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Proceedings
of the
BIOLOGICAL SOCIETY
of
WASHINGTON

Volume 93 6 November 1980 Number 3

THE BIOLOGICAL SOCIETY OF WASHINGTON

1980-1981
Officers

President: Richard Banks Secretary: Michael A. Bogan
Vice President: Raymond B. Manning Treasurer: David L. Pawson

Elected Council
Stephen D. Cairns Leslie W. Knapp

Anne C. Cohen Storrs L. Olson
Stanwyn G. Shetler

PROCEEDINGS
Editor: C. W. Hart, Jr.

Associate Editors

Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman
Plants: David B. Lellinger Vertebrates: Richard Banks

Insects: Robert D. Gordon

Membership in the Society is open to anyone who wishes to join. There are no prerequisites.
Annual dues of $10.00 include subscription to the Proceedings of the Biological Society of
Washington. Correspondence concerning membership should be addressed to the Treasurer,
Biological Society of Washington, National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C. 20560.

The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly
in February, May, August and November.

Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological
Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C. 20560.

Known office of publication: National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560.

Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044

Second class postage paid at Washington, D.C., and additional mailing office.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 515-522

TWO NEW SPECIES OF LIZZIA (HYDROZOA:
ANTHOMEDUSAE) FROM THE EASTERN
TROPICAL PACIFIC

Lourdes Segura

Abstract.—Two new planktonic species of hydromedusae from the East-
ern Tropical Pacific, Lizzia alvarinoae and Lizzia ferrarii, are described and
compared with the 5 other species of Lizzia, L. blondina, L. elizabethae,
L. fulgurans, L. gracilis, and L. octostyla. Morphological differences, as
well as differences in the number and characteristics of the oral tentacles,
are detailed.

Resumen.—Se describen dos especies nuevas de hidromedusas del planc-
ton del Pacifico Tropical Oriental, Lizzia alvarinoae y Lizzia ferrarii y se
comparan con las otras especies del género, L. blondina, L. elizabethae,
L. fulgurans, L. gracilis y L. octostyla de las cuales se diferencian en la
~morfologia, asi como en el numero y caracteristicas de los tentaculos orales.

Two new anthomedusas were found in the plankton collected on the series
of cooperative cruises made during the EASTROPAC expedition between
February 1967 and April 1968. A vast expanse of the Eastern Tropical Pa-
cific was surveyed on EASTROPAC, extending from 20°N to 20°S, and from
the American coasts offshore to 126°W. The plankton collections were taken
in oblique plankton hauls with a plankton net (0.5 m mouth opening) made
of 333 wm Nitex cloth. The collections were taken from different depths to
the surface in the epiplankton region. All drawings in this work were made
freehand from preserved material. Type-material of both species is depos-
ited in the National Museum of Natural History, Smithsonian Institution
(USNM).

Lizzia alvarinoae, new species
Figs. 1—2

Type-material.—Holotype, USNM 59270, collected 23 March 1968,
03°19.0'S;098°05.0’W, surface; 2 paratypes: USNM 58932, collected 19 Feb-
ruary 1968, 02°49.0’S,084°57.0’W, 0-209 m; USNM 58933, collected 7
March 1968, 02°31.5’S,105°03.5’W, 0—209 m.

Additional material.

Sta. 13.159; 10°46’S,098°00'W; 26-II-1967; 204—0 m; 3 specimens
Sta. 20.256; 12°45'’N ,098°22’W; 22-V-1967; 214-0 m; 1 specimen
Sta. 76.076; 04°42’S, 104°S8’W; 8-III-1968; 213-0 m; 2 specimens

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

516
1.0 mm

YIN 4, ot

pe
is
™
=
La}
y 1

Lizzia alvarinoae: 1, Drawn from preserved adult specimen showing shape of
the umbrella and the 4 marginal tentacles; 2, Stomach and peduncle, showing insertion of oral

Figs. 1-2.
tentacles above buccal margin and arrangement and shape of medusa buds.

S17

VOLUME 93, NUMBER 3

an

aya andes

a

pie ee

we og OW ae PRE OLS PM arn

2.5 mm

3, Detail of umbrella and stomach, showing different develop-

mental stages of medusa buds, one with 4 short marginal tentacles; 4, Drawn from preserved

adult specimen, showing mouthparts and size and shape of marginal tentacles.

Figs. 3-4. Lizzia ferrarii:

518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sta. 76.088; 10°53’S, 104°58’W; 10-III-1968; 210-0 m; 1 specimen
Sta. 77.144; 14°59'S,084°56’W; 15-II-1968; 202—0 m; | specimen
Sta. 77.159; 10°50’S,085°02’W; 16-II-1968; 198-0 m; | specimen
Sta. 77.438; 15°00'S,088°09’W; 8-IV-1968; 212—0 m; 1 specimen
Sta. 77.452; 15°00'S,093°06’W; 9-IV-1968; 194-0 m; | specimen

Diagnosis.—Hydromedusae with 4—20 unbranched oral tentacles; 4 radial
marginal tentacles, and oval-shaped medusa buds on stomach walls.

Description.—Bell of holotype 2.5 mm high, 2.0 mm in diameter. Um-
brella about 1.0-1.5 mm high and 1.5—2.0 mm wide, dome-shaped, usually
slightly higher than wide, with small apical projection (Fig. 1). Jelly mod-
erately thick, especially in apical region. Stomach large, barrel-shaped, at-
tached to a broad-based conical peduncle. Medusa buds oval-shaped,
stalked, located on interradial regions of stomach walls. Mouth simple with
round opening, surrounded by 4—20 unbranched oral tentacles arising above
mouth margin, each with | terminal nematocyst cluster (Fig. 2). Stomach
and mouthparts extend together beyond half the length of the bell cavity.
Four radial marginal tentacles, all of one kind, arise from bell margin. Basal
tentacle bulbs large, pear-shaped, with endoderm deeply pigmented. It was
not possible to determine the velum dimensions.

Remarks.—The species of Lizzia can be rather easily confused with Po-
docoryne or Rathkea in some stages of development (Russell, 1954). How-
ever, the latter genera can be distinguished by the form of the mouth-lips
which are expansions of the lips, each having a single cluster of nemato-
cysts. In Lizzia the oral tentacles are inserted a short distance above the
opening of the mouth, and each has one terminal nematocyst cluster.

The genus Lizzia was established by Forbes in 1846, and there are 5
species described at present. The major differences among them are sum-
marized in Table 1. Lizzia alvarinoae differs from the other species mainly
in two morphological characteristics. There are only 4 marginal tentacles,
all radial, and the number of oral tentacles varies between 4—20. In other
species the marginal tentacles are more than 4 and the oral tentacles do not
number more than 8.

Etymology.—Named in recognition of the prolific contributions of Dr.
Angeles Alvarino in the field of Marine Biology.

Lizzia ferrarii, new species
Figs. 3—4

Type-material.—Holotype, USNM 58929, collected 13 April 1968,
04°56.0’S ,095°13.0’W, surface; 2 paratypes: USNM 58930, collected 18 Feb-
ruary 1968, 05°41.0'S;085°03.0’'W, 0-220 m; USNM 58931, collected 8
March 1968, 04°42.0'S;104°58.0’W, 0-213 m.

Additional material.

VOLUME 93, NUMBER 3 SD)

Sta. 13.071; 04°16’S,092°05'W; 15-II-1967; 206-0 m; 1 specimen
Sta. 13.322; 11°47'N,095°08’W; 18-ITI-1967; 213-0 m; 2 specimens
Sta. 14.138; 07°55’S,088°46’W; —_1-ITI-1967; 214-0 m; 1 specimen
Sta. 14.255; 04°03’S,086°17’W; 22-III-1967; 207-0 m; 8 specimens
Sta. 14.287; 09°56’S,083°56’W; 26-III-1967; 210-0 m; 1 specimen
Sta. 45.365; 18°48’N,104°44’W; 10-[X-1967; 207-0 m; 4 specimens
Sta. 46.050; 02°50’N,105°16’W; 23-VITI-1967; 204-0 m; 1 specimen
Sta. 46.137; 11°48’N,092°02’W; 15-LX-1967; 206—0 m; 2 specimens
Sta. 46.147; 08°28’N ,092'05’W; 17-LX-1967; 170-0 m; 1 specimen
Sta. 46.155, 05°40’N ,091°58’W; 18-IX-1967; 224-0 m; 1 specimen
Sta. 47.049; 01°00’N ,082°00’W; 5-VIII-1967; 215-0 m; 1 specimen
Sta. 75.239; 05°00'N ,098°02’W; 4-IV-1968; 211-0 m; 1 specimen

Sta. 77.177; 05°41’S,085°03’W; 18-II-1968; 220-0 m; 1 specimen

Sta. 77.386; 02°40'S,088°04’W; 4-IV-1968; 206-0 m; | specimen

Sta. 77.394; 04°01'S,088°00'W; 4-IV-1968; 238-0 m; 2 specimens

Sta. 77.420; 10°22'S,087°57'W; 6-IV-1968; 215—O m; | specimen

Sta. 77.452; 15°00’S,093°06’W; 9-IV-1968; 194-0 m; 1 specimen

Sta. 77.482; 07°29'S,094°57'W; 12-IV-1968; 201-0 m; 1 specimen

Sta. 77.491; 04°56’S,095°13’W; 13-IV-1968; 217-0 m; 26 specimens

Diagnosis.—Hydromedusae with 12—24 unbranched oral tentacles; 8 long
marginal tentacles, 4 radial and 4 interradial in position, and pear-shaped
medusa buds developed on stomach walls.

Description.—Bell of holotype 2 mm high, 2 mm in diameter. Umbrella
about 1.0-2.5 mm high and 1.0-3.0 mm wide, dome-shaped, sometimes a
little wider than high, with a small apical projection. Jelly very thick, es-
pecially in apical region. Velum broad. Stomach cylindrical, attached to a
small conical peduncle. Medusa buds completely surrounding stomach,
pear-shaped, differing morphologically in different stages of development.
Some have only 4 short, equally developed, marginal tentacles (Fig. 3).
Mouth a simple round opening, surrounded by 12-24 unbranched, oral ten-
tacles arising above mouth margin, each with | terminal nematocyst cluster.
Stomach and mouthparts together extend beyond half the length of the bell
cavity. The 4 radial canals and ring canal narrow; with 8 long, similar,
marginal tentacles, 4 radial and 4 interradial (Fig. 4). Tentacles %4 longer
than bell height. Basal tentacle bulbs large, pear-shaped. Each tentacle bulb
with a dense mass of red endodermal pigment, which in radial tentacle bulbs
extends a short distance up radial canals.

Remarks.—Lizzia ferrarii differs from the related species in two morpho-
logical features. The number of oral tentacles, which in other species is no
greater than 20, varies between 12-24. L. ferrarii has 8 marginal tentacles.
All related species except L. gracilis have different numbers. However the
tentacles of L. gracilis are small and curve upwards over the margin of the
umbrella; the radial tentacles are twice as long as the interradial ones. In L.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

520

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MADAAA, “TT ADOULADAID *7 DIAISOJIO “T
so1oodsg

‘panunuoj— | IGRI,

SL, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ferrarii the marginal tentacles are large, equal in length and without any
curvature.

Etymology.—Lizzia ferrarii is named after Dr. Frank D. Ferrari who kind-
ly placed at my disposal the plankton collections of the EASTROPAC ex-
pedition.

Acknowledgments

I wish to express my appreciation to Dr. Frank Ferrari, Director of the
Oceanographic Sorting Center, Smithsonian Institution for the reasons cited
above. I am especially grateful to Dr. Angeles Alvarino for checking the
descriptions of these two new species and reviewing the manuscript.

Literature Cited

Agassiz, A. 1865. North American Acalephae. Illustrated catalogue of the Museum of Com-
parative Zoology at Harvard College, II—Mem. Mus. Comp. Zool. 1(2):1—234.
Bigelow, H. B. 1914. Fauna of New England. 12. List of the Medusae Craspedota, Sipho-

nophorae, Scyphomedusae, Ctenophorae.—Papers Boston Soc. Nat. Hist. 7:1-37.
Kramp, P. L. 1928. Papers from Dr. Mortensen’s Pacific Expedition 1914-16. Hydromedusae
and Anthomedusae.—Vidensk. Medd. dansk. Naturh. Foren. Copen. 85:27-62.

1959. The Hydromedusae of the Atlantic Ocean and adjacent waters.—Cana Rep.
46:3-283.
. 1961. Synopsis of the Medusae of the world.—J. Mar. Biol. Ass. U.K. 40:1—469.
. 1965. The Hydromedusae of the Pacific and Indian Oceans. Section I.—Dana Rep.
63: 1-162.
1968. The Hydromedusae of the Pacific and Indian Oceans. Sections II and III.—
Dana Rep. 13(72):1—200.
Mayer, A. G. 1900. Medusae of the world. Vol. I, II and IIJ.—Carnegie Foundation, Wash-
ington, D.C. 735 pp.
Russell, F. S. 1954. The Medusae of the British Isles: Anthomedusae, Leptomedusae, Lim-
nomedusae, Trachymedusae and Narcomedusae.—Cambridge University Press, 530 pp.

Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, Mex-
ico, 20 D.F.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 523-524

ORATOSQUILLA MEGALOPS, A NEW STOMATOPOD
CRUSTACEAN FROM TAIWAN

Raymond B. Manning

Abstract.—This new species is distinguished from O. solicitans Manning,
1978, by its unusually large eyes.

In a recent review (Manning, 1978) of some species of the Indo-West
Pacific stomatopod genus Oratosquilla, I pointed out the apparent variabil-
ity in material assigned to O. solicitans, and noted (p. 28) that three spec-
imens from Anping, Taiwan probably represented a distinct species.
Through the kindness of L. Tiefenbacher, Zoologisches Staatssammlung,
Munich, I was able to reexamine these specimens to verify that their eyes
are noticeably larger than those of other specimens assigned to O. solici-
tans. These three specimens are assigned to a new species herein.

I thank L. Tiefenbacher for the loan of the material and my wife Lilly for
preparing the illustrations.

Oratosquilla megalops, new species
Fig. 1

Material.—Anping [An-P’ing or Ampin, 23°00'N, 120°09’E], S Taiwan;
Haberer, leg.; June 1905: 3 6, total lengths 63 to 69 mm (largest male is
holotype, smaller males are paratypes; all specimens in Zoologisches Staats-
sammlung, Munich).

Diagnosis.—As in O. solicitans Manning, 1978:25, fig. 13, except: Rostral
plate (Fig. la, e, f) slightly broader than long, appearing elongate, tapering
anteriorly to convex anterior margin. Eye (Fig. 1b) moderate to large, cor-
neal indices 290-313 in specimens less than 70 mm long. Lobe between
basal prolongation of uropod (Fig. 1d) small, rounded, narrower than ad-
jacent spine.

Name.—The specific epithet is from the Latin and refers to the enlarged
eyes.

Remarks.—The eyes of O. megalops (Fig. 1b) resemble those of O. so-
licitans in shape but not in size. The corneal indices of O. megalops, 290-
313, do not overlap those of O. solicitans of similar size, 340—400 (see
Manning, 1978, table 1 on p. 42). The eyes of O. megalops also appear to
be larger than those of the similar O. hindustanica Manning, 1978, from
India. Only one of 27 specimens of the latter species, the smallest male

524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ieee)
* dallageas a
c¥ | d a
ste
oes |

Fig. 1. Oratosquilla megalops, male holotype, 69 mm: a, Anterior part of carapace; b, Eye;
c, Lateral processes of exposed thoracic somites; d, Basal prolongation of uropod. Anterior
part of carapace of: e, Male paratype, 63 mm; f, Smallest male paratype, carapace length
12.2 mm.

examined, 47 mm long, had a similar corneal index, 309. Specimens of O.
hindustanica 60 to 70 mm long had corneal indices ranging from 338 to 373.

The present material strongly suggests that each of the morphological
forms recognized in O. solicitans (see Manning, 1978:26—28, fig. 15a—d)
probably represents a distinct species.

Literature Cited

Manning, Raymond B. 1978. Further observations on Oratosquilla, with accounts of two new
genera and nine new species (Crustacea: Stomatopoda: Squillidae)—Smithsonian Con-
tributions to Zoology, No. 272:1-—44, figs. 1-25.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 525-535

REAPPRAISAL OF THE GASTROPOD GENUS
VARICOPEZA GRUNDEL (CERITHIIDAE:
PROSOBRANCHIA)

Richard S. Houbrick

Abstract.—The genus Varicopeza is described and its relationship to the
genera Bittium and Argyropeza is established. This monotypic genus, rep-
resented by Varicopeza pauxilla (A. Adams), 1s found in deep sea environ-
ments throughout the Indo-West Pacific. A synonymy and new description
which includes radular features and aspects of soft anatomy, and an histor-
ical review of the literature are presented. The ecology of the species and
the functional significance of the shell aperture are discussed.

Numerous lots of a small, Bittium-like cerithid dredged by the R.V. Al-
batross in the Philippine archipelago and deposited in the National Museum
of Natural History, Smithsonian Institution have prompted this review. This
small cerithid belongs to the monotypic genus Varicopeza Grundel, 1976
and has long been overlooked or misidentified in most museum collections
where it is frequently found mixed with other small cerithid species. Closely
related to the genera Argyropeza and Bittium, Varicopeza pauxilla (A.
Adams) has a long synonymic history and has been largely ignored in the
literature, except when mentioned incidentally.

In this review, I have examined material from the major American and
Australian museums and several other institutions. The sparse anatomical
material available for study was poorly preserved. I have surveyed the lit-
erature as completely as possible. Whenever possible, types of all nominal
species were examined. Scanning electron micrographs were made of the
radula, shell and operculum and preserved specimens examined for gross
anatomy.

The following abbreviations appear in the text: AMS, Australian Museum,
Sydney; ANSP, Academy of Natural Sciences of Philadelphia; BM, Berlin
Museum; BM(NH) British Museum (Natural History); NMV, National Mu-
seum, Victoria; TAU, Tel-Aviv University; USBF, United States Bureau
of Fisheries; USNM, United States National Museum of Natural History;
WAM, Western Australian Museum.

Family Cerithiidae Fleming, 1828
Genus Varicopeza Grundel

Varicopeza Griindel, 1976:46, plate 1, figures 11-13. Type-species: by tau-

526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tonomy, Varicopeza varicopeza Grundel, 1976 [=Varicopeza pauxilla (A.
Adams, 1854)].

Description.—Small, slender turreted shell characterized by flat-sided
whorls, deeply impressed suture and overall sculpture of spirally beaded
bands and axial riblets. Body whorl wide with distinctive thick varix op-
posite outer lip. Aperture wide with flaring outer lip and wide, deeply im-
pressed, anterior, inhalant, siphonal canal; wide, spoutlike, posterior ex-
halant canal in upper outer lip directly opposite anterior siphonal canal.
Protoconch smooth with convex whorls. Operculum ovate, paucispiral and
with subcentral nucleus. Radular ribbon taenioglossate (2+1+1+1+2) with
thin, sicklelike lateral teeth and marginal teeth serrated with tiny denticles.
Rachidian tooth with long, spadelike, basal plate, two tiny, basal denticles
and a long central tooth.

Etymology.—Varix-foot, from the Latin, varix, “‘dialation’’ or
and pes, “‘foot.’’ Varicopeza is a feminine Latin noun.

Remarks.—This monotypic genus is widely distributed throughout the
Indo-West Pacific geographic province where it is confined to the continen-
tal shelf and slope. The animal, shell, opercular and radular characters all
indicate a close relationship to the genera Bittium Gray, 1847, and Argy-
ropeza Melvill and Standen, 1901. The spoutlike exhalant siphonal canal in
the upper part of the outer lip of the type-species (see Fig. 1 a—e, j) is so
different from anything seen in Bittium or Argyropeza species that I believe
recognition of the genus Varicopeza Grundel, 1976 to accomodate the
species is justified, both on morphological and functional grounds.

The genus Varicopeza is not known from the fossil record. On the basis
of shell sculpture, Grundel (1976) believed the genera Varicopeza and Ar-
gyropeza Melvill and Standen, 1901 to be Recent representatives of the
family Procerithiidae Cossmann, 1905, subfamily Cryptaulinae Grundel,
1976, and placed Bittium in the same family. I find Grundel’s classification
too dependent on unimportant shell characters and topheavy with supra-
specific taxa; consequently, I prefer to regard Varicopeza as a genus in the
Cerithiidae until there is more solid information upon which to base su-
praspecific categories.

6

NTE *

Varicopeza pauxilla (A. Adams, 1854)
Figs. 1-2

Cerithidea (Pirenella) pauxilla A. Adams, 1854:86 (Holotype: BM(NH), not
registered; Type-locality: Burias, Philippines).

Cerithium pauxillum (A. Adams). Sowerby, 1855:889-890, pl. 186, fig. 294
(non Pease, 1860); Reeve, 1865, pl. 20, fig. 144; Tryon, 1887:141, pl. 34,
fig. 95; Kobelt in Martini Chemnitz, 1898:228, pl. 40, fig. 8; Schepman,
1909: 162; Oostingh, 1925:45.

VOLUME 93, NUMBER 3 S27.

Cerithium bicanaliferum Brazier, 1877:317 (Lectotype: AMS A-72, here se-
lected; 7 paralectotypes; Type-locality: 30 fathoms, Darnley Id., Torres
Straits, Queensland, Australia); Hedley, 1901:127, pl. 17, fig. 25; Ponder,
1972:45.

~Cerithium trigonostomum Melvill, 1910:10, pl. 1, fig. 15 (Lectotype:

BM(NH), not registered, largest from F. W. Townsend Collection, here

selected; 3 paralectotypes, two with Reg. No. 1912.8.16.97-8: Type-lo-

cality: 40 fathoms, Merkan Coast, Charbar [Pakistan]).

Argyropeza involuta Thiele, 1918:120, pl. 21, figs. 20, 20a (Holotype: BM,
not registered; Type-locality: sta. 245, 5°27.9’, 39°18.8’E at 463 meters
(near Zanzibar).

Varicopeza varicopeza Grundel, 1976:46—48, figs. 8, 11, 12, 13 (Holotype:
BM, not registered; Type-locality: 0°30’S, 107°5’'W, South China Sea).

Description.—Shell (Fig. 1 a—e, g—j): Shell small, about 7 mm long, 2.2
mm wide, thin, slightly transparent, slender and turreted, having apical an-
gle of 24 degrees and white to light tan in color. Teleoconch comprises 8-
13 straight-sided whorls that angle sharply inwards on each side at the deep-
ly impressed suture to form distinct sutural ramps. Each whorl sculptured
with raised spiral cords and broader, less distinct axial riblets. Where cords
cross ribs, small, sharp beads appear, giving an overall spiny appearance.
Early whorls of teleoconch have only 2 spiral cords; mid-whorls and pen-
ultimate whorl each with 3 dominant spiral cords and one weaker spiral
cord on the upper and lower sutural ramp. Protoconch of 3.5 concave,
smooth whorls the last 2 of which are sculptured with a centrally located
microscopic spiral thread. Lip of protoconch sinuous, thickened into spout-
like exhalant siphon at upper surface. Body whorl broad, with wide, prom-
inent varix opposite outer lip. Sculpture of body whorl comprises 4 raised,
primary spiral cords. Between each of these, is a thinner, raised spiral cord.
Base of body whorl constricted at siphonal area and sculptured with 6—7
thin, raised spiral cords. Outer lip crenulate, flaring and thickened on out-
side, due to lip varix. Aperture triangular-ovate with concave, smooth col-
umella and wide, anterior inhalant siphonal canal that is turned to the left
45 degrees to shell axis. A prominent posterior exhalant siphonal canal is
formed by spoutlike notch in upper, outer lip directly opposite to, and in
line with, anterior siphonal canal.

Operculum: (Fig. 1f) Operculum corneous, thin, ovate and paucispiral
with a subcentral nucleus. Operculum fits aperture snugly when animal is
retracted into shell, and has a very thin, flexible border.

Radula: (Fig. 2) Radular ribbon taenioglossate, tiny, about one-ninth the
length of the shell, and comprising about 20 rows of teeth. Rachidian tooth
Shield-shaped with spadelike basal plate upon which are 2 thickened, but
small basal cusps. Top of rachidian tooth straight with long, pointed, central

528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Varicopeza pauxilla, scanning electron micrographs showing shell physiognomy and
operculum. a—d, Apertural, side, dorsal and apertural views of adult shells, all USBF Station
5435, 80 meters, soft mud, off Nagubat Id., E. Mindanao, Philippines (USNM 276898), 30x;
e, Adult shell, 52 meters off Mersat Abu Samra, Gulf of Aqaba, Israel (USNM unaccessioned)
30x; f, Operculum of specimen from USBF Station 5426, 49 meters, off Mantaquin Id., E.
Palawan, Philippines (USNM 288300), 75x; g, Enlargement of protocon¢h showing sculptural
details and spoutlike notch in protoconch lip, 75x same data as in a—d; h, Detail of sculpture

VOLUME 93, NUMBER 3 529

Fig. 2. Scanning electron micrographs of radula of Varicopeza pauxilla. a, Radular ribbon
with some marginal teeth folded back to expose lateral teeth, 500; b, Enlargement of rachidian
and lateral teeth, 1,000; c-d, Marginal teeth showing cusp placement on inner and outer
marginals, 600x, 1,000.

<_—

on penultimate and adult whorls, 55x, same data as a—d; i, Close-up of aperture showing
exhalant siphonal canal, 200, same data as a—d; j, Detail of early whorls of teleoconch, 55x,
same data as a—d.

530 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tooth flanked by 3—4 small denticles on each side. Lateral tooth rhomboid
with straight top, serrated with one inner sharp denticle, a larger, triangular
pointed tooth, and with 6-7 smaller pointed denticles. Basal plate of lateral
tooth smooth, flat, with long, lower lateral extension onto basal radular
membrane. Inner marginal tooth long, hooklike and tapering to pointed tip,
serrated on both sides with 3-5 tiny denticles. Outer marginal tooth same,
only smooth on outer surface. Basal half of each marginal tooth becomes
wider and flatter where it is inserted onto the basal radular membrane.

Animal: Head-foot of preserved animals large and with deep groove
emerging from exhalant siphon and extending down right side of foot. Snout
long, bilobed at tip. Eyes very large in relation to animal. Mantle edge with
short, thick pallial tentacles. Organs of mantle cavity and internal anatomy
unknown. Eggs and larvae unknown.

Etymology.—From the Latin, pauxillus, ‘‘little,’’ *“small.”’

Discussion

Synonymy.—This species was first described by A. Adams (1854) as a
Cerithidea species, family Potamididae H. and A. Adams, 1854, but was
considered a Cerithium species by Sowerby (1855; 1865), Tryon (1887) and
Kobelt (1898). Shell sculpture, radular and opercular characters substantiate
its proper assignment to the Cerithiidae. Adams’ (1854) original diagnosis
was without a figure but described a “‘yellowish-brown shining species with
deeply channeled sutures, and with the whorls ornamented with 3 series of
rather acute tubercles.’’ His description also noted the unusually shaped
aperture with sinuous lip and the posterior siphonal canal. Examination of
the holotype shows it to be conspecific with recent material and the other
synonyms given above.

The nominal species, pauxilla, has been overlooked because of the poor,
tiny figures in Sowerby (1855; 1865) and the other classical iconographies
which do not clearly illustrate the sculpture or distinctive features of the
aperture. As a result, a number of names have been proposed, as can be seen
in the above synonymy. The homonym, Cerithium pauxillum Pease, 1860, is
a different species.

The earliest synonym is Cerithium bicanaliferum Brazier, 1877. This
name appropriately describes the unique siphonal canals. The names bican-
aliferum and pauxillum were missed by Melvill (1910), who named the
species, trigonostomum, “‘‘triangular mouth.’’ The type-material of C. tri-
gonostomum comprises 4 specimens, 2 of which were purchased by Sow-
erby and Fulton and have the registry number 1912.8.16.97-8. The other 2,
lacking registry numbers, are from the F. W. Townsend collection. I select
the best preserved specimen from the Townsend collection as the lectotype
of C. trigonostomum because Melvill’s (1910) paper was written about the
mollusks collected by Townsend.

VOLUME 93, NUMBER 3 531

Grundel (1976) proposed the genus Varicopeza to accomodate what he
thought was a new species and was apparently unaware of the other avail-
able specific names.

Examination of the holotype of Argyropeza involuta Thiele, 1918, has
convinced me that it is conspecific with Varicopeza pauxilla. Thiele’s spec-
imens are both immature individuals lacking outer lips and were erroneously
assigned by him to the genus Argyropeza Melvill and Standen, 1901. Al-
though related to the genera Bittium and Varicopeza, Argyropeza species
differ in having a distinctive protoconch sculpture of 2 spiral cords, a spiral
row of beads adjacent to the suture, and a sinusigera notch.

Remarks.—Varicopeza pauxilla is a common species, widely distributed
throughout the continental shelf and slope regions of the Indo-West Pacific
biogeographic province.

The color of this species is variable. Dead collected specimens tend to be
chalky white while freshly collected material is white or light tan and of
porcellaneous texture. Some shells are flesh colored or dark tan: a series of
specimens from Palau (ANSP 203855;203483) ranged in color from white to
brown, many being striped with a dark tan band on the lower half of each
whorl. A summary of the shell dimensions and characters is presented in
Table 1, below:

Table |.—Statistical summary of shell measurements (mm) of Varicopeza pauxilla.

Character Number Range Mean S.D.
Length 64 3.5—9.6 6.89 1533
Width 64 1.4—2.8 Dales 0.35
Aperture length 64 0.5-2.0 1.21 0335
No. whorls 64 8-14 te) [e239

Adults, with fully formed body whorl, varix, outer lip, and two siphonal
canals are so distinctive that they are unlikely to be confused with any other
small, deep-sea cerithid (Fig. la—e). Immature individuals or shells without
the outer lip may be difficult to identify without close examination of the
distinctive sculptural pattern. Varicopeza pauxilla may be confused with
young individuals of Rhinoclavis (Proclava) sordidula (Gould, 1849) which
have similar sculpture. The latter species differs by being much larger and
in having a median columellar plait in the aperture and a sutural ramp only
on the bottom portion of each whorl.

Cerithium elegantissimum Hedley, 1899 resembles Varicopeza pauxilla
in that the outer lip flares slightly, forming a weak exhalant posterior si-
phonal canal, but the shell is less slender and has different sculpture. The
former is clearly more like Bittium and forms a morphological link between
that genus and Varicopeza.

Cerithium scabriusculum Issel, 1869, may be a synonym of Varicopeza

SV PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pauxilla. | have not examined the type-material, but Issel’s (1869:pl. 3, fig.
11) illustration looks somewhat close to Varicopeza pauxilla.

Varicopeza pauxilla may also be mistaken for a turrid species because of
the similarity of the exhalant posterior siphonal canal to the “‘turrid notch.”’
Turrids however, normally have narrower apertures and slightly longer an-
terior siphonal canals.

Ecology.—The ecological information available suggests that this deep-
sea species is a microphagous detritivore and perhaps a filter feeder, as well.
It lives in great numbers on soft or fine sandy-muddy bottoms and has a
radula that is morphologically and functionally similar to those of Bittium
species. The numerous rod-shaped, ovoid fecal pellets of fine detrital par-
ticles are relatively large in comparison to the size of the animal, indicating
a continuous feeding habit. The deep groove leading from the exhalant si-
phon down the right side of the foot probably conveys mucous, fecal pellets
and other debris away from the animal. It is more pronounced than in other
cerithid species. The functional significance of the wide, exhalant, posterior
siphonal canal may be explained in terms of filter feeding on a soft substra-
tum. The inhalant and exhalant siphonal canals allow the animal to lie with
its aperture tangential to the surface of the substratum and may expedite
the flow of water and detrital particles through the mantle cavity. Normal
grazing activity can also take place. It does not appear that the animals are
partly buried in the sediment because none of the specimens examined were
eroded or worn on the dorsal or ventral surfaces of their shells. I thus
suggest that this small cerithid snail grazes and filter feeds at the water-
substratum interface.

Many specimens examined from the Philippines, Palau, and the Andaman
Sea were drilled, indicating heavy predation by naticid snails. Many empty
Shells from Palau contained sipunculids.

Although the protoconch is large, consisting of 3.5 whorls, the smooth
sculpture and spout-like notch (Fig. 1g) in the upper edge of the outer pro-
toconch lip and the absence of a sinusigera notch point to a direct or de-
mersal developmental mode rather than to an indirect, pelagic one.

Geographic distribution (Fig. 3).—The bathymetric range of this species
is summarized in Table 2. The geographic range is confined to the Indo-
West Pacific province. The one record from Bikini Atoll, Marshall Islands,
indicates that this species may yet be found in other areas of Micronesia
and possibly in Polynesia.

Specimens Examined

GULF OF AQABA: off Elat, 167 m; Elat, 121 m; Elat, 40-49 m; off
Mersat Abu Samra, 52 m; off Wadi Murach, 80-82 m; off Gesirat Fara’un,
64-73 m; Elat, from beach; off Nuweiba el Tarabin, 143-146 m; off Nuweiba

VOLUME 93, NUMBER 3 553

Fig. 3. Geographic distribution of Varicopeza pauxilla (A. Adams).

el Museina, 329-402 m (all USNM 783672-783678). ZANZIBAR: 5°27.9’/N,
39°18.8'E, 463 m, near Zanzibar (BM); 2 mi. W of Nyange Id., 20 m (ANSP
251215). INDIA: 10 mi. SE of Vizagapatnam, 58 m, Bay of Bengal (ANSP
294094). BURMA: 15°08’'N, 99°04’E, 50 mi. SW of Irrawaddy River, Pre-
paris N. Channel, 53 m (ANSP 293491). ANDAMAN SEA: 13°N, 97°4’E,
35 mi. W of Tavoy Id., Andaman Sea, 68 m (ANSP 292782); 08°29’N,
97°59’E, 25 mi. NNW of Phuket Id., W. Thailand, 42 m (ANSP 291467;
291826); 12°01’N, 92°5S’E, 14 mi. NNW of Port Blair, off S. Andaman Ids.,
49 m (ANSP 292348). WESTERN AUSTRALIA: Approximately 170 mi. W
of Port Hedland, W.A., 19°29’S, 116°01’E, 137 m (AMS). QUEENSLAND,
AUSTRALIA: off Endeavour Reef, N of Cooktown, 37 m (AMS); Dornley
Id., Torres Straits, 5S m (AMS A-72); Palm Id., N of Townsville, 27 m
(AMS C10188); Low Isles, near Port Douglas (AMS); Palm Id., (NMV).
NEW GUINEA: 1-2 mi. off Kaipoiri Village, Koeroedoe Id., Geel-
vink Bay, 1S m (ANSP 277870); 1 mi. S by E. Cape Dgaroewawoffi, Japen
Id., Geelvink Bay 12-29 m (ANSP 277723); W end Manubada (Local)
Id. off Port Moresby, Papua, 18-22 m (AMS). PALAU: 1-1.5 mi. off

Table 2.—Bathymetric range of Varicopeza pauxilla.

No. of stations Depth range (meters) Mean depth S.D.

48 11-686 90.92 122.36

534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Garakasan Point, E side of Babelthaup (ANSP 203855); off Malakal Harbor,
46-55 m (ANSP 236589; 203483). MARSHALL ISLANDS: 3 mi. SW of
Bikini Id., Bikini Atoll, 46 m (USNM 586688). PHILIPPINES: off SE Tawi-
tawi, 33 m (USNM 283634); Palawan Pass, off Cauayan Id., 95 m (USNM
258481); Linapacan Str., off Observatory Id., Palawan, 84 m (USNM
282437); SE off Bantayan Id., 59 m (USNM 281081); S off Corregidor Lt.,
55 m (USNM 263530); off Tocanhi Pt., Tawi-tawi, 90 m (USNM 274414);
off Nagubat Id., E. Mindanao, 80 m (USNM 276898); off Mantaquin Id., E.
Palawan, 49 m (USNM 288300); off Tacbuc Pt., E. Leyte, 88 m (USNM
282853); off Tacbuc Pt., E. Leyte, 104 m (USNM 283184); Linapacan Str.,
off Observatory Id., 84 m (USNM 282792); off Tinakta Id., Tawi-tawi Is.,
29 m (USNM 257689); Linapacan Str., off Observatory Id., Palawan, 84 m
(USNM 291621); off S. W. Tawi tawi, 33 m (USNM 312733); off Malavatuan
Id., W. Luzon, 33 m (USNM 278223); off Observatory Id., Palawan Pass,
79 m (USNM 282334); off Observatory Id., Palawan Pass, 79 m (USNM
282023); off Tinakta Id., Tawi tawi, 29 m (USNM 257690); Palawan Pass,
off Pt. Tabonan, 686 m (USNM 285856); off Sueste Pt., W. Luzon, 46 m
(USNM 262609); off S.W. Tawi tawi, 33 m (USNM 312727); Malampaya
Sound, Palawan, 11 m (USNM 286138); S.E. off Bantayan Id., 59 m (USNM
281061). BORNEO: off Sandakn Lt., 71 m (USNM 285765).

Acknowledgments

Within the Smithsonian Institution, I thank Dr. Joseph Rosewater and
Dr. Harold Rehder for critically reading the original draft of this manuscript.
June Jones kindly typed the final copy of the manuscript. I also thank Cathy
Lamb for her technical assistance and for helping to proofread the text.
Thanks are extended to the staff of the Scanning Electron Microscope Lab-
oratory and to Victor Krantz of Photographic Services for the photographs.
I gratefully thank the following individuals for the loan of material in their
charge: Dr. A. Barash, TAU; Dr. George Davis, ANSP; Dr. R. Kilias, BM;
Dr. Winston Ponder, AMS; Dr. Brian Smith, NMV; Dr. John Taylor,
BM(NH); and Dr. Fred Wells, WAM. Part of this work was done with the
aid of a Smithsonian Research Award and Smithsonian Fluid Research
Funds.

Literature Cited

Adams, A. 1854. A monograph of Cerithidea, a genus of Mollusca, with descriptions of
several new species, from the collection of Hugh Cuming, Esq.: To which are added,
descriptions of two new species of Colina, and one of Donax.—Proceedings of the
Zoological Society of London 22:83-87.

Brazier, J. 1877. Shells collected during the Chevert Expedition.—Proceedings of the Lin-
naean Society of New South Wales 1:311-321.

VOLUME 93, NUMBER 3 535

Cossmann, M. 1905. Mollusques éocéniques de la Loire-Inférieure.—Bulletin de la Société
des Sciences naturelles de l’ouest de la France 3(2):68—123 (135-189), pls. 1-6 (9-14).

Fleming, J. 1828. Mollusks. Encyclopedia Britannica, supplement to editions 4—6, volume 3,
part 1. Edinburgh.

Gould, A. A. 1849. Shells ‘‘brought home by the U.S. Exploring Expedition.’’—Proceedings
of the Boston Society of Natural History 3:106—-108.

Gray, J. E. 1847. The classification of the British Mollusca by N. E. Leach, M.D.—Annals
and Magazine of Natural History 20:267—273.

Grundel, J. 1976. Zur Taxonomie und Phylogenie der Bittium-Gruppe (Gastropoda, Cerithi-
acea).—Makakologische Abhandlungen Staatliches Museum fiir Tierkunde in Dresden
5(3):33—5S9. 2 pls., 17 figs.

Hedley, C. 1899. The Mollusca of Funafuti Atoll—Memoir of the Australian Museum

3(7):397-567, 80 figs.

. 1901. A revision of the types of the marine shells of the Chevert Expedition.—Records

of the Australian Museum 4(3):121—130, pls. 16-17.

Issel, A. 1869. Malacologia del Mar Rosso.—Biblioteca Malacologica. 387 pp., 5 pls. Pisa.
Kobelt, W. 1888-1898. Die Gattung Cerithium, 297 pp., 47 pls.—Jn Martini, F. H. S. and J.
H. Chemnitz, Neues systematisches Conchylien-Cabinet, etc. 1(26). Nurnburg.
Melvill, J. C. 1910. Descriptions of twenty-nine species of marine mollusca from the Persian
Gulf, Gulf of Oman, and North Arabian Sea, dredged by Mr. F. W. Townsend, of the
Indo-European Telegraph Service, 1901—1903.—Annals and Magazine of Natural His-

tory (8)6:1-17, pls. 1-2; 289-324, pls. 20-23.

Melvill, J. C. and R. Standen. 1901. The mollusca of the Persian Gulf, Gulf of Oman and Arabian
Sea, as evidenced mainly through the collections of Mr. F. W. Townsend, 1893-1900,
with descriptions of new species.—Proceedings of the Zoological Society of London
2:327—400, pls. 21-24.

Oostingh, C. E. 1925. Report on a collection of Recent shells from Obi and Halmahera (Mol-
uccas).—Mededeelingen van de Landbouwhoogeschool te Wageningen (Nederland)
29(1):3-362.

Pease, W. H. 1860. Descriptions of forty-seven new species of shells from the Island of Ebon,
Marshall’s Group, in the collection of H. Cuming.—Proceedings of the Zoological So-
ciety of London, pp. 431-438.

Ponder, W. F. 1972. Type specimens in the Macleay Museum, University of Sydney. VI.
Mollusks.—Proceedings of the Linnaean Society of New South Wales, vol. 97, part 1,
no. 429:42-55.

Schepman, M. M. 1909. The Prosobranchia of the Siboga-Expedition.—Siboga-Expeditie
49(43):100—231, 7 pls. Leiden: Brill.

Sowerby, G. B. 1855. Thesaurus Conchyliorum, or monographs of genera of shells.—Vol. 2,

Cerithium: 847-859, pls. 176-186. London.

. 1865. Cerithium, 20 pls. + index in Reeve, L. A., Conchologia Iconica: or illustrations

of the shells of molluscous animals. Vol. 15. London.

Thiele, J. 1918. Gastropoda der Deutschen Tiefsee-Expedition, pt. 2—Wissenschaftliche Er-
gebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer ‘‘Valdivia’’ 1898-1899,
17(2): 1-382. Jena.

Tryon, G. W. 1887. Manual of Conchology; structural and systematic; with illustrations of
the species.—First series: 9a, Cerithium, pp. 127-149, pls. 20-29. Philadelphia.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 536-550

PSEUDOCHIRELLA SQUALIDA GRICE &
HULSEMANN, 1967, FROM
CONTINENTAL SLOPE WATERS
OFF DELAWARE (COPEPODA:CALANOIDA)

Frank D. Ferrari

Abstract.—The original description of copepodid stage VI female Pseu-
dochirella squalida is augmented. Initial descriptions of stage VI male, stage
V female, and stage V male are presented. Sexually dirmorphic characters
in both stages are present in a variety of body segments, appendages, and
their armature. Some of their presumed functions are discussed. Bilateral
asymmetry is exhibited only in stage VI and is manifested in primary or
secondary sexual characters.

Introduction

The establishment of Pseudochirella squalida by Grice & Hulsemann
(1967) was based on a single copepodid stage VI female specimen collected
in the Indian Ocean on 28 June 1964 at ANTON BRUUN station 351B
(29°45'S, 64°58’E) between 350 and 1,710 m. Deevey & Brooks (1977) re-
ported 2 females in the Sargasso Sea collected at Station ‘“‘S’’ (32°10’N,
64°30’ W) on July 1969 and February 1970 between 500 and 1,000 m. To date
examination of calanoid copepods from midwater trawl samples of 2 cruises
(July 1975, ALBATROSS IV, and February 1976, OREGON II) in the area
of Deepwater Dumpsite 106 in continental slope waters off Delaware has
shown P. squalida to be the most abundant species of the genus. In this
paper I augment the description of the external morphology of copepodid
stage VI female (hereafter VI 2°), describe copepodid stage VI male (VI
36), copepodid stage V female (V ¢), and copepodid stage V male (Vd) and
discuss sexual dimorphism and bilateral asymmetry among the various
Stages.

Materials and Methods

Specimens were examined from midwater trawl samples collected during
cruises to Deepwater Dumpsite 106, an area encompassed by 70°31’—75°00’ W
and 36°00’-39°10’N. A more complete description of the gear and station
positions is given by Krueger et al. (1977). Table | gives collection data for
the samples containing P. squalida.

All specimens of P. squalida were removed from every sample and

VOLUME 93, NUMBER 3 Ss)

measured. They were prepared for examination by clearing in lactic acid
and staining with chlorazol black E. In the following presentation VI @ is
described and illustrated completely. VI 6, V 2, and V ¢ are compared to
this description and only those structures differing significantly from VI @
are illustrated. For clarity, illustrations of spines, setae and their ornamen-
tation are not complete. Generally for similar articles or elements in a series
only one is illustrated completely. In a few cases individual variation occurs
in the number of setae comprising large clusters, e.g. RiA2, RiMd or RiMx1.
These instances are not reported as it is impossible to determine whether
the setae were lost during preparation, not observed due to imperfect stain-
ing, or actually represent individual variation. Illustrations were made with
the aid of camera lucida or drawing tube. Anatomical terminology and ab-
breviations follow Ferrari and Bowman (in press) except for the mouthparts
where terminology and abbreviations of Giesbrecht (1892) are used.

Results
Pseudochirella squalida Grice & Hulsemann 1967
Pseudochirella squalida Grice & Hulsemann 1967:24, figs. 65—70.

Material.—71 VI 2, 10 VI 6, 18 V 2, 61 V ¢ from 27 midwater trawl
samples listed in Table 1.

Description.—VI 2: Length range 5.5—6.8 mm. Cph/Pg]1 articulation (Fig.
le) free laterally, fused dorsally. Pg4/Pg5 fused. Url (genital segment) (Figs.
la, b; 4d, e) with row of hairs across dorsal surface. Left side of Url pro-
duced midlaterally into slight bulge, accentuated by peripheral integumental
ridges; posterolaterally a small bump; series of integumental ridges across
ventrolateral face; small patch of anterolateral hairs. Right side 2 large pos-
terolateral bumps with long hairs; integumental ridges across ventrolateral
face; scattered hairs dorsally. Ventral surface slightly produced into genital
prominence with accentuating integumental ridges; large transverse ridge
posteriorly, with associated hairs. Ur2&3 (Fig. la) with thin hairs scattered
Over entire segment; distinctly thicker hairs along posterodorsal margin. CR
with hairs dorsally and laterally; first terminal setae reduced, originating
from ventral surface; remaining terminal setae and dorsal seta long, thick
and plumose; lateral seta small. Al (Figs. 2c—e) 24 free segments, 8—9 fused.
1A1 with tiny spines; esthetes on 2,3,5,6,9,13,18,25. B1A2 (Fig. 2a) proximal
comb of 6-7 setules; 1 distal seta. B2A2 2 distal setae. Re 7 segments; Re2—
6 1 seta each; Re7 1 seta midlength, 3 distally. Ril 2 distal setae; Ri2 separate
group of setae on 2 distal edges, lateral edge 8 setae, medial 9. BIMd as shown
(Fig. 3a); hairs of varying thickness on anterior surface. B2 3 medial setae.
ReMd 5 segments; Rel—4 1 seta each Re5S with 2. RilMd 3 medial setae,
thin hairs laterally; Ri2 10 distal setae. LeMx1 9 setae (Fig. 3c); Re 9 setae;

538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ri 16 setae. B2 5 distal setae. Li3 4 setae; Le2 5 setae. Lil 10 anterior spines
with long spinules; 4 posterior spines with short, thick spinules; hairs on
anterior surface. BIMx2 (Fig. 3e, f) with lateral spinules. LI-SMx2 with
small spines on posterior surface; each lobe 3 long spines with spinules or
plumes. Ri 5 segments; Ril—4 with large medial spine and reduced postero-
lateral spine; Ri5 3 well-developed spines. B1&2Mxp with rows of spinules
as illustrated (Fig. 4b). B1 3 groups of 3 spines. B2 2 medial spines with
spinules and 3 plumose setae. Ri 5 segments; with 0,0,0,1,1 lateral setae and
3,4,3,3,3 medial spines with spinules. BIP1 (Fig. 5a) row of medial hairs;
B2 row of medial hairs and strong curved distal spine extending over tu-
bercule of Ri. Re 3 segments; articulation between Re1&2 incomplete. Rel
with medial hairs and 1 Se with spinules. Re2 medial hairs, tiny spines on
anterodistal edge, 1 Se with spinules and 1 Si. Re3 hairs on anterior surface
and along lateral edge, Se with spinules, 1 St plumose medially, spinules
proximolaterally, serrate flange distolaterally, and 3 Si. RiP1l 1 segment,
forming medial tubercule on anterior face, with tiny spines (see Vaupel-
Kline, 1972, for a more complete discussion of this structure in Euchirella),
anterodistal hairs, 5 Si. B1P2 (Fig. 3h) medial hairs and 1 Si. B2 naked.
ReP2 3 segments. Rel medial hairs, 1 Se with serrate flange on both edges,
1 Si. Re2 similar to Rel but with row of tiny anterodistal spines, lateral
hairs, and lacking medial hairs. Re3 3 biflanged Se; St plumose medially,
serrate flange laterally, and 4 Si. P3 (Fig. 3i) similar to P2 except Ri 3
segments. Ril 1 Si. Ri2 lateral hairs and 1 Si. Ri3 posterior and lateral hairs,
1 Se, and 4 Si. P4 (Fig. 4a) similar to P3 except BI medially formed lobe
with comb of strong spines, 6—7 on left B1; 8—10 on right. B2 with posterior
hairs. Ri2 with posterior hairs. PS absent.

VI 6: Length range 5.1—-5.8 mm. Cph/Pg1 fused (Fig. 1d). Pg4/Pg5 fused;
small unarticulated, slightly curved, posterodistal spine on each side. Url
(Fig. lc) genital opening on left side. Ur2—-5 posterodistal serrate fringe;
without hairs on surface of segments. CR smaller than VI 9° with fewer
dorsal hairs; medial 2 setae longer than VI °. Al (Figs. 2f—-h) 24 free seg-
ments; 8&9 fused; articulation between 9&10 and 12&13 fused posteriorly;
many more esthetes especially on proximal segments. B1A2 (Fig. 2b) distal
seta reduced. B2 2 distal setae reduced. Re similar to VI 2 but seta on Re2
and | on Re7 absent. Ril naked. Ri2 mediodistal series with 7 setae; latero-
distal with 6. B1Md (Fig. 3b) gnathobase reduced to knob. B2 1 medial seta.
Re all segments fused, with 6 setae. Ril lateral hairs and 2 medial setae.
Ri2 9 distal setae. Mx1 (Fig. 3d) greatly reduced. Le 7 setae. Re 11 setae.
Ri&B2 fused. Ri 10 setae; B2 with 5. Li3 2 small setae. Lil&2 naked. Mx2
(Fig. 3g) greatly reduced, all segments fused. L1-5 small knobs with
1,2,2,2,2 setae; LS distal seta with swollen base. Ri 5 setae. Mxp (Fig. 4c)
slightly reduced. B1 1 medial and 2 distal setae; small spines absent. B2 5
medial setae; no small spines. Ri with 3,3,2,2,3 medial spines; 0,0,0,1,1
lateral setae. Pl (Fig. 5f) slightly reduced, similar to VI 2 but Rel &2 artic-

VOLUME 93, NUMBER 3 539

\
Hil
tt

\||
iM

)
!

SSS

SS

=>

aS

=

ee

Ss

==

=

=

Se

Fig. 1. Pseudochirella squalida: a, V1 2 Ur, dorsal; b, VI 2 Ur, ventral: Ga Vio Ur:
dorsal; d, VI 3 habitus, left lateral; e, VI 2 habitus, right, lateral; f, V 2° habitus, right lateral:
g, V ¢o habitus, right lateral.

540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Pseudochirella squalida: a, V1 2° A2;b, VI 3 We ey Wik @ AN, Meliseal, Wl 2 all,
(Mile o, Wl @ AN, Z2=W5e Te, WLS A N33 ep WIL) Ail, eile ns WAL Ss ANIL, 22 = 29.

VOLUME 93, NUMBER 3

a0)

z
z
z
5
2
é
4\ \\
4| {\
i
4} h\

‘\

Fig. 3. Pseudochirella squalida: a, VI 2 Md, posterior; b, VI ¢ Md, posterior; c, VI 9°

{x1, posterior; d, VI ¢ Mx1, posterior: e, VI 2 Mx2, posterior; f, VI 2 Mx2, anterior: g, VI
. Mx2, posterior; h, VI ? P2, posterior: i, VI 2 P3, posterior.

541

542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(
Ayn
Why

Za

Ziff
Vi
V/A

|
Wy
Noe:

HO NN

args —

\

Wl MW My

——
~~

\
(
(
(

Fig. 4. Pseudochirella squalida: a, V1 2 P4, posterior; b, VI 2 Mxp, posterior; c, VI ¢
Mxp, posterior; d, VI 2 genital segment, right lateral; e, VI 2 genital segment, left lateral; f,
VI 2 genital segment with spermatophore, left lateral; g, VI 2 genital segment with sper-
matophore, right lateral.

VOLUME 93, NUMBER 3 543

ulation complete; SeRel reduced and naked; SeRe2 thick and naked; SeRe3
naked. P2&3 similar to VI 2. P4 (Fig. 5c) also similar except B2 comb of
spines absent and posterior hairs on B2&Re2 absent. P5 (Figs. 5d&e) well-
developed, complicated and asymmetrical. Left and right B1 fused to cou-
pler. Left B2 larger than right. Left Re 3 segments. Re2 with medial blade
and 2 mediodistal tooth-like projections. Re3 with medial tooth-like projec-
tion and mediodistal patch of hairs. Left Ri 1 segment extending slightly
beyond Rel. Right Re 2 segments. Rel bent laterally, extending to middle
of left Re2. Right Re2 with proximomedial ridge and several ridges disto-
laterally. Right Ri 1 segment extending to end of left Ri; tip ovate; ridge on
anterior side.

V 2: Length range 4.3-4.7 mm. Cph/Pg1 (Fig. 1f) fused laterally. Pg4/5
fused. Pg5 with points similar to VI ¢. Ur 4 segments. Ur3&4 with dorsal
and ventral hairs and posteroventral serrate marginal fringe. CR similar to
VI 2. Appendages also similar except as noted. Ri2A2 1 seta. Ri3 8 medial
and 6 lateral setae. Ri2Md with 9 distal setae. ReMx!1 10 setae; Ri 15 setae.
RiMx2 incompletely fused at base. Ri2Mxp 3 spines; Ri3&4 with 2 each.
Ril&2P2 articulation partially fused. Ril&2P3 articulation partially fused.
B1P4 comb of spines absent. PS absent.

V o: Length range 4.6-5.0 mm. Cph/Pg1 separate (Fig. 1g). Pg4/5 sepa-
rate. PgS points similar to VI 6. Ur 4 segments. Ur2&3 with posteroventral
serrate marginal fringe. Appendages similar to VI 2 except as noted. Ri2A2
1 seta. Ri3 8 medial and 7 lateral distal setae. Re2&3Md fused. ReMx1 9
setae. RiMx2 fused to B2. Re2&3P1 without anterior hairs. Ril&2P2 par-
tially fused. Ril&2P3 partially fused. B1P4 without comb of spines. P5 (Fig.
5b) symmetrical. Left and right Bl fused. Re and Ri each as simple, undif-
ferentiated segment.

Discussion

As shown in Table 1, 3 other species of the genus have been collected
with Pseudochirella squalida: P. obtusa VI 2 (1 specimen); P. pustulifera
Mio? (Go), Vis (),eV"S ©): PP. scopularis VI -< (1). Most mature and
juvenile specimens of the 4 species were captured between 400 and 1,825
m, the depths to which most tows were confined. Capture data did not
indicate evidence of particular sex or stage occupying well-defined or ex-
clusive depth zones.

Twenty VI 2° of P. squalida were found with spermatophores attached
ventrally on the genital segment (16 with 1 spermatophore, 3 with 2, and |
with 3). Unlike some species of Pseudochirella and species of the related
genus Echirella, P. squalida does not have a distinctly well-sculptured area
on the genital segment (see descriptions and illustrations in Vervoort, 1949;
Park 1975, 1976a, 1976b, 1977). Areas such as the large dorsal lobe on
Euchirella messinensis and the large protrusion on E. truncata serve as
spermatophore attachment points away from the genital field. Fertilization

544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Pseudochirella squalida: a, V1 ° Pl, anterior; b, V do PS5, posterior; c, VI d P4,
posterior; d, VI 3 PS, posterior; e, VI 3 PS, anterior; f, VI ¢d Pl, anterior.

545

VOLUME 93, NUMBER 3

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546

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Characters which exhibit sexual dimorphism in stage V or stage VI and their state
in the remaining stage.

Character

Cph/Pgl

Pg4/Pg5

Pg5 points

Ur

position, genital
opening

Ur marginal hairs

Ur marginal fringe

CR setae

Al

A2 setae Re
Ri

Md gnathobase

setae B2
segments Re
setae Ri

Mx! size

spines Lil
setae Li2
setae Li3
setae Ri
setae Re
setae Le

Mx2 size

spines L1
spines L2
spines L3
spines L4
spines L5
spines Ri
segments Ri

Mxp size

VI °

fused dorsally
fused entirely
absent

4 segments

ventral

Ur 2&3

none

8&9 fused
9&10 free
12&13 free

9
19

well developed

3
all free
13

—= WWW WwW WwW

|
all free

VI ¢

fused entirely
fused entirely
present

5 segments

lateral (left)

none
Ur 2-4

longer
than VI °

8&9 fused
9&10 part free
12&13 part free

7
15

reduced teeth
absent

|
all fused
11

greatly reduced

greatly reduced

l
2
2
2
2
5)

all free

reduced
slightly

VQ

fused laterally
fused entirely
present

4 segments

absent

none
Ur 2&3
similar

to VI @

similar
to VI @

9
15

well developed

3
all free
12

similar
to VI @

14
5
4

15

10
9

similar
to VI @

WWW Ww

3
11
Ril/B2 fused
similar
to VI 2

Vo

free

free
present

4 segments

absent

none
Ur 2&3

similar
to VI 2

similar
to VI 2

9
16

well developed

3
2&3 fused
12
similar
to VI @
14
5
4
15
9
9
similar
to VI ¢
3
3
3
3
3
11
all free
similar
to VI °

VOLUME 93, NUMBER 3 547

Table 2.—Continued.

Character VI 2 VI 3 V?° Vo
spines B1 y) 3 9 9
spines Ri 18 I{5) 15 (5)
Pl size reduced similar similar

slightly to VI ° to VI 2

Ril & 2 partly fused free partly fused partly fused
margin Se spinules smooth spinules spinules
P4 Bl spine comb absent absent absent
P5 absent large complex absent small simple

tubes serve to connect the sperm sac to the genital opening (unpublished
observations).

Spermatophores of P. squalida were attached on the sternum to the left
or right of the genital plate. Multiple spermatophores were also attached in
this area. Figs. 4f, g show left and right lateral views of VI 2 genital segment
with the sperm sac narrowing to a short neck and attached to the left ster-
num by a simple plate. Several spermatophores including the one illustrated
were partly empty, the distolateral areas absent. A similar situation was
reported for Euchaeta norvegica and the implications discussed by Ferrari
(1977). In the illustrated example the genital plate appears flexed up into the
genital segment and a large mass of material seems to fill the vacated space
below it. This mass was not detected on specimens with full spermato-
phores. Due to the thick cuticle and opaque structures associated with the
spermatophore it was impossible to determine anything more of the relations
of the mass, genital plate and pore, and seminal receptacle.

Characters which exhibit sexual dimorphism in either stages V or VI and
their development in the remaining stage are described in Table 2. Very
little is known of calanoid behavior in general or even of the function of
various appendages, segments, or their armament. Nothing is known of the
behavior of P. squalida. However some suggestions about the function of
these sexually dirmorphic characters can be made based on studies of other
calanoids. The characters can be artificially grouped into: those for which
some information is available through behavioral observations of homolo-
gous morphological characters in other calanoids; those for which there is
no homologous behavioral information but information about comparative
morphology may permit some conjecture; those for which no statements
can be made.

In the first group the reduction in size and/or ornamentation or A2, Md,
Mx1, Mx2, and Mxp in VI ¢ indicates an inability of these animals to feed.
This is further substantiated by the degenerate condition of the gut (unpub-

548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Comparison of sexual dimorphism (sd) and bilateral asymmetry (ba) for both sexes
of stage V and stage VI.

ba sd

? None Cph/Pg1 articulation
Pg4/Pg4 articulation
V A2
Md
Mx!
Mx2
PS

6 None

Cph/Pg1 articulation
Pg4 points
2 Genital segment Ur-shape and number of segments
Ur-margin of segments
P4 comb of spines CR-length of setae
Al
A2
Md
Mx!
Mx2
VI Mxp
Pl
3 Genital segment P4

PS
PS

lished observations). Significant reductions occur in teeth of BIMd gnatho-
base and strong spines of LiMx1 and LI—5Mx2 which probably hold, tear,
and crush the food prior to injestion.

Concerning the VI ¢ PS and VI 2 Url (genital segment) much has been
written about the Centropagidae (Lee, 1972, and Blades, 1977), Labidocera
(Fleminger, 1975 and Blades & Youngbluth, 1979) and Euchaeta norvegica
(Hopkins et al., 1978). The VI ¢ uses his P5 to clasp the 2 and position
himself immediately prior to spermatophore transfer which is accomplished
by his left ReP5. The shape of the VI 2 genital segment may mirror the
clasp of her mate as well as the position of the spermatophore.

Regarding the second group of characters, sexual dimorphism in A1 of VI
2 and ¢ is manifested in greater number of esthetes of the ¢. Griffiths &
Frost (1976) state that VI ¢ esthetes on Calanus pacificus and Pseudoca-
lanus sp. may be chemoreceptors upon which dissolved organic matter pro-
duced by VI 2 accumulates. They suggest this organic matter is a phero-
mone.

CR setae of VI 2 are the initial contact point of VI ¢ during mating of
the heterarthandrid copepods Centropages typicus and Labidocera aestiva
(Blades, 1977, and Blades & Youngbluth, 1979). The VI ¢ grasps the VI

VOLUME 93, NUMBER 3 549

2 CR setae with his geniculate right Al. Obviously this behavior does not
occur during mating of P. squalida as VI ¢ lacks the geniculation. However
such as function for the CR setae should not be precluded since another
appendage could be used by the VI 4 in the initial contact.

No direct information is available about the function of the comb of spines
on BIP4 VI @. Considering the absence of P5 in VI 2 and the position of
the spines, it is possible that they function in an analogous manner to the
reduced PS in VI @ of other calanoid females. Lawson (1977) describes VI
2 PS of the Candaciidae as a ‘‘secondary sexual structure.’’ Fleminger
(1975) calls it a “‘sexually modified character’? of Labidocera. Blades &
Youngbluth (1979) have recently reported that Labidocera aestiva V1 2
uses P5 to remove the spermatophore after sperm discharge.

Pl is morphologically similar to P2—4 but differs not only by the presence
of the modified Ri and B2 setae but also in the ornamentation of SE Rel-3.
The further sexual dimorphism in Se Rel-—3 VI 6, reduction in size and
ornamentation (VI 2 morphology is virtually identical to V 2 and V 3@),
would appear to follow the progression of such reductions in the feeding
appendages anterior to it, as there are only minor morphological changes in
swimming appendages posteriorly.

The functional significance of such sexually dimorphic characters as Cph/
Pgl articulation, Pg4/5 articulation, points on Pg5, number of Ur segments,
position of genital opening, and posterior margins of Ur segments is not
understood.

Table 3 briefly lists those external morphological characters which exhibit
sexual dimorphism in either VI 2&6 or V 9&d and compares these to
characters exhibiting bilateral asymmetry. Bilateral asymmetry is apparent
only in the adult, is not manifested in one sex more than the other and
involves known primary or secondary sexual characters involved in repro-
duction.

Acknowledgments

I would like to thank Dr. Robert Gibbs, Jr. for the opportunity to study
these specimens and Drs. Thomas E. Bowman and Taisoo Park for review-
ing the manuscript.

Literature Cited

Blades, P. 1977. Mating behavior of Centropages typicus (Copepoda: Calanoida).—Marine

Biology 40:57-64.

, and M. Youngbluth. 1979. Mating behavior of Labidocera aestiva (Copepoda: Cal-

anoida).—Marine Biology 51:339-35S.

Deevey, G., and A. Brooks. 1977. Copepods of the Sargasso Sea off Bermuda: species com-
position, and vertical and seasonal distribution between the surface and 2000 m.—Bul-
letin of Marine Science 27(2):256—291.

550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ferrari, F. 1977. Spermatophore placement in the copepod Euchaeta norvegica Boeck 1872
from Deepwater Dumpsite 106.—Proceedings of the Biological Society of Washington
91(2):509-521.

, and T. Bowman (in press). Oithonidae (Copepoda:Cycopoida) from the east coasts of

Central and South America.—Smithsonian Contributions to Zoology, 312:1—27.

Fleminger, A. 1975. Geographical distribution and morphological divergence in American
coastal-zone planktonic copepods of the genus Labidocera.—Estuarine Research 1:392—
419.

Giesbrecht, W. 1892. Systematik und Faunistik der pelagischen Copepoden des Golfes von
Neapel und der angrenzenden Meeresabschnitte.—Flora und Fauna des Golfes von
Neapel, monograph. 19,831 pp., 541 pls.

Grice, G., and K. Hulsemann. 1967. Bathypelagic copepods of the western Indian Ocean.—
Proceedings of the United States National Museum 122(3583): 1-67.

Griffiths, A., and B. Frost. 1976. Chemical communication in the marine planktonic copepods
Calanus pacificus and Pseudocalanus sp.—Crustaceana 39(1):1—8, 1 pl.

Hopkins, C., J. Mauchline, and D. McLusky. 1978. Structure and function of the fifth pair of
pleopods of male Euchaeta norvegica (Copepoda:Calanoida).—Journal of the Marine
Biological Association of the United Kingdom 58:63 1-637.

Krueger, W., R. Gibbs, Jr., R. Kleckner, A. Keller, and M. Keene. 1977. Distribution and
abundance of mesopelagic fishes on Cruises 2 and 3 at Deepwater Dumpsite 106.—
NOAA Dumpsite Evaluation Report 77-1:377—422.

Lawson, T. 1977. Community interactions and zoogeography of the Indian Ocean Candaciidae
(Copepoda: Calanoida).—Marine Biology 43:71-92.

Lee, C. 1972. Structure and function of the spermatophore and its coupling device in the
Centropagidae (Copepoda:Calanoida).—Bulletin Marine Biology and Ecology 8:1-10,
10 pls.

Park, T. 1975. A new species of Euchirella and the male of Euchirella splendens Vervoort,

with notes on Euchirella truncata Esterly (Copepoda, Calanoida).—Crustaceana

28(3):291-297.

. 1976a. Calanoid copepods of the genus Euchirella from the Gulf of Mexico.—Con-

tributions in Marine Science 20:101—122.

. 1976b. A new species of Euchirella (Copepoda, Calanoida) from the pacific coast of

North America, with note on Euchirella pulchra (Lubbock).—Crustaceana 31(2):208-—

DNAs

. 1978. Calanoid copepods belonging to the families Aetideidae and Euchaetidae from

antarctic and subantarctic waters.—Jn David Pawson (ed.), Biology of the Antarctic

Seas VII, Antarctic Research Series 27:91—290.

Vaupel-Klein, F. 1972. A new character with systematic value in Euchirella (Copepoda,
Calanoida).—Zoologische Mededelingen 47:497—-5S11, 6 pls.

Vervoort, W. 1949. Some new and rare Copepoda Calanoida from East Indian Seas.—Zoolo-
gische Verhandelingen 5:3-53.

Smithsonian Oceanographic Sorting Center, Smithsonian Institution,
Washington, D.C., 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 551-562

LARVAL DEVELOPMENT UNDER LABORATORY
CONDITIONS OF THE TROPICAL SPIDER
CRAB MITHRAX (MITHRACULUS) CORYPHE
(HERBST, 1801) (BRACHYURA: MAJIDAE)

Liberta E. Scotto and Robert H. Gore

Abstract.—The larval development of Mithrax (Mithraculus) coryphe, a
tropical American shallow water marine spider crab is described and illus-
trated from stages cultured in the laboratory, and compared with larvae of
Mithrax (Mithraculus) forceps (A. Milne Edwards, 1875). Development
consists of two zoeal and a megalopal stage; a prezoeal stage is hypothesized
but was not observed. Temperature affects duration of development with
both zoeal stages lasting 2 days each at 30°C and 3 days each at 25°C. No
first crab stages were attained, but data obtained from the megalopal stages
at both temperatures allowed a planktonic duration of about 10 days to be
extrapolated. Larvae of M. coryphe are almost identical in most morpho-
logical features to those of M. forceps, so separation of the zoeae in the
two species will be difficult; megalopal stages are more easily distinguished.
Similarity in larval and postlarval stages corresponds with the general mor-
phological similarity exhibited by adults of the two species.

Mithrax (Mithraculus) coryphe 1s a tropical American marine spider crab
distributed from the Indian River region on the central eastern Florida coast,
southward throughout the eastern and southwestern Caribbean, to Sao Pau-
lo, Brazil (Powers, 1977; Velez F., 1977). It is associated with rocky or
coralline habitats, or seagrasses, from the intertidal zone to 55 m. The larval
development of members of this genus has been little studied (see Wilson
et al., 1979, for summary), in spite of the easy accessibility of their habitats
and the relative abundance of specimens. In a previous paper (Wilson ef
al., loc. cit.) we described the complete larval development for Mithrax
(Mithraculus) forceps, and compared morphological features in the larvae
of that species to those exhibited by larvae in the subgenus Mithrax (Mith-
rax). In this paper we continue our investigations on this genus, by providing
illustrations of the larval development of M. coryphe, and comparing it to
larvae of M. forceps. This is but the third report on larvae within the genus
Mithrax, and the second within the subgenus (Mithraculus).

Materials and Methods

Two ovigerous females were collected from Ninguange Lagoon, Santa
Marta, Colombia, from a milleporine patch reef in | m of water, on 23 May

SPY, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1979. The specimens were held in 8.5 cm covered glass laboratory dishes
in nonflowing seawater (35%.) until hatching occurred in one female on 3
June. A total of 96 larvae, divided evenly among 4, 24-compartmented plas-
tic trays, was cultured in controlled temperature units (CTU) at 25° and
30°C. A single tray (24 individuals) at 25°C was inadvertently starved on the
day of hatching; the remaining tray at 25°C, and both trays at 30°C received
Artemia nauplii daily; all trays received fresh seawater (35%) daily. Light
in the CTU’s was on a diel cycle, 12 hours on, 12 hours off.

Illustrations and measurements of larvae were made as in the previous
study (Wilson eft al., 1979). The descriptions and illustrations below are
based on zoeae and megalopae obtained in the fed series at 25° and 30°C.

Laboratory Culture Experiment

The larval development of Mithrax coryphe consists of two zoeal stages,
and a megalopal stage. Whether a prezoeal stage occurred is unknown; none
was observed the morning of the hatch. However, other members of the
genus begin their larval development with such a stage, albeit of very short
duration. It is thus possible that the stage took place but was passed before
the trays were examined. Duration of development in the zoeal and megal-
opal stage is presented in Table 1, and the percentage of larval survival is
illustrated in Figure 1; both show that the duration of larval development
is temperature-dependent. The minimum time spent in each zoeal stage was
2 days at 30°C and 3 days at 25°C. Regrettably, megalopal stage development
was not completed at any temperature. Consequently, no firm conclusion
can be made as to postlarval duration or total larval development. However,
if the duration of the megalopal stage is comparable to that in M. forceps,
then this stage probably lasts from 6-8 days. Using this value, the length of
planktonic existence in M. coryphe at 30°C may be about 10 days, or slightly
longer at 25°C. This duration is different from that observed in M. spino-
sissimus (Provenzano and Brownell, 1977), and M. forceps (Wilson et al.,
1979); the former required about 5—6 days, and the latter about 14 days, to
complete planktonic development.

_ The larval series at 25°C which was starved for one day showed the effects
of food deprivation on the early zoeal stage. The first zoeae required 5 days
(as compared to 3 days in the fed series at 25°C) to complete development.
Survival was poor, only 3 zoeae attaining stage II, and all dying as such.
By laboratory day 4 only 12 of the original 24 zoeae starved on day 1
remained alive in stage I, whereas 19 of 20 surviving zoeae in the 25°C fed
series had already molted to second stage. Three of the day-1-starved zoeae
did attain stage II, but were apparently too debilitated to complete this
stage; the greatest mortality in this series occurred prior to the molt to stage
II. Contrarily, nearly all their counterparts in the fed series reached megal-

VOLUME 93, NUMBER 3 5/3)

Table 1.—Duration of the larval stages of Mithrax (Mithraculus) coryphe at various tem-
peratures.

Duration (days)

Mini- Maxi- Total number

Temperature (°C) mum Mean Mode mum molting to next stage
Puce (Fed) Zoeae | 3 3 3 3 19
II 3 3.3) 3 4 7

Megalopa l — 4 4 All died in stage
25°C (Starved | day) Zoeae I 5) 5 5 5) 3

II 1 — — 3) All died in stage
30°C (Fed) Zoeae I 2 2 2 3 42
II 2 2 yy 2 26

Megalopa l — 3 4 All died in stage

opa, but it was throughout this stage that the greatest mortality occurred.
These data, admittedly brief, suggest that the early zoeal stages of Mithrax
coryphe probably have insufficient amounts of yolk, or none at all, subse-
quent to eclosion, so that survival through the first zoeal stage requires an
outside food source which must be available immediately upon hatching.
Best survival was seen in the 30°C series, a somewhat surprising result
because generally at this temperature the increased speed of development
and the shorter intermolt duration in the larvae often appear concomitant
with higher overall mortality than is seen at cooler temperatures where the
development is prolonged (e.g. Wilson et al., 1979). We believe that indi-
| vidual larval variation in development can be ruled out as a possible expla-
nation for the observed variance in mortality in M. coryphe because all the
'zoeae in this study came from the same hatching produced by a single
female. However, the fact that M. coryphe is more of a tropical-subtropical
species (based on distributional records), rather than a eurythermic-tropical
| species (sensu Briggs, 1974) may provide some explanation. If the species
_and its larvae were better adapted to warmer seawater temperatures, then
presumably the larvae would complete their development faster at the higher
temperatures, thereby settling out from the plankton sooner.

| Description of Developmental Stages

| The zoeae and megalopae of M. coryphe, while not completely identical
| to those of M. forceps, are sufficiently similar in most morphological char-
| acters to cause some difficulty in distinguishing between the two forms. But
because an extensive description of the zoeae and megalopae of M. forceps
, is available (Wilson et al., 1979), the following descriptions of larvae of M.
'coryphe will be limited to features which differ from those noted in M.
forceps, i.e., Structures not described are identical to those of M. forceps.

|

554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

100

80
30°C
60:

40 Zoea II

20 Megalopa

100

80

SURVIVAL

60

40

20

PERCENTAGE

100

60

25°C

40 Zoea Il

20

Megalopa
8 10

2 4 6
DAYS IN STAGE
Fig. 1. Percentage survival and stage duration of Mithrax coryphe larvae reared under
laboratory conditions. N = number of larvae cultured at each temperature; * = larvae starved
on first day at 25°C.

ee

VOLUME 93, NUMBER 3 555

= —
A 1.0mm
ie :
See B-1 0.25mm
[ee eee

Fig. 2. Mithrax coryphe, first zoea: A, Lateral view; B, Telson; C, Antennule; D, Antenna;
E, Mandibles; F, Maxillule; G, Maxilla; H, Maxilliped 1; I, Maxilliped 2.

556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SSW
SAN

A 10mm
Ss

B-J 0.25mm
el

Fig. 3. Mithrax coryphe, second zoea: A, Lateral view; B, Telson; C, Antennule; D, An-
tenna; E, Mandibles; F, Maxillule: G, Maxilla; H, Maxilliped 1; 1, Maxilliped 2; J, Maxilliped 3.

VOLUME 93, NUMBER 3 BS)//

A-B 1.0mm

C-J 0.25mm

Fig. 4. Mithrax coryphe, megalopal sensory and feeding appendages: A, Lateral view; B,
Dorsal view; C, Antennule; D, Antenna; E, Mandibles; F, Maxillule; G, Maxilla; H, Maxilliped
1; 1, Maxilliped 2; J, Maxilliped 3.

558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

In order to facilitate identification and allow comparison between the larvae
of the two species, all appendages in the zoeal and megalopal stages are
completely illustrated. As in M. forceps, the measurements of larvae are
based on the arithmetic averages of the specimens examined.

First Zoea.—(Carapace length: 0.67 mm; 10 specimens examined).

Carapace (Fig. 2A): As in M. forceps, but dorsal spine slightly more
recurved; rostral spine inserted at different angle than in M. forceps, so
that antennular rod in lateral view appears ventral to plane of rostral spine;
in M. forceps antennular rod lies in same plane as rostral spine, obscuring
latter. Setation number and position identical to M. forceps; thoracic ap-
pendages unsegmented, visible through carapace.

Telson (Fig. 2B): Slightly wider and longer than M. forceps, furcae slight-
ly longer.

Mandible (Fig. 2E): Processes slightly less robust.

Color: Differs from M. forceps in that the basipodites of maxillipeds 1
and 2 are orange-rose with spidery black chromatophores ventrally; in M.
forceps maxilliped 1 is transparent, with a single black chromatophore dis-
tally at junction of endopodite, whereas maxilliped 2 is colored similarly to
that of M. coryphe. The posterior margin of the eyes, and the dorsal cara-
pacial spine with orange-rose hue in M. coryphe; uncolored in M. forceps.

Second Zoea.—(Carapace length: 0.83 mm; 10 specimens examined).

Mandibles (Fig. 3E) remain stouter in M. coryphe, but otherwise identi-
cal.

Maxilla (Fig. 3G) nearly identical; scaphognathite in M. coryphe with 25
marginal setae (24 in M. forceps).

Maxilliped 3 (Fig. 3J) with epipodite bud, placed as illustrated; no bud
observed, but possibly overlooked in M. forceps.

Color: Maxilliped 1 and 2 retain first stage coloration: M. forceps lacks
chromatophores on maxilliped 1 in this stage; abdominal somites 1—4 lime-
green with orange and black chromatophores ventrally; M. forceps abdom-
inal color similar to stage I.

Megalopa.—(Carapace length =< width: 1.15 x 1.05 mm; 10 specimens
examined.

Carapace (Fig. 4A, B): Generally similar to M. forceps, although larger
in size. Differs in placement of additional tubercle on gastric region, making
total of 6 (instead of 5), placed 3,1,2 (instead of 3, 2); only 4 tubercles in
arch on cardiac region (instead of 5 in M. forceps); general carapacial se-
tation slightly more abundant in M. coryphe, placed as illustrated.

Abdomen (Fig. 4A, B; Fig. SA—C): Setation slightly different but probably
not significantly so.

Antennule (Fig. 4C): Identical in form and setation (but not in aesthetasc
positioning and number) to M. forceps; in Wilson et al. the aesthetascs were
stated to be placed in a V, progressing distally 2,2,2,1 but this should read,

VOLUME 93, NUMBER 3 559)

Fig. 5. Mithrax coryphe, megalopal locomotory appendages: A-C, Pleopods 1,4,5; D-F,
Pereiopods 1,2,5.

560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Comparison of male gonopods in Mithrax coryphe (left) and Mithrax forceps (right).
Sternal views on left, abdominal views on right of each figure.

progressing distally 1,2,2,2, instead; in M. coryphe 9 aesthetascs are posi-
tioned similarly, progressing distally 1,2,2,2,2.

Maxilla (Fig. 4G): Essentially similar; however, scaphognathite in M.
coryphe with 29-33, usually 32 marginal setae (in M. forceps 26-30 marginal
setae).

Maxilliped 1 (Fig. 4H): Nearly identical; epipodite differs slightly in hav-
ing usually 6, sometimes 7, marginal and terminal setae.

Maxilliped 3 (Fig. 4J): Similar; variation in epipodite setation ranges from
0-2 lateral, and 2—6 terminal setae in both species, thus both overlap.

Pereiopods (Fig. 5D—F): Chelipeds essentially similar in form, varying in
setation (probably not significantly); mean ratio (N = 5 specimens) of mov-
able finger to palm in M. coryphe = 1.37; this ratio in M. forceps = 1.24;
walking legs more or less identical in general appearance; dactyls slightly
shorter relative to propodus (0.78 in M. coryphe, 0.88 in M. forceps) on
first walking leg (pereiopod 2); coxal and ischial spines present both species,
figured, but inadvertently not discussed in our account of M. forceps; coxal
spine in M. coryphe with | seta, in M. forceps with 2.

Color: Differs from M. forceps. Carapace anteriorly rose-orange, poste-
riorly transparent, with paired black chromatophores only dorsally; gut
black interiorly; chelipeds and all walking legs rose-orange, darker on is-
chium, merus, carpus, lighter on propodus and dactyl. Abdominal somites
1-2 clear, 3-6 rose-orange; telson overall rose-orange, without individual
chromatophore. Peripheral ommatidia of eyes reflect rose-orange, corneas
dark. Rostrum, antennule and antennae clear, without chromatophores or
diffuse coloration.

VOLUME 93, NUMBER 3 561

Discussion

From the preceding description it is obvious that the zoeal and megalopal
stages of Mithrax coryphe will be extremely difficult to distinguish from
those of its closely related congener, M. forceps. Except for minor variation
in appendage armature, which may eventually prove to be insignificant, the
larval and postlarval stages are nearly identical. The most noticeable dif-
ference between the two species is in the coloration of live specimens on
the zoeal and megalopal stages, with the two species more easily separated
in the megalopal stage. In general, the carapace, telson and walking legs of
M. coryphe are rose-orange, whereas in M. forceps the colors are golden-
brown and greenish-gold, respectively, with nearly transparent walking legs.
Postlarvae of the two species also differ in positioning and number of chro-
matophores, with M. coryphe having but a single pair dorsally on the car-
apace (2 pairs in M. forceps), no chromatophores on abdominal somites 1
and 2 (a black, and several red chromatophores, respectively, on these
somites in M. forceps), and none on the mouthparts (several black chro-
matophores on the mandibles, labrum, and protopodites of maxilliped 3 in
M. forceps).

In addition to color, M. coryphe differs in the megalopal stage from M.
forceps in size (slightly larger), dorsal tuberculation on the carapace (1 more
gastric, 1 less intestinal tubercle), overall carapacial setation (slightly more
setose), in the number of antennular aesthetascs (9 instead of 7), and the
number of marginal setae on the scaphognathite (29-33, as against 26-30 in
M. forceps). The higher ratio of the movable finger to the palm length of
the chelipeds in M. coryphe may prove to be of some value, although too
few specimens were available to allow firm conclusions in this respect.

In a previous paper (Wilson et al., 1979) we compared the larval stages
of Mithrax (Mithraculus) forceps to the first zoea of Mithrax (Mithrax)
pleuracanthus, and to the larval and postlarvae of M. (Mithrax) spinosis-
simus. We found very little difference between M. forceps and M. pleu-
racanthus on the one hand, but considerable differences between M. for-
ceps and M. spinosissimus, on the other. This demonstrated that characters
defining subgenera at the adult level may not be useful at the larval level.
Moreover, the very great similarity between M. coryphe and M. forceps
larvae in this study, and thus to M. pleuracanthus larvae (at least in the
first stage), supports previous observations (e.g. Yang, 1976) that many of
the species in the various majid genera in which the larvae are known show
a remarkable consistency in larval characters, making them difficult to sep-
arate in the plankton. Mithrax coryphe thus becomes the second species in
the subgenus Mithraculus in which the larvae appear similar to at least one
species in the subgenus Mithrax (i.e. M. pleuracanthus), although differing
considerably from another (M. spinosissimus, Provenzano and Brownell,
ISD.

562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The great similarity exhibited by the larvae of M. coryphe and M. forceps
is carried over into the adult stage to some degree. Adults of both species
have oblique branchial sulci on the carapace dorsum, but are distinguished
chiefly by the number and armature of the anterolateral lobes (3, bluntly
rounded in M. coryphe; 4, spine-tipped in M. forceps), and whether the
cheliped carpus is dorsally nodose (M. coryphe) or smooth (M. forceps).
The gonopods in males of the two species show noticeable differences with
that of M. coryphe morphologically much simpler (Fig. 6). It would appear
that differentiation between the species is first observable at a very early
stage, probably the megalopa, based on our studies.

Acknowledgments

We thank Kim Allyn Wilson for her aid in collecting the ovigerous fe-
males, maintaining them in the laboratory, and for much other assistance.
We gratefully acknowledge the aid of Dr. Jose A. Lozano, Director, Insti-
tuto de Investigaciones Marinas de Punta de Betin, and Mr. Hernando Nes-
tor of the same institution for their gracious assistance and help during our
stay at “‘Invemar’’ in Santa Marta, Colombia.

Literature Cited

Briggs, J. S. 1974. Marine zoogeography.—McGraw-Hill, New York. ix + 475 pp., 65 text-
figs.

Powers, L. W. 1977. A catalogue and bibliography to the crabs (Brachyura) of the Gulf of
Mexico.—Contr. Mar. Sci. Univ. Texas 20(Suppl.): 1-190.

Provenzano, A. J., Jr., and W. N. Brownell. 1977. Larval and early postiarval stages of the
West Indian spider crab, Mithrax spinosissimus (Lamarck) (Decapoda: Majidae).—
Proc. Biol. Soc. Wash. 90:735—752, figs. 1-7.

Velez F., M. M. 1977. Distribucion y ecologia de los Majidae (Crustacea: Brachyura) en la
region de Santa Marta, Colombia.—An. Inst. Inv. Mar.—Punta Betin 9:109-140, figs.
1-28.

Wilson, Kim A., Liberta E. Scotto, and Robert H. Gore. 1979. Studies on Decapod Crustacea
from the Indian River region of Florida XIII. Larval development under laboratory
conditions of the spider crab Mithrax forceps (A. Milne Edwards, 1875) (Brachyura:
Majidae).—Proc. Biol. Soc. Wash. 92(2):307—327, figs. 1—S.

Yang, W. T. 1976. Studies on the western Atlantic arrow crab genus Stenorhynchus (Deca-
poda Brachyura, Majidae) 1. Larval characters of two species and comparison with
other larvae of Inachinae.—Crustaceana 31:157-177, figs. 1-13.

Smithsonian Institution, Fort Pierce Bureau, Fort Pierce, Florida 33450.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 563-572

A NEW SPECIES OF PONTOPHILUS (CRUSTACEA:
NATANTIA: CRANGONIDAE) FROM THE GULF OF
MEXICO AND THE WESTERN ATLANTIC!

Mike Dardeau

Abstract.—Pontophilus gorei, a new species of caridean shrimp, is de-
scribed from the Gulf of Mexico and the western Atlantic. It differs from
other Gulf of Mexico species in lacking an exopod on the first pereopod and
in the form of the rostrum. The synonymy of Philocheras with Pontophilus
is discussed.

In a review of the genus Pontophilus in the Gulf of Mexico, Pequegnat
(1970) listed three species, P. brevirostris Smith, 1881, P. gracilis Smith,
1882 and P. abyssi Smith, 1884. Specimens discussed as P. abyssi later
proved to be P. talismani Crosnier and Forest, 1973 (pers. comm., L. H.
Pequegnat). Examination of specimens collected during ‘*Project Hour-
glass,’ a biological sampling program on the continental shelf off central
western Florida (see Joyce and Williams, 1969), revealed that a fourth
species of Pontophilus, described herein, occurs in the Gulf of Mexico.

Pontophilus gorei, new species
| Figs. 1—4

Material examined.—Specimens are deposited in invertebrate collections
at the U.S. National Museum of Natural History, Washington, D.C.
(USNM); Rijksmuseum van Natuurlike Historie, Leiden, Netherlands
(RMNH); Florida Department of Natural Resources Marine Research Lab-
oratory, St. Petersburg, Florida (FSBC I); Texas A&M University, College
Station, Texas (TAMU); Indian River Coastal Zone Study Reference Mu-
seum, Fort Pierce, Florida, (IRCZS); and Marine Environmental Sciences
Consortium, Dauphin Island, Alabama (MESC). The abbreviation cl refers
to carapace length, measured in mm from the level of the posterior margin
of the orbits to the posterior margin of the carapace.

Holotype.—1| ovig. 2, cl 2.5; 26°24’N, 83°22’W, 55 m; 6 Aug. 1966, trawl;
USNM 172417.

Allotype.—1 6, cl 1.9; 27°37'N, 83°58’'W, 55 m; 2 Aug. 1966, trawl;
USNM 172418.

Paratypes.—West coast of Florida, off Tampa Bay: 1 6, cl 1.8; 2 2, cl
feSrled: Soytlvencl 0:6—122 27-3 /ONEos4 13) We (3.132 Dees 1966, dredge:
SINIMP 72419 Si cles ovie. 2. cl 2 27°37-N, 84> 13° W., 73°ms; 2

' Contribution No. 029 of the Marine Environmental Sciences Consortium, P.O. Box 386,
Dauphin Island, Alabama 36528.

564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Aug. 1966, trawl; RMNH D31979. 1 3, cl 1.9; 27°37'’N, 83°58’W, 55 m; 6
Feb. 1967, trawl; IRCZS 89:3474. 1 ovig. 2, cl 2.1; 27°37'N, 83°28'’W, 37
m; 2 June 1967, trawl; FSBC I 21454. West coast of Florida, off Sanibel
Island; | ovig. 2, cl 2.1; 26°24’N, 83°43’W, 73 m; 13 Oct. 1966, trawl;
TAMU 2-5944. 1 63, cl 1.8; 26°24’N, 83°43'’W, 73 m; 8 Aug. 1967, trawl;
TAMU 2-5943. 1 6, cl 1.8; 26°24'N, 83°22’W, 55 m; 12 Oct. 1967, trawl;
FSBC I 21455. 1 ovig. @, cl 2.0; 26°24’N, 82°58’W, 37 m; 7 Apr. 1967,
dredge; IRCZS 89:3475. 1 ovig @, cl 2.2; 26°24’N, 82°58’W, 37 m; 6 June
1967, dredge; IRCZS 89:3476. Coast of northwest Florida, off Cape San
Blas: 1 juv., cl 1.4; 28°2S’N, 85°15’W, 182 m; 29 Sept. 1977, trawl; MESC.
East coast of Texas, off Padre Island: 1 3, cl 1.9; 27°37'N, 96°33'W, 59 m;
5 June 1971, from fish stomach; TAMU 2-5942. Western Atlantic, off Geor-
gia: 1 5, cl 1.9; 30°55'N, 81°18’ W, 9 m; 9 Sept. 1973, small biological trawl;
USNM 172420.

Diagnosis.—Rostrum short, extending slightly beyond cornea, with pair
of lateral spines midway along length; tip spatulate with expanded rounded
apex. Carapace smooth with single, strong dorsomedial spine behind ros-
trum; hepatic spines lacking. Merus of first pereopod with strong spine
midway on flexor margin; subchelar spine simple.

Description.—A robust, smooth-bodied crangonid shrimp (Fig. la). Ros-
trum (Fig. Ic) short, depressed, extending slightly beyond cornea; 2 blunt
lateral spines midway along length; tip spatulate, expanded club shape. Car-
apace (Fig. 1b) slightly depressed, smooth, with 2 very faint longitudinal
carinae and single, strong, dorsomedial spine behind rostrum; suborbital
spine dorsal and posterior to antennal spine; pterygostomian spine strong,
equal to or extending slightly beyond eyes; a minute spine posteroventral
to pterygostomian spine, directed somewhat medially; hepatic spine lacking;
orbital notch present; eyes well developed.

Stylocerite (Fig. 2a) broad at base, tip acute, reaching nearly to end of
basal article of antennule; antennular article 1 longer than 2, with minute
distolateral spine; article 2 longer than 3; flagella overreach antennal scale,
inner slightly longer than outer, both ending in a tuft of setae. Antennal
blade (Fig. 2b) slightly broader proximally, tapering to subquadrate tip;
lateral border of antennal scale almost straight, ending in robust distolateral
spine, spine may or may not exceed antennal blade. Antennal articles 1—4
compressed; article 5 longer than combined lengths of articles 1-4, falling
just short of distal margin of antennal blade; flagellum tapering to slender
tip.

Mouthparts as illustrated (Figs. 2c—g).

Pereopod | (Fig. 3a) strong, subchelate; dactyl slender; propodus broad,
width 4 length, with serrate spinules proximally on flexor margin, subchelar
spine simple; carpus short, 5 length of propodus, small spine on distolateral
portion of extensor margin, large medial spine ventrally, overreaching prox-

VOLUME 93, NUMBER 3 565

pee Sa
2S

CZ
es

—
==

Fig. 1. Pontophilus gorei, paratype, ovigerous female (eggs not shown), cl 2.1, RMNH
D31979: a, Whole animal, lateral view. Paratype, ovigerous female, cl 2.1, TAMU 2-5944; b,
Carapace, dorsal view; c, Same, lateral view. Holotype, ovigerous female, cl 2.5, USNM
172417; d, Telson and uropods; e, Same, distolateral margin of exopod; f, Same, tip of telson.

566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

O
‘?)

eae one aes SS
SS Sow es

SN

SNINS

5

Fig. 2. Pontophilus gorei, holotype, ovigerous female, cl 2.5, USNM 172417: a, Left an-
tennule; b, Left antennal scale; c, Left mandible; d, Left maxillula; e, Left maxilla; f, Right
maxilliped 1; g, Left maxilliped 2; h, Left maxilliped 3.

VOLUME 93, NUMBER 3 567

q

,

Fig. 3. Pontophilus gorei, holotype, ovigerous female, cl 2.5, USNM 172417: a, Left per-
eopod 1; b, Left pereopod 2: c, Same, dactyl and propodus; d, Left pereopod 3; e, Left
pereopod 4; f, Left pereopod 5; g, Left pereopod 3, tip of dactyl.

568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.5 mm
(a,b,e)

Fig. 4. Pontophilus gorei, allotype, male, cl 1.9, USNM 172418: a, Left pleopod 1; b, Left
pleopod 2. Holotype, ovigerous female, cl 2.5, USNM 172417; c, Left pleopod 1; d, Left
pleopod 2. Paratype, ovigerous female, cl 2.1, FSBC I 21454; e, Left pleopod 5.

VOLUME 93, NUMBER 3 569

imal portion of propodus, rounded lobe with serrate spinules on flexor mar-
gin; merus slightly shorter than propodus, bearing two spines on distal ex-
tensor margin, distal flexor margin terminating in weak spine, strong spine
about midway on flexor margin; ischium and basis short, combined length
4 that of merus. Pereopod 2 (Fig. 3b) setose, short, reaching nearly to distal
margin of merus of first; chela minute, internal margins of fingers concave,
meeting only at tips (Fig. 3c). Pereopod 3 (Fig. 3d) long, slender, overreach-
ing antennal scale by length of dactyl and propodus; dactyl] with tuft of hairs
on flexor margin (Fig. 3g). Pereopod 4 (Fig. 3e) shorter and stouter than
third, only tip of dactyl overreaching antennal scale; dactyl with tuft of hairs
on flexor margin. Pereopod 5 (Fig. 3f) reaching as far as fourth when both
extended anteriorly; dactyl with tuft of hairs on flexor margin. All pereopods
lacking exopods and epipods.

Six branchiae (pleurobranchs) present; inferior apices directed posteri-
orly.

Abdomen (Fig. la) tapering to long, narrow sixth segment; pleura of all
segments broadly rounded ventrally; posterolateral angle of segments 1-4
bluntly rounded, that of segment 5 developed into a blunt tooth; all segments
lack sculpturing.

Pleopods as illustrated (Fig. 4a—e). Endopods of each unsegmented;
length ratio of latter to exopods in female holotype as follows: first 4;
second, */5; third, '2; fourth, 7; fifth, 5. Subapical appendix internae on
pleopods 2-5. For comparisons of relative lengths in the sexes see remarks
below.

Telson (Fig. 1d, f) long, narrow, truncate terminally, bearing 3 pairs of
terminal spines; lateral pair minute, intermediate pair +5 length of mesial
pair; 2 pairs of minute dorsal spines on lateral margins at about % and 45
length. Uropod long, narrow, setose; endopod slightly overreaching telson;
exopod falling just short of telson, lateral margin straight, terminating in
fixed tooth proximal to movable spine at distal end of margin (Fig. le).

Eggs 0.3-0.5 mm in diameter, depending on development.

Type-locality.—The ovigerous female holotype was taken 135 km due
west of Sanibel Island Light in the eastern Gulf of Mexico.

Distribution.—Most specimens were from the two ‘Project Hourglass”’
transects over the continental shelf of the west coast of Florida, off Sanibel
Island and Tampa Bay. A single specimen was taken 167 km off the north-
west coast of Florida in the central Gulf. The single specimen from a fish
stomach (Prionotus rubio) from the Texas coast extends the range of this
species into the western Gulf of Mexico. A single male specimen captured
off Georgia represents the only record for this species from the east coast
of the United States. Although this specimen was taken in 9 m, all Gulf
specimens were taken at greater depths ranging from 37 to 182 m.

570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology.—This species is named in honor of Dr. Robert H. Gore in
recognition of his many contributions to the field of carcinology and his
unselfish aid to others in the field.

Remarks.—The outer flagella of the antennules of males are broad, thick-
ened and distinctly shorter than the inner flagella. Outer flagella of females
are only slightly shorter and thicker than the inner flagella (Fig. 2a).

Sexual dimorphism is also exhibited in the form and development of the
pleopods. Margins of the endopod of the first pleopod of males (Fig. 4a) are
sinuous, terminating about %4 the length of the exopod, but in females (Fig.
4c) the margins are evenly curved and the tip is more lobate, the latter
reaching about + the length of the exopod. The second pleopod of males
bear a well developed appendix masculina and an appendix interna (Fig.
4b); females bear only an appendix interna (Fig. 4d). The relative lengths
of the endopods of pleopods 2—4 of both males and females are somewhat
variable but generally decrease in size sequentially from ¥5 to 4 the lengths
of the exopods in females and from “45 to */; the lengths of the exopods in
the males. The appendix interna arises near the base of the endopod in
males but has a more distal origin in females. The endopod of the fifth
pleopod of adult males is relatively long, varying from 4 to 2 the length of
the exopod; the appendix interna is in the usual basal position. In females,
however, the appendix interna is positioned subapically on the fifth pleopod,
and may extend beyond the endopod; the endopod is reduced, never ex-
ceeding % the length of the exopod (Fig. 4e).

Relative lengths of the endopods are a function of size as well as sex.
Ovigerous females, from 1.8 to 2.2 mm carapace length (cl), vary little. In
non-ovigerous females (recognizable at 1.5 mm cl), relative lengths of the
endopods increase in specimens of up to 1.8 mm cl, whereupon the relative
lengths remain consistent. Between 1.5 and 1.7 mm cl, the appendix interna
may be missing on the posterior pairs of pleopods. The appendix masculina
of the male is present at 1.5 mm cl, although it may be somewhat reduced.
Relative lengths of endopods show the same progressive increase as in fe-
males but continue to increase in specimens up to 2.2 mm cl. Juveniles, less
than 1.4 mm cl, may or may not possess an appendix interna.

Comparisons.—The strong spine on the flexor margin of the first pereopod
easily distinguishes this species from all other western Atlantic species of
Pontophilus. P. dimorphus Fujino and Miyake, 1971 from Sagami Bay,
Japan, seems to be the only described species with such a spine. A similar
spine is figured on the extensor margin by Bate (1888) for P. gracilis Bate,
1888 (=P. challengeri Ortmann, 1893) but Calman (1939) remarks in a foot-
note that none of the type-series of P. challengeri possess this spine. AI-
though P. dimorphus, like P. gorei, lacks an exopod on the first pereopod
the extensive spination on the carapace does not substantiate a close rela-
tionship.

VOLUME 93, NUMBER 3 571

Table 1.—Generic characters of Pontophilus and Philocheras exhibited by Gulf of Mexico
specimens (adapted from Kemp, 1911).

Characters typical of Characters typical of
Pontophilus Philocheras
no}
= 2g
=
3 ae z 23
na) ~ a) Va) -,
= eT a 5 D 2 As r=
= OH ~< ~”n “4o} vo
yz se 3°38 8B oe S44. &
2 SD Oe © Oi OG a Ee 6W 6 a
R Of Eg oe) ee af ©
oO bm he ox Oo iso} no} oe mo) iS} 3
2 DS 30 8S & & Ss Bn Ds So & =
4 ro) of =. = o > = cS = oD
o a ¢ a9 2 2 o ss ON et Ye = 2
s et ayaa) CS ie 5 x Ser Stat =
© aS ig Os oS Oy ie SYS Oe On oS
= © IS Seles sim Ee cy Vs va vo X
Oo no) QR OF & = RD we @O Or Oe cs
3) o & Qo 2 = @ fe fer fa fen IS
} a Oe Os OF ® © £29 99° Ov OF 9
= Jf be SUE Sine Boe
A OM@asMa MS <£ HF BAAS ME we <
Pontophilus gorei n. sp. RO ereX Pe LG) eee xen re OL) LAA
Pontophilus brevirostris Smith, 1881 KEP SEX weaker ERG? 2X oe ee ee ee
Pontophilus gracilis Smith, 1882 KGae eX EP Xe 01 aX NOS i ia a ES ye
Pontophilus talismani Crosnier &
Forest, 1973 OF VO aE Or Oe ae a

The present species is most closely related to a small group of extra-
American species which lack an exopod on pereopod |, possess a single
spine in the dorsal midline of the carapace and lack hepatic spines. These
species, however—Pontophilus monacanthus Holthuis, 1961 from Turkey,
P. opici Crosnier, 1971 from West Africa, P. megalocheir (Stebbing, 1915)
from South Africa and P. hendersoni Kemp, 1915 from South Africa and
India—all possess a bifid subchelar spine while that of P. gorei is simple.
P. gorei also differs from each in the form of the rostrum which is blunt,
expanded at the apex and possesses strong lateral spines midway along its
length.

Discussion.—Kemp (1911), using data on intergrades of the characters
shown in Table 1, concluded that in Asiatic waters it was impossible to
distinguish between the genera Pontophilus and Philocheras. More recently
European workers (Zariquiey Alvarez, 1968: Lagardere, 1971), using the
same characters, regarded the two genera as distinct in Atlantic and Med-
iterranean waters. In view of the ambiguous nature of some of the features
of P. gorei, expressed in Table 1, we can only follow Kemp (1911) and
consider Philocheras Stebbing, 1900 to be a synonym of Pontophilus Leach,
1817. As Kemp (1916) recognized, several natural groupings exist and one
solution to the systematic confusion in this unwieldy genus may lie in the
recognition of several genera.

Acknowledgments

I acknowledge William G. Lyons and David K. Camp, Florida Depart-
ment of Natural Resources Marine Research Laboratory, for the loan of

ye PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

specimens and for their unending patience. Dr. Thomas S. Hopkins (under
BLM Contract AA550-CT7-34), Dauphin Island Sea Lab, and Dr. John N.
Kraeuter, Virginia Institute of Marine Science, also contributed specimens.
I am particularly indebted to Dr. Linda H. Pequegnat, Texas A&M Uni-
versity, for the loan of comparative material and to Dr. L. B. Holthuis,
Ryksmuseum van Natuurlike Historie, for comments on the nature of the
Pontophilus/Philocheras problem. Finally, I thank Dr. Richard W. Heard
for his guidance, Ms. Linda B. Lutz for the illustrations and Dr. F. A.
Chace, Jr. for reviewing the manuscript.

Literature Cited

Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during
the years 1873—1876.—Report on the Scientific Results of the Voyage of H.M.S. Chal-
lenger, 1873-76, Zoology 24: xc + 942 pages, 76 figs., 157 pls.

Calman, W. T. 1939. Crustacea: Caridea.—The John Murry Expedition 1933-34 Scientific
Reports 6(4):183-224, figs. 1-8.

Joyce, E. A., Jr., and J. Williams. 1969. Memoirs of the Hourglass Cruises: Rationale and
pertinent data.—Florida Department of Natural Resources Marine Research Laboratory
1(1):1-50, 26 tables, figs. 1-12.

Kemp, S. 1911. Notes on Crustacea Decapoda. II. Descriptions of two new Crangonidae with

observations on the mutual affinities of the genera Pontophilus and Philocheras.—Rec-

ords of the Indian Museum 6:5-12, | table, pl. 2.

. 1916. Notes on Crustacea Decapoda. VI. Indian Crangonidae.—Records of the Indian

Museum 12:355—384, text-figs. 1-7, pl. 8.

Lagardere, J. P. 1971. Les crevettes des cotes du Maroc.—Travaux de |’Institut Scientifique
Cheérifien et de la Faculté des Sciences 36:7—140, figs. 1-325.

Pequegnat, L. H. 1970. Deep sea caridean shrimps with descriptions of six new species.—In
W.E. Pequegnat and F. A. Chace, Jr., eds., Contributions on the Biology of the Gulf
of Mexico. Texas A&M University Oceanographic Studies 1:59-123, figs. 4-1 to 4-17.

Zariquiey Alvarez, R. 1968. Crustaceos Decapodos Ibéricos.—Investigacion Pesquera 32:i-
xv, 1-510, figs. 1-164 (Barcelona).

Dauphin Island Sea Lab, P.O. Box 386, Dauphin Island, Alabama 36528.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 573-592

FOUR NEW SPECIES OF RENOCILA (ISOPODA:
CYMOTHOIDAE), THE FIRST REPORTED FROM
THE NEW WORLD

Ernest H. Williams, Jr. and Lucy Bunkley Williams

Abstract.—Renocila colini sp. n. is described from the flamefish, Apogon
maculatus (Poey), and the belted cardinal fish, Apogon townsendi (Breder),
from Mona Island, Puerto Rico; R. waldneri sp. n. from the harlequin bass,
Serranus tigrinus (Bloch), from La Caleta, near Santo Domingo, Domini-
can Republic; R. bowmani sp. n. from the harlequin bass from Saona Island,
Dominican Republic; and R. thresherorum sp. n. from Apogon retrosella
(Gill) from Loreto, Baja California Sur, Mexico.

Limited data suggests that members of this genus possess extremely re-
stricted geographic ranges, high rates of infestation, and a high degree of
host specificity. Oostegites of Renocila colini, and possibly other species
in the genus, are formed during a single molt.

Introduction

The genus Renocila is represented by 5 known species, R. dubia (Nier-
strasz, 1931) Barnard, 1936; R. heterozota Bowman and Mariscal, 1968; R.
indica Schioedte and Meinert, 1884; R. ovata Miers, 1880; and R. peri-
ophthalma Stebbing, 1900, from the Indopacific Region. Four new species
of the genus are described from the West Indies and the American Pacific
Regions.

Materials and Methods

Parasitized cardinalfishes were located at night using underwater lights
and scuba. They were collected with quinaldine or microprong spears pro-
pelled by miniature Hawaiian slings. Later in the work, fishes were captured
by forcing them into plastic bags with the face of underwater lights. Para-
sitized harlequin bass were collected during the day with a macroprong
spear and an elastic band speargun or with a microprong spear. Hosts were
immediately sealed in individual plastic bags, and stored in a dive bag for
no longer than 60 minutes. Location of isopods on each host was recorded
on an underwater slate.

Hosts were weighed to the nearest 0.1 g, measured for standard and total
lengths to the nearest mm, and examined for damage associated with the
isopod under 10 power of a dissecting microscope. Isopods were measured

574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

for total length and maximum width to the nearest 0.1 mm, while alive.
They were preserved in 70% ethanol. Eggs and larvae were removed from
female isopods, counted, and a random sample of 10 measured for total
length and maximum width to the nearest 0.01 mm. Mouthparts and ap-
pendages were mounted in glycerine jelly. Drawings of appendages were
made with the aid of a Bausch and Lomb Trisymplex microprojector, whole
specimens were drawn from projections of 35 mm slides made with a Nikon
F2, 55 mm Nikon macro lens and bellows. Telsons of the illustrated species
were drawn in a natural or some what depressed position; therefore, the
length of telsons in the dorsal views do not represent the actual total lengths.

Renocila colini, sp. n.
Figs. 1-27 and 105

Type-host and locality (date and depth).—Flamefish, Apogon maculatus
(Poey), Carmelita, Mona Island (23 April 1976) (10 m.).

Additional hosts and localities (date and depths).—Belted cardinal fish,
Apogon townsendi (Breder), east of Playa Carabinero, Mona Island (20
April 1976) (15 m); Playa Sardinera, Mona Island (22 April 1976) (15 m);
flamefish, Playa Carabinero (27 December 1975) (30 m), east of Playa Car-
abinero (20 April 1976) (15 m), Playa Sardinera (21, 22 and 23 April 1976;
25 May 1977) (17 m); and Carmelita (23 April 1976) (10 m).

Location.—Male-female pair or occasionally female, male, or transitional
specimens attached to dorsal surface along side of the dorsal fin. Male an-
terior to female (Fig. 105).

Specimens studied.—42 (all type-material).

Type-specimens.—Holotype (female), USNM 173920; allotype (associated
male) USNM 173921; 8 paratypes, USNM 173922-25; 32 paratypes in au-
thors’ collection.

Diagnosis.—Anterior margin of head inflexed, not produced into lobe
between bases of antennae 1. Posteroventral angle of pereonites 5—7 pro-
duced, that of pereonite 7 overlapping only pleonite |. Telson '/7 to % wider
than long. Antennae | slightly broader and slightly shorter than antennae 2.
Pereopods 1-3 without swelling in dactyl and without lobe at posterodistal
corners of basis. Pereopods 6—7 subequal in length. Inner ramus of uropod
more than half as long as outer ramus.

Further details.—Antennae | 8-merous. Antennae 2 8-merous. First seg-
ment of mandibular palp expanded, 3rd segment with 7 to 15 stout setae
with broad distal ends along outer margin, and 2 similar, but longer apical
setae; 2nd segment with 3 closely spaced setae on inner margin similar to
marginal setae of 3rd segment. Incisor process of mandible broad pointed
with a fine tip. Maxilla 1 with 4 recurved apical spines. Distal lobes of

VOLUME 93, NUMBER 3 575

Figs. 1-16. Renocila colini, female: 1, Apex of maxilla 1; 2, Maxilla 1; 3, Antennae and
anterior margin of head, ventral; 4, Right uropod, dorsal; 5, Dorsal view; 6, Lateral view; 7,
Apex of mandible; 8, Mandible; 9, Apex of maxilla 2; 10, Maxilla 2; 11, Apex of 3rd segment,
left mandibular palp; 12, Left mandibular palp; 13, Seta of maxilliped; 14, Apex of palp,
maxilliped; 15, Maxilliped; 16, Scales on maxilla 2. (Whole mouthparts and pereopods 28x;
enlargements of portions of mouthparts 280.) (Scales in mm.)

576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

maxilla 2 each with 2 blunt spines, occasionally 1 (Fig. 9). Distal segment
of maxillipedal palp with 3 stout recurved spines.

Penis lobes of male separate. Appendix masculina of male pleopod 2
linear, with unmodified apex (terminal setae on apex of juvenile male). Fe-
males possess a reduced appendix masculina 33 to 67% the length found in
the male.

Color.—Dorsal and appendages of living specimens uniform yellowish
brown.

Variation.—Posteroventral angle of pereonite 4 occasionally slightly pro-
duced. Pereonite 7 occasionally extends slightly beyond pleonite 1. Telson
averages '4 wider than long with a range of '/7 to %. Antennae | occasionally
equal in length to antennae 2, but occasionally much longer than antennae
De

Remarks.—Of the 42 specimens of Renocila colini collected from 29
hosts, 20 were females, 18 males, and 4 transitionals; no juveniles were
collected. Nine females with oostegites were 12.0 to 17.5 mm in length,
mean 16.2 mm; 5.9 to 9.0 mm in width, mean 8.0; lacked penis lobes; and
possessed reduced appendix masculina ranging from approximately 42 to
67% of the length found in the male. Eleven females lacking oostegites were
10.2 to 16.2 mm long, mean 13.2 mm; 4.5 to 9.0 mm in width, mean 6.6 mm;
lacked penis lobes; and possessed reduced appendix masculina ranging from
approximately 33 to 67% the length found in the male. Four transitionals
with reduced penis lobes were 8.0 to 13.0 mm long, mean 10.7 mm; 4.5 to
5.5 mm in width, mean 4.8 mm, and possessed reduced appendix masculina
ranging from approximately 67 to 75% of the length found in male. Males
were 7.5 to 13.0 mm long, mean 9.9 mm, and 2.8 to 4.5 mm in width, mean
3.6 mm.

Only | of the 9 females with oostegites possessed an empty brood pouch.
Numbers of eggs or young in the other females varied from 46 to 175 and
averaged 115. The smallest and apparently least developed brood numbered
157 and were spherical to subspherical embryos 0.83 to 1.08 mm (averaging
0.95 mm) long by 0.74 to 0.88 mm (averaging 0.81 mm) wide. Broods of 3
females, numbering 46 to 152, were oblong embryos 0.99 to 1.18 mm (av-
eraging 1.08 mm) long by 0.81 to 0.91 mm (averaging 0.89 mm) wide. One
female contained 74 oblong embryos with a cephalic end formed, but not
possessing eyes, 1.22 to 1.37 mm (averaging 1.27 mm) long by 0.74 to 0.83
mm (averaging 0.78 mm) wide. Broods of 3 females, numbering 94 to 175,
were larvae with 6 pereonites and a loose cuticle, apparently ready to molt,
2.60 to 2.78 mm (averaging 2.69 mm) long by 0.86 to 0.95 mm (averaging
0.93 mm) wide.

The average feminity index (Legrand, 1951-width/length x 100) of the
males associated with females (N = 13) is 36.7; males not associated with

VOLUME 93, NUMBER 3 S)//7/

Figs. 17-27. Renocila colini: 17, Dorsal view, male; 18, Right uropod, male; 19, Pereopod
1, female; 20, Pereopod 7, female; 21, Distal end, outer ramus of uropod, male; 22, Pereopod
1, male; 23, Pereopod 7, male; 24, Seta of pleopod 2, male; 25, Pleopod 2, male; 26-27, Setae
of pleopod 2, male. (Whole mouthparts and pereopods 28x; enlargements of portions of mouth-
parts 280.) (Scale in mm.)

578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

females (N = 5) 33.6; transitionals (N = 4) 46.4; females not associated with
males (N = 7) 48.8; females associated with males (N = 13) 50.5.

The 27 infested flamefish were 3.8 to 7.1 cm in standard length, and av-
eraged 5.1. Male-female pairs of isopods were equally abundant on all sizes
of hosts. The belted cardinalfish infested were 4.0 and 6.3 cm in standard
length. Damage to the host was minor to unnoticeable, occasionally occur-
ring at the attachment point of the female isopod.

The position of the male and female on the host as discussed above and
shown in Figure 105 was consistent in all the specimens collected and in
numerous other specimens observed in the field. Pairs or individual isopods
occurred as frequently on the left as on the right side of the dorsal fin. Charles
Arneson (pers. comm.) observed a cardinalfish at Mona Island which pos-
sessed 3 Renocila sp. on 1 side of the dorsal fin and 2 on the opposite side.
Unfortunately this specimen was not captured.

Four abnormalities were noted. The first left pleopod of a female with
oostegites was reduced to less than % normal size; and the first right pleopod
of female lacking oostegites was reduced to approximately ’% normal size.
The left pereopod 7 of a female lacking oostegites was reduced to approx-
imately '/s of the size of the normal right pereopod 7. The right uropod of
a transitional specimen was reduced and did not extend to the posterior end
of the telson; the outer ramus was the most reduced portion, the basis the
least reduced portion.

All females with oostegites were associated with males, which suggests
either the presence of a male is necessary to form oostegites in the female,
or the presence or process of producing oostegites attracts males. However,
the one female in the process of molting with 2 of the oostegites exposed
was associated with a male. Oostegite formation seems to be independent
of the size of the female as some of the largest and smallest collected lacked
these structures.

The prolonging of the male stage by the associated female as noted in
Anilocra physodes (Linnaeus) by Legrand (1951) and in Lironeca puhi Bow-
man by Bowman (1959), apparently occurs in Renocila colini. Some of the
males associated with females were longer (lengths 7.5 to 13.0 mm) than
any of singly occurring males (lengths 8.0 to 10.1 mm); which indicates
males associated with females tend to retain male characters to a larger size
than non-associated males.

The specific name is in honor of the discoverer and collector of the first
specimen of this isopod, Dr. Patrick L. Colin.

Renocila waldneri, sp. n.
Figs. 28-52 and 106
Type-host and locality (date and depth).—Harlequin bass, Serranus ti-
grinus (Bloch), La Caleta, near airport, Santo Domingo, Dominican Repub-
lic (23 November 1978 and 17 May 1979) (17 m).

VOLUME 93, NUMBER 3 579

‘

37

Figs. 28-43. Renocila waldneri, female holotype: 28, Apex of maxilla 1; 29, Maxilla 1; 30,
Antennae and anterior margin of head, ventral; 31, Right uropod, dorsal; 32, Dorsal view; 33,
Lateral view; 34, Apex of mandible; 35, Mandible; 36, Apex of maxilla 2; 37, Maxilla 2; 38,
Apex of 3rd segment, left mandibular palp; 39, Left mandibular palp; 40, Seta of maxilliped;
41, Apex of palp, maxilliped; 42, Maxilliped; 43, Scales on maxilla 2. (Whole mouthparts and
pereopods 28x; enlargements of mouthparts 280.) (Scales in mm.)

580 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Location.—Male-female pair or single male, transitional, or female spec-
imens attached to dorsal surface alongside of the dorsal fin. Male in contact
with female along lower anterior side (Fig. 106).

Specimens studied.—18 (all type material).

Type-specimens.—Holotype (female) USNM 173926; allotype (associated
male) USNM 173927; 8 paratypes USNM 173928-32; 8 paratypes in authors’
collection.

Diagnosis.—Anterior margin of head inflexed, not produced into lobe
between bases of antennae |. Posteroventral angle of pereonite 5 moderately
produced, of pereonites 6—7 produced, that of pereonite 7 overlapping ple-
onites 1 and 2. Telson 4% to % wider than long. Antennae 1 much broader
and slightly shorter than or equal in length to antennae 2. Pereopods 1-3
without swelling in dactyl and without lobe at posterodistal corner of basis.
Pereopods 6—7 subequal in length. Outer ramus of uropod slightly longer
than inner ramus.

Further details.—Antennae | 8-merous. Antennae 2 8-merous. First seg-
ment of mandibular palp expanded; 3rd segment with 16 stout setae with
broad distal ends along outer margin, and 3 similar, but longer apical setae;
2nd segment with 3 closely spaced setae on outer margin similar to longer
apical seta of 3rd segment. Incisor process of mandible broad pointed. Max-
illa 1 with 4 recurved apical spines. Dorsal lobes of maxilla 2 each with 2
blunt spines. Distal segment of maxillipedal palp with 3 stout recurved
spines occasionally 4 (Fig. 41).

Penis lobes of male separate. Appendix masculina of male pleopod 2
linear, with unmodified apex, arising near base of endopod, about as long
as endopod. Female possessed a reduced appendix masculina 33 to 67% the
length found in the male.

Color.—Dorsal of living specimens uniform brown, appendages yellowish
brown.

Variation.—Posteroventral angle of pereonite 5 occasionally not pro-
duced. Telson averages 2 wider than long with a range of 14 to %.

Remarks.—Of the 18 specimens of Renocila waldneri collected from 12
hosts, 9 were females, 8 males, and | a transitional; no juveniles were
collected. Four females with oostegites were 15.3 to 19.3 mm in length mean
17.4 mm; 8.0 to 9.3 mm in width, mean 8.6 mm; lacked penis lobes; and
possessed reduced appendix masculina ranging from approximately 33 to
50% of the length found in the male. Five females lacking oostegites were
12.7 to 15.7 mm long, mean 14.6 mm; 6.8 to 8.1 mm in width, mean 7.6 mm;
lacked penis lobes; and possessed reduced appendix masculina ranging from
approximately 33 to 67% the length found in the male. A transitional 10.8
mm long and 5.8 mm wide with reduced penis lobes, possessed a reduced
appendix masculina approximately 63% of the length found in the male.

VOLUME 93, NUMBER 3 581

Males were 5.0 to 10.8 long, mean 8.1 mm; 1.5 to 4.9 mm in width, mean
3.3 mm.

Two females possessed 221 and 279 spherical to subspherical embryos
0.93 to 1.08 mm (averaging 0.98) long by 0.85 to 0.99 mm (averaging 0.93)
wide. One female contained 155 oblong embryos with a cephalic end formed,
but not possessing eyes, 1.34 to 1.54 mm (averaging 1.44) long by 0.72 to
0.82 mm (averaging 0.77) wide. The marsupium of one female contained 135
larvae with 6 pereopods and 13 embryos with eyes, but no appendages.
Internal segmentation was apparent in the embryos, which were evidently
ready to molt. Embryos were 1.10 to 1.20 mm (averaging 1.15 mm) long by
0.72 to 0.82 mm (averaging 0.77 mm) wide. Larvae were 2.11 to 2.26 mm

_ (averaging 2.16) long by 0.77 to 0.86 mm (averaging 0.82 mm) wide.

The average feminity index of the males associated with females (N = 6)
was 38.0; males not associated with females (N = 2) 44.9; transitionals (N =
1) 53.7; females not associated with males (N = 3) 51.7; females associated
with males (N = 6) 49.7.

The 12 infested harlequin bass were 5.7 to 8.1 cm in standard length and
averaged 7.0 cm. Damage to the host was very slight to unnoticeable.

Approximately 10% of the dorsal surface of two gravid female R. waldneri
were covered with growths of algae. Twelve species of algae occurred in

| these growths (David L. Ballantine, pers. comm.). Numerous mites (Ar-

achnoidea: Acarina) were associated with either the algae or these two iso-
pods.

The distribution of R. waldneri seemed extremely limited. Extensive ob-
servations along the 17 m depth contour approximately 800 m to the north

and south of the type locality indicated general habitat and abundance of

the harlequin bass which were very similar to the type-locality, but no R.
waldneri were observed. Renocila waldneri were not observed at 6 other

_ localities on the south coast of the Dominican Republic.

The specific name is in honor of Raymond E. Waldner, who first noted

_ this isopod during a scuba dive with the authors.

Renocila thresherorum, sp. n.
Figs. 53-79

Type-host and locality (date and depth).—Apogon retrosella (Gill), Lo-
reto, Baja California Sur, Mexico (13 October 1978) (4.6 m).

Location.—Male-female pair (and one singly occurring specimen; not col-
lected), attached to dorsal surface alongside of the dorsal fin. Male and
female on either side of the dorsal fin.

Specimens studied.—2.

582 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 44-52. Renocila waldneri: 44, Dorsal view, male allotype; 45, Pereopod 1, female
holotype; 46, Pereopod 7, female holotype; 47, Right uropod, male allotype; 48, Right uropod,
male paratype; 49, Pereopod 7, male allotype; 50, Pereopod 1, male allotype; 51, Pleopod 2,
male allotype; 52, Distal end, outer ramus of uropod, male paratype. (Whole mouthparts and
pereopods 28x; enlargements of portions of mouthparts 280.) (Scale in mm.)

VOLUME 93, NUMBER 3 583

Figs. 53-68. Renocila thresherorum, female holotype: 53, Apex of maxilla 1; 54, Maxilla
1; 55, Antennae and anterior margin of head, ventral; 56, Right uropod, dorsal; 57, Dorsal
view; 58, Lateral view; 59, Apex of mandible; 60, Mandible; 61, Apex of maxilla 2; 62, Maxilla
2; 63, Apex of 3rd segment, left mandibular palp; 64, Seta of maxilliped; 65, Apex of palp,
maxilliped; 66, Maxilliped; 67, Scales on maxilla 2; 68, Left mandibular palp. (Whole mouth-
parts and pereopods 28x; enlargements of portions of mouthparts 280.) (Scales in mm.)

584

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Yj QJ J

DM

Figs. 69-79. Renocila thresherorum: 69, Dorsal view, male allotype; 70, Right uropod,
male allotype; 71, Pereopod 1, female holotype; 72, Pereopod 7, female holotype; 73, Pereopod
1, male allotype; 74, Pereopod 7, male allotype; 75, Seta of pleopod 2, male allotype; 76,
Pleopod 2, male allotype; 77, Distal end outer ramus of uropod, male allotype; 78-79, Setae
of pleopod 2, male allotype. (Whole mouthparts and pereopods 28x, enlargements of portions
of mouthparts 280.) (Scale in mm.)

VOLUME 93, NUMBER 3 585

Type-specimens.—Holotype (female) USNM 173933; allotype (associated
male) USNM 173934.

Diagnosis.—Anterior margin of head inflexed, not produced into lobe
between bases of antennae 1. Posteroventral angle of pereonite 5 not pro-
duced, of pereonite 6 moderately produced, and of pereonite 7 produced,
that of pereonite 7 overlapping pleonite |. Lateral margins of pereonites 2-3
somewhat notched, of 4—7 notched. Telson 4 wider than long. Antennae |
much broader and slightly shorter to equal in length with antennae 2. Pe-
reopods 1-3 without swelling in dactyl and without lobe at posterodistal
corner of basis. Pereopods 6—7 subequal in length. Outer ramus longer than
inner ramus of uropod.

Further details.—Antennae 1 8-merous. Antennae 2 8-merous. First seg-
ment of mandibular palp slightly expanded; 3rd segment with 14 stout setae
with broad distal ends along outer margin and 2 similar, but longer apical
setae. Incisor process of mandible broad pointed with a fine tip. Maxilla 1
with 4 recurved apical spines. Distal lobes of maxilla 2 with 2 and 3 blunt
spines. Distal segment of maxillipedal palp with 3 stout recurved spines.
Penis lobes of allotype male fused medially, forming a short bilobate pro-
cess. Appendix masculina of male pleopod 2 linear, with unmodified apex,
arising near base of endopod, about as long as endopod. The female (ho-
_lotype) possessed a greatly reduced appendix masculina.

Color.—Dorsal of living specimens dark brown, more intensely marked
on margins of pereonites and pleonites. Telson light brown centrally and on
distal end. Appendages light brown.

Remarks.—The female was 16.7 mm in length and 9.5 mm in width, the

associated male was 12.0 mm in length and 4.3 mm in width, and the host
_ was 5.2 cm in standard length.
The marsupium of the female contained 147 larvae with 6 pereopods and
33 embryos with eyes, but no appendages. Internal appendages and seg-
mentation were apparent on specimens of the later group of young when
cleared in glycerine jelly. Apparently the group of young were in the process
of molting when preserved. Embryos were 1.47 to 1.81 mm long and aver-
aged 1.70 mm, and 0.78 to 0.88 mm wide and averaged 0.85 mm. Larvae
_ were 2.55 to 2.84 mm long and averaged 2.67 mm, and 0.83 to 0.98 mm wide
_and averaged 0.88 mm. The female possessed a rudimentary appendix mas-
culina.

The specific name is in honor of the discoverers and collectors of this
isopod Dr. Ronald E. and Ann G. Thresher.

Renocila bowmani, sp. n.
Figs. 80-104 and 107

Type-host locality (date and depth).—Harlequin bass, Serranus tigrinus

586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 80-95. Renocila bowmani, female holotype: 80, Apex of maxilla 1; 81, maxilla 1; 82,
Antennae and anterior margin of head, ventral; 83, Right uropod, dorsal; 84, Dorsal view; 85,
Lateral view; 86, Apex of mandible; 87, Mandible; 88, Apex of maxilla 2; 89, Maxilla 2; 90,
Apex of 3rd segment, left mandibular palp; 91, Seta of maxilliped; 92, Apex of palp; maxilliped;
93, Maxilliped; 94, Scales on maxilla 2; 95, Left mandibular palp. (Whole mouthparts and
pereopods 28x; enlargements or portions of mouthparts 280.) (Scales in mm.)

VOLUME 93, NUMBER 3 587

Figs. 96-104. Renocila bowmani: 96, Dorsal view, male allotype; 97, Right uropod, male
allotype; 98, Pereopod 1, female holotype; 99, Pereopod 7, female holotype; 100, Pereopod 1,
male allotype; 101, Pereopod 7, male allotype; 102, Pleopod 2, male allotype; 103-104, Setae
_ of pleopod 2, male allotype. (Whole mouthparts and pereopods 28x; enlargements of portions
of mouthparts 280.) (Scale in mm.)

588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Bloch), south central coast of Saona Island, Dominican Republic (18 May
1979) (10.5 m).

Location.—Male-female pair attached to dorsal surface alongside of the
dorsal fin. Male in contact with female along lower anterior side (Fig. 107).

Specimens studied.—2.

Type-specimens.—Holotype (female) USNM 173935; allotype (associated
male) USNM 173936.

Diagnosis.—Anterior margin of head inflexed, not produced into a lobe
between bases of antennae 1. Posteroventral angle of pereonites 5—7 pro-
duced, that of pereonite 7 overlapping pleonites 1-3. Telson 34 wider than
long. Antennae 1 much broader and slightly longer than antennae 2. Per-
eopods 1-3 without swelling in dactyl and without lobe at posterodistal
corner of basis. Pereopods 6—7 subequal in length. Outer ramus of uropod
longer than inner ramus.

Further details.—Antennae | 8-merous. Antennae 2 7-merous. First seg-
ment of mandibular palp expanded; 3rd segment with 16 stout setae with
broad distal ends along outer margin, and 2 similar, but longer apical setae.
Incisor process of mandible broad pointed with a fine tip. Maxilla | with 4
recurved apical spines. Dorsal lobes of maxilla 2 each with 2 blunt spines.
Distal segment of maxillipedal palp with 3 stout recurved spines.

Penis lobes of male separate. Appendix masculina of male pleopod 2
linear, with unmodified apex, arising near base of endopod, about as long
as endopod. Female (holotype) possessed a reduced appendix masculina
approximately 4 the size occurring in the male.

Color.—Dorsal of living specimens and appendages uniformly black.

Remarks.—The female was 18.0 mm in length and 9.2 mm in width; the
associated male was 11.5 mm in length and 4.8 mm in width; and the host
was 7.8 cm in standard length. The female did not possess a marsupium.

Eight of 48 harlequin bass observed on a uniform rock bottom 10.0 to
10.5 m deep with dense growths of soft corals and scattered small coral
heads were infested with R. bowmani. All infestations consisted of male-
female pairs. Density of harlequin bass was low averaging 5 to 7 along 100
m by 5 m transects. The fish seemed to be somewhat clustered in some
areas and absent in others. Infested hosts were normally isolated with only
2 parasitized fishes occupying adjacent territories.

=>

Fig. 105 (top): Male and female Renocila colini on the flamefish, Apogon maculatus (Poey),
underwater photograph; Fig. 106 (middle): Male and female Renocila waldneri, sp. n., on the
harlequin bass, Serranus tigrinus (Bloch), photograph in ship laboratory; Fig. 107 (bottom):
Male and female Renocila bowmani, sp. n., on the harlequin bass, Serranus tigrinus (Bloch),
underwater photograph.

VOLUME 93, NUMBER 3

590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The specific name is in honor of Dr. Thomas E. Bowman and his work
with cymothoid isopods of fishes.

Discussion

Bowman and Mariscal (1968) found Renocila heterozota only in Port Vic-
toria, Seychelles, although Mariscal had examined numerous other popu-
lations of Amphiprion spp. in the Pacific and Indian Oceans. In the present
study R. colini was only observed at Mona Island and R. waldneri and R.
bowmani were only observed at the separate locations off the south coast
of the Dominican Republic, although extensive collections and examinations
were conducted throughout Puerto Rico, Desecheo Island, Caja de Muertos,
Culebra Island, Vieques Island; St. Thomas; St. John; St. Croix; Virgin
Gorda; Anagada; Chub Cay, Eleuthera, Cat Island, Conception Isle, Rum
Cay, Crooked Island, Long Island, Great Exuma, Great Inagua, Little Ina-
gua, Acklins Island, and Long Island, Bahamas; and Santa Marta and Car-
tagena, Colombia. Renocila heterozota occurred on 5 to 10% of the single
species of host infested (Bowman and Mariscal, 1968); R. colini occurred
on 25 to 33% of 2 host species; R. waldneri on 40 to 50% of one host species
and R. bowmani on 16.7% of one host species. On the 4 species Renocila
for which any biological data is available, a pattern of extremely limited
geographic range, high levels of infestation, and strong host specificity is
indicated. More data is needed for the remaining species to determine the
extent of this pattern in other members of the genus.

Another genus of external parasitic isopods in the West Indies, Anilocra,
contrasts markedly with the 3 species of Renocila described from this area.
The 8 species of Anilocra possess wide geographic ranges, low levels of
infestation, and usually specificity to several species in 1 or more genera or
families of hosts. They also differ by being generally much larger and by
infesting much larger hosts (Williams and Williams, unpubl. data). The
known populations of Renocila bowmani and R. waldneri are separated by
less than 100 km; and R. bowmani and R. colini by less than 50 km.
Possibly other isolated species of Renocila will be discovered when the
remainder of the West Indies is thoroughly examined for external isopods of
fishes.

Females of Renocila colini, R. waldneri, R. thresherorum and R. bowmani
possess a reduced appendix masculina. Retention of this structure has not
been noted in members of the genus Renocila, but has been discussed by
Trilles (1964).

Menzies, Bowman, and Alverson (1955) suggested that oostegites were
produced during a single molt in Lironeca convexa Richardson, 1905, be-
cause they failed to find incompletely developed oostegites. In Renocila
colini one female specimen possessed fully formed oostegites on pereonites

VOLUME 93, NUMBER 3 591

5 through 7 which had molted, but not on pereonites 1 through 4 which had
not molted. Apparently the oostegites of Renocila colini are formed during
a Single molt, and possibly this may be the case for other, if not all, members
of the genus.

Juveniles of Renocila colini and R. waldneri have not been collected from
or observed on their host species. These isopods may not settle on their
final host until they developed into males. Also lack of bone deformation
suggests that these isopods do not become associated with juveniles of their
final host as juvenile isopods. Possibly intermediate hosts are involved,
because a prolonged planktonic existence is not compatible with the very
restricted distributions of these isopods.

Key to the Species of Renocila

The key is modified from a previous key by Bowman and Mariscal (1968)
prepared when only 5 species were known in the genus.

la. Dactyls of pereopods 1-3 with swelling on outer margin......... 2
lb. Dactyls of pereopods 1—3 without swelling ..................... 4
Pe AMtennaculasiOnteh enanvalntemMac 2 ics 45min. 55 6c dubia
PD ntennacslwlongen than amtemmac 2.) sce s..4 seb e osicuut tees ancy ks 3
3A, Telsom wir ey ieee eee Cee eee cence Bee ec eee rete oe indica
Bbahel Sonu oneenthanawiGer nu. soc ccncee4 sei es acke oF periophthalma
4a. Posteroventral angle of pereonite 7 reaching pleonite 1 .......... 5

4b. Posteroventral angle of pereonite 7 reaching pleonite 2 or beyond 6
5a. Posteroventral angle of pereonites 5 produced, coxae of pereonites
GTO AG POLS Oeste cei see Pace Sei en cg nes, es SR aes aks colini
5b. Posteroventral angle of pereonite 5 not produced, coxae of pere-
OMiKeSsO—/ MAnhOwW-POlNled mas pe ae ee cians one thresherorum
6a. Posteroventral angle of pereonite 7 reaching base of telson .... ovata
6b. Posteroventral angle of pereonite 7 reaching pleonite 2 or 3...... 7/
7a. Outer ramus of uropod more than twice as long as inner ramus .
515: Sig SPSS DOU o O°G ee Oe Eo ce heterozota

7b. Outer ramus of uropod only slightly longer than inner ramus..... 8
8a. Brown in color, antennae 2 8-merous, antennae | slightly shorter

fiANeAMTeniINnacIort. was wees ete, OS Site. ore ee. eee waldneri

8b. Black in color, antennae 2 7-merous, antennae | slightly longer

LMA AMICK ete rele 28k OP SOP EEE. SIE, Ah ie, bowmani

Acknowledgments

Thanks are expressed to Michael J. Dowgiallo, Raymond E. Waldner,
Cecilio Diaz Carela, Adolphus G. Bunkley, Jr., Dr. Patrick L. Colin, A.
Charles and Deborah A. Arneson, Tracey J. Wolfe, Dr. Ronald E. and Ann

592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

G. Thresher for help in collecting isopods; and to Charles Arneson for one
of the underwater photographs. Appreciation is expressed to Dr. Thomas
E. Bowman, U.S. National Museum of Natural History for reviewing the
manuscript. This project was supported in part by a grant from Dr. Ismael
Almodovar, President of the University of Puerto Rico.

Literature Cited

Bowman, T. E. 1959. Description and notes on the biology of Lironeca puhi n. sp. (Isopoda:
Cymothoidae), parasite of the Hawaiian moray eel, Gymnothorax eurostus (Abbott).—
Crustaceana 1:83-91.

, and R. N. Mariscal. 1968. Renocila heterozota, a new cymothoid isopod, with notes
on its host, the anemone fish, Amphiprion okallopisos, in the Seychelles.—Crustaceana
14:97-104.

Legrand, J. J. 1951. Etude statisque et expérimentale de la sexualité d’ Anilocra physodes L.
(Crustac, Isopode, Cymothoidé).—Bull. Soc. Hist. Nat. Toulouse 85:176—183.

Menzies, R. J., T. E. Bowman, and F. G. Alverson. 1955. Studies of the biology of the fish
parasite Livoneca convexa Richardson (Crustacea, Isopoda, Cymothoidae).—Wassmann
Jour. Biol. 13:277-295.

Trilles, J. P. 1964. Sur la présence et l’evolution régressive de l’appendix masculina chez les
Isopodes Cymothoidae.—C. R. Hebd. Séanc. Acad. Sci., Paris 258:5739-5741.

University of Puerto Rico, Department of Marine Sciences, Mayaguez
00708.

|

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 593-596

A NEW STROBILOPS (MOLLUSCA: PULMONATA:
STROBILOPSIDAE) FROM BAJA
CALIFORNIA SUR, MEXICO

Walter B. Miller and Carl C. Christensen

Abstract.—Strobilops californica, a new species of pulmonate land snail
inhabiting the Sierra de la Victoria of Baja California Sur, Mexico, is de-
scribed.

In December 1973, while searching in the high mountains of the Cape
Region of Baja California, Mexico, for live specimens of Rabdotus beldingi
(Cooper, 1892), we discovered a single dead shell of an unfamiliar species
of small snail. We were successful in collecting living R. beldingi and other
terrestrial mollusks but were unable to find additional specimens of the
unknown small species. Subsequent examination by Alan Solem, Field Mu-
seum of Natural History, revealed that this unique shell is that of a species
of Strobilops, a genus only sparsely distributed in western North America
and not previously known to inhabit Baja California. Comparison with other
west Mexican Strobilops demonstrated the Baja California form to be an
undescribed species.

We collected this snail in the higher elevations of the Sierra de la Victoria,
a region only rarely visited by malacologists; it is therefore unlikely that
additional material of this new species will soon become available for study.
While we are reluctant to describe a new species on the basis of a single
specimen, we believe that the relative inaccessibility of the habitat of this
species and the desirability of documenting its occurrence in Baja California
justify such action at this time.

Strobilops californica, new species
Fig. 1

Description.—Shell large for genus, solid, trochiform, sharply carinate,
much wider than high, finely and closely ribbed above periphery, with about
140 riblets on body whorl and 120 on penultimate whorl, convex and finely
ribstriate below. It is widely umbilicate, the umbilicus contained about 3
times in diameter. Parietal callus thick, prominent where the strong parietal
lamella joins it. No infraparietal or interparietal lamellae visible from the
aperture, and none can be seen by transmitted light. Nearly a half whorl
behind the aperture, a series of 4 basal folds can be detected by transmitted
light. The innermost 3 are relatively short, subequal, parallel and equally

594 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sm, “
Zan)

Alp

RSF
lit 244

4.0 mm

Fig. 1. Holotype of Strobilops californica, USNM 799595: A, Dorsal view; B, Side view;
C, Umbilical view, with basal folds seen in transparency.

spaced, running for about one sixth of a whorl; the outer fold is longer,
running for about one fourth of a whorl. Whorls 6%, gradually increasing
in size, first 2 worn and smooth, last descending abruptly to aperture. Ap-
erture is rounded-lunate, peristome thick, only very slightly expanded.

Holotype.—USNM No. 799595; diameter 3.95 mm, height 1.85 mm, width
of umbilicus 1.3 mm.

Type-locality.—La Laguna, Sierra de la Victoria, Baja California Sur,
Mexico; along slope above creek immediately behind La Laguna shelter.
Elevation ca. 6,500 ft., 31 December 1973, W. B. Miller and C. C. Chris-
tensen, collectors.

Taxonomy

The subgeneric classification of Strobilops was summarized by Pilsbry
(1927-1935, and 1948). He recognized three subgenera, namely Strobilops
Pilsbry, 1893, Discostrobilops Pilsbry, 1927, and Enteroplax Gude, 1899.

Zilch (1959) raised Enteroplax to generic rank, and Solem (1968) reviewed
this genus in detail. Among the characteristics of Enteroplax delineated by
Solem are the low conic shell, prominent radial ribs above threaded periph-
ery and in umbilicus, sculpture absent on body whorl below periphery,
strongly elevated edge of parietal callus which fuses with the upper parietal
lamella, and the presence of 2 parietal lamellae. Enteroplax is currently
known only from the Philippine Islands and New Guinea.

In the genus Strobilops, the subgenus Discostrobilops is strongly de-
pressed, subdiscoidal, and widely umbilicate, while Strobilops s.s. has tro-
chiform shells with a narrow to moderate umbilicus.

Although S. californica has many characters of Enteroplax, namely the
low conic shell, the strongly elevated edge of the parietal callus, and a wide
umbilicus, it does not have a threaded periphery and it does have a sculpture
of fine radial striae on the body whorl below the periphery. Because of these

VOLUME 93, NUMBER 3 595

distinctions, we consider S. californica to belong properly in the genus
Strobilops. However, it does not appear to fit perfectly in either of the
described subgenera of Strobilops. Since we do not feel justified in describ-
ing a new subgenus on the basis of a single shell without anatomical data,
we believe that subgeneric classification should be deferred until additional
specimens, including anatomies, can be obtained.

Differential Diagnosis

Other species of Strobilops inhabiting western Mexico are found on So-
corro Island about 400 km southwest of the southern tip of Baja California,
and, presumably, in Sinaloa.

The species presumed from Sinaloa, Strobilops sinaloa Morrison, 1953,
was described from material found by U.S. Department of Agriculture in-
spectors on a shipment of plants said to have originated in Sinaloa, Mexico.
It is a Discostrobilops and has the depressed shell of that subgenus.

Specimens of Strobilops from Socorro, one of the Revillagigedo Islands,
have been cited in print under three names. Dall (1926) identified specimens
from a single locality (CAS Loc. 24,782) as S. labyrinthica (Say) and S.
strebeli (Pfeiffer). Pilsbry (1927-1935) described material from the same So-
corro locality as S. hannai, stating that this was the species identified by
Dall as S. labyrinthica and that Dall’s S. strebeli was probably also this
species. We have examined the holotype of S. hannai in the collection of
the Academy of Natural Sciences of Philadelphia (ANSP 256587a) and the
Socorro Island specimens in the California Academy of Sciences collection
identified by Dall as S. labyrinthica and S. strebeli. We found that the two
specimens identified as §. labyrinthica were unquestionably that species;
the two specimens identified as S. strebeli, however, were more widely
umbilicate and more depressed than typical S. strebeli and are probably S.
hannai, as Pilsbry suggested. Unfortunately, we did not have the holotype
of S. hannai at the same time that we were examining Dall’s specimens so
that a direct comparison could not be made. Nevertheless, it was immedi-
ately obvious that Dall’s “‘S. strebeli’’ from the CAS and Pilsbry’s S. hannai
from the ANSP were both only about two-thirds the diameter of S. califor-
nica; they were also more coarsely ribbed and narrowly umbilicate.

Discussion

At the type-locality the vegetation consisted primarily of oaks (Quercus
devia and Q. tuberculata) and pinon pines (Pinus cembroides). We found
many live specimens of R. beldingi as well as many shells of R. levis (Dall,
1893), R. montezuma (Dall, 1893), Pseudosubulina eiseniana (Cooper,
1893), Glyphyalinia cf. paucilirata (Morelet, 1851), and live animals of Der-
oceras cf. laeve (Miiller, 1774). This faunal association of the high oak-pine

596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

forest was understandably different from the lower desert communities of
Baja California (Christensen, 1979).

The significance of finding S. californica in that locality, however, was
not apparent to us at the time as we did not recognize it in the field as a
Strobilops and because we were intent on finding R. beldingi. Our failure
to find additional specimens of S. californica was due to our concentration
of collecting effort in the vicinity of rock outcrops (typical Rabdotus habitat)
rather than under fallen logs and leaf mold, where Strobilops would be more
likely to occur.

The single specimen of S. californica is an adult shell, solid, and recent,
as indicated by fresh periostracum in the umbilicus and on the base of the
body whorl.

Acknowledgments

We are grateful to Barry Roth for the loan of the California Academy of
Sciences specimens, to Joseph Rosewater for facilitating our examination
of specimens at the United States National Museum, and to George Davis,
Robert Robertson, and the staff of the Academy of Natural Sciences of
Philadelphia for facilitating our examination of the ANSP collection. The
visit to Philadelphia by one of us (CCC) was supported by the Jessup Fund
of the ANSP.

Literature Cited

Christensen, C. C. 1979. A preliminary analysis of the land mollusks of Baja California Sur,
Mexico.—Annual Report of the Western Society of Malacologists (for 1978) 11:15—20.

Dall, W. H. 1926. Land shells of the Revillagigedo and Tres Marias Islands, Mexico.—Proc.
Calif. Acad. Sci. (4) 15(15):467-491, pl. 35-36.

Hanna, G. D. 1922. Notes on the anatomy of Strobilops labyrinthica (Say).—Nautilus 35:91—
93.

Morrison, J. P. E. 1953. Two new American species of Strobilops.—Nautilus 67:53—55.

Pilsbry, H. A. 1927-1935. Manual of Conchology (Strobilopsidae), 28:1—63.—Acad. Nat. Sci.

Philadelphia, PA.

. 1948. Land Mollusca of North America (north of Mexico).—Acad. Nat. Sci. Phila-

delphia Monogr. 3, 2(2): i—xlvii + 521-1113.

Solem, A. 1968. ‘‘Ptychodon’’ misoolensis Adam and van Benthem Jutting, 1939, a New
Guinea strobilopsid land snail and review of the genus Enteroplax.—Veliger 11(1):24—
30.

Zilch, A. 1959. Handbuch der Palaozoologie. 2(1):1—200. Berlin, Germany (Bebr. Borntrae-
ger).

(WBM) Department of General Biology, University of Arizona, Tucson
85721; (CCC) Department of Zoology, Bernice P. Bishop Museum, Hono-
lulu, Hawai.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 597-608

A GIANT EXTINCT INSECTIVORE FROM CUBA
(MAMMALIA: INSECTIVORA:SOLENODONTIDAE)

Gary S. Morgan, Clayton E. Ray, and Oscar Arredondo

Abstract.—A femur, identified as that of a previously unknown giant so-
lenodontid insectivore, is reported from a fossil deposit in Cuba containing
a typical Greater Antillean late Pleistocene mammalian fauna. The fossil is
closest in morphology to Solenodon cubanus among known insectivores
Lack of adequate material precludes description of a new taxon, althougt.
the femur probably represents an undescribed species of Solenodon. Based
on measurements of the femur, the giant Cuban solenodontid would have
been considerably larger than any living member of the Insectivora. Addi-
tion of a new very large insectivore suggests a substantial radiation of in-
sectivores in the Greater Antilles, similar to that of capromyid rodents and
of megalonychid sloths.

This report is based on a partial femur (USNM 299480) from western
Cuba belonging to a previously unknown mammal. The specimen was col-
lected on 15 March 1959 by Oscar Arredondo and César Garcia del Pino
from the Abra de Andrés, Las Alturas de Esperon, Mesa de Anafe, Sierra
del Rosario, near the city of Guanajay, Pinar del Rio Province, Cuba. Ac-
cording to the new geographic subdivision of Cuba, this locality is now in
Habana Province, but because maps showing the new Cuban provinces are
not generally available at present, we will use the more conventional bound-
aries and names of the Cuban provinces. The femur was collected from a
reddish-colored breccia deposited in a crevice in a rock wall of Miocene
age. A late Pleistocene age is suggested for the breccia based on the ver-
tebrate fossils collected from it. The associated vertebrate fauna includes
three species of small megalonychid ground sloth, Megalocnus rodens,
Mesocnus sp. and Neocnus gliriformis (for use of Neocnus rather than its
synonyms Microcnus and Cubanocnus see Varona, 1976), and two species
of capromyid rodent, Geocapromys columbianus and Capromys sp. (either
C. pilorides or C. prehensilis). All of these species have been recovered
from late Pleistocene cave deposits elsewhere in Cuba and, with the excep-
tion of Capromys, all are now extinct. Although much paleontological field
work has been conducted throughout Cuba during the past 20 years, no

additional specimens referable to this unique mammal have yet come to
light.

Description of femur.—The fossil femur which is the subject of this study
lacks the head, much of the lesser trochanter, and the medial condyle (Fig.

598 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1). It is relatively short and massive with a prominent greater trochanter,
third trochanter, trochanteric fossa, and intertrochanteric crest. The femoral
shaft is straight for most of its length, but is flexed slightly anteriorly near
the proximal end. The shaft is nearly hemicircular in cross section, convex
on the anterior surface and almost flattened on the posterior surface. The
greater trochanter appears to have projected approximately 4-5 mm above
the femoral head and is separated from the head by a deep groove on the
anterior surface. In anterior aspect, the greater trochanter is broad at its
base narrowing proximally to a relatively sharp prominence. On the anterior
surface of the greater trochanter just distal to the proximal end there is a
small, transversely elongate protuberance, presumably for attachment of
the gluteus minimus. In lateral view, the greater trochanter is flexed some-
what anteriorly and is broadly rounded proximally. The trochanteric fossa
is well developed, forming a very deep pit in the posterior surface of the
greater trochanter. The intertrochanteric crest is prominent and composed
of two portions; the anterior part is a thin ridge of bone arising at the tip of
the greater trochanter and forming the posterior border of the trochanteric
fossa and the distal portion of the crest is transverse to the shaft and is
gently concave. The vertical and horizontal portions of the intertrochanteric
crest meet at nearly a right angle (85°) just proximal to the third trochanter.
Although broken off near its base, the lesser trochanter appears to have
been strongly developed and to have met the medial edge of the shaft at
approximately a right angle. The lesser trochanter is located slightly higher
on the shaft than is the third trochanter. The third trochanter is a very
prominent triangular-shaped process which extends about one-sixth the
length of the femur. On the distal end, the patellar groove, although partially
missing, is relatively narrow, slightly concave, and projects anteriad of the
femoral shaft. Proximal to the patellar trochlea there is a deep pit for re-
ception of the patella during strong extension of the leg. The distal end of
the femur is deep anteroposteriorly, but is not particularly broad. The ar-
ticular surface of the lateral condyle is vertical and relatively narrow, where-
as the intercondylar notch is comparatively broad.

Comparison with other mammals.—Although incomplete, this specimen
retains enough diagnostic features to permit detailed comparisons with other
mammalian groups. We have compared the fossil femur with femora of
representative genera of all orders of native mammals known from the late

Baas

Fig. 1. A, C, E, G.—Posterior, anterior, lateral, and medial views of giant solenodontid
femur from the Abra de Andrés, near Guanajay, Pinar del Rio Province, Cuba (USNM 299480);
B, D, F, H.—Posterior, anterior, lateral, and medial views of the left femur of Solenodon
cubanus (USNM 49508). All views x7/e.

599

VOLUME 93, NUMBER 3

600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pleistocene and modern fauna of the West Indies. Only the Marsupialia,
Insectivora, Chiroptera, and Rodentia are represented in the contemporary
fauna of the West Indies. Primates and Edentata are added when late Pleis-
tocene faunas are included. Specimens from the following collections were
used in this study: National Museum of Natural History (USNM), Carnegie
Museum of Natural History (CM), Florida State Museum (UF), and the
personal collection of Oscar Arredondo (OA).

The femora of Chiroptera can be excluded from further consideration for
obvious reasons such as their small size, very slender shaft, and highly
derived morphology of the proximal end. The only marsupials in the West
Indies are the didelphid genera Didelphis and Marmosa, both known in this
region only from the Lesser Antilles. The femur of these marsupials is quite
different from that of the fossil. The femoral shaft is much longer and more
slender, the third trochanter is virtually absent, the greater trochanter ex-
tends only slightly proximal to the head, and the patellar trochlea is very
broad, as is the articular facet of the lateral condyle. Many features of the
femora of New World monkeys, the elongate shaft, reduced third trochanter
located in a more proximal position, and the comparatively broad patellar
trochlea, preclude placement of the fossil in the Primates. In the rodent
femora examined, the lesser trochanter is located primarily on the posterior
surface of the femoral shaft, the third trochanter is vestigial and located far
distal to its position in the fossil, and the patellar groove is much deeper.
Small, extinct ground sloths of the family Megalonychidae are the only
representatives of the Edentata in the West Indies. The femur in these small
sloths differs strikingly from that of the fossil in such features as the relative
proportions of the shaft, the weak development of the greater and lesser
trochanters, trochanteric fossa, and intertrochanteric crest, and the shape
of the patellar groove. Only in the femora of certain genera in the order
Insectivora can a reasonably close match be found for the Cuban fossil
femur.

Femora of all living insectivore families were available for study, with the
number of genera examined for each family following it in parentheses:
Solenodontidae (1), Tenrecidae (7), Chrysochloridae (4), Soricidae (11), Tal-
pidae (7), and Erinaceidae (7). Following Butler (1972) and McKenna (1975)
the tree shrews (Tupaiidae) and elephant shrews (Macroscelididae) are ex-
cluded from the Insectivora (sensu stricto, 1.e. restricted to the Lipotyphla),
but in any case the fossil bears little resemblance to the femora of either
group. In addition to living insectivores, the fossil was compared to the
femur of Nesophontes, the only genus in the extinct Greater Antillean in-
sectivore family Nesophontidae and to the femora of several fossil Solen-
odon-like forms and the extinct solenodontid Antillogale (regarded as So-
lenodon by Van Valen, 1967, and Varona, 1974). The fossil Cuban
insectivore was also compared to the femur of Deinogalerix koenigswaldi

)
|
|
|

VOLUME 93, NUMBER 3 601

from the Miccene of Italy, the largest known insectivore, living or extinct
(Freudenthal, 1972).

With the exception of one genus of tenrec, the fossil differs considerably
from the femora of all tenrecids and chrysochlorids, all of which have the
greater trochanter approximately even with or lower than the head, lack a
well defined trochanteric fossa and intertrochanteric crest, and have a small-
er third trochanter located farther distally on the shaft. The femur of Mi-
crogale, one of the smallest genera of tenrecs, is similar to the fossil in
morphology, differing only in its considerably smaller size and elongate,
flange-shaped third trochanter. The fossil bears little resemblance to the
femur of any living talpid or soricid. Besides the obvious discrepancy in
size, the femur in moles and shrews lacks a well developed trochanteric
fossa and intertrochanteric crest, has the third trochanter located higher on
the shaft, and is relatively broader distally. Modern erinaceids are divisible
into two subfamilies, the Erinaceinae and Echinosoricinae, which differ sig-
nificantly in their femoral morphology. Erinaceine femora differ in a number
of ways from the Cuban fossil, including having a less pronounced greater
trochanter, the trochanteric fossa not as concave, and the third trochanter
developed as an elongate flange extending nearly one-third the length of the
shaft. The patellar groove in hedgehogs does not extend nearly as far proxi-
mally as it does in the fossil and the lateral condyle is somewhat reduced.
Among living echinosoricines, the fossil femur resembles most closely the
femur of Echinosorex, especially in its overall proportions and in the strong
development of the greater trochanter, third trochanter, and trochanteric
fossa. However, the morphology of the distal end of the femur in Echino-
sorex is like that of other erinaceids and quite unlike the Cuban fossil. The
femur of the gigantic extinct echinosoricine Deinogalerix is very similar in
most respects to the femur of Echinosorex, except, of course, for its tre-
mendous size. As with Echinosorex, several fundamental differences in the
distal end of the femur argue against a close relationship between Deino-
galerix and the Cuban specimen. Nonetheless the femora of these two giant
insectivores are quite similar in gross morphology, particularly in their long,
relatively gracile (for an insectivore) overall form and the strongly developed
greater trochanter, third trochanter, and trochanteric fossa.

As was suspected on the basis of geography, the fossil bears a closer
resemblance to the femora of the West Indian insectivores Solenodon and
Nesophontes than it does to those of any other living insectivores. Although
there are some minor differences, the femora of these two genera are more
similar to one another than either is to that of any other living insectivore.
The differences in postcranial osteology between these two genera are less
pronounced, for instance, than the differences observed between certain
genera within the Tenrecidae or Erinaceidae. McDowell (1958) summarized
the many similarities in the cranial and postcranial osteology of Solenodon

602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and Nesophontes and hypothesized a close phylogenetic relationship be-
tween them, placing both genera in the Solenodontidae. However, the pro-
found differences in dentition between Solenodon and Nesophontes seem
to preclude their placement in the same family, at least in the present state
of our knowledge of insectivore relationships. The possibility does exist that
the Solenodontidae and Nesophontidae are closely related within the sori-
comorph insectivores and may even have been derived from a single **in-
vasion’’ of the West Indies or “‘proto-Antilles’’ by a late Cretaceous or early
Tertiary soricomorph (MacFadden, in press).

Comparison with Solenodon, Nesophontes, and Antillogale.—To deter-
mine if the fossil femur is closest in morphology to Solenodon, Antillogale,
or Nesophontes, the fossil was compared to the femora of all living and
extinct West Indian insectivores for which the femur is known. These in-
clude, Solenodon cubanus and several large Solenodon cubanus-like forms,
all from Cuba, S. paradoxus and Antillogale marcanoi from Hispaniola,
and six species of Nesophontes, three from Hispaniola and one each from
Puerto Rico, Cuba, and the Cayman Islands.

The most striking feature of the fossil femur is its large size. It is 27%
longer than the longest modern Solenodon femur measured (Table 1) and
is considerably larger than the femur of any living insectivore. Aside from
the obvious difference in size, the largest Nesophontes being barely half the
size of Solenodon or Antillogale, the femur of Nesophontes differs consis-
tently from those of the latter two genera in several features. Antillogale
and Solenodon are certainly closely related, if not congeneric, and in fem-
oral morphology they are quite similar. In the following comparison of fem-
oral characters in Nesophontes and Solenodon, the characters ascribed to
Solenodon apply also to Antillogale, unless noted otherwise. In Solenodon
the lateral condyle is transversely flattened, in contrast to its convexity in
Nesophontes. The patellar groove of Solenodon projects anteriad of the
shaft, unlike Nesophontes in which the patellar trochlea is in line with the
axis of the femoral shaft. Solenodon has a deep pit on the anterior surface
of the shaft just proximal to the patellar groove, for reception of the patella
in strong extension of the lower leg. This pit is very weakly developed in
the large species of Nesophontes, N. edithae, and is absent in the smaller
species. In all Solenodon specimens examined, the third trochanter is more
strongly developed than in any species of Nesophontes and is located slight-
ly higher on the shaft. The lesser trochanter is of slightly different shape in
the two genera, pointed and projecting at a right angle from the shaft in
Nesophontes and triangular, relatively broader, and projecting somewhat
proximally in Solenodon. The femur of Solenodon has a longer neck and
a slightly oblong head, whereas the neck is shorter in Nesophontes (owing
at least in part to the relatively larger head) and the head is almost perfectly
hemispherical. The greater trochanter in Solenodon exhibits a strong an-

603

VOLUME 93, NUMBER 3

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604 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

terior flexion which is not nearly as pronounced in the smaller genus. Fi-
nally, there is a difference in the angle formed by the intertrochanteric crest,
acute in Solenodon, right to obtuse in Nesophontes.

Difficulties arise when the fossil is compared to all species of Solenodon
and Nesophontes, rather than to generalized characters for the individual
genera. Comparisons reveal that there are notable differences between the
femora of the species within each of these two genera. For instance, the
femur of Nesophontes edithae differs considerably from the femora of the
smaller species of the genus, particularly in its relatively larger head, broad-
er patellar groove, and comparatively broader shaft. The index of robustness
(minimum shaft width/total length of femur) reveals that the femur of the
N. edithae specimens measured is as robust as the femur of Solenodon
paradoxus, an animal twice its size. The remaining species of Nesophontes
have more slender femora.

Although only one skeleton of modern Solenodon cubanus was available
for study, we did examine two fossil femora from western Cuba of a form
very close to the modern Cuban solenodon. Solenodon cubanus appears to
differ in several important femoral characters from S. paradoxus. The index
of robustness demonstrates clearly that the femur of S. cubanus is of more
slender build than that of S. paradoxus. Unlike S. paradoxus and like Ne-
sophontes, the greater trochanter in S. cubanus is not flexed anteriorly to
a marked degree. The patellar groove in §. cubanus is narrower and less
concave than in §. paradoxus and the pit proximal to this groove is shal-
lower in the former.

The femur of Antillogale marcanoi differs from that of S. cubanus and
S. paradoxus in several features. The most striking feature of Antillogale
is the relative massiveness of its limb elements. Like the humerus and ulna
described by Patterson (1962), the femur of Antillogale is similar to that of
S. paradoxus in the breadth of the proximal and distal ends, but has a
noticeably shorter shaft, giving it a much stouter, more massive appearance.
In addition, the femoral shaft of Antillogale has a distinct curvature not
observed in other West Indian insectivores. The pit on the anterior surface
of the shaft proximal to the patellar groove is deeper and the greater tro-
chanter is flexed anteriorly to a greater degree than in either species of
Solenodon. In these last two features and in the relative robustness of the
shaft and broad, concave patellar trochlea, the femur of Antillogale more
closely resembles that of S. paradoxus than it does the femur of S. cubanus,
Nesophontes, or the Cuban fossil.

In almost every aspect of its morphology, the Cuban fossil femur resem-
bles the femur of Solenodon cubanus more closely than it does the femur
of any other West Indian insectivore (Fig. 1). Although the giant fossil femur
does resemble Nesophontes more closely in several characters than it does
S. paradoxus, particularly in the slenderness of the shaft and the reduced

VOLUME 93, NUMBER 3 605

anterior flexion of the greater trochanter, it also shares these characters
with S. cubanus. In characters such as the stronger development and the
more proximal location of the third trochanter, the angle formed between
the shaft and the lesser trochanter, the length of the femoral neck, the angle
formed by the intertrochanteric crest, the shape of the lateral condyle, the
anterior projection of the patellar groove, and presence of a well defined pit
proximal to the patellar groove, the fossil resembles Solenodon more closely
than Nesophontes and in particular, resembles S. cubanus more closely
than S. paradoxus or Antillogale.

Although the fossil femur is most similar to the femur of Solenodon cu-
banus among known insectivores, several differences are apparent, the most
obvious being one of size. The fossil solenodontid femur is 1.4 times longer
than that of modern S. cubanus. The ratio of femur length/head and body
length was calculated for three modern individuals of Solenodon paradoxus,
the species most closely related to the fossil for which these data are avail-

able. The resulting ratio (x = .14) suggests a head and body length for the
fossil solenodontid of approximately 470 mm. This is about the size of a
large adult male opossum (Didelphis virginiana), and is considerably larger
than any living member of the Insectivora. There are several fossil Solen-
odon cf. cubanus femora from western Cuba which are intermediate in size
between modern S. cubanus and the giant solenodontid (Table 1). These
_ specimens are very similar to S. cubanus, differing primarily in their larger
_ size. Based on the available fossil material, it is not clear if these interme-
| diate-sized specimens represent a third late Pleistocene solenodontid species
_or are representative of a late Pleistocene population of S. cubanus which
_was larger than the modern form. Other characters which distinguish the
| giant solenodontid from S. cubanus are the more prominent greater tro-
_chanter, deeper trochanteric fossa, better developed groove separating the
_head from the greater trochanter, larger third trochanter, and slightly con-
_ cave lateral condyle. Most of these characters are not unique to the giant
_solenodontid, but rather are characters found developed to a lesser degree
/in Solenodon cubanus. It is possible that the observed differences are re-
| lated to the large size of the fossil, but this cannot be determined from the
limited material available.
: In summary, the giant insectivore femur can be assigned confidently to
_the Solenodontidae. In morphology it agrees closely with Solenodon, in
particular with S$. cubanus, to which the fossil appears to be most closely
| related among known insectivores. This specimen almost certainly repre-
sents an undescribed species, tentatively assignable to the genus Solenodon,
but the incomplete femur described here does not provide adequate material
} for the formal description of a new taxon.
_ Discussion.—The presence in the late Pleistocene fauna of Cuba of a giant
| solenodontid, larger than any living insectivore, raises some intriguing ques-

)
i

|)
{

606 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tions regarding its ecological position in that fauna. Based on the carnivo-
rous habits of its closest living relatives, it seems likely that the giant Cuban
solenodontid was also carnivorous, at least in part. Certainly Solenodon
paradoxus eats small mammals, lizards, and frogs, in addition to various
invertebrates (based on observations of captive S. paradoxus by C. E. Ray
and others; see also Allen, 1910; Pena, 1977; Verrill, 1907), so it is not
inconceivable that a very closely related animal of considerably larger size
would have preyed on small to medium-sized vertebrates. Taken in the
context of the entire Cuban fauna, the carnivorous habits of Solenodon and
possibly the giant solenodontid may reflect the absence of other mammalian
predators. In the absence of members of the Carnivora, the majority of
carnivorous niches in Cuba are filled by nonmammalian predators. The larg-
est native carnivorous vertebrates on Cuba today are the boa, Epicrates
angulifer, several species of medium-sized raptorial birds, and Solenodon
cubanus. The inclusion of late Pleistocene faunas would add the gigantic
flightless owl, Ornimegalonyx oteroi, two species of giant barn owl, the
extinct eagle, Aquila borrasi, and possibly the giant solenodontid.
Absence of members of the Carnivora from Cuba invites comparison with
other well known faunas in which carnivores (i.e. Carnivora) are absent. In
the near absence of Carnivora in the Tertiary of South America (with the
exception of procyonids in the Neogene) an impressive array of carnivorous
marsupials and large flightless predaceous birds is found (Marshall, 1977).
In Australia there evolved a full complement of marsupial carnivores and
the titanic varanid lizard, Megalania, in addition to several other medium-
to large-sized carnivorous varanids (Hecht, 1975). There are, however, no
members of the Insectivora in either of these faunas (with the minor excep-
tion of small shrews of the genus Cryptotis in the modern fauna of north-
ernmost South America). Olson (1978) noted the striking parallei between
the Miocene fauna of the Gargano Peninsula of Italy (Freudenthal, 1972)
and that of the late Pleistocene of Cuba. The Gargano Peninsula was ap-
parently an island in the Miocene and during that time its faunas are char-
acterized by the virtual absence of carnivores (with the exception of an otter
of remarkably large size) and by the presence of several species of large
raptorial birds, the largest known insectivore, Deinogalerix koenigswaldi,
and rodents of very large size (Freudenthal, 1972). The virtual absence of
mammalian carnivores and the presence of a giant insectivore and very large
raptorial birds mirrors the situation in Cuba during the late Pleistocene.
Giant rodents are unknown from Cuba, but the small Cuban ground sloths
may have occupied a similar niche. It is probably no coincidence that the
giant Cuban solenodontid and Deinogalerix, the largest of known insecti-
vores living or extinct, both occur in faunas devoid of other mammalian
predators. Competition from more advanced predators of the Carnivora has

VOLUME 93, NUMBER 3 607

probably prevented modern insectivores from attaining the large size of
these two fossil forms.

The giant solenodontid described here, in addition to recently discovered
species of Nesophontes from Puerto Rico, Vieques, Cuba, and the Cayman
Islands, provides evidence to support our conviction that the Quaternary
vertebrate fauna of the West Indies is at present incompletely known. Fur-
thermore, the lack of terrestrial vertebrate fossils in the West Indies older
than the late Pleistocene strongly indicates that there is still much to learn
about the Greater Antillean insectivore fauna. If the Greater Antillean in-
sectivores were derived from North American early Tertiary soricomorphs
as suggested by most authors (MacFadden, 1980; Matthew, 1918; Pat-
terson, 1962; Simpson, 1956), we might justifiably expect an insectivore
radiation similar to that of another soricomorph group apparently isolated
on an island since the early Tertiary, the Tenrecidae of Madagascar. Rather

extensive radiations of capromyid rodents, including eight endemic genera

ee eee —_— —-

—

and at least 15 species, and megalonychid ground sloths, including seven
endemic genera and eight species, have been documented in the Greater
Antilles. Both of these radiations have probably taken place since the early
Miocene. With the addition of a new species of solenodontid we now know
of three genera and at least a dozen species of endemic Antillean insecti-
vores. These figures compare with 12 genera and approximately 25 species
of tenrecs on Madagascar, an island about five times larger than Cuba and
of considerably greater ecological diversity than any West Indian island.

| The discovery of a giant solenodontid on Cuba raises the possibility that an
even more extensive radiation of insectivores remains to be discovered in

the West Indian fossil record.

Acknowledgments

John E. Guilday and Allen D. McCrady of the Carnegie Museum of Nat-
ural History kindly loaned us the femur of Antillogale. The figure was drawn

_ by Lawrence B. Isham. For helpful comments on the manuscript we thank

Jon A. Baskin, Susan M. Ford, Luis S. Varona, and Charles A. Woods.

Literature Cited

Allen, G. M. 1910. Solenodon paradoxus.—Mem. Mus. Comp. Zool. 40:1—54.

Butler, P. M. 1972. The problem of insectivore classification. Pp. 253-265 in K. A. Joysey
and T. S. Kemp, eds. Studies in vertebrate evolution.—Oliver and Boyd, Edinburgh.

Freudenthal, M. 1972. Deinogalerix koenigswaldi nov. gen., nov. spec., a giant insectivore
from the Neogene of Italy.—Scripta Geol. 14:1-19.

Hecht, M. K. 1975. The morphology and relationships of the largest known terrestrial lizard,
Megalania prisca Owen from the Pleistocene of Australia.—Proc. Roy. Soc. Victoria
87:239-249.

608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

MacFadden, B. J. 1980. Rafting mammals or drifting islands?: Biogeography of the Greater
Antillean insectivores Nesophontes and Solenodon.—Jour. Biogeog. 7:11—22.

Marshall, L. G. 1977. Evolution of the carnivorous adaptive zone in South America. Pp. 709-
721 in M. K. Hecht, P. C. Goody, and B. M. Hecht, eds. Major patterns in vertebrate
evolution.—Plenum Press, New York.

Matthew, W. D. 1918. Affinities and origin of the Antillean mammals.—Bull. Geol. Soc.
Amer. 29:657—666.

McDowell, S. B., Jr. 1958. The Greater Antillean insectivores.—Bull. Amer. Mus. Nat. Hist.
115(3):113-214.

McKenna, M. C. 1975. Toward a phylogenetic classification of the Mammalia. Pp. 21-46 in
F. S. Szalay and W. P. Luckett, eds. Phylogeny of the Primates.—Plenum Press, New
York.

Olson, S. L. 1978. A paleontological perspective of West Indian birds and mammals. Pp. 99—
117 in F. B. Gill, ed. Zoogeography in the Caribbean. The 1975 Leidy Medal Sympo-
sium.—Proc. Phila. Acad. Sci., Spec. Publ. 13:1-128.

Patterson, B. 1962. An extinct solenodontid insectivore from Hispaniola.—Mus. Comp. Zool.,
Breviora 165:1—11.

Pena Franjul, M. 1977. Habitos nutritivos del Solenodon paradoxus Brandt y el programa de
reproduccion en captividad.—Zoodom 1:11-25.

Simpson, G. G. 1956. Zoogeography of West Indian land mammals.—Amer. Mus. Novitates
1759: 1-28.

Van Valen, L. 1967. New Paleocene insectivores and insectivore classification.—Bull. Amer.
Mus. Nat. Hist. 135(5):217—284.

Varona, L. S. 1974. Catalogo de los mamiferos vivientes y extinguidos de las Antillas.—Acad.

Cien. Cuba, Habana, Cuba, 139 pp.

. 1976. Reemplazo de Cubanocnus por Neocnus (Mammalia: Edentata).—Inst. Zool.,

Acad. Cien. Cuba, Misc. Zool. 2:4.

Verrill, A. H. 1907. Notes on the habits and external characters of the Solenodon of San
Domingo (Solenodon paradoxus).—Amer. Jour. Sci., Ser. 4, 20:55—57.

(GSM) Dept. of Vertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560; (CER) Dept. of Paleobiology, Smithsonian Institution,
Washington, D.C. 20560; (OA) Sta. No. 27, esquina a E, Reparto Capri,
Arroyo Naranjo, Habana 19, Cuba.

ADDENDUM

Arredondo and Varona (1974. Poeyana 131:1—12) described a new genus
and species of extinct canid, Cubacyon transversidens, based on a partial
maxilla with two teeth from a cave deposit in Cuba. Olson (1978) questioned
the validity of this species and Hall (in press. The Mammals of North Amer-
ica, 2nd Ed., The Ronald Press, New York) synonymized it with the domes-
tic dog, Canis familiaris. Morgan and Ray follow Hall in regarding Cubacyon
as a domestic dog. Arredondo and Varona (1974; in litt.), however, believe
that C. transversidens is a valid species based on the configuration of the P*
and M2 and by its association in a fossil deposit with extinct mammals. There-
fore, any comments in this paper regarding the absence of endemic Carniv-
ora on Cuba are the opinion of Morgan and Ray, whereas Arredondo
regards Cubacyon as part of Cuba’s late Pleistocene fauna.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 609-654

NEW AMPHIPODA FROM THE SOUTHERN OCEAN,
WITH PARTIAL REVISIONS OF THE
ACANTHONOTOZOMATIDAE AND

PARAMPHITHOIDAE

Les Watling and Heather Holman

Abstract.—Six new species and 1 new genus of Acanthonotozomatidae,
2 new species of Paramphithoidae and | new species of Stegocephalidae are
described from Antarctic waters, chiefly of the Scotia Sea region. New
‘revisions are offered for the acanthonotozomatid genera Acanthonotozo-
mella, Iphimedia, Iphimediella, and Pseudiphimediella and the paramphi-
thoid genera Epimeria and Parepimeria.

) With this paper we begin a series of studies on the Amphipoda from the
Scotia Sea region of the Southern Ocean. This and the following papers will
be directed toward the redescription of poorly known or incorrectly de-
scribed species as well as the description of new species. To facilitate the
redescription of older taxa and the revision of generic concepts we have
borrowed much type-material and would like to thank the following persons
_and institutions for making specimens available to us: Dr. Denise Bellan-
Santini, Station Marine d’Endoume; Ms. Joan Ellis, British Museum (Nat-
‘ural History); Ms. Elizabeth Louw, South African Museum; Dr. H.-E. Gru-
ner, Zoologisches Museum der Humboldt—Universitat zu Berlin; and Dr.
Roy Olerod, Swedish Museum of Natural History. Additional material was
sent to us for examination by Dr. R. Y. George and Dr. L. D. McKinney.
We would like to express our gratitude to Ms. Patrice Rossi whose pencil
and ink illustrating capabilities saved us much time and effort, and to Dr.
J. L. Barnard for his critical examination of our manuscript.

_ This study was funded by the Smithsonian Oceanographic Sorting Center
| under contract No. PC-801851. We would like to thank Ms. B. Landrum for

: her interest in this project.

|

|
Acanthonotozomatidae

| Acanthonotozomella Schellenberg
Acanthonotozomella Schellenberg, 1926:332.
Paracanthonotozoma Bellan-Santini:1972:177 (new synonymy).

_ Type-species.—Acanthonotozomella alata Schellenberg, 1926 (original
| designation).

|

610 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Diagnosis {modified from Schellenberg (1926) and J. L. Barnard (1969) ].—
Body with numerous processes; telson wide, entire to emarginate; antenna
| shorter than antenna 2, accessory flagellum absent; upper lip distally
grooved; mandible incisor multi-toothed, molar poorly developed, with setal
row extending between incisor and molar; inner plate of maxilla 1 not mi-
nute, few setae distally, palp biarticulate, reaching end of outer plate; max-
illiped well-developed, palp 4-articulate, article 2 not expanded or produced;
gnathopods 1 and 2 simple, alike; coxae | and 3 subrectangular, coxa 2
shorter than either coxa | or 3; coxae 5—7 rounded.

Remarks.—The genus Acanthonotozomella, as diagnosed above, is con-
sidered here to contain the following species: A. alata, Schellenberg 1926;
A. trispinosa (Bellan-Santini 1972), originally described as the type-species
of the genus Paracanthonotozoma; and A. barnardi n. sp., described below.
Recently, Bushueva (1978) described the species A. pushkini, but as will be
discussed below, we believe this species represents a distinct genus, inter-
mediate in position to Acanthonotozomella and Acanthonotozomoides, to
which we will give the name Acanthonotozomoposis. The genera, Acan-
thonotozoma, Acanthonotozomoides, Acanthonotozomella and Acanthon-
otozomoposis are united by their common possession of the following fea-
tures: gnathopods | and 2 simple; maxilla | palp 2-articulate; maxilliped
palp exceeds outer plate, 4-articulate, article 2 not produced or expanded.
The characters which distinguish these genera are given in Table |. Acan-
thonotozomoides and Acanthonotozomella can be readily characterized by
features of the body as well as mouthparts. Bushueva (1978) used the shape
of coxae 1-3, maxilliped palp article 2 not produced along article 3 and
pereopod 7 larger than 5 and 6 to assign her species pushkini to Acanthon-
otozomella. However, the presence of a minute inner plate and expanded
palp article 2 on maxilla 1, lack of acute body processes, form of mandible
incisor and similarity of coxae | and 2 distinguish Bushueva’s species from
all others in Table 1 and warrants, in our opinion, the erection of a new
genus.

Acanthonotozomella trispinosa (Bellan-Santini), new combination
Paracanthonotozoma trispinosum Bellan-Santini, 1972:177, pl. 6.

Diagnosis. —Coxae 1 and 3 anteroventral corners produced; antenna |
peduncle article | with distal tooth; mandible seta row short; pereopods 5,
6 and 7 alike, all with excavate posterior margins, lobe above excavation
subacute.

Remarks.—Comparison of the type-specimen of A. alata with the illus-
trations of A. trispinosa revealed many similarities, especially in dorsal
armature. They were found to differ in the following features: coxa 2 is
distally rounded in A. alata and is distally subacute in A. trispinosa; antenna

611

VOLUME 93, NUMBER 3

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612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1 peduncle article | bears a dorsal tooth in A. trispinosa but such a tooth
is absent in A. alata; the hind margin of pereopods 5-7 basis is rounded
above the excavation in A. alata but forms an acute lobe in A. trispinosum.
Since the specimens also differed strongly in size (A. alata, 3 mm; A.
trispinosa, 12 mm) it is possible that the above differences are size-related
and the two species are synonymous. Also see comparisons in Table 1.

Acanthonotozomella barnardi n. sp.
Figs. 1-3

Material.—Holotype: Eltanin Cruise 9, Sta. 740, 18 Sept. 1963, 56°06’S,
66°19-30'W, 384-494 m, 1 6 (USNM 173583).

Diagnosis.—Pleonite 3 with single, dorsal, upright tooth; coxa | rectan-
gular distally, not produced acutely at anterodistal corner; coxa 2 rectan-
gular distally; mandible with multidenticulate incisor and poorly developed
molar with row of more than 30 setae between incisor and molar; bases of
pereopods 5 and 6 similar, anterior and posterior margins almost parallel
with posterodistal corner extending downward in a smooth lobe; pereopod
7 distinct from pereopods 5 and 6, posterior margin of basis excavate dis-
tally, posterodistal corner extended downward as a slender lobe.

Description.—Head, without rostrum, shorter than pereonite 1. Integu-
ment of body covered with small conelike papillae giving rough appearance.
Pereonite | to urosomite | each with single dorsal projection extending
upward and posteriorly. Projection on pereonite | forming upright, truncate
keel with slight posterior extension. Projections on pereonites 2—6 in form
of upright, posteriorly projecting teeth on posterior part of each somite,
increasing in size with successive somite. Pleonite 3 bearing single upright
keel, posterior part of which forms subacute apex without projecting over
following somite. (Exact structure of tooth on urosomite | uncertain as it
was broken on this specimen.) Pleurae of pereonites 1-7 all projecting out-
ward and posteriorly. Epimeral plates 1-3 each with lateral tooth on pos-
terior margin. Distal margin of epimeral plate 1 forming subacute point.
Distal margin of epimeral plates 2 and 3 rounded anteriorly, more angular
posteriorly. Coxae 1-3 rectangular distally. Coxa | slightly excavate ante-
riorly. Coxa 2 more slender distally and shorter than coxa | or 3. Coxa 4
lower margin crescentic. Coxae S—7 rounded posteriorly.

Rostrum with raised lateral edges, shorter than first peduncle article of
antenna |. Ocular bulge pronounced. Antenna | peduncle article 1 equal in
length to combined lengths of articles 2 and 3, accessory flagellum absent.
Article 4 of antenna 2 subequal to article 5. Upper lip distinctly grooved
with fine hairs along edge to either side of groove. Mandible multidenticu-
late; toothed accessory plate present on right mandible; molar poorly de-
veloped with row of approximately 34 setae between incisor and molar; palp

VOLUME 93, NUMBER 3 613

Fig. 1. Acanthonotozomella barnardi, male: a, Body, side view; b, Head; c, Antenna 1; d,
Antenna 2; e, Upper lip; f, Mandible; g, Mandible palp; h, Lower lip; i, Maxilla 1; j, Maxilla
2:

articles 2 and 3 equal in length. Lower lip tapering distally and unnotched.
Palp of maxilla 1 biarticulate and extending past outer plate; inner plate
slender with 2 short spines and 4 longer setae on distal margin. Maxilla 2
inner and outer plates subequal in length, serrate setae on distal margins of
both, somewhat longer on outer plate. Maxilliped palp 4-articulate, articles
1-3 equal in length, article 4 slightly shorter and bearing approximately 5

614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Acanthonotozomella barnardi, male: a, Maxilliped; b, Gnathopod 1; c, Gnathopod
2; d, Telson.

setae on tip. Gnathopods simple and alike; carpus slightly longer than propo-
dus, dactyl half length of propodus. Pereopods 5 and 6, anterior and pos-
terior margins of basis parallel with posterodistal corner, extending down-
ward as rounded lobe past distal margin of ischium. Pereopod 7 basis
excavate behind with posterodistal corner extending downward as a sub-
acute lobe past distal margin of ischium. Telson appears to be entire, poor
condition of distal margin makes this uncertain. No uropods were intact.

Etymology.—This species is named in honor of Dr. J. L. Barnard.

Distribution.—Known only from the type-locality.

Remarks.—This species differs from A. alata and A. trispinosa primarily
by having a single dorsal tooth on pleonite 3 (vs. 2 teeth), a distally truncate
coxa 2 (vs. round or subacute), pereopods 5 and 6 hind margin not excavate —
(vs. excavate) , and a long seta row (ca. 30 setae) on the mandible (vs. 8— |
10 setae).

Acanthonotozomopsis n. gen.

Type-species.—Acanthonotozomella pushkini Bushueva 1978.

Diagnosis.—Body with blunt dorsal and lateral processes; coxa 2 shorter
than but similar in shape to coxae | and 3; coxa 4 triangular, without pos-
terior protuberance; mandible incisor broadened, seta row long; maxilla 1

VOLUME 93, NUMBER 3 615

Fig. 3. Acanthonotozomella barnardi, male: a, Pereopod 5; b, Pereopod 6; c, Pereopod 7.

inner plate minute, palp article 2 expanded; maxilliped palp article 2 broad,
not produced along article 3; gnathopods simple, of similar shape; telson
entire.

Remarks.—The distinctiveness of this genus is discussed in the remarks
for Acanthonotozomella and its features listed in Table 1.

Iphimedia Rathke

Iphimedia Rathke, 1843:85.
Panoploea Thomson, 1880:2.—Karaman and Barnard, 1979:110.
Cypsiphimedia K. H. Barnard, 1955:87 (new synonymy).

Type-species.—l. obesa Rathke 1843.

Diagnosis.—Upper lip entire or slightly emarginate; maxilla 1 palp biar-
ticulate, of variable length; maxilliped palp exceeding outer plate, 3-artic-
ulate, article 2 produced medially along article 3; gnathopod 1 chelate;
gnathopod 2 chelate or subchelate; telson variably emarginate or incised.

Remarks.—At their extremes, [phimedia and Panoploea were signifi-
cantly different from each other. Karaman and Barnard (1979), when
synonymizing the two genera, showed there was a continuous variation in
shape and size of upper and lower lips, mandible and maxilla | palp. Krapp-

616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Iphimedia gibba: a, Body, side view; b, Mandible; c, Lower lip; d, Maxilla 1; e,
Maxilliped; f, Gnathopod 1.

Schickel (1976) noted that the palp of maxilla 1 of J. minuta changed in
length with size of the individual. Karaman and Barnard (1979) further dis- |
tinguished /phimedia from the closely related Cypsiphimedia on the basis
of the latter possessing an enlarged first pereonite and lacking dorsal body
processes (but see later). As with the palp of maxilla | and the strength of
the mandible, it can be shown that there is also a gradation in size of per- _
eonite 1, ranging from equal to pereonite 2 in J. obesa Rathke, as long as __
pereonites 2 and 3 in J. spinosa (Thomson), to as long as pereonites 2-4 in _

VOLUME 93, NUMBER 3 617

IM)
St All
hf)
{\ Hi

Fig. 5. Iphimedia magellanica, female: a, Body, side view; b, Head; c, Antenna | peduncle
articles 1 and 2: d, Antenna 2 peduncle; e, Upper lip; f, Right mandible; g, Mandible palp; h,
Lower lip; i, Maxilla 1.

I. eblanae (Bate). Additionally, the number and shape of dorsal processes
range from 4 pairs of short teeth (J. obesa), to 3 pairs of long and slender
teeth (J. joubini), to 3 pairs of short processes (J. spinosa), to a single
minute pair of processes (J. excisa). Thus, the species assigned to Cypsi-
phimedia by Karaman and Barnard (1979) are not outside the concept of
Iphimedia as diagnosed above. This genus, more than any other in the

618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ee : - \
R N | \N

=

Fig. 6. Iphimedia magellanica, female: a, Maxilla 2; b, Maxilliped without palp; c, Max-
illiped palp; d, Gnathopod 1; e, Gnathopod | propodus fixed finger and dactyl; f, Gnathopod
2; g, Gnathopod 2 propodus fixed finger and dactyl.

family, is characterized by a series of gradational characters. In our opinion
the genus needs a further re-examination and perhaps should be subdivided
into more discrete units. It is also one of the few acanthonotozomatid genera
to be extensively represented outside polar waters. The genus as presently
constituted contains the following species: J. capicola K. H. Barnard, I.

VOLUME 93, NUMBER 3 619

discreta Stebbing, I. eblanae (Bate), I. excisa (K. H. Barnard), I. gibba
(K. H. Barnard), J. gladiolus K. H. Barnard, J. grossimana Ledoyer, I.
haurakiensis Hurley, I. imparilabia n. sp., I. joubini (Chevreux), I. jugos-
lavica Karaman, I. macrocystidis (K. H. Barnard), I. magellanica n. sp.,
I. minuta (Sars), I. multidentata (Schellenberg), J. obesa Rathke, J. or-
chestimana Ruffo, I. pacifica Stebbing, I. rickettsi (Shoemaker), I. spinosa
(Thomson), J. stegosaura (Griffiths).

Iphimedia gibba (K. H. Barnard), new combination
Fig. 4

Cypsiphimedia gibba K. H. Barnard, 1955:88, fig. 43.

Material.—Holotype: South African Museum No. 10318.

Diagnosis (emended from K. H. Barnard, 1955).—Body with minute
paired dorsal processes on pleonite 3, otherwise smooth dorsally; pereonite
1 expanded, head directed ventrally; lower lips apically notched; maxilla 1
palp longer than outer plate; gnathopod 2 propodus ovate.

Description.—The following supplements the description given by K. H.
Barnard (1955). Body with pair of minute dorsal processes on pleonite 3.
Epimeral plate 3 posteroventral corner with 2 teeth. Mandible accessory
plate minute, feebly bidentate. Maxilla 1 palp longer than outer plate. Max-
illiped palp 3-articulate; article 2 expanded and produced along article 3.

Remarks .—Our re-examination of the holotype has shown the mouthparts
of this species to be typical of Iphimedia. Also, contrary to the statement
of Karaman and Barnard (1979) the palp of maxilla 1 is in fact longer than
the outer plate. The size of pereonite | is not beyond the range otherwise
seen for [phimedia and its ties to the genus appear to be strengthened by
the presence of minute dorsal processes on pleonite 3.

Iphimedia magellanica n. sp.
Figs. 5-7

Material.—Holotype. Eltanin Cruise 11, Sta. 977, 13 Feb. 1964, 52°32'S,
63°53’W, 299 m, 1 2 with eggs (USNM 173584), 6 mm. Paratypes: same
station, 4 juveniles (USNM 173585).

Diagnosis.—Body with short, paired dorsal processes on pereonite 7 and
pleonites | and 2; pleonites 1-3 with mid-dorsal keel; pereopod 7 basis
posterior margin with 3 teeth, coxa with single posterodistal tooth.

Description.—Body with 3 pairs dorsal processes, all short and somewhat
curved downward; pleonites 1 and 2 with short mid-dorsal keel; pleonite 3
mid-dorsal keel enlarged. Pleurae on pereopods 5-7 extend as posteriorly-
directed acute processes. Epimeral plates 1 and 2 with small posterodistal
tooth. Hind margin of epimeral plate 3 with 2 teeth. Head lateral margin

620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Iphimedia magellanica, male: a, Pereopod 5; b, Pereopod 6; c, Pereopod 7; d, |
Telson. |

VOLUME 93, NUMBER 3 621

slightly produced as acute process distally; rostrum extends nearly length
of first peduncle article of antenna 1. Ventral tooth of antenna | peduncle
article 1 extends to end of peduncle article 2, dorsal tooth small. Antenna
2 peduncle article 3 with short lateral and dorsal apical teeth. Upper lip
broadly rounded, with paired ventrolateral groups of fine setae. Left man-
dible with blunt, smooth incisor, accessory plate weakly toothed; right man-
dible without accessory tooth; palp article 2 twice length of article 3, armed
with only 2 distal setae; third palp article with setae on ventral margin of
distal third of article. Lower lip variably incised. Maxilla 1 palp biarticulate,
not reaching end of outer plate; inner plate small, with 5 long setae; outer
plate with 11 stout setae apically. Maxilla 2 slender, inner and outer plates
subequal in width; distal setae on outer plate only slightly longer than on
inner plate. Maxilliped palp 3-articulate, article 2 produced medially along
article 3, article 1 longer than article 2, third article elongate, distally blunt,
two-thirds the length of article 2; inner plate narrow, armed with plumose
setae along distal half of inner margin; outer plate apically subacute.

Gnathopod | chelate; coxa distally subacute, with 4 short setae; basis
with scattered short setae along inner margin; dactyl equal in size to fixed
finger of propodus, with 2 long, blunt setae; propodus fixed finger with 4
long setae, 2 of which are distally forked. Gnathopod 2 chelate; coxa distally
quadrate; article 6 longer than article 5. Pereopods 5-7, basis posterior
margin with 3 teeth, | located dorsally and 2 ventrally. Uropod 1 rami
subequal; uropod 2 outer ramus shorter than inner. Telson shallowly cleft.

Etymology.—Name derived from Magellanic zoogeographic province.

Distribution.—Known only from locality listed above.

Remarks.—This species is distinctive among species of [phimedia in its
possession of only 3 pairs of short, downwardly curving dorsal processes
and single, upright keel on pleonite 3 in combination with the form of the
posterior pereopods and coxae. The general body form suggests affinities
with J. multidentata and I. macrocystidis; however the absence of paired
posterior teeth on coxa 7 and the number of teeth on pereopod 7 basis hind
margin readily distinguish J. magellanica from both species.

Iphimedia imparilabia n. sp.
Figs. 8, 9

Material.—Holotype: Eltanin Cruise 6, Sta. 340, 3 Dec. 1962, 53°07—08’S,
59°21—23'’W, 567-578 m, 1 6, 7 mm (USNM 173586).

Diagnosis.—Body with short, paired dorsal processes on pereonite 7 and
pleonites 1 and 2, and with mid-dorsal keel on pleonite 3; pereopod 7 coxa
posterior margin rounded, basis posterior margin slightly concave distally,
without conspicuous teeth; maxilla | palp biarticulate, short; maxilliped palp
article 3 very short.

622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Iphimedia imparilabia, male: a, Body, side view; b, Head; c, Upper lip; d, Mandible;
e, Lower lip; f, Maxilla 1; g, Maxilla 2; h, Maxilliped.

Description.—Head interantennal angle subacute, anteroventral corner
with short subacute process. Body with short, paired dorsal processes on
pereonite 7 and pleonites 1 and 2; mid-dorsal keel strongly developed on
pleonite 3. Pleurae on pereonites S—7 not conspicuously produced. Epimeral
plates 1 and 2 without teeth at posterodistal corners; epimeral plate 3 with
lateral and distal teeth on posterior margin. Coxa 2 subrectangular; coxa 7
rounded posteriorly.

VOLUME 93, NUMBER 3 623

Fig. 9. Iphimedia imparilabia, male: a, Gnathopod 1; b, Gnathopod propodus fixed finger
and dactyl; c, Gnathopod 2; d, Gnathopod 2 propodus fixed finger and dactyl; e, Pereopod 5;
f, Pereopod 6; g, Pereopod 7; h, Telson.

Antenna | peduncle article 1 without major teeth on distal margin; pe-
duncle article 2 with short, blunt dorsal tooth distally. Upper lip truncate
distally, with paired groups of distolateral setae. Mandible incisor elongate,
slightly spooned, accessory plate on left mandible elongate; palp article 2
with distal group of 3 setae; palp article 3 half length of article 2, with 9

624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distal setae. Lower lip lobes unequal in length distally, both incised. Maxilla
1 palp biarticulate, not reaching end of outer plate, articles subequal in
length, outer plate 10 stout setae apically; inner plate broken. Maxilla 2
inner and outer plates subequal in width; outer plate bearing setae equal to
three-fourths length of plate, much longer than those on inner plate. Max-
illiped palp 3-articulate, article 2 produced medially along most of length of
short article 3; outer plate without setae along outer margin; inner plate
truncate distally.

Gnathopod | subchelate; article 6 shorter than 5; dactyl and propodus
fixed finger shortened. Gnathopod 2 chelate; articles 5 and 6 subequal in
length; article 6 with setae along distal half of ventral margin; fixed finger
of propodus with dense cover of setae. Pereopod 5 and 6 basis posterodistal
corner narrowly rounded. Pereopod 7 basis posterior margin distally con-
cave, with few small serrations. Uropod | rami subequal, armed with short
setae. Uropod 2 outer ramus shorter than inner, both with a few short setae.
Uropod 3 rami sublanceolate, armed with short setae. Telson slightly emar-
ginate, with terminal pair of setae.

Etymology.—The species name is derived from the Latin impar, meaning
unequal and /abium meaning lip, referring to the unequal lobes of the lower
lip.

Distribution.—Known only from the type-locality.

Remarks.—The form of the dorsal body processes on this species closely
resembles those of several other species such as I. spinosa and I. capicola;
however, the following mouthpart features distinguish it from all other mem-
bers of the genus: lower lip with unequal lobes; very short palp article 3 and
lack of setae on outer plate outer margin on the maxilliped.

Iphimediella Chevreux, new synonymy

Iphimediella Chevreux 1911.
Pariphimediella Schellenberg 1931 (part).

Type-species.—l. margueritei Chevreux 1912.

Diagnosis [modified from Barnard (1969)].—Antenna | accessory flagel-
lum uniarticulate; epistome not broad, less than 3 times as wide as high;
upper lip entire or weakly incised; mandible incisor elongate, not exces-
sively thickened or broadened, oriented to cut in transverse plane; lower lip
without inner lobes, distal apices generally subacute; maxilla | palp biartic-
ulate, reaching end of outer plate; maxilliped palp 4-articulate, article 2
scarcely or not produced; gnathopods 1 and 2 chelate; telson slightly cleft.

Remarks.—Karaman and Barnard (1979) noted that only minor differ-
ences in the mandible and lower lip had been used by Schellenberg (1931)
to separate Pariphimediella and Iphimediella. We do not agree with Kar-
aman and Barnard, however, in including Pseudiphimediella nodosa Dana

VOLUME 93, NUMBER 3 625

and Pariphimediella glabra Schellenberg in the genus [phimediella as pres-
ently defined. These two species have mouthparts which are more broad-
ened and have mandible incisors which cut in the frontal plane as opposed
to the transverse-plane orientation seen in [phimediella. According to the
above diagnosis, the genus [phimediella is here considered to consist of the
following species: I. bransfieldi K. H. Barnard, J. cyclogena K. H. Barnard,
I. margueritei Chevreux, I. rigida K. H. Barnard, J. imparidentata (Bellan-
Santini), 7. microdentata (Schellenberg), 7. octodentata (Nicholls), I. ser-
rata (Schellenberg). The latter two may be synonymous as Nicholls (1938)
evidently mistook Schellenberg’s (1926) drawing of J. serrata to show only
a single dorsal process on pleonite 3. In his text however, Schellenberg
clearly stated: ‘“‘Die 2 letzten Meso- und die 3 Metasomsegmente tragen
dorsal je ein Paar grosse, nach ruchwarts gerichtete, annahernd gerade
Zahne’’ (1926:328). We have not yet examined specimens of either species
and thus are hesitant to synonymize the two at this time.

Key to species of [phimediella

1. Pleonites with enlarged mid-dorsal carinae, paired dorsal processes

MOM OROMMNENC man. shpe cea dnasitas dea ceexs I. imparidentata (Bellan-Santin1)
— Pleonites with distinct paired dorsal processes ................. 2
2. Without paired dorsal processes on pleonite 3 ................. 3
SVG pale dudorsal processes On, pleOnite sean see reine ee 5
3. Paired dorsal processes meet to form a ‘“‘U”’ dorsally, pleonite 3

Write: Sie nGh” ROSS INGE Bs ss 65 coke ob coo cco bose ene I. georgei
— Paired dorsal processes meet to form a “‘V”’ dorsally; pleonite 3

WA MOUES AUK CCL cy co: a ReNn ores, ees mice Cr aed re Oeil a 4

4. Coxa | smoothly rounded anteroventrally, pereopod 7 basis
broadly rounded behind, hind margin smooth; upper lip incised
OES Gee eS BORE as cee en nae I. bransfieldi K. H. Barnard
— Coxa | serrate or dentate below; pereopod 7 basis hind margin
serrate and nearly parallel to anterior margin; upper lip entire J. dis-
coveryl Nn. Sp.
5. Paired dorsal processes on pereonites 6 and 7 and on pleonites 1-

a re ee er Re ce th Meee eh ES Set as 6
— Paired dorsal processes on pereonite 7 and on pleonites 1-3 .... 7
6. Coxa | ventral margin concave, coxae 4—7 ventral and posterior
margins coarsely serrate, almost dentate .. J. serrata (Schellenberg)
— Coxa | ventral margin straight, coxae 4—7 ventral and posterior
MAoins ile WaSciialeur. : sem win taters cr I. octodentata (Nicholls)
7. Coxa7 posterior margin produced as an elongate process; pereopod
TESS WN Z OCSISHOR (ESN see es deeb nee oboe I. acuticoxa N. sp.

— Coxa 7 smoothly rounded behind, at most with small tooth; per-

626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

eopod 7 basis smoothly rounded with single small tooth at postero-

distal:CORME tae. sarrcriettien pens nee acdsee! bens «mete pa 8
8. Prominent medial tooth on antenna 1 peduncle article 1 extends

well past third peduncle article; coxa | tapering distally, ventral

MAKIN EN ARON ye Wty eee eae I. margueritei Chevreux
— Teeth on antenna | peduncle article | not extending past third ar-
ticle; coxa | not tapering, smooth at anteroventral corner ....... 9

9. Paired dorsal teeth form a distinct *‘V’’ shape; mid-dorsal keel pres-
ent on pleonites 1-3, additional upright keel present on urosomite

Tinayiaties till anernaiatid od ek at Rates Pe a8 ve ecan aaa I. rigida K. H. Barnard
— Paired dorsal teeth do not form a ‘‘V”’ shape, no mid-dorsal keel

PreSenth asec wa Ee aes ew awiowan eels Be i Eee 10
10. Dorsal processes short, stubby; telson only shallowly emarginate;

UPPekalip renee eo eee: I. microdentata (Schellenberg)
— Dorsal processes elongate, slender; telson distinctly cleft; upper lip

LINC ESC Ce i oireetigtitet leh Mae Scere i nah atly ce Ouest I. cyclogena K. H. Barnard

Iphimediella georgei n. sp.
Figs. 10, 11

Material.—Holotype: Islas Orcadas Cruise 19, Sta. 7, 8 Mar. 1979,
62°18.9’S, 55°13’W, 610 m, | 2 (28 mm) with eggs (USNM 173587). Para-
types: Eltanin Cruise 27, Sta. 1924, 27 Jan. 1967, 75°11’S, 176°13'W, 728-732
m, 2 juveniles (USNM 173588).

Diagnosis.—Body with paired dorsal processes on pereonite 7 and pleo-
nites 1 and 2; pleonite 3 with keel only, produced slightly backward into
acute tooth; coxa 1 anteroventral margin rounded, posteroventral margin
drawn downward into anteriorly directed hook.

Description.—Three pairs dorsal processes meet mid-dorsally to form
smoothly rounded ‘‘U.’’ Urosomite | with prominent mid-dorsal carina.
Head anterolateral sinus with acute tooth above and longer acute tooth
below. Coxa | anteroventral margin smoothly rounded; ventral margin be-
comes distinctly concave before forming anteriorly directed hook (easily
broken) at posteroventral margin. Coxae 2 and 3 similar in shape to coxa
1 but more slender, less rounded anteriorly and less concave ventrally.
Antenna | peduncle article 1 with prominent tooth projecting ventrally past
distal margin of peduncle article 2, with 2 smaller teeth projecting laterally
along midline; peduncle article 2 with teeth projecting along each lateral
margin past distal margin of article 3; accessory flagellum uniarticulate.
Antenna 2 longer than antenna | in female. Upper lip distinctly incised.
Mandible tapering, incisor toothed; molar prominent and setose; palp
3-articulate; accessory plate on right mandible toothed, left accessory plate
not as well developed. Lower lip entire, apices smoothly rounded, inner

VOLUME 93, NUMBER 3 627

a
\ \ |

| : i J Zo \\\

Fig. 10. Iphimediella georgei, female: a, Body, side view; b, Antenna | peduncle and basal
flagellar articles; c, Upper lip; d, Mandible; e, Mandible palp; f, Lower lip; g, Maxilla 1; h,
Maxilla 2; i, Maxilliped without palp; j, Maxilliped palp.

lobes absent. Maxilla 1 palp biarticulate, exceeding length of outer plate,
setae half way along inner margin; inner plate subtriangular with 18 plumose
setae on inner margin. Maxilla 2 inner and outer plates subequal in width;
setae extend along medial margin of inner plate. Maxilliped palp 4-articulate
with fourth article minute, covered by setose, hoodlike extension of article

628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Iphimediella georgei, female: a, Gnathopod 1; b, Gnathopod | propodus fixed
finger and dactyl; c, Gnathopod 2; d, Telson.

3; palp article 2 distinctly expanded but not produced along article 3; palp
article 3 longer than broad with plumose setae giving way to shorter, stouter
setae along distal margin; inner plate with plumose setae along distal and
medial margins. Gnathopods | and 2 chelate. Gnathopod | article 3 subequal
to article 4 but less than % length of article 6; article 5 subequal to article
6. Gnathopod 2 article 3 longer than article 4, 34 length of article 6; article
5 subequal to article 6. Pereopods 5 and 6 basis posterior margin smoothly
rounded proximally, drawn into a small tooth at posteroventral corner; pos-
terior margin slightly serrate. Telson longer than wide, shallowly cleft; distal
margin minutely irregular. Uropods 1 and 2 biramous. Uropod 1 peduncle
longer than rami; outer ramus slightly shorter than inner. Uropod 2 peduncle
slightly shorter than inner ramus; outer ramus shorter than inner. Uropod
3 either uniramous or missing inner ramus.

Etymology.—Named in honor of Dr. Robert Y. George.

Remarks.—This species can be easily distinguished by its dorsal arma-
ture. The only other [phimediella species with 3 pairs of dorsal processes
are I. bransfieldi and I. discoveryi. I. georgei differs from the latter 2 species
in possessing a dorsal keel on pleonite 3. Additionally, the dorsal processes
come together at the midline as a broadened “‘U”’ in J. georgei and as a
“Vv? in I. bransfieldi and I. discoveryi. The mouthparts of J. georgei are

VOLUME 93, NUMBER 3 629

typically iphimediellan with the expanded but not produced maxilliped palp
article 2 being similar to that seen in J. bransfieldi.

Iphimediella acuticoxa Nn. sp.
Figs. 12, 13

Material.—Holotype: Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S,
54°43’'W, 210-220 m, 1 juvenile, 5 mm (USNM 173589).

Diagnosis.—Body with paired elongate dorsal processes on pereonite 7
and pleonites 1—3; lower lip notched apically; maxilla 1 palp as long as outer
plate; pereopod 7 posterior margin excavate between 2 large teeth; coxae
5-7 hind margins acutely produced.

Description.—Head anteroventral corner with shallow sinus bounded by
pair of subacute processes. Body with paired elongate dorsal processes on
pereonite 7 and pleonites 1-3, last 2 pairs being somewhat upturned. Pleo-
nite 3 with slight mid-dorsal keel. Pleurae of pereonites 5—7 extend outward
as acute processes. Epimeral plates 1-3 each with lateral and distal teeth
on posterior margins. Coxae | and 2 distally rounded; coxae 5-7 postero-
distal corners acutely produced.

Antenna | peduncle article 1 with 2 short dorsal teeth on distal margin,
without ventral tooth; peduncle article 2 with dorsal, ventral and lateral
teeth distally. Upper lip narrowly rounded, entire. Mandible elongate, with
accessory plate on both left and right sides; molar a short, setose, flat-topped
protuberance; palp article 2 with single distal seta, article 3 with covering
of fine setae, longer setae on distal third of article. Lower lip with apical
notch. Maxilla 1 palp biarticulate, exceeding outer plate, article 2 twice
length of article 1; inner plate more than half length of outer plate, with 7
strong, plumose setae distally; outer plate with 5 stout setae at apex. Maxilla
2 inner and outer plates subequal in width; outer plate with long, simple
setae at apex slightly longer than those on inner. Maxilliped palp 4-articu-
late, article 2 expanded but not produced along article 3, latter long and
slender; palp article 1 only slightly longer than article 2, article 4 minute;
outer and inner plates apically subacute; inner plate with setae only on distal
half of medial margin.

Gnathopod | chelate, generally devoid of setae; dactyl: partly fused to
propodus; propodus finger bears 5 long plumose setae with backwardly-
directed setules. Gnathopod 2 subchelate; article 6 longer than article 5,
both articles sparsely setose; dactyl and fixed finger of propodus equal in
size and shape. Pereopods 5-7, basis posterior margin excavate between
dorsal and ventral teeth. Uropod | rami weak, outer ramus unarmed. Uro-
pod 2 outer ramus shorter than inner. Telson shallowly V-cleft, with 2 ter-
minal pairs of setae.

Etymology.—The name refers to the shape of coxa 7.

630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

———=
SS S=

L—
ZZ

SZ

:

Hacer ITY

Fig. 12. Iphimediella acuticoxa, juvenile: a, Body, side view; b, Head; c, Antenna 1 pe-
duncle articles 1 and 2; d, Antenna 2 peduncle articles 2—4; e, Upper lip; f, Left mandible with
palp; g, Right mandible; h, Lower lip; i, Maxilla 1; j, Maxilla 2.

Distribution.—Known only from locality listed above.

Remarks.—This species is unique in the genus with respect to the acutely
produced hind margins of coxae 5—7. Maxilliped palp article 2 is expanded
medially and slightly produced; however the overall aspect of the palp is
more closely related to several other [phimediella species than it is to

species of I[phimedia.

VOLUME 93, NUMBER 3 631

1
MY HN

b

Fig. 13. Iphimediella acuticoxa, juvenile: a, Maxilliped; b, Maxilliped distal end of article
3 and minute article 4, seta omitted; c, Gnathopod 1; d, Gnathopod 1 propodus fixed finger
and dactyl; e, Gnathopod 2; f, Gnathopod 2 propodus fixed finger and dactyl; g, Pereopod 5;
h, Pereopod 6; i, Pereopod 7; j, Telson.

Iphimediella bransfieldi Barnard 1932
Figs. 14, 15

Iphimediella bransfieldi Barnard, 1932:119 (part).—Nicholls, 1938:70, fig.
37J.

632 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 14. Iphimediella bransfieldi, female: a, Body, side view; b, Head; c, Antenna | peduncle
and uniarticulate accessory flagellum; d, Upper lip; e, Mandible; f, Lower lip; g, Maxilla 2.

Material.—Discovery Sta. 175, 2 Mar. 1927, 63°17'S, 59°48’W, 200 m
(BMNH No. 1936. 11. 2. 1080-1081) 1 2 ovigerous, 22 mm (lectotype).
Eltanin Cruise 51, Sta. 5761, 8 Feb. 1972, 76°01.6’S, 179°49.9’E, 388-399
m, 1 2, with eggs, 22 mm.

Diagnosis.—Body with short, paired dorsal processes on pereonite 7 and
pleonites | and 2; antenna | peduncle article 2, ventrodistal tooth longer

VOLUME 93, NUMBER 3 633

i

Fig. 15. Iphimediella bransfieldi, female: a, Maxilla 1; b, Maxilliped; c, Maxilliped distal
end of article 3 and minute article 4, seta omitted; d, Telson.

than dorsal tooth; coxae 1—4 anteroventral margins smooth; upper lip emar-
ginate.

Description.—The following supplements the description given by Bar-
nard (1932). Body with short, wide dorsal processes on pereonite 7 and
pleonites | and 2, each pair forms distinct *‘V’’ dorsally; urosomite | without
mid-dorsal carina. Head anterolateral margin with narrow sinus bounded
below by short tooth; rostrum shorter than antenna | first peduncle article.
Coxa | anteroventral corner broadly rounded.

Antenna | peduncle short, ventrodistal tooth on first article with 2 short
accessory teeth; peduncle article 2, ventral tooth longer than dorsal; acces-
sory flagellum uniarticulate. Upper lip slightly incised, ventrolateral margins
with dense covering of short hairs. Right mandible incisor and accessory
plate multidentate; molar conical, strong, with apical tuft of short hairs.
Lower lip apically narrow, without inner lobes. Maxilla 1 palp article 2
elongate, armed along distal half of inner margin; outer plate subrectangular,
distal edge with strong spines, medial edge setose distally; inner plate sub-
acute, distal half of medial margin with short plumose setae. Maxilla 2 inner
and outer plates longer than wide. Maxilliped outer plate subovate, unarmed
along proximal half of lateral margin; palp with 4 articles, the last minute,

634 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\
i al We y

Fig. 16. Iphimediella discoveryi, female: a, Body, side view; b, Head; c, Antenna | peduncle
articles and accessory flagellum; d, Upper lip; e, Right mandible; f, Left mandible; g, Lower
lip; h, Maxilla 1; i, Maxilla 2; j, Maxilliped.

detectable only at high magnification, partly covered by hoodlike extension
of article 3; palp article 3 longer than article 2, subequal in length to article
1. Telson with broad V-shaped cleft, apices acute.
Distribution.—South Shetland Islands, approximately 200 m.
Remarks.—K. H. Barnard (1932) briefly described, but did not illustrate,

VOLUME 93, NUMBER 3 635

d

Fig. 17. Iphimediella discoveryi, female: a, Gnathopod 1; b, Gnathopod 2; c, Pereopod 5;
d, Telson distal margin.

this species from 2 ovigerous syntypes collected off the South Shetland
Islands. In his description he noted that the smaller of the 2 specimens
differed somewhat from the larger. We have examined both specimens and
found them to differ considerably, especially in their mouthpart morphol-
ogy. Since the smaller of the 2 specimens seemed to represent to K. H.
_ Barnard exceptions to the typical J. bransfieldi, we have chosen it as the
holotype of a new species, J. discoveryi, which is described below. The
larger specimen becomes the lectotype of J. bransfieldi.

Iphimediella discoveryi n. sp.
Figs, 16,17

~Iphimediella bransfieldi Barnard, 1932:119 (part).

Material.—Holotype: Discovery Sta., 175, 2 Mar. 1927, 63°17'S, 59°48’W,
200 m (BMNH No. 1936. 11. 2. 1080-1081), 1 2 ovigerous, 14 mm. Para-
type: Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S, 54°43’W, 210-220
mz, | juvenile.

Diagnosis.—Body with short, paired dorsal processes on pereonite 7 and
_ pleonites | and 2 meeting to form ‘‘V”’ dorsally; antenna 1 peduncle article
_ 2 dorsodistal tooth longer than ventral; coxae 1—4 anteroventral margins and
pereopods 5-7 basis hind margin serrate; upper lip entire.

636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Description.—Body with short, paired dorsal processes on pereonite 7
and pleonites 1 and 2; processes meet mid-dorsally to form a ‘‘V’’; urosomite
| without mid-dorsal carina. Head anterolateral margin with shallow sinus
bounded below by long, acute tooth. Coxa | tapering, anteroventral corner
not rounded, ventral margin slightly concave, serrate.

Antenna | peduncle article 2 with dorsal tooth longer than ventral. Upper
lip broadly rounded with setae along ventral margin. Mandible stout relative
to I. bransfieldi; molar a slight, setose protuberance; accessory plate on left
mandible toothed, on right mandible smooth. Lower lip apically truncate,
without inner lobes. Maxilla 1 palp article 2 armed with hairlike setae along
most of inner margin; outer and inner plates similar to J. bransfieldi. Max-
illiped palp with minute article 4; palp article 2 nearly as wide as long; article
3 longer than article 2, article | longer than article 3; outer plate as broad
as long, distal margin with elongate plumose setae. Pereopod 5 basis anterior
and posterior margins parallel, posterior and ventral margins serrate. Telson
shallowly cleft, apices rounded.

Etymology.—The species is named in honor of the R.R.S. Discovery.

Distribution.—Antarctic Peninsula, 200-220 m.

Remarks.—The form and number of paired dorsal processes on the body
gives this species the appearance of J. bransfieldi. The following features,
however, readily distinguish the two: 1) coxal plates 1-4 anteroventral margins
and pereopod 5-7 basis hind margins serrate in J. discoveryi and smooth in
I. bransfieldi; 2) pereopod 5 basis much less expanded in J. discoveryi than
in I. bransfieldi; 3) antenna 1 peduncle article 2 dorsal tooth longer than
ventral in J. discoveryi whereas the reverse is the case in I. bransfieldi; 4)
upper lip ventral margin entire in J. discoveryi and emarginate in I. brans-
fieldi; 5) telson apices rounded in /. discoveryi and subacute in J. bransfieldi.

Iphimediella cyclogena Barnard 1930
Figs. 18, 19

Iphimediella cyclogena Barnard, 1930:349, fig. 23.
Iphimediella intermedia Nicholls, 1938:71, fig. 37 (new synonymy).

Material.—Eltanin Cruise 12, Sta. 1002, 15 Mar. 1964, 62°40’S, 54°44—
45'W, 265 m, | 2; Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S,
54°43'W, 210-220 m, 3 3d. Terra Nova Sta. 4 (194?) (BMNH No. 1930.
8.1: 180-182) (Oates Land7?).

Diagnosis.—Body with 4 pairs elongate, smooth dorsal processes; coxa
1 anteroventral corner narrowly rounded, ventral margin concave; coxae 2
and 3 tapering distally, posteroventral corner produced ventrally; antenna
1 peduncle article | ventrodistal tooth extends beyond distal margin of pe-
duncle article 2; head lateral margin with narrow notch bounded above by
sharp point and below by blunt lobe.

VOLUME 93, NUMBER 3 637

Fig. 18. Iphimediella cyclogena, Terra Nova specimen, male: a, Head and coxa 1; b,
Pleonites 2, 3 and urosome; c, Antenna | peduncle and accessory flagellum; d, Upper lip; e,
Mandible; f, Mandible palp; g, Lower lip; h, Telson. Male, Eltanin Sta. 1003; i, Telson.

Description.—The following supplements the description given by K. H.
Barnard (1930). Antenna 1 with minute accessory flagellum. Upper lip
slightly incised. Mandible molar with small apical tuft of short hairs. Lower
lip without inner lobes. Maxillae | and 2 similar to J. bransfieldi. Maxilliped
palp article 4 minute, as wide as high, covered by hoodlike extension of
article 3; article 2 shorter than articles | and 3, only slightly produced me-

638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 19. Iphimediella cyclogena, Terra Nova specimen, male: a, Maxilla 1; b, Maxilla 2;
c, Maxilliped; d, Gnathopod 1; e, Gnathopod | propodus fixed finger and dactyl; f, Gnathopod
2; g, Gnathopod 2 propodus and dactyl, setae omitted.

dially. Gnathopod | chelate, sparsely setose; dactyl overhangs fixed finger |
of propodus. Gnathopod 2 chelate, setose along ventral margin of articles
4, 5 and 6; dactyl shorter than fixed finger of propodus. Telson shallowly
Cleft

Distribution.—Shelf off tip of Antarctic Peninsula, Oates Land, Ross Sea,
Adelie Coast, 329-540 m.

VOLUME 93, NUMBER 3 639

Remarks.—We compared the illustrations of J. cyclogena given by K. H.
Barnard (1930:350) with specimens borrowed from the British Museum and
found the following variations: 1) coxa | as illustrated appeared to be sharply
angled at the anteroventral corner, however, all specimens we examined
had a rounded anteroventral corner; 2) the length of the dorsal processes
on pleonite 3 were proportionately longer and slightly upturned in smaller
versus larger specimens; 3) the head lateral margin as illustrated appeared
to be very diagrammatic since the lobe below the sinus is actually much less
rounded; 4) the shape of the telson differed from specimen to specimen.
Nicholls (1938:73) distinguished J. intermedia from I. cyclogena using ‘‘the
shape of the head, of the first coxal plate, outline of peraeon segment 7 and
_ pleon segments 1-3, in proportions and other details of peduncular joints of
first and second antennae and the presence of secondary cutting plates on
both mandibles.’’ For all specimens of J. cyclogena we examined, the above
characters had to be rejected either on the basis of overall variability or
because of incorrect interpretation of the illustrations. Thus we conclude
that J. intermedia is a synonym of I. cyclogena.

Pseudiphimediella Schellenberg

| Pseudiphimediella Schellenberg, 1931:119.
Pariphimediella Schellenberg, 1931:121 (part) (new synonymy).

Type-species.—Amphitoe nodosa Dana, 1853 (original designation).

Diagnosis.—Upper lip incised or sinuous; epistome not more than 3 times
as wide as high; mandible incisor broad, thin-edged, multi-toothed, and
curved toward midline so as to cut in the frontal plane; maxilliped palp
article 2 broadened, not produced, outer plate broad.

Remarks.—Schellenberg (1931) distinguished Pseudiphimediella from
Pariphimediella on the basis of the former possessing a deeply incised upper
lip and broadened articles 1 and 2 on the maxilliped palp. As discussed
under the remarks for [phimediella, Schellenberg’s P. glabra differs from
other [phimediella species (sensu Karaman and J. L. Barnard, 1979) but is
similar to Pseudiphimediella nodosa in possessing mouthparts with a broad-
ened aspect and, more significantly, in the orientation of the mandible in-
cisor which cuts in the frontal plane. The genus Pseudiphimediella as pres-
ently defined thus contains the 2 species P. nodosa (Dana) and P. glabra
(Schellenberg).

Pseudiphimediella glabra (Schellenberg, 1931) new combination
Fig. 20
Pariphimediella glabra Schellenberg, 1931:121, fig. 65; pl. 1, fig. B.
Material.—Eltanin Cruise 9, Sta. 740, 18 Sept. 1963, 56°06— 07'S, 66°19-

640 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

“yy,

|

|

:

AMY:
jG |

duncle ventral view; d, Antenna | peduncle dorsal view; e, Mandible; f, Mandible palp; g,
Lower lip; h, Maxilla 1; i, Maxilliped; j, Gnathopod 2 propodus fixed finger and dactyl; k,

Fig. 20. Pseudiphimediella glabra, female: a, Body, side view; b, Head; c, Antenna | pe- |
Pereopod 6; 1, Pleonite 3 and urosomites | and 2; m, Telson.

30’W, 384-494 m, 1 2 with eggs, 2 dd; Eltanin Cruise 11, Sta. 977, 13 Feb.
1964, 52°32’S, 63°53’W, 229 m, 1 2 with eggs, 2 dd, 1 juvenile.
Diagnosis.—Body with paired dorsal processes on pereonites 6 and 7 and
pleonites 1—3; urosomite 1 with strong mid-dorsal keel; maxilla 1 inner plate |
with 9-10 stout plumose setae; maxilliped inner plate medial margin armed |

with several setae.

VOLUME 93, NUMBER 3 641

_ Fig. 21. Pseudiphimediella nodosa, female: a, Mandible; b, Maxilla 1; c, Maxilliped.

Description.—The following supplements the description given by Schel-

lenberg (1931). Antenna | peduncle article | with 3 teeth along distal margin,
: 2 ventral and 1 lateral. Mandible shortened, with broad, thick incisor; ac-

cessory plate with elongate cutting edge; palp article 3 half length of article
2, armed along distal half of ventral margin. Lower lip distal apex broad-

ened, sinuous. Maxilla 1 palp biarticulate, reaching beyond distal half of
medial margin; outer plate subrectangular; inner plate armed with short,
stout plumose setae. Maxilliped palp 4-articulate, article 3 apex extended
: slightly covering minute fourth article (seen using SEM); article 2 expanded
medially but only slightly along article 3, outer plate broadly ovate, armed
with stout plumose setae. Gnathopods 1 and 2 chelate; fixed finger of gnath-
-opod 2 propodus much wider than dactyl, heavily armed with setae.
— Distribution.—Burdwood Bank, Falkland Islands and Magellanic area, 2—
494 m.

Remarks.—The specimens examined by us differed from the illustrations
of Schellenberg (1931) in the following features: mid-dorsal keel on uroso-
mite | varied considerably in size; pereopod 6 basis hind margin lower
corner was without a tooth; coxa 1 was more rounded distally.

Pseudiphimediella nodosa (Dana, 1853)
Fig. 21

Iphimedia nodosa Dana, 1853—55:928, pl. 63, figs. 3A, B.
Pseudiphimediella nodosa: Schellenberg, 1931:119, fig. 64, pl. 1, fig. A.
Iphimediella nodosa: K. H. Barnard, 1932:119, fig. 67.

642 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Material.—William Scoresby Sta. 85, 25 March 1927, 8 mi. S, 66°E of
Lively Island, East Falkland Island, 79 m, 1 2 with young (BMNH 1936.
11. 2. 1082).

Diagnosis.—Body with paired dorsal processes on pereonite 7 and ple-
onites 1-3; urosomite 1 with mid-dorsal keel; maxilla 1 inner plate with few,
elongate plumose setae; maxilliped inner plate medial margin unarmed.

Description.—The following supplements the descriptions given by Schel-
lenberg (1931) and K. H. Barnard (1932). Right mandible with small acces-
sory plate; incisor spoon-shaped, multitoothed; mandible oriented such that
incisors move in frontal plane; palp article 2 elongate, with 2 setae on distal
corner. Maxilla 1 inner plate weakly armed; palp article 2 broad. Maxilliped
palp article 4 minute, covered by hoodlike extension of article 3; palp article
2 broadened, not produced; inner plate medial margin weakly armed.

Distribution.—Falkland Islands, Magellanic region; low tide to 150 m.

Remarks.—The illustrations given by Schellenberg (1931) for this species
are extremely diagrammatic and do not contain the information critical for
evaluating its relationships. On examination of British Museum material we
found that the maxilliped palp articles 1 and 2 were not appreciably broad-
ened compared to some species of I[phimediella and Gnathiphimedia. P.
nodosa was, however, very distinct in the shape and orientation of the
mandible. We have looked at mandible orientation in several genera of
Acanthonotozomatidae and conclude that several generic complexes can be
discerned using this feature. The mandible incisors of Pseudiphimediella
and Maxilliphimedia are oriented such that they meet and thus cut or pinch
along the frontal plane of the head, which is the condition typical of most
gammaridean anphipods. Guathiphimedia, Iphimediella and most species
of Iphimedia have mandible incisors that are oriented to cut along the trans-
verse plane of the head. It is this orientation which has led to the description
of acanthonotozomatid mouthparts as being arranged in a “‘conical bundle.”’
Within the transversely biting mandible group there is some variation in the
form of the incisor, for example, in Gnathiphimedia the thickened, smoothly
rounded incisor edge apparently functions in a ‘‘crushing’’ mode while the
blade-like incisor of Iphimediella probably functions in a cutting mode.

Paramphithoidae

Epimeria Costa

Epimeria Costa in Hope, 1851:46.
Pseudepimeria Chevreux, 1911:1167 (new synonymy).
Subepimeria Bellan-Santini, 1972:225.

Type-species.—Gammarus corniger J. C. Fabricius 1779.
Diagnosis (emended from J. L. Barnard, 1969).—Rudimentary accessory
flagellum; mandible molar large, ridged; lower lip lacking inner lobes; max-

VOLUME 93, NUMBER 3 643

illiped palp 4-articulate; gnathopods simple or subchelate, dactyls often spi-
nose, much shorter than propodus; coxae 4—5 together forming a more or
less crescentic curve below.

Remarks.—As pointed out by Karaman and Barnard (1979) the characters
used by Bellan-Santini (1972) to establish the genus Subepimeria (presence
of uniarticulate accessory flagellum, simple gnathopods) were not distinctive
at the generic level. All species of Epimeria examined in this study have
been found to have an uniarticulate accessory flagellum. Karaman and Bar-
nard (1979) also suggested that E. geodesiae (Bellan-Santini) showed very
weak gnathopodal palms and thus were not, in fact, simple. Weak palms are
also found in E. puncticulata K. H. Barnard. These latter species intergrade
to the true simple gnathopod seen in Pseudepimeria Chevreux, leaving no
characters by which Pseudepimeria and Epimeria can be distinguished.

Epimeria rimicarinata n. sp.
igs. 22, 23

| Material.—Holotype, Eltanin Cruise 27, Sta. 1875, 15 Jan. 1967, 72°32'S,
171°26-28’E, 337-329 m, 1 non-ovigerous °, 35 mm (USNM 173590); para-
type, Cruise 32, Sta. 2080, 31 Jan. 1968, 75°50-52’S, 173°08’W, 468-474 m,
.1 3, 35 mm (USNM 173591).
| Diagnosis.—Body with mid-dorsal carinae on pereonites 3-7, pleonites
1-3 and urosomites 1 and 2, those on pereonites 5-7, pleonites 1-3 and
-urosomite 1 distinctly cleft into anterior and posterior teeth; pereonites 5—
7, pleonites 1-3 and urosomite | with dorsolateral carinae; coxa 4 broadly
quadrate ventrally, posteroventral corner slightly produced; coxa 5 pos-
teroventral corner slightly, bluntly produced.
| Description.—Body bluntly processiferous. Mid-dorsal carinae on per-
eonites 3-7, pleonites 1-3 and urosomites | and 2, those on pereonites 5-7,
| pleonites 1-3 and urosomite | distinctly cleft into 2 consecutive posteriorly-
| directed teeth. Pereonites S—7, pleonites 1—3 and urosomite | with small
dorsolateral carinae. Pleonites 1-3 and urosomite 1 each bear one additional
lateral protuberance anteriorly. Coxae 1-3 narrowly subrectangular with
anteroventral corners rounded, posteroventral corners angular. Coxa 4 ven-
tral margin broadly quadrate, posteroventral corner slightly produced. Coxa
_5 posteroventral margin bluntly produced. Rostrum long, strongly curved
ventrally, extending beyond antenna | peduncle article 1. Antenna | shorter
than antenna 2 in both sexes, with uniarticulate accessory flagellum; pe-
| duncle article 1 twice length article 2, with several small teeth on mediodistal
margin. Upper lip incised, lobe apices setose. Mandible molar large, crush-
_ing (‘‘triturative’’); incisor multidentate; left lacinia mobilis multidentate,
right lacinia mobilis bidentate; palp articles 2 and 3 subequal, heavily setose.
_Lower lip entire, without inner lobes. Maxilla | inner plate subtriangular
_with 9 apical plumose setae; outer plate with 14 slender spines on distal

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

h

‘

I

\

\

Ay
x

:

hy

WS
\Y \\

KE \
SOA QMA, \
S

Fig. 22. Epimeria rimicarinata, female: a, Body, side view; b, Antenna 1; c, Upper lip; d,

Mandible; e, Mandible palp; f, Lower lip; g, Maxilla 1; h, Maxilla 2.

margin; palp article 2 distally armed with short, stubby spines extending

proximally along 44 inner margin, grading into fine setae. Maxilla 2 inner
and outer plates approximately equal in width, heavily setose along distal
margins. Maxilliped inner plate with plumose setae along medial and distal
margins; outer plate with nonplumose setae distally, short submarginal
spines medially; palp 4-articulate, article 2 longest; palp article 4 with 8
slender spines on inner margin. Gnathopods subchelate, similar, gnathopod

VOLUME 93, NUMBER 3 645

Fig. 23. Epimeria rimicarinata, female: a, Maxilliped, without palp; b, Maxilliped palp; c,
Gnathopod 1; d, Gnathopod 2; e, Gnathopod 2 propodus and dactyl; f, Pereopod 5; g, Pereopod
6; h, Pereopod 7; i, Telson.

/

| 1 slightly smaller than gnathopod 2, articles 5 longer than articles 6; dactyls
| with numerous spines on inner margin. Pereopod 5 basis with small proximal
| lobe on posterior margin, distal 24 posterior margin parallel to anterior mar-
/ gin. Pereopods 6 and 7 proximal lobes of basis posterior margins succes-
sively enlarged, producing incised appearance. Telson cleft 4, setae on
| apices.

|

4

646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON |

Etymology.—The name is derived from the Latin rima = cleft and cari-
na — keels |

Distribution.—Known only from the Ross Sea. |

Remarks.—This species is most easily distinguished by the cleft mid-dor- |
sal carinae, a feature thus far unique in the genus. |

Parepimeria Chevreux

Parepimeria Chevreux, 1912:1168.
Parepimeriella Schellenberg, 1931:165 (new synonymy).

Type-species.—Parepimeria crenulata Chevreux, 1912. |

Diagnosis.—Antenna 1 with untarticulate accessory flagellum; mandible |
with large, ridged molar; lower lip with inner lobes; maxilliped palp 4-artic- |
ulate, article 3 not produced distally, inner plate not reaching palp article |
2; gnathopods similar; telson entire. |

Remarks.—The generic diagnosis has been modified to include a 4-articled |
palp on the maxilliped, a feature which Chevreux evidently did not see due |
to the dense cover of setae distally on article 3. Schellenberg (1931) distin- |
guished Parepimeriella from Parepimeria on the basis of its rudimentary |
rostrum, widely separated outer lobes of the lower lip, and broadly rounded
distal margin of gnathopod 2 coxa. Barnard (1932) pointed out the variable |
nature of the rostrum length; the other 2 characters are differences only of |
degree as evidenced by the new species described herein. The genus as now |
constituted contains the following species: P. crenulata Chevreux 1912, P. |
bidentata Schellenberg 1931, P. irregularis (Schellenberg 1931); P. major }
K. H. Barnard 1932; and P. minor n. sp.

Key to species of Parepimeria

1. Rostrum deflexed, not extending along article 1 of antennal ..... 2 ||
— Rostrum extends out in front of head, at least to half length of antenna |

WArtiClen noose eee OS ee 4
2. Epimeral plate 3 posterior margin tooth ...................200-- 3 |i

Epimeral plate 3 posterior margin smooth ..P. major K. H. Barnard 1932
3, Coxa 2 distal margm broadly rounded -ee 5. ..5-5- 1.) eee
Pe ey I A neh ch sa he | P. irregularis (Schellenberg 1931) |

— Coxa 2 distal margin narrowly rounded .............. P. minor n. sp. |
4. Last 3 pereon segments with mid-dorsal and pair of sub-dorsal cari- |

TNC ieee tec uop Sch oe doit Miia See Ack LS re: P. crenulata Chevreux 1912 |
— Last 3 pereon segments lacking mid-dorsal carina ...............
HP Ordre 5 SN erg ae ets ene Ne Raa es HR Woe, P. bidentata Schellenberg 1931

|
|
)
|
;

-

VOLUME 93, NUMBER 3 647

Parepimeria bidentata Schellenberg
Fig. 24

_Parepimeria bidentata Schellenberg, 1931:164.
_ Parepimeria crenulata K. H. Barnard, 1932:179 (part).

Material.—Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41'S, 54°43’W,
210-220 m, 5 spec.; Discovery Sta. 42, 1 April 1926, off mouth of Cumber-

_jand Bay, South Georgia, 120-204 m, 1 6, 12 22 ; Isla Orcadas Cruise 19,

|
:
|
|
|
)
|

Sta. 27, 29 Mar. 1979, 53°57'S, 36°08’W, 180 m, 1 @ with eggs.

Diagnosis.—Rostrum long, extending horizontally outward from head;
body with paired dorsolateral processes on pereonites 3 to 7; pleonites 1
and 2 with backwardly-directed mid-dorsal carinae; pleonite 3 with acute
mid-dorsal tooth; epimeral plate 3 posterior margin serrate; maxilliped palp
article 4 strong; gnathopods | and 2 dactyl shorter than propodus.

Description.—Body with paired sub-dorsal carinae on pereonites 3-7, in-
creasing in length posteriorly; pleural tubercles pronounced on pereonites
4-6; pleonites 1—3 with mid-dorsal carinae, the first two posteriorly-directed,
the last recurved anteriorly; posterior margin of epimeral plates 2 and 3
strongly serrate; coxa | anteroventral corner subacutely produced anterior-
ly, posteroventral corner rounded, ventral margin slightly serrate; coxa 3
anteroventral corner bluntly produced downward; coxa 4 narrowly rounded
distally. Rostrum long, extending beyond distal end of antenna | peduncle
article 1, often beyond distal end of peduncle article 2; head angle subacute;
eyes large, protruding from side of head. Antenna | with uniarticulate ac-
cessory flagellum; peduncle articles 1-3 subequal in length; peduncle and
flagellar articles bear ventrally-directed long, thin setae. Antenna 2 longer
than antenna 1; peduncle article 5 slightly longer than article 4. Mandible
with strong, triturative molar; left lacinia mobilis with approximately 4 teeth,
right lacinia mobilis bifid; palp articles 2 and 3 elongate, approximately 5
times as long as wide; both articles armed ventrally throughout their length
with long stiff setae. Maxilla 1 palp with subapical group of setae; inner
plate with 3 terminal setae, outer plate with 7. Maxilla 2 inner and outer
plates terminally rounded, outer plate without setae along medial margin.
Maxilliped with 4-articulate palp; palp terminal article slender, spinelike,
three-fourths length of article 3; article 3 with dense terminal cluster of
setae; outer plate reaches less than half length of palp article 2, not falciform;
inner plate short, reaching only to end of first palp article.

Gnathopod 1 simple; dactyl greater than half length of article 6, slender,
spiniform; articles 5 and 6 subequal in length; article 5 proximally expanded,
with 2 rows of ventrally-directed, long, stiff setae, one medial, the other
along ventral margin. Gnathopod 2 similar to gnathopod 1, but articles 5
and 6 more elongate. Pereopods 5 and 6 basis subrectangular, posteroventral

a, catty /»
| Wy

Fig. 24. Parepimeria bidentata, female: a, Body, side view; b, Maxilliped; c, Gnathopod 1.

corner extended as a lobe. Pereopod 7 basis tapers distally. Uropods 1-3, |
outer ramus shorter than inner. Telson entire, broadly rounded. |

Distribution.—Shelf off Joinville Island (Antarctic Peninsula), Shag |
Rocks, South Georgia. |

Remarks.—In most details, especially with respect to the mouthparts, P. |
bidentata and P. crenulata are very similar. Chevreux (1912) did not record |
an article 4 on the maxilliped palp but since it is difficult to see in other }
species, it is likely that it escaped Chevreux’s notice. The major difference |
between P. bidentata and P. crenulata then, becomes the presence of a |
mid-dorsal carina on pereonites 2-7 in P. crenulata which is lacking in P. |
bidentata. We re-examined the specimens from Discovery Station 42 |
(Barnard 1932) and found them to correspond to the description of P. bi- |
dentata rather than P. crenulata as determined by K. H. Barnard. It is not |
known whether the remainder of the material examined by K. H. Barnard |
also is P. bidentata. !

Parepimeria minor n. sp.
Fig. 25

Material.—Eltanin Cruise 6, Sta. 410, 31 Dec. 1962, 61°18—20’'S, 56°09-
10’W, 220-240 m, 3 spec., 5.0 mm holotype (USNM 173592); 4.0 mm and |
2.0 mm paratypes (USNM 173593). |

‘’ VOLUME 93, NUMBER 3 649

Fig. 25. Parepimeria minor, female: a, Head and pereonite 1; b, Pereonites 6 and 7 and
pleonites 1-3; c, Mandible; d, Lower lip; e, Maxilla 1; f, Maxilliped; g, Gnathopod 1; h,
| Gnathopod 2; i, Pereopod 4.

/

| Diagnosis.—Rostrum short, blunt; head with dorsal keel; body with
paired dorsolateral processes of variable strength on peronites 2—7; mid-
| dorsal carinae on pleonites 1 and 2, pleonite 3 dorsally keeled; epimeral
| plate 3 posterior margin serrate; maxilliped palp article 4 strong; gnathopod
| 1 articles 6 and 7 subequal in length.

: Description.—Head, rostrum short, blunt; dorsal keel extends length of

650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 26. Parandaniexis dewitti, female: Body, side view.

head; antennal angle subacute; eye slightly protruding, ommatidia not con-
tiguous. Body with paired, low, sub-dorsal processes on pereonites 2-7.
Pleonites 1 and 2 with mid-dorsal carinae; pleonite 3 dorsally keeled; epi-
meral plate 3 posterior margin serrate. Mandible molar strong, triturative;
incisor multidentate; lacinia mobilis of right mandible with 3 teeth; palp
articles 2 and 3 elongate, subequal, with dense rows of setae along ventral
margin; palp article 3 with proximal group of setae dorso-medially. Lower
lip with nearly coalesced inner lobes; outer lobes broadly rounded, 1-3 blunt
setae medially near apex. Maxilla 1 palp apex with 6 stout setae; inner plate
with 4 short plumose setae terminally. Maxilliped palp 4-articulate; article
4 strong, as long as article 3; outer plate extends halfway along palp article
2; inner plate reaches base of palp article 2. Gnathopod 1 simple; coxa
subacutely produced anteriorly; article 7 as long as article 6; article 5 proxi-
mally expanded. Gnathopod 2 similar to gnathopod | but slightly larger;
coxa narrowly rounded distally. Coxa 4 broadly rounded distally, posterior
margin excavate proximally. Telson entire.

Etymology.—The name refers to the small size of the species.

Distribution.—Known only from the locality listed above.

Remarks.—This species differs from the other species which have a short
rostrum by the serrate posterior margin of epimeral plate 3 (smooth in P.
major) and the narrow distal margin of coxa 2 (broadly rounded in P. ir-
regularis). K. H. Barnard (1932) described an aberrant form and a variety
(miothele) of P. crenulata which seem to be very similar to P. minor.
Barnard’s var. miothele differs from P. minor in having no pleural tubercles
on the pereonites and a short non-deflexed rostrum which extends halfway
along antenna | peduncle article 1. The aberrant form is described as having
a short rostrum and, on pleonite 3, only a mediodorsal keel. Barnard does
not indicate whether or not the rostrum is deflexed so it is not possible to
determine its affinities with P. minor.

VOLUME 93, NUMBER 3 651

Stegocephalidae
Parandaniexis Schellenberg
Parandaniexsis Schellenberg, 1929.

Type-species.—Parandaniexis mirabilis Schellenberg, 1929.

Diagnosis (from J. L. Barnard, 1969).—Mandible incisor smooth; maxilla
1 palp biarticulate; maxilla 2 outer plate not geniculate or gaping; maxilliped
palp article 2 not produced; pereopods 5 and 6 article 2 slender; pereopod
7 article 2 broad; pereopod 4 subchelate; telson entire.

Parandaniexis dewitti n. sp.
Figs. 26, 27

Material.—Islas Orcadas Cruise 575, Sta. 38, 22 May 1975, 57° 00.4’S,
26°10.1’W, 2,740-2,757 m, 2 2°: holotype 42 mm, with eggs, USNM
173594; paratype 32 mm, USNM 173595.

Diagnosis.—Body with strong mid-dorsal carinae on pleonites 1-3, elon-

- gate, thin, mid-dorsal tooth on urosomite 1; antenna 1 basal flagellar article
only slightly longer than peduncle; pereopod 4 subchelate, propodus with
- slight proximal lobe on ventral margin.
Description.—Ovigerous female. Head submerged in pereonite |. Eye
_ lacking. Body with strong mid-dorsal carinae on pleonites 1-3. Urosomite
1 with mid-dorsal elongate curved tooth. Epimeral plates 2 and 3 poster-
odistal corner acutely extended. Coxa | anteroventral corner subacute, pos-
_ terior margin rounded. Coxa 2 subacute distally; coxae 3 and 4 anterodistal
_ margins convex distally, posterodistal margins concave distally, both plates
ventrally subacute.

Antenna 1 peduncle slightly shorter than basal flagellar article; uni-
articulate accessory flagellum extends more than halfway along basal fla-
gellar article; main flagellum of 6 articles. Antenna 2 peduncle article 5 as
long as flagellum; article 4 very short, as long as wide. Upper lip twice as
wide as high, emarginate. Mandible incisor smooth, accessory tooth sub-
triangular, present in left mandible only. Lower lip lobes broadly rounded,
with dense covering of fine setae. Maxilla 1 palp biarticulate, extending
beyond outer plate; palp second article armed with short, stout setae along
apex and distal half of medial margin; outer plate armed with elongate heavy
spines; inner plate with 11 elongate, less robust setae. Maxilla 2 inner plate
3 times as wide as outer plate; outer plate distal setae non-plumose. Max-
illiped palp 4-articulate, article 2 not produced, articles successively de-
creasing in length and width, outer plate broadly ovate, armed with short
setae; inner plate short, reaching base of palp, medial margin armed with
elongate setae. Gnathopods 1 and 2 simple; gnathopod | articles 5 and 6
subequal in length; gnathopod 2 article 6 longer than article 5. Pereopod 2
subchelate; propodus ventral margin excavate, bearing small proximal pro-

652 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Bf
VEZ
=

Hh

s

Fig. 27. Parandaniexis dewitti, female: a, Antenna 1; b, Antenna 2; c, Mandible; d, Lower
lip; e, Maxilla 1; f, Maxilla 1 palp; g, Maxilla 2; h, Maxilliped; i, Gnathopod 1; j, Gnathopod
2; k, Pereopod 2; 1, Pereopod 2 distal end; m, Pereopod 5; n, Pereopod 6; 0, Pereopod 7; p,
Telson.

tuberance with continuous row of short setae; articles 5 and 6 subequal.
Pereopods 5 and 6, article 2 slender; pereopod 6 article 6 siightly longer
than article 4. Pereopod 7 article 2 broad, tapering distally. Telson entire,
apically subacute.

Etymology.—The species is named in honor of Dr. H. H. DeWitt, Chief
Scientist for the cruise during which the specimen was collected.

VOLUME 93, NUMBER 3 653

Distribution.—South Sandwich Islands.

Remarks.—The subchelate pereopod 4 and the morphology of the mouth-
parts unquestionably places this species in the genus Parandaniexis. It dif-
fers from the only other known species, P. mirabilis Schellenberg by the
shorter basal flagellar article of antenna 1, curved and acutely tapering coxae
3 and 4, large carinae on pleonites 1-3, and small ventral lobe on pereopod
4 propodus. The outline of the body is remarkably similar to that illustrated
tor Andaniexis spinescens (Alcock 1894). The latter species, originally
placed in the genus Andania by Alcock, was moved to the genus Andaniexis
by Stebbing (1906). It was so incompletely described, however, that its
precise affinities are impossible to establish.

Literature Cited

Alcock, A. 1894. Natural history notes from H.M. Indian Marine Survey Steamer “‘Investi-
gator’, Commander R. F. Hoskyn, R. N., Late Commanding.—ser. II. no. 1. On the
results of the deep-sea dredging during the season 1890-91 (concluded).—Ann. Mag.

| nat. Hist. (6) 13:411.

- Barnard, J. L. 1969. The families and genera of marine gammaridean Amphipoda.—U:S.
Natl. Mus. Bull. 271:1—535.

Barnard, K. H. 1930. Crustacea. Part IX. Amphipoda.—British Antarctic (‘Terra Nova’)

Expedition, 1910, Zoology 8:307-454.

. 1932. Amphipoda.—Discovery Reports 5:1—326.

. 1955. Additions to the fauna-list of the South African Crustacea and Pycnogonida.—

Ann. S. Afr. Mus. 43:1—107.

Bellan-Santini, D. 1972. Invertébrés marins des XII et XV Expédition Antarctiques Frangaises
en Terre Adelie 10,—Amphipodes Gammariens.—Tethys 4:683-702.

Bushueva, I. V. 1978. A new amphipod species (Amphipoda, Gammaridea) from the Davis
Sea (eastern Antarctic).—Zool. Zh. 57:450-453.

Chevreux, E. 1911. Sur les amphipodes des Expédition Antarctique Frangaise.—C. R. Acad.

Sci. Paris 153:1166—-1168.

. 1912. Deuxiéme expédition dans L’antarctique, dirigée par le Dr. Charcot 1908-1910,

Diagnoses d’amphipodes nouveau.—Bull. Mus. Nat. Paris 4:208-218.

Dana, J. D. 1853-55. Crustacea. Part I].—United States Exploring Expedition 14:689-1618
(1853), pls. 1-96 (1855).

Hope, F. W. 1851. Catalogo dei Crostacei Italiani e di molti altri del Mediterraneo, Napoli,
48 pp.

Karaman, G. S., and J. L. Barnard. 1979. Classificatory revisions in Gammaridean Amphipoda
(Crustacea), Part I.—Proc. Biol. Soc. Washington 92(1) :106—165S.

Krapp-Schickel, G. 1976. Marine amphipods from Pantelleria and Catania (Sicily).—Bull.
Zool. Mus. Univ. Amsterdam 5(5):31-—45.

Nicholls, G. E. 1938. Amphipoda Gammaridea.—Australasian Antarctic Expedition 1911-14,
Sci. Rep. ser. C, 2(4):1-145.

Rathke, H. 1843. Bertrage zur Fauna Norwegens.— Verhandl. Kaiserl. Leopolinish - Caro-
linischen Akad. Naturforsch. Breslau 20(1): 1-264, 264b, 264c.

Schellenberg, A. 1926. Die Gammariden der Deutschen Sudpolar-Expedition 1901—1903.—

Deutsch. Sudpolar Expedition 10, Zool. 18:233-414.

. 1929. Die abyssale und pelagische Gammariden. Reports on the scientific results of

654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz on the
‘‘Albatross . . . 1899-1900 and 1904-1905. —Bull. Mus. Comp. Zool. 69:191—201, 1 pl.
. 1931. Gammariden und Caprelliden des Magellangebietes, Sudgeorgiens und der Wes-
tantarktis.—Further Zool. Res. Swedish Antarctic Exped. 1901-1903, 2(6): 1-290.
Stebbing, T. R. R. 1906. Amphipoda I, Gammaridea.—Das Tierreich, 21:1 —806.

Thomson, G. M. 1880. New species of Crustacea from New Zealand.— Ann. Mag. Nat. Hist.,

(5) 6: 1-6.

Department of Oceanography, Ira C. Darling Center, University of
Maine, Walpole, Maine 04573.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 655-661

THE CALLS AND TAXONOMIC POSITIONS OF HYLA
GIESLERI AND OLOLYGON OPALINA
(AMPHIBIA: ANURA: HYLIDAE)

W. Ronald Heyer

Abstract.—The advertisement calls of Hyla giesleri, H. microps, Ololy-
gon catharinae and O. opalina are described. The call data together with
morphological differences indicate that Hyla giesleri is a species distinct
from H. microps, with which it is currently placed, and that Ololygon opal-
ina 1s a full species, not a subspecies of Ololygon catharinae as originally
described. The calls in each species pair differ in several species specific
coding features; small behavioral and morphological adjustments can ac-
count for these differences.

Field recordings of advertisement calls of frogs from southeastern Brazil

clarify the taxonomic positions of two species of hylid frogs, Hyla giesleri

and Ololygon opalina. Duellman (1977) and Lutz (1973) included Hyla gies-
leri Mertens in the synonymy of Hyla microps Peters, following the syn-

- onymy of Bokermann (1966). Lutz (1968) described opalina as a new sub-

species of Hyla catharinae. Lutz (1973) and Duellman (1977) maintained
this relationship. I follow Fouquette and Delahoussaye (1977) and consider
catharinae a member of the genus Ololygon. The data presented in this
paper demonstrate that Hyla giesleri is specifically distinct from Hyla mi-
crops and that Ololygon opalina is a distinct species within the Ololygon
catharinae complex.

Methods and Materials

Field recordings made with an Uher CR-134 stereo cassette recorder were
analyzed using a Kay Sonagraph model 6061 B, a Hewlett Packard 7402A
strip chart recorder and a Bruel and Kjaer 2121 frequency analyzer. Spec-
imens are deposited in the Museu de Zoologia, Sao Paulo, and National
Museum of Natural History, Washington, D.C. (USNM).

Hyla giesleri

The advertising call is intensity modulated, beginning relatively quietly
and ending loudly. The call (Fig. 1A) consists of about 30 discrete pulses
(two calls analyzed have 27 and 32 pulses) at an average pulse rate of about
100 per second. The duration of the entire call is about 0.3 s, and the du-

656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

I

r

ah See ee

|

Fig. 1. Oscillographic strip chart recordings of portions of advertisement calls of (A) Hyla
giesleri and (B) Hyla microps. Scale bars = 0.01 s. Hyla giesleri call from specimen USNM
208170 recorded at an air temperature of 20.5°C, Santa Teresa, Espirito Santo, Brazil. Hyla
microps call from specimens USNM 208412-16 recorded at an air temperature of 19.5°C, Ter-
esopolis, Rio de Janeiro, Brazil.

ration of individual pulses ranges from about 0.003 s at the beginning of the
call to about 0.01 s towards the end of the call. The call has a complex
spectral pattern, but does have a harmonic structure (Fig. 2A); the low
frequency spectra presumably the fundamental (200-800 Hz range), and the
high frequency spectra (3,000—3,800 Hz range) some multiple of the pre-
sumed fundamental. The dominant frequency at a temperature of 20.5°C
ranges from a peak of about 3,000 Hz for the initial pulses in the call to a
peak of about 3,600 Hz for the later pulses in the call.

The call of H. giesleri is similar to the call of H. microps in being pulsatile
and having harmonic structure (Fig. 3). The calls differ distinctively in du-
ration and dominant frequencies.

The call of Hyla microps is intensity modulated, starting out somewhat
softly but quickly becoming and staying loud. The call consists of about 55-
60 discrete pulses (Fig. 1B) at an average pulse rate of about 50 per second.
The duration of the entire call is about 1 s, and the duration of individual
pulses ranges from about 0.005 s for the initial pulses to about 0.01 s for
pulses throughout the rest of the call. The call has a harmonic structure
with a complex spectral pattern (Fig. 2B); it appears likely that the funda-
mental is in the 600-700 Hz range, with the dominant being the sixth har-
monic. The dominant frequency at a temperature of 19.5°C ranges from
about 4,600-5,300 Hz.

The recorded specimen of Hyla giesleri is part of a series taken at Santa

VOLUME 93, NUMBER 3 657

INTENSITY

RELATIVE

Seewe \

T T ol
6

FREQUENCY IN~ kHz

Fig. 2. Spectral analyses for advertisement calls of (A) Hyla giesleri (two calls) and (B)
Hyla microps (three calls). The dots represent peak RMS meter readings at discrete frequency
settings on the Brtiel and Kjaer frequency analyzer, using a tape loop as the sound source.
Specimen data same as for Fig. 1.

Teresa, Espirito Santo, Brazil. These specimens were compared with a to-
potype of Ayla giesleri at the USNM (taken by the collector of the type-
specimens). Previously, Hyla giesleri had been reported only from the type-
locality in the State of Rio de Janeiro, Brazil. Preserved specimens of Hyla
giesleri are easily distinguished from those of Hyla microps by the presence
in H. microps of a distinct white subocular spot or area on the upper lip.
Hyla giesleri specimens lack a distinct white spot or area on the upper lip
below the eye.

Ololygon opalina, new rank

The advertisement call begins quietly and builds in intensity. The call is
pulsatile (Fig. 4A), but the pulses are not discrete. Because of the contrast
in intensity between the beginning and ending of the call, only the louder
ends of the calls appear on sonagrams (Fig. 5A). The entire call lasts about
0.7 s. Each pulse lasts about 0.03 s and is partially pulsed. The call has
harmonic structure (Fig. 6A), with the sixth harmonic emphasized as the

658 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

FREQUENCY IN kHz

|
TIME IN SECONDS

Fig. 3. Sonagrams (narrow band) of advertisement calls of (A) Hyla giesleri and (B) Hyla
microps. Specimen data same as for Fig. 1.

dominant frequency. The dominant frequency at 15.8° C varies from 3,300
to 4,100 Hz.

The call of O. opalina is distinct from the call of O. catharinae from
Santa Catarina. The calls have a general similarity in starting out quietly

Fig. 4. Oscillographic strip chart recordings of advertisement calls of (A) Ololygon opalina
and (B) Ololygon catharinae. Scale bars = 0.1 s. Oloygon opalina call from specimen USNM
208473 recorded at an air temperature of 15.8°C, Teresopolis, Rio de Janeiro, Brazil. Ololygon
catharinae call from specimen USNM field no. 7359 recorded at an approximate air temperature
of 22°C, Pirabeiraba, Santa Catarina, Brazil.

VOLUME 93, NUMBER 3 659

IN kHz
to)

>

1 8]

FREQUENCY
hb
i
‘

- = Se ee

I 2
TIME IN SECONDS

Fig. 5. Sonagrams (narrow band) of advertisement calls of (A) Ololygon opalina and (B)
Ololygon catharinae. Specimen data same as for Fig. 4.

and ending relatively loudly, in being pulsatile, and in having harmonic
structure. In several details, the calls differ markedly as can be seen when
they are compared (Figs. 4, 5, 6).

The call of O. catharinae begins very quietly and ends much more loudly

97 A
3
>
é
Ww
Zz
uJ
e
z
|
9
WwW B
>
=
<x
a
u
ce
3

2 4 6

FREQUENCY IN kHz

Fig. 6. Spectral analyses for advertisement calls of (A) Ololygon opalina (two calls) and
(B) Ololygon catharinae (two calls). The dots represent peak RMS meter readings at discrete
frequency settings on the Briiel and Kjaer frequency analyzer, using a tape loop as the sound
source. Specimen data same as for Fig. 4.

660 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Fig. 4B). In the field, the first part of the call was not heard above the
background of other chorusing frogs. Some frogs called from plastic bags
after capture; only then was the first part of the call heard. The call consists
of a number of separate pulses (Fig. 5B); each of the pulses has a relatively
uniform intensity envelope, not distinctly pulsatile as in O. opalina. The
entire call sequence lasts somewhat more than two seconds. The duration
of a given pulse within the call lasts about 0.04 s. The pulses, particularly
the latter, are frequency modulated (Fig. 5B). The call has harmonic struc-
ture (Fig. 6B), with either the fifth or sixth harmonic emphasized as the
dominant frequency. The dominant frequency at about 22°C varies from
2,200 to 3,100 HZ.

The nominate subspecies of O. catharinae was described from the State
of Santa Catarina, Brazil, where the recordings reported here were made.
Lutz (1968) described a number of subspecies of O. catharinae, one of
which (simplex = obtriangulata), she later (1973) raised to a full species.
The call data presented here demonstrate that the subspecies Lutz (1973)
defined as catharinae and opalina differ at the species level. There remain
three subspecies as defined by Lutz (1973) for which the status is still un-
clear: O. c. bocainensis, O. c. brieni, O. c. trapicheiroi.

Specimens of O. catharinae and O. opalina are easily distinguished by
color pattern. The dorsum of O. opalina is rather uniform; the groin region
of bold dark and light markings contrasts markedly with the surrounding
sides and dorsum (for a color photo, see plate V in Lutz 1973). The dorsum
of O. catharinae consists of a series of alternating dark and lighter brown
transverse bands (similar to those seen in O. c. brieni and O. humilis as
photographed in plate V of Lutz 1973); the groin area has an intensification
of the dorsal pattern, but there is no contrast between the dorsal and lateral
pattern with the groin pattern.

Discussion

The species discussed represent morphologically similar species pairs.
The morphological similarity presumably indicates close phyletic relation-
ships. For each species pair, a common call pattern can be derived, yet
there are at least three aspects of the calls for both species pairs where
differences are sufficient to code species specific information. These differ-
ences are: (1) call duration, (2) pulse rate, and (3) dominant frequency chan-
nel. In addition, there is a pulsatile difference within notes for O. catharinae
and opalina that could code species specificity. All of these call differences
can be accounted for by relatively small adjustments in the calling appara-
tus. For example, differences in call duration and pulse rate are under neu-
romuscular control; small changes in behavior of this system could lead to
the call differences seen. The frequency differences and pulsatile structure

VOLUME 93, NUMBER 3 661

within note differences could be accounted for by small morphological
changes in the vocal cords and/or vocal sac. Thus it appears that at least
one premating isolating mechanism in frogs can evolve through fine tuning
of a functional system; major changes are not required.

Acknowledgments

The following persons helped in various aspects of the field work: Ronald
I. Crombie, Maria Christina Duchéne, Annelise Gehrau, and Francisca Car-
olina do Val. George R. Drewry and Roy W. McDiarmid critically read the
manuscript. This is a contribution to the research project, ‘‘Evolutionary
Zoogeography of the Atlantic Forest System of Brazil: The Anuran Ex-
ample,’ supported jointly by the Museu de Zoologia da Universidade de
Sao Paulo and the Smithsonian Institution’s Amazon Ecosystem Research
Program.

Literature Cited

Bokermann, W. C. A. 1966. Lista anotada das localidades tipo de anfibios brasileiros.—
Servico de Documentagao, Sao Paulo. 183 p.

Duellman, W. E. 1977. Liste der rezenten Amphibien und Reptilien. Hylidae, Centrolenidae,
Pseudidae.—Das Tierreich 95:1—225.

Fouquette, M. J., Jr., and A. J. Delahoussaye. 1977. Sperm morphology in the Hyla rubra
group (Amphibia, Anura, Hylidae), and its bearing on generic status.—J. Herpetology

11:387-396.

Lutz, B. 1968. Geographic variation in Brazilian species of Hyla.— Pearce-Sellards Ser. 12:
1-14.
. 1973. Brazilian species of Hyla.—University of Texas Press, Austin & London. xix
ae YASS) [Ds

Division of Amphibians and Reptiles, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 662-673

TWO NEW LAMIPPID COPEPODS PARASITIC ON
GORGONIANS FROM HAWATI AND THE BAHAMAS

Mark J. Grygier

Abstract.—Descriptions are presented of Linaresia magna, new species,
and Sphaerippe caligicola, new genus and species, gorgonian-inhabiting
lamippid copepods parasitic within galls on a Hawaiian Placogorgia and a
Bahaman Callogorgia respectively. L. magna is the first lamippid described
from the Pacific. It differs from the two previously known species of Lin-
aresia in its greater size, in the shape of the lateral arms and lack of ven-
tromedial knobs in the female, and in details of setation. It lives in galls
formed from individual host polyps, with a density of 0.42 individuals per
centimeter of host skeleton. The sex ratio is nearly even; spermatophores
are attached in pairs to the females and eggs are found in loose bundles
within the galls. Some specimens are associated with the ascothoracican
cirriped Gorgonolaureus. The new genus Sphaerippe is distinguished by a
globular female, bands of papillae on the male, no maxillipeds, broad, weak-
ly segmented second antennae, a short furca with spines, and multidigitate
thoracic limbs with acicules. S. caligicola lives as a male-female pair within
a boot-shaped gall. Several egg masses containing about 40 eggs each are
found within the gall.

The Lamippidae are a small family of poecilostome copepods parasitic
within various Octocorallia. Those found in the vicinity of Banyuls-sur-Mer,
France, are the best known, having been studied by de Zulueta (1908, 1910,
1911), Bouligand & Delamare-Deboutteville (1959a, b), and Bouligand
(1960a, b, 1961, 1965). The most extensive recent review is that of Bouligand
(1966). Here he recognized two genera, Lamippe (subgenera Lamippe,
Lamippula, Lamippina, Lamippella, Enalcyonium, and Isidicola) and Lin-
aresia. Except for Lamippula, which was newly defined, the subgenera of
Lamippe were previously of generic rank. Stock (1973) reestablished all
these taxa as genera and presented a key for distinguishing them. A repre-
sentative of a new genus, Magnippe, has recently been described by Stock
(1978).

The two new lamippids described in this paper were discovered fortu-
itously while I was examining gorgonians for gallicolous ascothoracican cir-
ripeds. Ms. Katherine Muzik of the Museum of Comparative Zoology at
Harvard University sent me dried branches of a new Placogorgia (n. sp.
A) from off Makapuu Point, Oahu, Hawaii, where they had been collected

VOLUME 93, NUMBER 3 663

by Boh Bartko in the Star IJ submersible at 366 meters. This dried material
was reconstituted overnight in a trisodium phosphate solution. Some as-
cothoracicans (Gorgonolaureus sp.) were present in a few large galls, but
much more numerous smaller galls contained peculiar stellate animals that
could be assigned to the lamippid genus Linaresia. The two species of this
genus previously known are L. mammillifera de Zulueta (1908) and L. bou-
ligandi Stock (1979) from the Mediterranean coast of Spain and the west
coast of Florida, respectively. This is the first lamippid known from Hawaii,
and, aside from undescribed species present at La Jolla, California (Taylor,
unpublished) and at Corona del Mar, California (Illg, personal communi-
cation), it is the first from anywhere in the Pacific.

Dr. Frederick Bayer of the Smithsonian Institution sent a collection of
gorgonian-associated ascothoracican material including a stalk of Callogor-
gia (USNM Acc. no. 295887) from off Grand Bahama Island with a small,
boot-shaped growth on it. This gall contained a pair (male and female) of
lamippid copepods, rather than ascothoracicans; these specimens are de-
scribed herein as representatives of a new genus, Sphaerippe. A number of
lamippids are already known from the West Atlantic and the Caribbean
(Patton, 1963: Dudley, 1973: Stock, 1973, 1978, 1979).

All figures were drawn by the author with the aid of a camera lucida from
whole specimens mounted and cleared in lactic acid, or in some cases for
Linaresia, dissected specimens mounted in glycerol or Turtox CMC-10 with
acid fuchsin.

Linaresia de Zulueta

Linaresia magna, new species
Fig. 1

Material.—Holotype, °, at U.S. National Museum of Natural History,
USNM 173711; Paratypes: 15 22, 15 66, and a Placogorgia branch with
unopened galls, USNM (for the lot) 173712; 15 22, 15 66, at Museum of
Comparative Zoology, Harvard University; numerous 2 2 and 64 and un-
opened galls retained at Scripps Institution of Oceanography.

Type-locality.—About 10 kilometers east of Makapuu Point, Oahu, Ha-
wail (21°19’N, 157°34'’W), at 366 meters depth. Taken by Boh Bartko in Star
IT submersible in 1979.

Distribution.—Known only from type-locality.

Host.—Endoparasitic in Placogorgia sp.

Etymology.—From Latin magnum (=large), referring to the animal’s size
relative to its congeners.

Description.—Female (Fig. la—g): Length varying due to uneven shrink-
age in dried state, 1.4-2.3 mm, averaging 1.9 mm (n = 30) not including
furca or lateral processes. Body (Fig. la) yellow to amber in color, outline

664 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

| 0.1mm idef
0.02 bol jk!

k.

Fig. 1. Linaresia magna: a—g, Female; h-l, Male: a, Ventral aspect of female, reconstruc-
tion, anterior end uppermost, showing pair of spermatophores with duct leading to genital
apparatus; b, First antennae, ventral aspect, including the 3 most anterior papillate tubercles;
c, Second antenna, ventromedial aspect; d, First pair of thoracopods, ventral aspect; e, Second
pair of thoracopods, ventral aspect; f, Female genital apparatus (genital plate and vulvae),
ventral aspect, genital plate displaced somewhat to left during mounting; g, Same as f with
genital plate removed to show clavate projections between vulvae and depression which ac-
comodates convex upper surface of plate; h, Ventral aspect of male, anterior end at left,
showing spermatophores within body and terminal swelling at base of furca; i, First antenna,
median aspect; j, Second antenna, median aspect; k, First thoracic limb, lateral aspect; 1,
Second thoracic limb, lateral aspect.

VOLUME 93, NUMBER 3 665

variable, but typically roughly rectangular, twice as long as wide, tapering
slightly posteriorly to a furca. Each furcal ramus about 1 mm long with
annulate ridges, tapering to a spine. Sixteen lateral processes (arms) along
sides of body arranged into 5 sets along each side; all arms about 1 mm
long, cylindrical with annulate ridges, tapering to blunt ends. First 3 pairs
each with biramous ventral and uniramous dorsal element. Posterior tip of
biramous arm slightly narrower and longer than anterior, bifurcation occur-
ring in distal quarter of arm. Last 2 pairs with only biramous ventral ele-
ment, last pair wider than the others. Arms produced laterally, but usually
bent ventrally. Other external features including 2 pairs of antennae, oral
cone, 2 pairs of thoracic limbs, and genital plate (‘terminal swelling’’ of
Bouligand, 1966) with associated structures. Ventral surface with several
transverse folds or ridges, which, together with intervening cuticle, bear
minute slits and depressions, triangular in side view, and more common
anteriorly. Antennae and oral cone between first pair of arms; first antennae
on anteriormost end of animal. Five papillae-laden tubercles anteriorly, 1
dorsal of first antennae, 2 lateral between pairs of antennae, and 2 lateral
posterior to second antennae. First thoracic limbs equidistant between oral
cone and second thoracic limbs, latter between second pair of arms. First
antennae (Fig. 1b) 0.05 mm long, directed laterally. Five setae on anterior
border and 5 more terminal, short but not spinelike. Segmentation obscure,
possibly 2 segments. Second antennae (Fig. Ic) slightly larger and conical,
segmentation obscure, single hooked claw at tip. Oral cone a very small
projection antero-posteriorly elongate. Thoracic limbs (Fig. 1d—e) reduced
to nipplelike structures lacking setae. Genital apparatus (Fig. If) just ante-
rior to last pair of arms. Plate roughly pentagonal, 0.3 mm wide and slightly
longer, with convoluted anterior edge. Plate prehensile, with rear edge an-
chored and upper side convex. Vulvae at anterior corners of plate, bulbous
part of each partly overlain by plate; tapered lateral part of each free. A pair
of narrow, clavate projections pointing posteriorly from beneath anterior
edge of plate (Fig. 1g).

Male (Fig. 1h—1): Length 0.9-1.6 mm, averaging 1.3 mm (n = 27) not
including furca. Body (Fig. 1h) yellow to amber in color, fusiform, tapering
more anteriorly than posteriorly; widest point (0.5—0.6 mm) at midlength.
A posterior furca, each ramus stout and tapering, with a terminal and ex-
ternal subterminal spine. Furcal rami 20-25% as long as rest of body. Other
appendages including 2 pairs of antennae and 2 pairs of thoracic limbs an-
teriorly; also spherical ventral protrusion (terminal swelling) just anterior to
furca. Transverse genital slit ventrally, just posterior of widest point of
body, bowed slightly forward in middle. Papillae covering integument, but
Sparse posteriorly and on ventral surface near thoracic and cephalic ap-
pendages. Five papillae-covered tubercles at anterior end arranged as in
female. First antennae (Fig. 11) 0.06 mm long, with no apparent segmenta-
tion, conical with 1 baso-medial seta, 2 long and 2 short setae in center of

666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

medial side, 1 seta on middle of anterior side or on medial face, and 2 long
and | short terminal setae. All setae stout and tapering to a point. Second
antennae (Fig. 1j) of same dimensions as first, with indistinct basal segment,
spherical segment with sclerites, and slightly bulbous distal segment bearing
stout, hooked claw. Mouth posterior to second antennae, reduced to minute
indented oval. Mandibles, maxillae, and maxillipeds absent. First and sec-
ond thoracic limbs (Fig. 1k—l) about same size as antennae. Protopodites
with sclerites. Exopodites with 3 claws, 2 terminal and smaller one between
and proximal to these on posterior side. Endopodites absent. Interpodal
plate connecting each pair of limbs across midline.

Remarks.—The 5 papillate cephalic tubercles, the simple tapering furcal
rami, and the pronounced development and precise arrangement of the fe-
male lateral processes (arms) are diagnostic of the genus Linaresia de Zu-
lueta (1908). Two other species are known, L. mammillifera de Zulueta
(1908), the type-species of the genus, and L. bouligandi Stock (1979).

The female of L. mammillifera is more rotund than that of L. magna. Its
length is only 1.1 mm, 40% shorter that the new species. The arms of L.
mammillifera are capitate, unlike L. magna, where they taper gradually.
On bifurcate arms, the posterior branch is clearly subordinate in L. mam-
millifera, being acapitate and set off at a right angle. In L. magna there is
a slight asymmetry, but no obvious subordination between the bifurcate
tips. The first antennae of the female L. mammnillifera lack a subterminal
seta found in L. magna, and the thoracic limbs are slightly less degenerate
in the former species. Three enigmatic structures are arranged serially on
the ventral side of the L. mammnillifera female posterior to the thoracopods.
The anterior 2 are round knobs situated medially, while the posterior one
may also be on the midline, or be manifested in other individuals as a paired
structure. Stock (1978) considers these to be rudimentary limbs. These
structures are absent in L. magna. The furcal rami of the male L. mam-
millifera are proportionately longer than those of L. magna, and its main
body is only half as long as the present species. There are 2 extra setae on
the medial side of the first antennae in L. magna males. No mention is
made of a terminal swelling in the males of L. mammnillifera.

Only the female is known of L. bouligandi. Its main body is 0.5—0.6 mm
long, much smaller than L. magna. The body outline is also more rotund.
The arms are proportionately longer and narrower than in L. magna, and
have wider annulations. The bifurcations of the ventral arms are very long
compared to L. magna. The first antennae are more reduced in L. bouli-
gandi, with only 0-3 setae, compared to 10 in L. magna. The claw on the
second antenna is bifid in L. bouligandi, but simple in L. magna. Two
medial swellings on the ventral surface, similar to those in L. mammnillifera,
are found in L. bouligandi, but not in L. magna. According to Stock (1979),
L. bouligandi has 2 arcuate sclerites connected by a straight, transverse

VOLUME 93, NUMBER 3 667

sclerite some distance anterior of the genital plate (‘terminal swelling’’).
He tentatively labelled this structure the vulva, citing Bouligand & Dela-
mare-Deboutteville (1959a). The structure on L. mammnillifera referred to
by Stock was in later papers by Bouligand redescribed as the third (single
or paired) ventral knob, discussed above. Bouligand (1960a) showed that
the oviducts open at the true vulvae, the structures at the anterior corners
of the genital plate. Stock’s ‘‘vulva’’ is probably homologous to the third
ventral knob in L. mammillifera, and is probably not a reproductive struc-
ture. The structure is lacking in L. magna. The genital plate is situated at
the extreme posterior end of the body in L. bouligandi, rather than farther
forward as in the other 2 species. In all 3 species the position of the genital
plate is fairly constant relative to the last pair of arms. There is apparently
no armanent of the furcal rami in L. bouligandi.

Life cycle.—Many males have a pair of spermatophores developing within
chambers that are one-third the length of the animal (0.8 mm). The sper-
matophores must exit through the ventral transverse slit. They are attached
in pairs by a cord to the female. Attachment may be at various places on
the underside, but usually just in front of the genital plate. This cord is over
1 mm long and is twisted; it contains the ducts leading from each sper-
matophore. These may join distally, or they may remain separate for most
of the length of the cord. The duct narrows greatly at the point of attach-
ment. No sperm were seen. Spermatophores attached to the female (Fig.
la) are 0.6 mm long and reniform.

Most females are associated with eggs, either loosely attached within the
gall or confined in thin membranous sacs on the underside of the female.
Egg sacs are not reported from either of the other species of Linaresia. The
eggs are about 0.14 mm in diameter. One was seen with a developing nau-
plius inside. It was impossible to see much detail, but the larva did have 3
pairs of appendages with 3—S terminal setae per ramus. Stages intermediate
between the male and female are known for L. mammnillifera (Bouligand,
1960a), but not in L. magna or L. bouligandi. It is possible, though, that
all 3 species are protandric hermaphrodites.

Ecological notes.—Each gall corresponds to a single polyp. L. magna
apparently does not live within the polyp, for a membrane separates the
gastro-vascular cavity of the polyp from the space within which the copepod
is living. The dorsal side of the female is appressed to the gorgonian’s axis,
with the body lined up longitudinally, producing a lenticular swelling of the
gorgonian’s bark. Unlike L. mammillifera, there are no cases of the arms
of the parasite being exposed and possibly simulating a missing polyp.

Each gall contains a female and, usually, a male (rarely 2 or 3) and eggs.
Solitary males were found, but not within galls. Often there were one or
more females associated with galls formed by an ascothoracican cirriped
(Gorgonolaureus sp.). Here there were no enlarged polyps; the parasitic

668 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

copepod lived between the valves of the ascothoracican and the gorgonian
tissue covering it. In order to estimate the degree of infestation, branches
of the gorgonian totalling 139 mm in skeletal length were thoroughly
searched for copepods; 59 individuals were found, corresponding to a den-
sity of 0.42 copepods per centimeter of host. Thirty-one of these were fe-
males and 28 males, suggesting a 1:1 sex ratio.

The host of L. mammillifera is Paramuricea clavata of the family Para-
muriceidae (potentially confusing name change of this gorgonian explained
in Stock, 1979). Both L. bouligandi and L. magna are found in Placogorgia
species, also in the Paramuriceidae. It is too soon to tell, however, whether
Linaresia is confined to this one family of gorgonians. Both L. mammnillifera
(subtidal) and L. bouligandi (76 m) are from shallow waters, while L.
magna (366 m) is from bathyl depths. The 3 species of Linaresia currently
encompass the Mediterranean, Western Atlantic, and Indo-West Pacific bio-
geographical provinces. This may be construed as a Tethyan distribution,
in which case it is reasonable to expect that more species will be discovered
at similar latitudes in the future.

Sphaerippe, new genus

Diagnosis.—Lamippid copepods endoparasitic within a gall on a gorgo-
nian (Callogorgia sp.). Sexually dimorphic, female much larger than male
and with slightly different armament on appendages. Female globose, with
slight anterior appendage-bearing protrusion; male less rotund, with more
prominent appendage-bearing protrusion and 5 bands of papillae around
trunk. Both sexes with biarticulate first antennae (single spine on basal
segment, many on terminal segment); broad, weakly biarticulate (possibly
triarticulate in female) second antennae terminating in hooked claw; 2 pairs
of thoracopods with uniarticulate protopodites, exopodites, and endopo-
dites; and separated furcal rami, conical, with a subterminal and 4 terminal
spines. Maxillipeds absent. Thoracic endopodites of both sexes well-devel-
oped with palmately arranged digitations, each terminating in an acicule.
Exopodites similar to endopodites, but with more digitations. Sometimes
inexact symmetry of numbers of digitations on left and right sides of animal,
and males possessing more digitations than females.

Remarks.—Sphaerippe differs from Linaresia and Magnippe Stock
(1978) in not having the lateral arms that give the females of these two
genera stellate appearances. Lamippe Bruzelius (1859) and Isidicola Gra-
vier (1914) have well-developed maxillipeds, which are lacking in Sphaer-
ippe. Lamippina Bouligand (1960) has well-developed thoracic endopodites
like Sphaerippe, but also has rudimentary maxillipeds, furcal acicules, and
a Clearly articulated second antenna with 3 segments in both sexes, condi-
tions found wanting in Sphaerippe. Lamippula Bouligand (1966) lacks max-
illipeds but has furcal acicules and long, clearly segmented first and second

VOLUME 93, NUMBER 3 669

antennae, contrary to the condition found in Sphaerippe. Enalcyonium Ols-
son (1869) is similar to Lamippula, though lacking furcal acicules, but oth-
erwise distinguishable from Sphaerippe by the same criteria as Lamippula.
Lamippella Bouligand & Delamare-Deboutteville (1959) is the closest of the
known genera to Sphaerippe in lacking maxillipeds and furcal acicules and
in having circular rows of papillae and poorly segmented antennae, but it
has reduced, unarmed endopodites on its thoracic limbs, contrasting with
the well-armed endopodites of Sphaerippe. Sphaerippe is much more ro-
tund than any other genus, and, aside from /sidicola, is the only relatively
unmodified lamippid to live in a gall (Gravier, 1914). Pronounced sexual
dimorphism has previously been known only in Linaresia and Lamippina,
through there is a size difference between the sexes in Lamippe (Bouligand,
1966).

Etymology.—From Latin sphaera (=sphere) and the ending of Lamippe,
the type-genus of the family Lamippidae, referring to the globular shape of
the female. Gender feminine.

Type-species.—Sphaerippe caligicola, new species.

Sphaerippe caligicola, new species
Fig. 2

Material.—Holotype, ¢, USNM 173709; Paratype, | 2 with eggs, USNM
173710.

Type-locality.—Off Southwest Point, Grand Bahama Island (26°31'N,
98°51’'W). Taken at 366 meters on 14 February 1965; Gerda Station 503.

Distribution.—Known only from type-locality.

Host.—Endoparasitic within a gall on Callogorgia sp.

Etymology.—From Latin caliga (=boot) and Latin -icola (=an inhabit-
ant), referring to the boot-shaped gall within which the copepod lives.

Description.—Male (Fig. 2a—d): Body (Fig. 2a) ellipsoidal, 0.62 mm long
and 0.32 mm wide, with anterior end protruding. Color white. Cuticle
smooth except for 5 bands of hemispherical papillae encircling trunk. Two
pairs of antennae, oral cone, and 2 pairs of thoracic limbs near anterior end,
ventral transverse genital slit just anterior to fourth band of papillae, and
pair of short furcal rami also present. First antennae (Fig. 2b) 0.05 mm long,
biarticulate. Basal segment with | short, median seta, terminal segment with
8 short setae. Second antennae (Fig. 2b) 0.06 mm long, with bulbous base
and short, conical second segment terminating in strong, hooked claw. Oral
cone round, covered by labrum with straight posterior edge. From posterior
opening, oral cavity leading forward to center of cone, then straight into
body; this cavity distally a transverse slit, but proximally, bifurcation at
each end of slit may reveal pair of mouthparts not protruding past posterior
edge of labrum. Maxillipeds absent. First thoracic legs (Fig. 2c) biramous,

670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Sphaerippe caligicola: a—d, Male; e-i, Female: a, Ventral aspect of male (Holotype)
showing developing spermatophores and bands of papillae (more papillae present than shown
here); b, First (above) and second (below) antennae, antero-lateral aspect; c, Right first (above)
and second (below) thoracopods, ventral aspect; d, Furca, ventro-posterior aspect; e, Lateral
aspect of female (Paratype), showing vulvae below and anterior to furca; f, Furca, ventro-
lateral aspect; g, Right (above) and left (below) second thoracopods, ventro-lateral aspect, with
exopodites oriented vertically, showing extended acicules; h, Right first thoracopod, medial
aspect, showing acicules; i, First (left) and second (right) antennae on anterior end of body;
j, Gall in place of Callogorgia, ovals representing stylized polyps.

VOLUME 93, NUMBER 3 671

with uniarticulate protopodites, exopodites, and endopodites. Interpodal
plate connecting this pair across midline. Endopodite and exopodite ter-
minating in digitations (integral parts of segment, not spines or setae) that
are slightly bulbous basally, presumably with a retracted acicule at tip of
each. Right endopodite of first pair of legs with 4 digitations, right exopodite
with 11, left endopodite with 4, and left exopodite with 12. On second pair
(Fig. 2c) components similar to those in first, except interpodal plate more
pronounced. Right endopodite with 5 digitations, right exopodite with 10,
left endopodite with 5, and left exopodite with 9. Digitations arranged pal-
mately on all limbs, 2 rows on exopodites. Genital slit bowed slightly for-
ward in middle. Furcal rami (Fig. 2d) separated on buttockslike lobes, each
ramus a flabby cone with 1 large and 2 small terminal spines and 1 subter-
minal lateral spine as large as largest terminal one.

Internal structure including 4 longitudinal muscle bands, many circular
muscles, and pair of developing spermatophores anterior to genital slit.
Muscles present in antennae and limbs. Each row of papillae on slight an-
nulus probably representing a segment, but perhaps only reflecting a state
of arrested peristalsis.

Female (Fig. 2e—-i): Body (Fig. 2e) globose, 1.24 mm long with anterior
end protruding slightly. Surface smooth. Other features arranged as in male
except for genital apparatus. First antennae (Fig. 21) 0.05 mm long, cylin-
drical with median seta on first segment and many very short setae on
second. Second antennae (Fig. 2i) twice as long as first, apparently biartic-
ulate (suture around cylindrical basal segment perhaps indicating that 3 seg-
ments are present), with hooked claw at tip. Oral cone and mouthparts not
observed due to difficulty of obtaining ventral view. Thoracic limbs similar
to males except for number of digitations and presence of an extended
acicule on many of them. Distribution of digitations as follows: on first pair
(Fig. 2h), exopodites with 7 and endopodites with 2; on second pair (Fig.
2g), exopodites with 6 and endopodites with 5. Furca (Fig. 2f) as in male.
Vulvae anterior to furca, but still on posterior side of sphere, hemispherical
knobs on each side of midline with 2 arcuate sclerites surrounding opening
on each.

Remarks.—Sphaerippe caligicola is distinguished from other lamippids
because of its rotundity (other lamippids are sausage-shaped to vermiform),
and the acicule-bearing digitations of the thoracic limbs are quite unusual.
Lamippina aciculifera, as its name implies, bears many acicules, including
some on digitations of its thoracic limbs (Bouligand, 1961), but there are
also normal setae and claws on its limbs, as well as additional acicules on
the furca. Many other lamippids bear acicules, either on the furca (for ex-
amples, see de Zulueta, 1908) or over the body surface (Enalcyonium chat-
toni), but on these, the endopodite is reduced and naked, or, at most, armed
with a few normal setae. The combination of acicule-laden thoracic limbs

672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and furcal rami armed only with spines is unique to this new species of
lamippid.

The asymmetry in the number of digitations, at least in the male, may
limit the value of this character in distinguishing §. caligicola from any
undiscovered species in the genus Sphaerippe.

Effect on host.—The axis of the Callogorgia host has polyps arranged in
rings of 4, the rings spaced 2 mm apart. The gall containing the pair of
copepods took the place of one of these rings. The gall (Fig. 2j) was boot-
shaped, with the “‘sole’’ distal and parallel to the axis. It extended 1.3 mm
away from the axis, was 3 mm long, and was covered with spicules like the
rest of the gorgonian. No external opening was present. The inside of the
gall was lined with a membrane with a 6-fold symmetry in cross-section at
the ‘‘toe.’’ It is not known whether this membrane was secreted by the host
or the parasite.

Life cycle.—Five egg clusters totalling 198 eggs were found within the
gall distal to the axis along the ‘‘sole’’ and within the “‘toe’’ of the boot.
Dimensions of the eggs averaged 110 x 80 wm. The female had a large
number of oocytes within its ovaries, and a pair of spermatophores was
being produced by the male. No empty spermatophores were found; nor
were any larvae, developing or hatched. It may be that this species of lam-
ippid mates for life, since the male and female are enclosed in a common
gall. Alternatively, the male’s small size and more formidable armament
may allow it to break out of the gall and reinvade the same or another
gorgonian as a female. Such a system of protandric hermaphroditism has so
far been demonstrated only in Linaresia mammillifera (Bouligand & Dela-
mare-Deboutteville, 1959a; Bouligand, 1960a). Presumably the gall must
rupture at some time to release newly hatched larvae, though a small hole
bored by one would also allow the others egress.

Acknowledgments

I thank Ms. Katherine Muzik of the Museum of Comparative Zoology
and Dr. Federick Bayer of the U.S. National Museum for supplying the
copepod-infested gorgonian material. Thanks also go to Dr. William A.
Newman of Scripps Institution of Oceanography for allowing me the use of
his laboratory, for reading an earlier draft of this paper, and for many helpful
comments and suggestions. This work was done during the tenure of a
National Science Foundation Graduate Fellowship. This paper is a Contri-
bution of Scripps Institution of Oceanography, new series.

Literature Cited

Bouligand, Yves. 1960a. Sur l’organisation des lamippides, copépodes parasites des octocor-
alliares (premiére note).—Vie Milieu 11:335-380.

VOLUME 93, NUMBER 3 673

—. 1960b. Notes sur la famille des Lamippidae, premiére partie.—Crustaceana 1:258-

278.

. 1961. Notes sur la famille des Lamippidae, 2e partie. —Crustaceana 2:40-52.

—. 1965. Notes sur la famille des Lamippidae, 3e partie.—Crustaceana 8: 1-24.

. 1966. Recherches récentes sur les copépodes associés aux anthozoaires. Jn Rees, W.

J. (ed.), The Cnidaria and Their Evolution.—Zool. Soc. London Symp. 16 (Academic

Press, London): 267-306.

Bouligand, Yves, and Delamare-Deboutteville, Claude. 1959a. Le dimorphisme sexuel de

Linaresia mammillifera Zulueta, 1980, copépode parasite de |’octocoralliare Muricea

chamaeleon van Koch.—C. R. Acad. Sci. Paris 248:286—288.

, and . 1959b. Lamippzlla faurei n. g. n. sp. Considérations morphologiques sur

la famille des lamippides, copépodes parasites des octocoralliares.—C. R. Acad. Sci.

Paris 249:1807—1809.

Dudley, Patricia L. 1973. Enalcyonium carrikeri, a new species of lamippid copepod from
Alcyonium carneum Agassiz in New England.—Crustaceana 25:75-87.

Gravier, C. 1914. Sur un type nouveau de crustacé parasite d’alcyonaires de |’ Antarctique
sud-americaine.—C. R. Acad. Sci. Paris 158:354—356.

Patton, W. K. 1963. Animal associates of the gorgonian coral Leptogorgia virgulata at Beau-
fort, N. C_—Amer. Zool. 3:522 (Abstr. 203).

Stock, Jan H. 1973. Copepoda of the family Lamippidae from the western Atlantic and the
Caribbean.—Stud. Fauna Curacao Other Caribb. Isl. 43(140): 22-41.

—. 1978. Magnippe caputmedusae n. gen., n. sp. (Copepoda: Lamippidae), a highly

transformed endoparasite in octocorals of the genus Thesea from the Gulf of Mexico.—

Mem. Hourglass Cruises 3(5): 1-11.

. 1979. A new species of Linaresia (Copepoda, Lamippidae) endoparasitic in the oc-

tocoral Placogorgia from the Gulf of Mexico.— Mem. Hourglass Cruises 5(1):1-7.

de Zulueta, Antonio. 1908. Note préliminaire sur la famille des Lamippidae, copépodes par-

asites des alcyonaires.—Arch. Zool. Exp. Gén. 9:1-30.

1910. Deuzieme note sur la famille des lamippides.—Arch. Zool. Exp. Gen. 6:137-
148.
. 1911. Los copepodos des los celentereos.—Mem. Soc. Esp. Hist. Nat. 7:5-58.

Scripps Institution of Oceanography, A-008, University of California, San
Diego, La Jolla 92093.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 674-696

NEW AND LITTLE-KNOWN NEOTROPICAL
KINNARIDAE (HOMOPTERA: FULGOROIDEA)

R. G. Fennah

Abstract.—The genera Southia and Oeclidius are redefined, and Bytrois
is synonymised with the former and Paroeclidius is reduced to a subgenus
of the latter. Two new species of Southia and eight new species of Oeclidius
are described, two of the latter being wholly cavernicolous.

The kinnarine Kinnaridae of the New World have hitherto been divided
between four genera, Oeclidius Van Duzee, Paroeclidius Myers, Southia
Kirkaldy, and Bytrois Fennah, the last three of which are known from only
One or two species. A re-evaluation of the suitability of certain characters
for generic definition has suggested that all the material studied can more
appropriately be accommodated in two genera that are separable as follows.

Key to the New World genera of Kinnarinae

1. Adults flightless; vertex at anterior margin more than twice as broad
as long in middle; eyes absent; tegmina not surpassing abdomen;
aedeagus bulbous, either entirely or basally ...... Oeclidius Van D.
— Adults not flightless; vertex at anterior margin not more than twice
as broad as long in middle; eyes present; tegmina much surpassing
abdomen; aedeagus bulbous basally, or otherwise ............... 2
2. Vertex with anterior margin usually distad of level of middle of eyes;
lateral margins of frons sinuately incurved at union with lateral ca-
rinae of post-clypeus; rostrum with apical segment longer than sub-
apical; tegmina with transverse veinlet between M and Cul at apex
of basal cell; aedeagus bulbous at base, and usually with slender
VerticalscylindricalerOds ccs su oe cows oe mre anee Oeclidius Van D.
— Vertex with anterior margin not or scarcely surpassing level of mid-
dle of eyes; lateral margins of frons uniting with laieral carinae of
post-clypeus in an even uninterrupted curve; rostrum with apical
segment shorter than subapical; tegmina without transverse veinlet
between M and Cul at apex of basal cell; aedeagus very broadly
tubular, or with ventral half shallowly basin-like .. Southia Kirkaldy.

In the descriptions that follow, the length of the post-tibia is the dorsal
length from the base to the base of the metatarsus, and the width is the
width at the middle as seen in dorsal view.

VOLUME 93, NUMBER 3 675

Genus Southia Kirkaldy

Kirkaldy, 1904:279. Type-species, Delphax opposita F., 1803:84.
Bytrois Fennah, 1945:450. Type-species, Bytrois nemoralis Fennah. New
synonym.

Key to species of Southia

1. Tegmina not more than 3.5 mm long; genae distinctly tumid below
Sven conounducolomotoodya dank iUSCOUS) 4se5. 464560 pecs 0 ae 2

— Tegmina distinctly more than 3.5 mm long; genae at most only feebly

tumid below eye; ground color of body yellowish, or light orange-
DOWLING fc ties cle drac oka Beeu aia le Cay BSS aoiow Wala wore 4

2. Lateral margins of frons not elevated; lateral and median ocelli
small; pronotum creamy-white except on lateral lobes .. pisenor, sp. n.

— Lateral margins of frons elevated; lateral and median ocelli relatively
large; pronotum mostly reddish-brown or olive-brown

3. Tegmina with a fuscous spot in first apical (stigmatic) cell; pronotum

reddish-brown, only slightly paler along carinae and margins......
ee artes a oirycs. beteay) prngesnnt eh aide tate, oe siretely scene airy ach opposita (F.)

— Tegmina without a fuscous spot in first apical cell; pronotum olive-

brown, greenish-white along carinae and margins ................
ek ite trerheen. sprivet Let oe past, so ukera aati. nemoralis (Fennah)

4. Vertex broader at anterior margin than long in middle line; lateral

pronotal carinae and lateral mesonotal carinae obscure or absent;

pregenital sternum of female laterobasally granulate and with a pair

of tumescences; third valvulae of ovipositor each with two distinct
LODSSHCISCAI ewan oes oases scutes eee cle tak oe ees capnorhina, sp. n.

— Vertex not broader at anterior margin than long in middle line; lateral

pronotal carinae and lateral mesonotal carinae distinct; pregenital

sternum of female smooth, devoid of tumescences; third valvulae of
Ovipositor each with a single lobe distally ........ iridescens Fennah

Southia opposita (Fabricius)
Figs. 1-9

Delphax opposita Fabricius, 1803:84.

The type, and only known specimen, of this species is a female, not a
male as given by Stal (1869:95), in the Naturhistoriska Riksmuseet, Stock-
holm. The shrivelled genitalia (which superficially resemble the genital cap-
sule of a male) are like those of nemoralis, as far as can be ascertained
without dissection. The median carina of the mesonotum is fine, clearly-
defined and percurrent to the apex of the mesoscutellum; the lateral carinae

676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

© as

Figs. 1-9. Southia opposita (F.): 1, head and thorax, dorsal view; 2, vertex, dorsal view;
3, head and pronotum, left side; 4, face and lateral lobes of pronotum; 5, right antenna; 6,
tegmen; 7, basal cell of tegmen, showing diffuse thickening of membrane between Sc+R+M
and Cul; 8, pregenital sternum (of dried specimen); 9, third valvulae of ovipositor, posterior
view (of dried specimen).

are obsolete. The pronotum is reddish-brown, and only slightly paler along
the carinae and margins.

Southia nemoralis (Fennah), new combination
Figs. 10-14

Bytrois nemoralis Fennah 1945:450.

In this species, the type of which is in the U.S. National Museum of
Natural History, the median carina of the mesonotum is fine and moderately
distinct as far as the base of the mesoscutellum, where it disappears. The
pronotum in the typical (Trinidadian) population is olivaceous, and greenish-
white along the carinae and margins. A female from British Guiana (Ma-
baruma, N. W. District, 11.1931, J. G. Myers) in the British Museum (Nat-
ural History) which appears to belong to this species has a pronotum that is
reddish-brown and has margins and carinae that are stramineous.

Southia pisenor, new species
Figs. 15=23

Vertex shorter in middle line than broad at posterior margin (about 1:1.3),
almost horizontal, lateral margins straight, strongly converging distad, api-
cal margin obscure, shorter than width at base (about 1:1.5), disc rather
deeply hollowed; frons in profile straight in distal two-thirds, strongly
curved in basal third, smoothly rounding into vertex, in anterior view longer
than broad (about 2.5:1), wider at apex than at base (about 3.5:1), basal
margin transverse, lateral margins feebly sinuate, diverging distad, moder-

VOLUME 93, NUMBER 3 677

11

Figs. 10-14. Southia nemoralis (Fennah): 10, head and thorax, dorsal view, 11, head and
pronotum, left side; 12, tegmen; 13, male genitalia, posterior view; 14, same, right side, with
part of aedeagus visible in transparency.

ately elevated in basal half, weakly so in distal half, apical margin slightly
concave, median ocellus present, post-clypeus tricarinate, anteclypeus me-
dially carinate, rostrum slightly surpassing post-trochanters; eyes and lateral
ocelli present, blemma above each lateral ocellus, antennae with basal seg-
ment not as long as broad, subcylindrical, widening distad, second segment
longer than first (3:1), and longer than broad (2:1). Pronotum medially car-

Figs. 15-23. Southia pisenor, n. sp.: 15, face and lateral lobes of pronotum; 16, head and
thorax, dorsal view; 17, head and pronotum, left side; 18, tegmen; 19, male genitalia, posterior
view; 20, same, posterolateral view from right; 21, same, right side, with aedeagus partly seen
in transparency; 22, aedeagus, right side; 23, apical portion of aedeagus, posteroventrolateral
view from right.

678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inate, fine carina along anterior margin from middle to near level of lower
margin of eye, then bent obliquely towards tegula; mesonotum broader than
long (1.3:1), medially carinate, lateral carinae obsolete; legs moderately
slender, post-tibiae longer than wide (19.6:1), with seven teeth apically,
distinctly separated into two groups, basal metatarsal segment and second
segment each with six teeth, tarsal claws small, each relatively stout basally.
Tegmina normal, costal margin very feebly produced at base in a shallow
convex lobe, wing-tucking apparatus between Cul and Cu2 in basal cell
developed as stout triangular lobe; Sc+R+M stalk as long as basal cell, five
subapical cells present and nine apical cells including stigmatic cell.

Dark fuscous; lateral margins of clypeus, frons, vertex and mesoscutel-
lum, light reddish-brown; rostrum, all tibiae and post-tarsi, light brownish
yellow; pronotum creamy-yellow, except on middle portion of lateral lobes.
Tegmina hyaline, faintly yellowish, an oblique band from basal cell across
clavus to posterior margin, dilute fuscous, veins reddish brown. Wings hya-
line, veins fuscous.

Anal segment of male moderately long, shortly tubular in basal half, la-
teroapical angles each strongly produced laterocaudad and slightly deflexed,
apical margin broadly transverse. Pygofer very short, dorso-lateral angles —
only slightly produced, lateral margins in side view concave, ventral margin ©
rather short, slightly concave. Aedeagus scoop-shaped, with dorsal surface
much shorter than ventral surface, the latter submembranous. Styles mod-
erately long, stout, narrow basally, strongly widening beyond middle with |
inner margin in posterior view concave, with small lobe at middle, in profile
with stout vertical process, bluntly rounded at its apex, on dorsal margin at
middle.

Male.—Length, 2.7 mm; tegmen, 3.5 mm. |

Holotype 6—Panama: Patino, 19.viii.52, F. S. Blanton, in U.S. National |
Museum of Natural History. |

This species stands apart from others in the much less elevated lateral |
frontal carinae and more acutely pointed mesoscutellum, and is also well |
distinguished by the form of the male anal segment, aedeagus and styles. In |
the single specimen available, all ocelli are relatively smaller than in other |
species, and are deep red in color. The specific name is a classical proper |
name, and is a noun in apposition. |

Southia iridescens Fennah
Figs. 24-29, 36
Fennah 1945:124.

Vertex shorter in middle line than broad at posterior margin (about 1:1.2),
lateral margins straight, converging distad, apical margin transverse, shorter |
than width at base (about 1:1.4), disc deeply hollowed; frons in profile con- |

' VOLUME 93, NUMBER 3 679

aD

{8 &

Figs. 24-29. Southia iridescens Fennah: 24, head and thorax, dorsal view; 25, head and
pronotum, left side; 26, tegmen; 27, male genitalia, right side; 28, same, posterolateral view
from right; 29, same, posterior view.

vex, more strongly so in basal half, smoothly rounding into vertex, in an-
terior view longer than broad (measured between outer edges of lateral
margins) (about 3.7:1), wider at apex than at base (about 2.0:1), basal margin
slightly concave, lateral margins sinuately convex, diverging to distal fifth,
foliate, apical margin transverse, median ocellus present, post-clypeus deep-
ly inserted into frons, tricarinate, anteclypeus medially carinate, rostrum
reaching to middle of abdomen; eyes and lateral ocelli present, a single
obscure blemma above each lateral! ocellus, antennae with basal segment
slightly longer than broad (about 1.2:1), cylindrical, widening distad, second
segment longer than first (2.6:1) and longer than broad (2.2:1). Pronotum
medially carinate, fine carina along anterior margin from middle to near
level of lower margin of eye, then bent obliquely towards tegula; mesonotum
broader than long (1.6:1), finely and rather obscurely tricarinate, with ca-
rinae parallel; legs slender, post-tibiae longer than wide (24.0:1), with seven
teeth apically, separated into two groups, basal metatarsal segment and
second segment each with six teeth, tarsal claws small. Tegmina normal,
costal margin slightly produced at base in shallow convex lobe, wing-tuck-
ing apparatus between Cul and Cu2 in basal cell developed as stout shallow
flange; Sc+R+M stalk scarcely as long as basal cell, five subapical cells
present and nine apical cells.

Light orange-brown; frons, genae, protarsi and mesotarsi, fuscous, basal
segment of antennae, carinae of pronotum and femora, pale orange-yellow.

680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Tegmina hyaline, faintly yellowish, diffuse spot in basal half of stigmatic
cell, and small suffusion just beyond claval apex, fuscous. Wings hyaline,
veins fuscous.

Anal segment of male rather short, stirrup-shaped, ring-like in basal half,
lateroapical angles and apical margin moderately produced caudad, apical
margin transverse, thickened on each side of middle. Pygofer short, dor-
solateral angles not at all produced, lateral margins produced in rounded
lobe at middle, ventral margin moderately broad, slightly concave. Aedea-
gus relatively large, scoop-like, with lower margin strongly produced cau-
dad, apical margin thin. Styles moderately long, stout, broad basally, nar-
rowed medially and expanding distally, apically slightly inflected mesad.
Pregenital sternum of female smooth, with surface evenly convex.

Male.—Length, 3.2 mm; tegmen, 4.5 mm.

Female.—Length, 3.1 mm; tegmen, 5.5 mm.

1 6 2 9, Trinidad: undergrowth of hill forest, 18.xi1.1928; 6.1.1929; Blue
Basin, 4.xi.1929, J. G. Myers, in British Museum (Natural History).

This species is distinguished by its relatively large size, the profile of the
head, the structure of the pygofer, aedeagus and styles and that of the
pregenital sternum.

Southia capnorhina, new species
Figs. 30-35

Vertex shorter in middle line than broad at posterior margin (about 1:1.4),
almost horizontal, lateral margins straight, markedly converging distad, api-
cal margin transverse, shorter than width at base (about 1:1.4), disc deeply
hollowed; frons in profile convex, more strongly so in basal half, curving
evenly into vertex, in anterior view longer than broad (about 4:1), wider at
apex than at base (about 2.7:1), basal margin transverse, lateral margins
Straight, diverging distad, foliate, apical margin transverse, median ocellus
present, a pair of blemmata above each lateral ocellus, post-clypeus tricar-
inate, anteclypeus medially foliately carinate, rostrum reaching to middle of
abdomen; eyes and lateral ocelli present, antennae with basal segment
slightly longer than broad (1.1:1), subcylindrical, widening and slightly
curved distad, second segment longer than first (2.5:1) and longer than broad
(2.4:1). Pronotum medially carinate, fine carina along anterior margin from
middle to near level of lower margin of eye, but becoming obsolete at this
point; mesonotum a little broader than long, medially carinate except on
mesoscutellum, lateral carinae obsolete; legs slender, post-tibiae longer than
wide (21.3:1) with 7-8 teeth apically in two groups, basal metatarsal segment
and second segment each with 6 teeth, tarsal claws small, each relatively
stout basally. Tegmina normal, costal margin slightly produced ventrad in
basal third in shallowly convex lobe, wing-tucking apparatus between Cul

VOLUME 93, NUMBER 3 681

ay
\

35

US

Figs. 30-36. Southia capnorhina n. sp.: 30, head and thorax, dorsal view; 31, same, left
side; 32, tegmen; 33, anal segment of female, dorsal view; 34, pregenital sternum postero-
ventral view, with posterior margin uppermost, and paired tumescences near anterior margin
below; 35, third valvulae of ovipositor, posterior view. Southia iridescens Fennah. 36, third
valvulae of ovipositor, posterior view, for comparison for fig. 35.

ae

33

and Cu2 in basal cell developed as stout narrowly triangular lobe; Sc+R+M
Stalk scarcely as long as basal cell, M slightly thickened or granulate near
base, five subapical cells present and ten apical cells.

Stramineous; lateral margins of frons, clypeus except laterally, post-tibiae
apically and metatarsal segments apically, dark fuscous; middle part of disc
of pronotum, and mesonotum, light orange-brown with faint greenish-golden
sheen, metanotum, abdominal tergites in posterior half, pale yellowish
brown. Tegmina milky-hyaline, faintly yellowish, suffusion narrowly over-
lying R-M transverse vein, Sc and basal quarter of stigmatic cell, fuscous,
veins pale brownish yellow, transverse veinlets brown. Wings hyaline, veins
yellowish-brown.

Anal segment of female with lateroapical angles each produced and api-
cally truncate, in posterior view with slight papilla near each inner angle.
Pregenital sternum broader than long, strongly curving dorsad at middle,
with pair of weak tumescences sublaterally towards anterior margin, integ-
ument laterad of these finely granulate in oblique rows.

Female.—Length, 3.1 mm; tegmen, 5.0 mm.

Holotype ° .—British Guiana: New River, 750 ft., 21—23.11.1938, C. A.
Hudson, in British Museum (Natural History).

Other material.—1 ¢°, Brazil: Mato Grosso, 12°49’S—51°45’W, 9.x1.1968,
W. J. Knight.

This species, which is nearest to S. iridescens, differs from it in the less

682 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

prominent margin around the antennal foramen, in the proportions of the
vertex, the carination of the pronotum and mesonotum, the shape and sculp-
turing of the pregenital sternum and the shape of the third valvulae. It also
differs in tegminal marking, there being no infuscate spot in the membrane
just distad of the claval apex. The most distinctive superficial feature is the
infuscation of the lateral margins of the frons in their distal half, and of the
carinae and intercarinal areas of the clypeus. The specific name, the Greek
roots of which refer to this feature, is regarded as a noun in apposition.

Genus Oeclidius Van Duzee

Van Duzee, 1914:40. Type-species, Oeclidius nanus Van Duzee, 1914:40.
Paroeclidius Myers, 1928:19. Type-species, Paroeclidius luizi Myers,
1928:20. Syn. n.

Key to species of Oeclidius of Cuba, Jamaica, and Haiti

1. Vertex at anterior margin more than twice as broad as long in mid-
dle; eyes absent; tegmina not surpassing abdomen; integument un-
pismentedy ek. Sh A OE oe OO 2
— Vertex at anterior margin not broader than long in middle; eyes
present; tegmina much surpassing abdomen; integument pigmented

oe SQA RTE CEE SSR 2 OO) sto? a AD Sa Ae ee 3

2. Disc of frons ecarinate; mesonotum without a median carina, lateral
Carinae ODSCULE§ “OcNe4, Caer. tnt SUM, ALE tnt CAE eae ee minos Sp. N.

— Disc of frons with two arcuate carinae; mesonotum distinctly tri-
Carinates Wit COU CUE TU. Ciba fir) Cat Tie ye antricola, sp. n.

3. Vertex surpassing eyes by length of an eye; frons in profile straight,
IMECUING Vertex ACUTCIY/ "ea. ay. Whee, ee ata ee a ee eee luizi (Myers)

— Vertex relatively shorter; frons in profile convex, more strongly so
in‘ basal "hal ee, SPA SE MARE ty Ee) ase ie 4

4. Mesonotum bright green, with apex of scutellum white ..........

AR umibearen eam Seta. ait rion Ag RAN | Neneh Res ni iio a fulgidus (Van D).
— Mesonotum not green, apex of scutellum not white ............ 5
5. Tegmina with a fuscous spot at apex of each apical vein.........

PE 2a RSC EE, VAR REE Ree EBSaaee Re eaeehd eee fuscosus (Van D.)
— Tegmina without a fuscous spot at apex of each apical vein ..... 6

6) Vertex in) profile evenly rounding intomnons= 7. > 4. nae eee 7

— Vertex in profile meeting frons abruptly, the point of junction dis-
SUAS) eee rice a a eA eR eae tM ts bs er Fs oo 8

7. Tegmina with posterior subapical cell (Cula) much smaller than
cell M3+4, and only about half as long ........... trinitatis Myers

— Tegmina with posterior subapical cell not much smaller than cell
MB--4and almose asvlong? 44.) toe eee es ee aboraca, sp. Nn.

VOLUME 93, NUMBER 3 683

8. Tegmina with subapical cell Cula about twice as long as broad .. 9
— Tegmina with subapical cell Cula not twice as long as broad .... 11
9. Post-tibiae with apical teeth evenly spaced in a single line ...... 10
— Post-tibiae with apical teeth in two unequal groups, not in a single

NERS oer peice ee AR an ear oe en pelagon, sp. n.

10. Tegmina with cell Cula scarcely twice as long as broad, virtually
touching commissural margin; no fuscous spot at stigma; median
frontalrocellstalbsent= ne. ojesa oe co. persephone, sp. n.

— Tegmina with cell Cula more than twice as long as broad, remote
from commissural margin; fuscous spot present at stigma; median
imonealvocellusypresent - ooh 4. kes ne sees ec te hanabanillae Myers

11. Apex of vertex distad of anterior margin of eyes; frons longer than
broad, 5.1:1; distal venation of tegmina reddish-brown .........

ee ete aay VPI EE, WEI SIEERES BOP ME AL II ee 2 princeps, sp. Nn.

— Apex of vertex not distad of anterior margin of eyes; frons longer
than broad, 4.2:1; distal venation of tegmina dark fuscous ......

5 6% OS ESE Bi eS pe ee ey ode eee ee nie a ae conopa, Sp. Nn.

Oeclidius persephone, new species
Figs. 37—42

Vertex longer in middle line than broad at posterior margin (about 2.3:1),
almost horizontal, lateral margins straight, converging distad, apical margin
transverse, shorter than width at base (about 1:1.7), disc deeply hollowed
longitudinally; frons in profile convex, more strongly so in basal half, meet-
ing vertex at obtuse angle, in anterior view longer than broad (about 5.4:1),
wider at apex than at base (about 2.3:1 ), basal margin transverse, lateral
margins straight, diverging distad, foliate, apical margin transverse, median
ocellus absent, post-clypeus tricarinate, anteclypeus medially carinate, ros-
trum reaching to middle of abdomen; eyes and lateral oceili present, anten-
nae with basal segment about as long as broad, subcylindrical, widening distad,
second segment longer than first (3:1) and longer than broad (2:1). Pronotum
medially carinate, fine carina along anterior margin from middle to near
level of lower margin of eye, then bent obliquely towards tegula; mesonotum
little broader than long, tricarinate, with carinae parallel; legs slender, post-
tibiae longer than wide (21.9:1), with seven teeth apically, basal metatarsal
segment and second segment each with seven teeth, tarsal claws small, each
relatively stout basally. Tegmina normal, costal margin slightly produced at
base in shallow convex lobe, wing-tucking apparatus between Cul and Cu2
in basal cell developed as a stout triangular lobe; Sc+R+M< stalk scarcely
longer than basal cell, M slightly thickened or granulate near base, five
subapical cells present and nine or ten apical cells.

Creamy white; lateral margins of frons and vertex, disc of pronotum and

684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

42

Figs. 37-42. Oceclidius persephone n. sp.: 37, head and pronotum, dorsal view; 38, head
and thorax, left side; 39, tegmen; 40, Male genitalia, posterior view; 41, same, posterolateral
view from left; 42, same, right side.

mesonotum, except for carinae and posterolateral margins, and legs, dilute
fuscous, metanotum, abdominal tergites in posterior half, anal segment of
male and inner edge of genital styles, fuscous or reddish brown, suffusion
under translucent parts of abdominal terga in posterior half of abdomen,
dull red. Tegmina hyaline, faintly yellowish, more distinctly so in first apical
cell, veins fuscous. Wings hyaline, veins fuscous.

Anal segment of male moderately long, subtubular in basal half, lateroap-
ical angles each strongly produced laterocaudad and deflexed at tip, apical
margin broadly concave. Pygofer short, dorsolateral angles only slightly
produced, flaring, lateral margins appearing tumid, ventral margin short,
transverse or slightly concave. Aedeagus almost bulbous basally, with nar-
row tubular process at apex recurving dorsad. Styles moderately long, stout,
narrow basally, strongly widening and twisted beyond middle with inner
margin strongly sinuate, outer margin shallowly concave, apical margin tri-
cuspidate, with two outer cusps closely approximated.

Male.—Length, 3.2 mm; tegmen, 3.8 mm.

Holotype 3.—Jamaica: Clarendon, Portland Ridge, Portland Caves,
15.vi.74, S. Peck, in the Canadian National Insect Collection.

This species, which is almost certainly endemic, differs from the known
Jamaican species of Oeclidius, O. fulgidus Van D. and O. fuscosus Van D.
in the absence of a median ocellus. From the former it also differs in the
color of the mesonotum, and from the latter in the color of the frons and
the lower part of the body.

The specific name is a classical proper name and is a noun in apposition.

VOLUME 93, NUMBER 3 685

Figs. 43-50. Oececlidius minos, n. sp.: 43, head and thorax, dorsal view; 44, face and lateral
lobes of pronotum; 45, head and pronotum, left side; 46, third segment of antenna; 47, tegmen;
48, male genitalia, posterior view; 49, same, posterolateral view from left; 50, same, right
side.

Oeclidius minos, new species
Figs. 43-50

Head in dorsal view broader than long, with basal part of frons amply
visible. Vertex broader at hind margin than long in middle line (2.7:1). An-
terior margin carinate, concave, lateral margins carinate, concave, converg-
ing distad, posterior margin transverse, sinuate, disc slightly transversely
depressed; frons in middle line longer than broad, widest at about four-
sevenths from base, convex transversely and in profile, basal margin shorter
than apical margin, lateral margins markedly convex, subfoliately produced
anterolaterad, disc smooth, ecarinate, frontoclypeal suture weakly im-
pressed; clypeus relatively small, tricarinate, rostrum long, slender, reach-
ing to middle of abdomen, eyes and ocelli absent, antennae with basal seg-
ment cylindrical, slightly widening distad, only slightly longer than broad,
second segment cylindrical, longer than broad (about 2.4:1), rounded-trun-
cate apically, third segment subovoid, about twice as long as broad, flagel-
lum long, about four-fifths as long as tegmen. Pronotum relatively large,
anterior margin convex, not carinate, not overlapping vertex medially, pos-
terior margin shallowly angulately excavate, median carina distinct, lateral
carinae obscure, sinuately diverging from near middle line to lateral angles;
tegulae present; mesonotum broader than long (about 1.3:1), slightly convex
transversely, almost ecarinate, with only feeble indication of a pair of lateral
carinae that strongly diverge caudad; post-tibiae longer than wide (20:1),

686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

laterally unarmed, apically with six teeth, basal metatarsal segment with
four teeth, second segment with four teeth, tarsal claws slender, small,
pulvilli small. Tegmina relatively small, reaching scarcely beyond middle of
abdomen, broadest near base, tapering distad and narrowly rounding at
apex, Sc+R, M and Cul present, each forking at transverse line of veinlets,
claval suture distinct, common claval vein apparently not attaining margin.
Wings each reduced to a small lobe on metapleuron.

Creamy white; all sclerites of thorax and abdomen faintly suffused yel-
lowish brown, lateral margins of frons, median carina of pronotum, legs and
male genitalia, brown. Tegmina yellowish hyaline, veins and margin of teg-
men yellowish brown.

Anal segment of male short, relatively deep, lateroapical angles rectan-
gulate. Pygofer short, dorsolateral angles a little produced, convex, slightly
inflected, ventral margin entire. Aedeagus tubular, broad at base, rapidly
narrowing into slender tube directed dorsad. Styles rather broad, each di-
rected dorsad and strongly bent cephalad in distal third, widest at about
two-thirds from base, rounded-truncate apically.

Male.—Length, 2.7 mm; tegmen, 1.7 mm.

Holotype 3.—Jamaica: Clarendon, Jackson Bay, Jackson Bay Cave,
2.vill./4, S. Peck, on roots on moist clay floor, in Canadian National Insect
Collection.

This species is distinguishable from both normally-developed species of
Oeclidius and the wholly subterranean O. hades Fennah (Fennah, 1973:442)
by the shape of the head and the weaker carination of the mesonotum, and
from O. hades it differs also in the more slender legs. The male genitalia
are quite distinctive, but conform in broad structure to the pattern common
in the genus.

The specific name is a classical proper name, and is a noun in apposition.

Oeclidius antricola, new species
Figs. 51-59

Head in dorsal view broader than long, with basal part of frons amply
visible. Vertex broader at hind margin than long in middle (3.5:1), anterior
margin shallowly concave, weakly carinate, lateral margins carinate,
straight, converging distad, posterior margin transverse, disc not depressed,
medially carinate; frons longer than broad (up to 1.2:1, according to view-
point), widest at middle, transversely subconvex, with median third flat,
moderately convex in profile, basal margin little shorter than apical margin,
lateral margins distinctly convex, pair of arcuate carinae, united little before
base of frons, percurrent to apex or nearly so, not meeting distally, fron-
toclypeal suture slightly impressed, clypeus relatively small, laterally cari-
nate, postclypeus obscurely medially carinate, but distinctly so at junction

VOLUME 93, NUMBER 3 687

Figs. 51-59. Oeclidius antricola, n. sp.: 51, body and right tegmen, dorsal view; 52, face
and lateral lobes of pronotum; 53, head and pronotum, left side; 54, tegmen; 55, male genitalia,
posterior view; 56, same, posterolateral view from left; 57, same, right side; 58, anal segment
of male, posterior view; 59, aedeagus, posterior view.

with anteclypeus, which also is medially carinate, rostrum long, slender,
slightly tapering near apex, reaching to basal third of abdomen; eyes and
ocelli absent, antennae with basal segment barrel-shaped, slightly longer
than broad, second segment cylindrical, longer than broad (3:1), rounded-
truncate apically, third segment rather elongate, flagellum longer than third
segment (about 2.5:1). Pronotum relatively large, anterior margin strongly
carinate, broadly convex, not overlapping vertex medially, posterior margin
sinuately concave, median carina weak, lateral lobes of pronotum flat; teg-
ulae present; mesonotum broader than long (about 1.7:1), moderately con-
vex transversely, distinctly tricarinate, with lateral carinae strongly diverg-
ing basad; post-tibiae longer than wide (21.7:1), laterally unarmed, apically
with 6—7 teeth, basal metatarsal segment with five teeth, second segment
with four teeth, tarsal claws slender, small, pulvillus moderately well de-
veloped. Tegmina reaching to apex of abdomen, broadest near base, taper-
ing distad and narrowly rounding at apex, Sc+R, M and Cul present, each
forking at transverse veinlets, position of furcation of veins rather irregular,
claval suture distinct. Wings each reduced to small broadly rounded scale.

688 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Creamy white; legs pale stramineous, carinae of head and thorax fuscous.
Tegmina hyaline, pallid stramineous, veins concolorous.

Anal segment of male short, in dorsal view widening distad in basal two-
thirds, then narrowing in apical third, apical margin shallowly excavate.
Pygofer with dorsolateral angles convex, slightly produced, markedly flaring
laterocaudad, ventral margin produced in slight lobe on each side of middle.
Aedeagus short and broad, vertical, with a circular orifice in middle at wid-
est part. Styles relatively long and broad in side view with dorsal margin
deeply concave, apical margin narrowly rounded, in posterior view gradu-
ally widening distad, moderately curved, rounded apically.

Male.—Length, 3.0 mm; tegmen, 2.0 mm.

Holotype 3.—Jamaica: Clarendon, Jackson Bay, Jackson Bay Cave,
2.vill.74, S.Peck, on roots on moist clay floor, in Canadian National Insect
Collection.

Other material.—6 3, 25 nymphs of different stages, with same data as
type.

This greatly modified species can only be compared with the Mexican O.
hades Fenn. From this it differs in the proportions of the vertex and of the
second antennal segment, the shape and venation of the tegmina, and the
shape of every element of the male genitalia.

The specific name is from the Latin antrum, cave, and suffix -cola, in-
habitant, and is a noun in apposition.

Oeclidius aboraca, new species
Figs. 60-66

Vertex longer in middle line than broad at posterior margin (about 3.5:1),
almost horizontal, lateral margins straight, converging distad, apical margin
concave, shorter than width at base (about 1:2.0), disc deeply hollowed
longitudinally; frons in profile shallowly convex, rounding smoothly into
vertex, in anterior view longer than broad (about 3.5:1), wider at apex than
at base (about 3.6:1), basal margin transverse, lateral margins slightly sin-
uate, diverging distad to one-sixth from apex, foliate, apical margin trans-
verse, median ocellus small, postclypeus tricarinate, anteclypeus medially
carinate, rostrum reaching to middle of abdomen; eyes and lateral ocelli
present and pair of blemmata on each side, antennae with basal segment
about as long as broad, subcylindrical, widening distad, second segment
longer than first (3:1) and longer than broad (2.7:1), third segment with short
arista in addition to flagellum. Pronotum medially carinate, fine carina
along anterior margin from middle to near level of lower margin of eye, then
bent obliquely towards tegula; mesonotum a little broader than long, tricar-
inate, with carinae almost parallel; legs slender, post-tibiae longer than wide
(25:1), with seven teeth apically in two unequal groups, basal metatarsal

VOLUME 93, NUMBER 3 689

66

Figs. 60-66. Oececlidius aboraca, n. sp.: 60, head and thorax, dorsal view; 61, head and
pronotum, left side; 62, base of third segment of antenna; 63, tegmen; 64, male genitalia,
posterior view; 65, same, left side; 66, same, posterolateral view from right.

segment and second segment each with six teeth, tarsal claws small, each
relatively stout basally. Tegmina normal, costal margin slightly produced at
base in shallow convex lobe, wing-tucking apparatus between Cul and Cu2
in basal cell developed as stout triangular lobe; Sc+R+M stalk about as
long as basal cell, five subapical cells present and ten apical cells. Last
subapical cell (cell Cula) longer than broad at base (2.7:1).

Pale yellow; lateral margins of frons and vertex very narrowly, basal
segment of antennae, disc of mesonotum, except mesoscutellum, dark fus-
cous, fore- and middle legs striped with fuscous, abdominal terga and anal
segment of male, dilute fuscous, suffusion under translucent parts of ab-
dominal terga in posterior half of abdomen, dull red. Tegmina hyaline, faint-
ly yellowish, veins dull yellow. Wings hyaline, veins fuscous.

Anal segment of male short, ring-like, lateroapical angles each broadly
and weakly produced ventrad, apical margin transverse. Pygofer short, dor-
solateral angles only slightly produced, flaring, lateral margins weakly sin-
uate, ventral margin short, transverse or slightly concave. Aedeagus almost
bulbous basally, with stout tubular process at apex directed caudad, slender
process a little below this, also directed caudad. Styles moderately long,
stout, widening distad and bent mesodorsad in apical third, acute apically,
moderately long vertical peg-like process on dorsal margin towards base.

Male.—Length, 2.8 mm; tegmen, 4.0 mm.

Holotype 3.—Jamaica: Howard Gap, vii.1961, J. Maldonado Capriles, in
U.S. National Museum of Natural History.

690 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

This species, which is almost certainly endemic, differs from O. fulgidus
Van D. in the colour of the mesonotum, and from O. fuscosus Van D. in
that of the frons.

The specific name is a classical proper name, and is a noun in apposition.

Oeclidius pelagon, new species
Figs. 67-72

Oeclidius fuscosus Myers 1928:18 (misidentification).

Vertex longer in middle line than broad at posterior margin (nearly 4.0: 1),
almost horizontal, lateral margins almost straight, parallel, apical margin
transverse, only little shorter than width at base, disc deeply hollowed lon-
gitudinally; frons in profile convex, more strongly so in basal half, meeting
vertex almost rectangulately, in anterior view longer than broad (about
5.4:1), wider at apex than at base (about 2.3:1), basal margin transverse,
lateral margins straight, diverging distad, foliate, apical margin transverse,
median ocellus absent, post-clypeus tricarinate, anteclypeus medially cari-
nate, rostrum reaching to middle of abdomen; eyes and lateral ocelli present,
antennae with basal segment about as long as broad, subcylindrical, wid-
ening distad, second segment longer than first (3:1) and longer than broad
(2:1). Pronotum medially carinate, fine carina along anterior margin from
middle to near level of lower margin of eye, then bent obliquely towards
tegula; mesonotum little broader than long, tricarinate, with carinae sub-
parallel; legs slender, post-tibiae longer than wide (18:1), with seven teeth
apically, basal metatarsal segment and second segment each with seven
teeth, tarsal claws small, each relatively stout basally. Tegmina normal.
costal margin slightly produced at base in shallow convex lobe, wing-tucking
apparatus between Cul and Cu2 in basal cell developed as stout triangular
lobe; Sc+R+M stalk scarcely longer than basal cell, five subapical cells
present and nine or ten apical cells.

Dark reddish brown; lateral margins of frons and vertex, postclypeus,
disc of pronotum and outer third of lateral lobes and tegulae dorsally, pale
ochraceous, basal segment of rostrum, carinae and posterolateral margins
of mesonotum, mesotibiae and post-tibiae, anal segment of male and third
valvulae of female, pale brownish yellow, metanotum and abdominal terga
in posterior half, dark fuscous. Tegmina milky hyaline, faintly yellowish,
veins yellowish brown. Wings hyaline, veins fuscous.

Anal segment of male exceptionally long, subtubular in basal half, latero-
apical angles each greatly dilated into broad, apically-rounded lobe that is
produced ventrad and slightly overlaps its counterpart apically. Pygofer
moderately long, dorsolateral angles broadly produced, lateral margins ap-
pearing tumid, ventral margin short, transverse or slightly concave, notched
medially. Aedeagus almost bulbous basally, with a narrow tubular process

VOLUME 93, NUMBER 3 691

UL

Figs. 67-72. Oececlidius pelagon, n. sp.: 67, head and thorax, dorsal view; 68, head, prono-
tum and mesonotum, left side; 69, tegmen; 70, male genitalia, posterolateral view from left:
71, same, posterior view; 72, same, right side.

at apex directed dorsad. Styles moderately long, stout, narrowest at base,
strongly widening beyond middle with inner and outer margins sinuately
convex in posterior view, apical margin strongly oblique, with outer angle
acute.

Male.—Length, 3.2 mm; tegmen, 3.8 mm.

Holotype 3.—Cuba: Soledad, 28.111.1925, J. G. Myers, BM 1937-789, in
British Museum (Natural History).

This species, which is almost certainly endemic, differs from the Jamaican
O. fuscosus VanD. in its smaller size, in the color of the basal antennal
segment and in the absence of infuscation on the apical veinlets of the
tegmen where they join the apical margin.

The specific name is a classical proper name and is a noun in apposition.

Oeclidius princeps, new species
Figs. 73-78

Vertex longer in middle line than broad at posterior margin (about 3.0:1),
almost horizontal, lateral margins straight, converging distad, apical margin
transverse, shorter than width at base (about 1:2.4), disc deeply hollowed
longitudinally; frons in profile convex, more strongly so in basal half, meet-
ing vertex at obtuse angle, in anterior view longer than broad (about 5.0:1),
wider at apex than at base (about 2.5:1), basal margin transverse, lateral
margins straight or feebly concave, diverging distad, foliate, apical margin
concave, median ocellus distinct, post-clypeus tricarinate, anteclypeus me-
dially carinate, rostrum reaching to middle of abdomen; eyes and lateral
ocelli present, blemmata apparently absent, antennae with basal segment

692 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

76

Figs. 73-78. Oececlidius princeps, n. sp.: 73, head and thorax, dorsal view; 74, head and
pronotum, left side; 75, tegmen; 76, male genitalia, right side; 77, male genitalia, posterolateral
view from left; 78, same, posterior view.

short, not as long as broad, cylindrical, second segment longer than first
(3:1) and longer than broad (2:1), third segment without arista. Pronotum
medially carinate, fine carina along anterior margin from middle to near
level of lower margin of eye, then bent obliquely towards tegula; mesonotum
little broader than long, tricarinate, with carinae parallel; legs slender, post-
tibiae longer than wide (23:1), with seven teeth apically, not separated into
two groups, basal metatarsal segment and second segment each with seven
teeth, tarsal claws small. Tegmina normal, costal margin slightly produced
ventrad at base in shallow convex lobe, wing-tucking apparatus between
Cul and Cu2 in basal cell developed as stout triangular lobe; Sc+R+M stalk
as long as basal cell, five subapical cells present and ten apical cells.

Pallid ochraceous; mesonotum, except for carinae and posterolateral mar-
gins, orange-brown, metanotum, abdominal tergites in part, reddish brown.
Tegmina hyaline, faintly yellowish, veins dull yellow. Wings hyaline, veins
fuscous.

Anal segment of male moderately long, subtubular in basal half, lateroap-
ical angles each strongly produced laterocaudad and deflexed at tip, apical
margin broadly concave. Pygofer short, dorsolateral angles only slightly
produced, not flaring, ventral margin relatively long, transverse or slightly
concave. Aedeagus almost bulbous basally, with narrow tubular process at
apex recurving dorsad. Styles moderately long, stout, rather narrow basally,
strongly widening beyond middle with inner margin convex, outer margin
shallowly concave, subcrenulate or trilobed, with two outer lobes closely
approximated.

VOLUME 93, NUMBER 3 693

Figs. 79-85. Oceclidius conopa, n. sp.: 79, head and thorax, dorsal view; 80, head and
pronotum, left side; 81, tegmen; 82, right styles, dorsal view; 83, male genitalia, posterior
view; 84, same, posterolateral view from left; 85, same, right side.

Male.—Length, 3.4 mm; tegmen, 4.5 mm.

Holotype 3 .—Haiti: Port au Prince, 11.1925, G. N. Wolcott, in British
Museum (Natural History).

This species, which is almost certainly endemic, is apparently a Haitian
counterpart of the Jamaican O. persephone, but differs markedly from this
species in the shape of the styles.

The specific name is the Latin term for prince, and is used as a noun in
apposition.

Oeclidius conopa, new species
Figs. 79-85

Vertex longer in middle line than broad at posterior margin (about 2.1:1),
almost horizontal, lateral margins straight, converging distad, apical margin
transverse, shorter than width at base (about 1:2.0), disc deeply hollowed
longitudinally; frons in profile convex, more strongly so in basal half, meet-
ing vertex at an obtuse angle, in anterior view longer than broad between
outer edges of lateral margins (about 4.0:1), wider at apex than at base
(about 2.2:1), basal margin transverse, lateral margins shallowly sinuate,
diverging to distal sixth, foliate, apical margin between carinae transverse,
median ocellus present, post-clypeus tricarinate, anteclypeus medially car-
inate, rostrum reaching to middle of abdomen; eyes and lateral ocelli pres-
ent, blemmata apparently absent, antennae with basal segment about as long

694 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 86-92. Oececlidius trinitatis Myers: 86, head and thorax, dorsal view; 87, face; 88,
head and pronotum, left side; 89, tegmen; 90, male genitalia, posterior view; 91, same, pos-
terolateral view from left; 92, same, right side.

as broad, subcylindrical, widening distad, second segment longer than first
(3:1) and longer than broad (2:1). Pronotum medially carinate, fine carina
along anterior margin from middle to near level of lower margin of eye, then
bent obliquely towards tegula; mesonotum little broader than long (about
1.3:1), tricarinate, with carinae subparallel; legs slender, post-tibiae longer
than wide (23:1), with seven teeth apically, not clearly separated into two
groups, basal metatarsal segment and second segment each with seven
teeth, tarsal claws small, each relatively stout basally. Tegmina normal,
costal margin slightly produced mesad at base in a shallow convex lobe,
wing-tucking apparatus between Cul and Cu2 in basal cell developed as
stout narrowly-triangular lobe; Sc+R+M stalk about as long as basal cell,
five subapical cells present and ten apical cells including stigmatic cell.

Pallid ochraceous; disc of mesonotum, except for carinae, and postero-
lateral margins and mesoscutellum, fuscous, more dilute in lateral fields,
metanotum darker fuscous, abdominal terga 3—5 except in posterior half,
and ceriferous segments 6-8 of female, fuscous. Tegmina hyaline, faintly
yellowish, a little more distinctly so in stigmatic cell, veins fuscous. Wings
hyaline, veins fuscous.

Anal segment of male short, almost ring-like, lateroapical angles each
feebly produced ventrad in broadly rounded lobe, apical margin concave.
Pygofer moderately long, dorsolateral angles only slightly produced, mod-
erately inflected mesad, lateral margins straight, ventral margin short, trans-

VOLUME 93, NUMBER 3 695

verse or slightly concave. Aedeagus almost bulbous basally, with a narrow
tubular process at apex recurving dorsad. Styles long, stout, broad basally,
slightly narrowed at middle, where inner margin is strongly concave, outer
margin straight, apical margin strongly trilobate, with outer lobe narrow,
acute, directed dorsomesad and strongly pigmented.

Male.—Length, 2.5 mm; tegmen, 4.1 mm.

Female.—Length, 2.6 mm; tegmen, 5.0 mm.

Holotype 6, Haiti: Port au Prince, vii.1961, J. Maldonado Capriles, in
U.S. National Museum of Natural History.

This species, which is almost certainly endemic, differs from all known
Caribbean species of Oeclidius in the form of the styles.

The specific name is a classical proper name and is a noun in apposition.

Oeclidius trinitatis (Myers)
Figs. 86-92

Myers, 1928:18.

The opportunity is taken to figure the male genitalia of this Cuban species,
the type of which is in the British Museum (Natural History).

Subgenus Paroeclidius Myers

Oeclidius (Paroeclidius) luizi (Myers), new combination
Paroeclidius luizi Myers 1928:20.

In view of the great variability between species of Oeclidius in the length
of the vertex and the degree of development of the mesonotal carinae, there
appears to be insufficient ground for maintaining Paroeclidius, which in-
cludes only one species, as a separate genus on the basis of differences
associated with these structures.

Literature Cited

Fabricius, J. C. 1803. Systema Rhyngotorum: 1-314.

Fennah, R. G. 1945. The Fulgoroidea, or lanternflies, of Trinidad and adjacent parts of South

America.—Proc. U.S. Nat. Mus. 95:411-520.

. 1973. Three new cavernicolous species of Fulgoroidea (Homoptera) from Mexico and

Western Australia.—Proc. Biol. Soc. Washington 86 (38):439- 446.

Kirkaldy, G. W. 1904. Bibliographical and nomenclatorial notes on the Hemiptera. No. 3.—
Entomologist 37:279-383.

Myers, J. G. 1928. Notes on Cuban Fulgoroid Homoptera.—Studies on Cuban Insects
1928: 13-28.

Stal, C. 1869. Hemiptera Fabriciana 2.—Handl. Svenska Vet. Akad. 8(1):1—130.

Van Duzee, E. P. 1907. Notes on Jamaican Hemiptera: a report on a collection of Hemiptera
made on the island of Jamaica in the spring of 1906.—Bull. Buffalo Soc. nat. Sci. 8(5):
3-79.

696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. 1914. A preliminary list of the Hemiptera of San Diego County, California.—Trans.
San Diego Soc. Nat. Hist. 2:1-57.

% Commonwealth Institute of Entomology, British Museum (Natural His-
tory), Cromwell Rd., London SW7 S5BD, England.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 697-707

A SIPHONOSTOME COPEPOD ASSOCIATED WITH A
VESTIMENTIFERAN FROM THE GALAPAGOS RIFT
AND THE EAST PACIFIC RISE?

Arthur G. Humes and Masahiro Dojiri

Abstract—Ceuthoecetes aliger, new genus, new species, a siphonostome
copepod belonging to the new family Dirivultidae Humes and Dojiri (in
press) occurs on the tentacular crown of a large vestimentiferan living in
warm water near hydrothermal vents on the Galapagos Rift and the East
Pacific Rise.

Two species of copepods are known as associates of Vestimentifera
Webb, 1969, a group of tubicolous marine worms placed in the Pogonophora
by Webb (1969) but among the Annelida by van der Land and N@rrevang
(1977). The clausidiid Tychidion guyanense Humes, 1973, is associated with
Lamellibrachia luymesi van der Land and N@¢grrevang, 1977, in 500 m off
Guyana. Dirivultus dentaneus Humes and Dojiri, in press (new family Di-
rivultidae), occurs on the tentacular crown of Lamellibrachia barhami
Webb, 1969, in 1125 m, off southern California. This paper includes the
description of a third species of copepod, this time from the tentacular
crown of a large vestimentiferan from the Galapagos Rift and the East Pa-
cific Rise. The worms from these two areas, tentatively identified as the
Same species, are presently under study by Dr. Meredith L. Jones, National
Museum of Natural History, Smithsonian Institution.

The vestimentiferans, living in warm water near hydrothermal vents on
the Galapagos Rift (about 380 km northwest of the Galapagos and 1,000 km
west of Ecuador) and the East Pacific Rise (about 250 km south of the tip
of Baja California), were collected during dives by the submersible Alvin,
operated by the Woods Hole Oceanographic Institution (see Corliss et al.,
ISSD))s

Three preserved vestimentiferans were washed gently in 70 percent ethyl
alcohol, making sure that the alcohol passed freely between the many la-
mellae on the tentacular crown. The copepods were recovered from the
sediment obtained after passing the solution through a fine net (openings
about 100 wm square). The first vestimentiferan yielded 34 adults and 17

' Contribution no. 7 of the Galapagos Rift Biology Expedition, supported by the National
Science Foundation.

698 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

copepodids; the second | adult and 6 copepodids; and the third 2 adults and
18 copepodids. The total number was 37 adults and 41 copepodids.

The study of the copepods was aided by grant DEB 77 11879 from the
National Science Foundation.

All measurements were made on specimens in lactic acid. The figures
were 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 and MXPD = maxilliped.

Dirivultidae Humes and Dojiri, in press
Ceuthoecetes, new genus

Diagnosis.—Siphonostomatoida, Dirivultidae. Body unmodified. Uro-
some 5-segmented in female, 6-segmented in male. Caudal ramus with six
setae. Rostrum not well defined. First antenna 10-segmented, with aesthete
on next to last segment. Prehensile second antenna with minute exopod and
2-segmented endopod with one terminal claw.

Oral cone short. Mandible a long slender blade with serrate tip, lacking
palp. First maxilla with small outer lobe. Second maxilla prehensile, second
segment merging with long slender claw. Maxilliped prehensile and 4-seg-
mented with terminal claw.

Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 2-
segmented endopod. Formula for endopod 0-0; I,1. Legs 1—4 alike in both
sexes.

Leg 5 sexually dimorphic, in female placed laterally with four setae on
free segment and dorsal seta absent, in male placed ventrally with five setae
on free segment and ‘‘dorsal’’ seta present. Leg 6 in female lacking identi-
fiable setae or spines but perhaps represented by two small processes on
genital area, in male with two setae on posteroventral flap on genital seg-
ment.

Other features as in species described below.

Associated with Vestimentifera.

Type-species.—Ceuthoecetes aliger, new species.

Etymology.—The generic name is a combination of KkevOos, the depths,
and ouxy7ys, an inhabitant. Gender masculine.

Ceuthoecetes aliger, new species
Figs. 1-28

Type-material.—8 2° 2, 26 66, and 17 copepodids from one large vesti-
mentiferan in 2,595 m, Alvin dive no. 915, East Pacific Rise, 20°51’N,
109°4.9'W, 22 April 1979. Holotype °, allotype, and 25 paratypes (4 2°,
21 366) deposited in the National Museum of Natural History, Smithsonian

699

VOLUME 93, NUMBER 3

ra Ac.
fog eee
7 EE
——— pt LOL a ag
—- << ee

—~ oe
= —<. tp LLLE
———— A hd Ae
~ SSS S555" RE
S Wit eecee
DDS

xe
Wee

Genital

dorsal (B); 3,

’

Dorsal (A); 2, Urosome

Figs. 1-6. Ceuthoecetes aliger, female: 1,
area, dorsal (C); 4, Genital area, ventral (D); 5, Segment of leg 5 and genital segment, lateral

(E); 6, Caudal ramus, ventral (F).

700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

iit

10

Figs. 7-12. Ceuthoecetes aliger, female: 7, Oral area, ventral (F); 8, First antenna, ventral
(F); 9, Second antenna, antero-inner (D); 10, Mandible, posterior (D); 11, First maxilla,
posterolateral (C); 12, Second maxilla, posterior (C).

VOLUME 93, NUMBER 3 701

Institution, Washington, D.C.; the remaining paratypes (dissected) and the
copepodids in the collection of the first author.

Other specimens.—1 2, 6 copepodids from part of the tentacular crown
of 2 large vestimentiferans, in 2,482 m, Alvin dive no. 884, Garden of Eden
site, Galapagos Rift (see Corliss et al., 1979), 0°48.1'N, 86°7’W, 25 January
1979; 1 2, 1 6, and 18 copepodids from one entire large vestimentiferan,
same locality and date.

Female.—Body (Fig. 1) cyclopoid. Length 1.02 mm (0.94—1.15 mm) and
greatest width 0.36 mm (0.36—0.37 mm), based on 10 specimens. Segment
of leg 1 not separated from cephalosome. Epimeral areas of segments of
legs 1-4 rounded. Ratio of length to width of prosome 1.40:1. Ratio of length
of prosome to that of urosome 1.08:1.

Segment of leg 5 (Fig. 2) 59 x 84 um. Genital segment 130 x 111 um,
only slightly expanded laterally in anterior third where segment bears a pair
of prominent digitiform dorsoposteriorly directed lobes approximately 62 x
16 um (Figs. 3, 4, 5). Genital areas (Figs. 3, 4, 5) lacking identifiable setae
or spines but with two small conical processes about 5 wm and 11 um (Fig.
4). Three postgenital segments from anterior to posterior 92 x 92, 70 x 84,
and 59 x 81 wm. Anal segment with posteroventral row of small spines on
each side (Fig. 6).

Caudal ramus (Fig. 6) elongate, 122 x 32 um, ratio 3.81:1. Outer lateral
seta 59 um, dorsal seta 46 wm, outermost terminal seta 68 wm, all smooth.
Innermost terminal seta 132 wm, with inner hairs. Two median terminal
setae 167 wm (outer) and 232 um (inner), both bilaterally barbed. Ventral
surface of ramus near lateral seta with two rows of minute spinules.

Body surface with few small hairs (sensilla) as in Figure 1.

Egg sac unknown.

Rostral area (Fig. 7) not well defined.

First antenna (Fig. 8) 10-segmented and 288 um long. Lengths of segments
(measured along their posterior nonsetiferous margins): 8 (20 wm along an-
terior margin), 43, 27, 41, 27, 27, 27, 30, 22, and 27 wm respectively. Formula
for armature: 1, 13, 9, 4,2,2,2,2,2 + 1 aesthete, and 12. All setae smooth.
Second antenna (Fig. 9) 4-segmented, 124 um long. Protopod 2-segmented

but coxa subdivided, giving appearance of three segments. Exopod a minute
process. Endopod 2-segmented, first segment slightly swollen with patch of
_ small spines distally on antero-inner surface; second segment bearing three
naked setae and a recurved claw approximately 30 um long.

Labrum (Fig. 7) broad with posteroventral margin weakly indented. Oral
cone short with incomplete dentate inner ring (Fig. 7).
| Mandible (Fig. 10) a long slender blade 65 «wm with finely serrate tip.
| Paragnath not seen. First maxilla (Fig. 11) with small outer lobe having
| three setae, large inner lobe with four setae, all setae smooth. Second max-
|
|

|
|

702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SS

SSS
SSSSS5

Ss
=
—= << —

Figs. 13-17. Ceuthoecetes aliger, female: 13, Maxilliped, posterior (F); 14, Leg | and
intercoxal plate, anterior (G); 15, Leg 2, anterior (G); 16, Leg 3, anterior (G); 17, Leg 4,
anterior (G).

VOLUME 93, NUMBER 3 703

illa (Fig. 12) with elongate first segment 78 x 25 um including proximal
blunt spinelike process. Second segment merging with long slender claw,
forming functional claw 157 um long provided distally with small spinules.
Maxilliped (Fig. 13) 4-segmented. First segment small with long spinulose
inner seta. Second segment elongate, 189 x 54 um, with short inner seta.
Third segment 51 um long with two very unequal setae. Fourth segment 27
jum, with one seta and two minute spinules. Claw nearly straight, 135 um,
obscurely subdivided and bearing short spinules along inner side.

Ventral area between maxillipeds and first pair of legs without special
sclerotization.

Legs 1—4 (Figs. 14, 15, 16, 17) biramous, with all rami 3-segmented except
for 2-segmented endopod of leg 4. Formula for armature as follows (Roman
numerals indicating spines, Arabic numerals representing setae):

P, coxa 0-0 basis 1-1 exp I-1; I-11; _ II,I,4

enp O-1; 0-2; 1,2,3

P, coxa 0-0 basis 1-0 exp I-1; le JOUL IA!
enp O-1; 0-2; 1,2,3
P, coxa Q-O basis 1-0 exp I-1; I-1; I,I,5
enp 0-1; 0-2; 1,1,3
P, coxa 0-0 basis 1-0 exp I-1; eI

enp 0-0; I,1

Leg 1 (Fig. 14) with intercoxal plate bearing pair of processes on ventral
margin. Basis with smooth inner seta 43 ~m long. Outer spines on exopod
weak and setiform. Proximal five inner setae on endopod with short spinules
near tips instead of long hairs. Outer margin of first endopod segment
smooth (as in legs 2—4 also). Leg 2 (Fig. 15) and leg 3 (Fig. 16) with bifurcate
distal outer process on second endopod segment and with several setae
showing short distal spinules as in leg 1. Leg 4 (Fig. 17) with exopod 216
wm. Endopod with smooth unarmed first segment 49 x 21 wm; second seg-
ment 76 X 21 wm, with terminal barbed spine 146 wm and inner seta 151
ym, segment with small spinules along outer margin.

Leg 5 (Fig. 5) irregular in form, unornamented, greatest dimensions 43 x
22 um (16 wm wide in distal part), bearing four smooth setae from dorsal
to ventral 32, 22, 32, and 46 wm. Dorsal seta absent.

Leg 6 perhaps represented by pair of small conical processes on genital
area (Fig. 4).

Color in living specimens unknown.

Male.—Body (Fig. 18) with cephalosome a little broader than in female.
Length (excluding setae on caudal rami) 1.14 mm (1.03—1.29 mm) and great-
est width 0.42 mm (0.37—0.45 mm), based on 10 specimens. Ratio of length
to width of prosome 1.39:1. Ratio of length of prosome to that of urosome
fe Sls

704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Boe PN
‘
77 N
1 1
‘ 1
‘ 1
N ”
yor
.
y .
UJ 1
‘
. , !
SQ, ’

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a

DN),

———
ae

pmb
CO

Figs. 18-23. Ceuthoecetes aliger, male: 18, Dorsal (A); 19, Urosome, dorsal (B); 20, Uro-
some, lateral (B); 21, First antenna, ventral (E); 22, Second antenna, antero-outer (D); 23,
Maxilliped, posterior (F).

VOLUME 93, NUMBER 3 705

ae

ie AT
SS

LZ

ZEEE

Figs. 24-28. Ceuthoecetes aliger, male: 24, Leg 1 and intercoxal plate, anterior (G); 25,
Leg 2 and intercoxal plate, anterior (G); 26, Leg 3, anterior (G); 27, Legs 5 and 6, ventral (E);
28, Spermatophore, as attached to genital area of female (E).

706 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Segment of leg 5 (Fig. 19) 81 x 81 um. Genital segment 118 x 113 um,
almost quadrate in dorsal view, but with prominent dorsal ridge in lateral
view (Fig. 20). Four postgenital segments from anterior to posterior 76 x
86, 76 x 81, 59 x 70, and 65 x 70 um.

Caudal ramus resembling that of female but a little smaller, 113 x 27 um,
ratio 4.2:1.

Body surface and rostrum as in female.

First antenna (Fig. 21) 10-segmented, recurved, length if straightened ap-
proximately 318 wm. Lengths of segments (measured along their posterior
nonsetiferous margins): 19 (27 wm along anterior margin), 41, 22, 22, 46, 30,
30, 46, 35, and 30 um respectively. Armature: 1, 14, 7, 4, 4, 2,2, 2 + minute
knob, 1 + aesthete + 2 small knobs, and 11 + bifid knob.

Second antenna (Fig. 22) resembling that of female but small exopod bifid.
First endopod segment with inner ridge and few small outer spinules, and
second endopod segment with one seta bifurcate. Labrum, oral cone, man-
dible, first maxilla, and second maxilla as in female. Paragnath not seen.
Maxilliped (Fig. 23) with formula for armature as in female. Seta on first
segment spinulose. Second segment stouter than in female. Claw 84 um,
much shorter than in female and relatively stouter and more unguiform.

Legs 1-4 having same segmentation and formula for armature as in fe-
male. First segment of endopod in all four legs with outer hairs (these hairs
absent in female). Leg | (Fig. 24) with inner seta on basis 78 wm long. Leg
2 as in Figure 25. Exopod of leg 3 (Fig. 26) with outer spines 30, 103, 59,
65, and 89 wm from proximal to distal. (In female these spines 13, 28, 27,
29, and 82 um). Leg 4 as in female.

Leg 5 (Fig. 27) placed ventrally. Free segment without fine ornamentation,
43 x 32 um, with five smooth setae, two inner setae broad and 34 wm long,
three outer setae slender, from outer to inner 31, 40, and 61 wm. Seta near
insertion of free segment (corresponding to usual dorsal seta, but here
moved ventrally) 78 wm.

Leg 6 (Fig. 27) a posteroventral flap on genital segment bearing two un-
equal smooth setae 51 wm and 70 um.

Spermatophore (Fig. 28) oval, 57 x 41 um, with long neck.

Color in living specimens unknown.

Etymology.—The specific name aliger, Latin meaning winged, alludes to
the pair of winglike lobes on the genital segment of the female.

Remarks.—Ceuthoecetes aliger, new genus, new species, appears to be
related to Dirivultus dentaneus Humes and Dojiri, in press, described from
the vestimentiferan Lamellibrachia barhami Webb, 1969. The nature of the
first maxilla, the prehensile character of the second maxilla, and the seg-
mentation and armature of legs 1—4 provide evidence for this affinity. Fur-
thermore, the general appearance of the mandible, a very conservative ap-
pendage, reflects this presumed relationship.

There are, however, fundamental differences between Dirivultus denta-

VOLUME 93, NUMBER 3 707

Table 1.—Comparison between Dirivultus dentaneus Humes and Dojiri, in press, and Ceu-
thoecetes aliger, new genus, new species.

Dirivultus Ceuthoecetes
First antenna
Female 13-segmented 10-segmented
Male 12-segmented 10-segmented
Exopod of second antenna l-segmented Small process
Third segment of exopod
of leg 3 III,1,5 RIES
Ieepy5
Female Minute free segment Large free segment
with | seta with 4 setae
Male Distinct free segment Free segment with
with 2 setae 5 setae

neus and Ceuthoecetes aliger (Table 1). Although the differences which
exist between the two species concerning the first antenna and leg 5 are
thought to be important, we believe at this time that Ceuthoecetes can be
accommodated within the family Dirivultidae.

The two collections of Ceuthoecetes aliger are separated by about 3,700
km. This suggests that other populations are likely to be discovered along
the intervening oceanic ridge system as other vent communities are studied.

Acknowledgments

We wish to thank Dr. J. Frederick Grassle of the Woods Hole Oceano-
graphic Institution for allowing us to examine the vestimentiferans from the
Alvin dives and for helpful suggestions during the preparation of this paper.

Literature Cited

Corliss, J. B., J. Dymond, L. I. Gordon, J. M. Edmond, R. P. von Herzen, R. D. Ballard, K.
Green, D. Williams, A. Bainbridge, K. Crane, and T. H. van Andel. 1979. Submarine
thermal springs on the Galapagos Rift.—Science 203: 1073-1083.

Humes, A. G. 1973. Tychidion guyanense n. gen., n. sp. (Copepoda, Cyclopoida) associated

with an annelid off Guyana.—Zool. Med. 46:189-196.

, and M. Dojiri. in press. A new siphonostome family (Copepoda) associated with a

vestimentiferan in deep water off California.—Pac. Sci.

van der Land, J., and A. Ngrrevang. 1977. Structure and relationships of Lamellibrachia
(Annelida, Vestimentifera)—K. Danske Vid. Selsk., Biol. Skr. 2:1—102.

Webb, M. 1969. Lamellibrachia barhami, gen. nov., sp. nov. (Pogonophora) from the north-
east Pacific.—Bull. Mar. Sci. 19:18—47.

Boston University Marine Program, Marine Biological Laboratory,
Woods Hole, Massachusetts 02543.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 708-713

A NEW SPECIES OF SCLEROBREGMA (POLYCHAETA:
SCALIBREGMATIDAE!) FROM OFF THE
SOUTHEASTERN UNITED STATES

Rodney D. Bertelsen and Donald P. Weston

Abstract.—Sclerobregma stenocerum, a new polychaete species of the
family Scalibregmatidae, is described from the continental shelf from Cape
Lookout, North Carolina to Daytona Beach, Florida. The placement of S.
stenocerum in the genus Sclerobregma and its affinity with the only other
member of the genus, S. branchiatum, are discussed.

The scalibregmid genus Sclerobregma previously included only Sclero-
bregma branchiatum Hartman, 1965, as S. branchiata. This species has been
collected at slope and abyssal depths (400 to 2,500 m) off New England.

During two recent studies, a new species of Sclerobregma was collected
at shelf depths throughout the South Atlantic Bight (Cape Hatteras to Cape
Canaveral). As part of a U.S. Bureau of Land Management sponsored
study, Texas Instruments, Inc. sampled the benthos of seven cross-shelf
transects from Cape Fear, North Carolina to Daytona Beach, Florida. Sam-
ples were taken seasonally from February to November, 1977 using a 0.06
m? box corer. Forty-six specimens of this new Sclerobregma species were
collected at locations indicated in Figure 1.

An additional specimen was found off Cape Lookout, North Carolina
during a cooperative University of Wisconsin—Virginia Institute of Marine
Science study of continental shelf benthos. Samples were taken seasonally
from May, 1977 through January, 1978 in an area extending from northeast
of Oregon Inlet to east of Cape Lookout, using a 0.1 m? Smith MacIntyre
grab.

Sclerobregma stenocerum, new species
Figs. 2-3

Holotype.—ENE of Daytona Beach, Florida; 29°34'N, 80°22’W; 44 m;
Sta. 899-1 (7D); 2 September 1977; USNM 58955.

Paratypes.—1 specimen; NE of Jacksonville, Florida; 31°03’N, 80°26’ W;
34 m; Sta. 230-1 (SE); 26 February 1977; USNM 58956.

1 specimen; E of Jacksonville, Florida; 30°23’N, 80°36’ W; 35 m; Sta. 262-
2 (6D); 1 March 1977; USNM 58957.

! The Greek word bregma has as its stem bregmat-. Therefore, the family name should be
Scalibregmatidae (G. Steyskal, pers. comm.)

VOLUME 93, NUMBER 3 709

Q
NORTH OREGON INLET
CAROLINA a

HATTERAS

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FLORIDA

Fig. 1. Map of the South Atlantic Bight of the eastern United States showing sampling
locations. Texas Instruments’ stations indicated by small circles; Univ. of Wisconsin—Virginia
Institute of Marine Science collection areas within open blocks. Sclerobregma stenocerum
collection sites shown by darkened triangles.

710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Sclerobregma stenocerum: A, Prostomium, lateral view; B, Anterior end, dorsal
view; C, Anterior end, ventral view.

1 specimen; E of Charleston, South Carolina; 32°40'N, 78°47’W; 37 m;
Sta. 488-4 (2E); 13 May 1977; USNM 58958.

1 specimen; E of Cape Lookout, North Carolina; 34°34.5'N, 76°13.1'W;
38 m; Sta. 002-1; 31 May 1977; USNM 58959.

Description.—Holotype 8.3 mm in length, 1.2 mm in width, with 46 seg-—
ments. Other specimens up to 23.1 mm long, 2.8 mm wide, with up to 49 |
segments. Body inflated anteriorly between setigers 6 and 17. Posterior cy-
lindrical and slender. Segmental divisions obscurred by transverse annula-
tions and longitudinal striations (Fig. 2b, c).

Prostomium (Fig. 2a—c) T-shaped with long, slender frontal horns. Single
pair of posterolateral eyes present on prostomium. Each eye V-shaped with
points directed anteriorly (Fig. 2a). Eyes partly hidden by buccal segment
in some specimens. Nuchal organs branched, originating at posterolateral —
margins of prostomium. Proboscis a soft, eversible pouch. Nuchal organs |
and proboscis generally retracted in preserved material. Buccal segment
achaetous and apodous.

Branchiae 3 pairs; inserted posterior to notopodia of setigers 3—5 (Figs.

VOLUME 93, NUMBER 3 711

0.01 mm

- Ge

RG

Fig. 3. Sclerobregma stenocerum: A, Fourth parapodium, anterior view; B, Posterior para-
podium, anterior view; C, Acicular seta from anterior setiger; D, Furcate seta.

Vile PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Distinguishing features of Sclerobregma branchiatum and Sclerobregma steno-
cerum.

Sclerobregma branchiatum Sclerobregma stenocerum
Eyes Absent Present
Frontal horns Short Elongate
Acicular setae Long, pointed, hirsute Very short, blunt, smooth
Branchiae 4 pairs (setigers 2-5) 3 pairs (setigers 3—S)
Ventral cirri Some with slender, pale Without terminal process

process at tip

2b, 3a). Branchiae basically pinnate in form; each pinna with additional
bifurcations.

Anterior 15 setigers with reduced parapodial lobes, lacking both dorsal
and ventral cirri (Fig. 3a). Middle and posterior parapodia with inflated
dorsal and ventral cirri, and small, rounded interramal organs (Fig. 3b). Cirri
becoming more elongate in far posterior setigers. Both dorsal and ventral
cirri usually retain reddish-brown pigmentation in alcohol.

Setae of 3 types: 1) smooth capillaries in both fascicles of all setigers; 2)
short, curved, blunt acicula (Fig. 3c) in both rami of first 2-3 setigers (2-6
acicula per fascicle); 3) furcate setae (Fig. 3d) in both rami of all setigers
from the third. Tines of furcate setae of unequal length and spinous along
inner margins. Up to 14 furcate setae per fascicle, inserted in row anterior
to capillary setae.

Ventral and ventrolateral anal cirri present; exact number indeterminable
as cirri are easily detached. Number of anal cirri ranging from 0-3 on spec-
imens examined.

Distribution.—Continental shelf, North Carolina to Florida; 17 to 218 m.

Etymology.—Steno, Greek for narrow; Keras, Greek for horn; referring
to the long, narrow frontal horns on the prostomium.

Discussion

Following Ashworth (1901) and Kudenov and Blake (1978), Sclerobregma
stenocerum is referable to the genus Sclerobregma on the basis of the ar-
enicoliform body and the presence of dorsal and ventral cirri, acicular setae
and branchiae. S. stenocerum differs from the generic description (Hart-
man, 1965) in that eyes are present. While the presence or absence of eyes
is a valid specific character, their presence in S. stenocerum is insufficient
justification to exclude it from the genus (J. Kudenov, pers. comm.). S.
stenocerum may be readily distinguished from S$. branchiatum by the char-
acteristics given in Table 1.

Examination of the type-material of Sclerobregma branchiatum, provided

VOLUME 93, NUMBER 3 718

by the Allan Hancock Foundation, revealed two errors in the original de-
scription. Hartman (1965; p. 185) stated that the ‘‘upper cirri are longest,
terminating in slender pale tips’’ (see also Hartman’s Plate 42, Fig. C). In
the type-material, the ventral, rather than the dorsal cirri, bear slender, pale
tips. Only the six anteriormost ventral cirri have this terminal structure,
while the remaining ventral cirri and all dorsal cirri are blunt and inflated as
in S. stenocerum.

Secondly, Hartman (1965; Plate 42, Fig. C) illustrated a small cirrus-like
projection beneath the “‘neuropodia’’ (actually the notopodia). Though not
discussed in the text, the figure caption refers to this structure as a ventral
cirrus. The structure could not be found on the holotype (AHF Poly 0484)
or the 17 paratypes (AHF Poly 0485) examined and is presumed to be an
error in the figure.

Acknowledgments

The authors wish to thank Dr. Jerry Kudenov of the University of South-
ern California for his helpful comments and loan of type-material from the
Allan Hancock Foundation. Valuable advice provided by Dr. Kristian Fau-
chald of the U.S. National Museum, and his critical review of the manu-
script is also gratefully acknowledged.

This material is based on research partially supported under Contract No.
AAS550-CT7-2 from the U.S. Bureau of Land Management to Texas Instru-
ments, Inc. Additional support was received through the National Science
Foundation under Grant No. OCE77-08531 and by the National Oceanic
and Atmospheric Administration’s office of Sea Grant, Dept. of Commerce
through an institutional grant to the University of Wisconsin.

Contribution No. 930 from the Virginia Institute of Marine Science.

Literature Cited

Ashworth, J. H. 1901. The anatomy of Scalibregma inflatum Rathke.—Quart. J. Micro. Sci.,
London 45:237-309.

Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda
and other North Atlantic areas.—Allan Hancock Foundation Occasional Paper 28:1-
378.

Kudenov, J. D., and J. A. Blake. 1978. A review of the genera and species of the Scalibreg-
midae (Polychaeta) with descriptions of one new genus and three new species from
Australia.—J. Nat. Hist. 12:427—444.

(RDB) Normandeau Associates, 15 Pickering Ave., Portsmouth, New
Hampshire 03801; (DPW) Department of Invertebrate Ecology, Virginia
Institute of Marine Science, and School of Marine Science, College of Wil-
liam and Mary, Gloucester Point, Virginia 23062.

PROC. BIOL. SOC. WASH.

93(3), 1980, pp. 714-724 |

TWO NEW SPECIES OF CENTROLENELLA FROM
BOLIVIA (ANURA: CENTROLENIDAE)

David C. Cannatella

Abstract.—Two new species of glass-frogs, Centrolenella bejaranoi and

C. bergeri, are described from the cloud forest of the Amazonian slopes of |

Bolivia. One species is allied to the C. fleischmanni group; the other is
apparently a member of the C. prosoblepon group. This is the first record
of the family Centrolenidae from Bolivia.

A recent checklist (Duellman, 1977) of the centrolenid frogs shows that
they are present in most countries in South America; notable exceptions are
Chile, Paraguay, Uruguay, and Bolivia. The apparent absence of these glass-
frogs from Bolivia is surprising, because the yungas region of the Amazo-
nian slopes certainly offers suitable habitat, and Centrolenella are known
from southern Peru (Duellman, 1976). In fact, the absence of Bolivian Cen-
trolenella is an artifact of collections; two recently discovered glass-frogs
are described in this paper.

In January 1979, Thomas J. Berger and I collected in the yungas of the
Chapare region, northeast of Cochabamba, Bolivia (Fig. 1). From the city
of Cochabamba (2,600 m) the road winds northeasterly to a pass (3,370 m,
km 40.5) and descends through the town of Colomi (3,240 m, km 47.7) in
the drainage of the Rio Corani. At about 3,000 m the upper edge of the
pristine forest is reached; tree ferns and viney bamboo are present. The
road heads east and enters the drainage of the Rio Santa Isabel. The cloud
forest on the upper slopes of the valley is virtually uncut (Fig. 2); numerous
streams drain the slopes.

There is a short tunnel through the mountainside at km 86.8, 1,950 m.
From km 101 to km 122 the road descends rapidly from 1,950 m to 860 m.
The small village of Villa Tunari is at km 160 (300 m). One of our collection
sites was a tiny rivulet at km 102, 1,980 m, 15.2 km NE (downhill) of the
tunnel. Alternatively, the locality can be described as 58.1 km SW Villa
Tunari. Our efforts yielded a small collection of frogs, including two new
species of Centrolenella. This is the first record of the family Centrolenidae
from Bolivia.

Methods.—In the following descriptions, the diagnoses follow the format
of Lynch and Duellman (1973). The webbing formulae are described as in
Savage and Heyer (1967); snout-vent length is abbreviated SVL. For spec-
imen comparisons I have examined the material in The University of Kansas

]

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VOLUME 93, NUMBER 3 Tis)

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Fig. 1. Map indicating type locality (arrow) of Centrolenella bergeri and Centrolenella
bejaranoi, 58.1 km SW Villa Tunari (by road), 1,980 m, Departamento Cochabamba, Bolivia.

Museum of Natural History (KU) listed by Lynch and Duellman (1973) and
Duellman (1976). Color notes from life are taken from the field notes of
William E. Duellman and the author. The color of the peritoneum in pre-
served specimens was determined by cutting into the abdomen and record-
ing from direct observation. Fig. | was redrawn from Mapa de Bolivia,
1:1,000,000, 1973, Instituto Geografico Militar, Bolivia.

Centrolenella bejaranoi, new species
Fig. 3

Holotype.—KU 182369, adult male, 23.8 mm SVL, from 58.1 km SW
Villa Tunari (by road), 1,980 m, Departamento Cochabamba, Bolivia
(65°50’W, 17°11'S), obtained 21 January 1979, by Thomas J. Berger and
David C. Cannatella.

Paratypes.—KU 182370-1, same data as the holotype.

Diagnosis.—1) Prevomerine teeth absent; 2) bones pale green in life; 3)
parietal peritoneum mostly clear, the portion immediately ventral to the

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mm. Bottom: View of cloud forest at type locality.

heart white; pericardial peritoneum ventral to the heart white; visceral peri-
toneum clear; 4) color in life dark green with minute off-white spots on
dorsal surfaces; in preservative, lavender with minute white spots; 5) web-
bing formula of fingers III(3—3~)IV; 6) webbing formula of toes I(2—

VOLUME 93, NUMBER 3

Centrolenella bergeri, male, paratype, KU 182364, SVL 23.8 mm.

717

718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2.5)II(1.25—2.5)IN(1.25—2.5)1IV(2.5—1.5)V; 7) snout round in dorsal view;
subtruncate in lateral profile; 8) dorsal skin spiculate; 9) forearm and tarsus
lacking conspicuous dermal folds; 10) humeral spine absent in males; 11)
lower three-fourths of tympanum visible, oriented dorsolaterally, with slight
posterior inclination.

The presence of a spiculate dorsum which is lavender in preservative
distinguishes this species from most other Andean species of Centrolenella.
Centrolenella buckleyi, C. spiculata, C. truebae, and C. johnelsi have spic-
ules on the dorsal surfaces. The much larger C. buckleyi differs by having
humeral spines in the males and lacking white spots; C. truebae differs by
having dark flecks on the dorsum and a denser arrangement of spicules than
C. bejaranoi. Centrolenella spiculata is similar to C. bejaranoi in the gen-
eral appearance and density of spicules, but has prevomerine teeth and
much more extensive webbing. The spicules of Centrolenella bejaranoi are
heterogenous in size; those of C. buckleyi, C. truebae, and C. spiculata are
rather homogenous in size.

Cochran and Goin (1970) described Centrolenella johnelsi from a single
male specimen from Medellin, Colombia. I have not examined specimens
of C. johnelsi, but have relied on the description and also notes on the
holotype provided by William E. Duellman for the following comparisons:
Centrolenella bejaranoi and C. johnelsi have spiculate dorsa and lack pre-
vomerine teeth. However, C. johnelsi is a much larger frog (29.3 mm vs.
23.8—-24.4 mm in males) than C. bejaranoi. Also, C. johnelsi possesses
humeral spines; these are absent in C. bejaranoi.

Description.—Adult males of moderate size for centrolenid frogs; SVL
23.8-24.4 mm (xk = 24.1, n = 3); females unknown. Head slightly wider than
body; snout short, round in dorsal view, subtruncate in lateral view; canthus
round and indistinct, loreal region slightly concave; lips not flared; nostrils
nearly terminal on snout, not flared, directed laterally, internarial region
concave. Eye moderately large, directed anterolaterally. Supratympanic
fold obvious; lower three-fourths of tympanum visible, directed anterolat-
erally with slight posterior inclination. Prevomerine dentigerous processes
and teeth absent; choanae oval; tongue broadly cordiform, slightly free pos-
teriorly; short vocal slits present in males, extending from angle of jaw
almost to midlateral edge of tongue.

Humeral spine absent; conspicuous dermal folds lacking; order of fingers
from shortest to longest 1-2-4-3; webbing absent between first three fingers
(Fig. 4); fingers lacking lateral fringes; finger discs broad, subtruncate; sub-
articular tubercles small, round, single; supernumerary tubercles absent;
palmar tubercle moderately large, round, single; thenar tubercle elliptical.

Hind limbs slender; tibia length 53.3—56.6% of SVL; conspicuous tarsal
fold absent; inner metatarsal tubercle small, flattened, elliptical; outer meta-
tarsal tubercle absent; subarticular tubercles small, round; supernumerary

VOLUME 93, NUMBER 3 TAS

tubercles absent; toes about one-half webbed (Fig. 5); discs rounded to
slightly truncate, smaller than those on fingers.

Skin on dorsal surfaces bearing spicules of heterogenous sizes; spicules
present on upper eyelids and upper lip. Skin of belly and ventral thighs
granular; chin, forelimbs, flanks, and shanks smooth. Anal opening directed
posteriorly at upper level of thighs; anal tubercles and ornamentation ab-
sent.

Color in preservative.—Dorsal surfaces lavender with minute white spots,
each white spot enclosing a large spicule. Ventral surfaces white or trans-
parent.

Color in life.—Dorsal surfaces dark green with minute off-white spots.
Digits orange-green; chest white; parietal peritoneum generally clear, that
portion ventral to the heart is white; pericardial peritoneum covering ventral
surface of the heart white; visceral peritoneum clear; bones and vocal sac
pale green. Iris metallic yellow with black flecks.

Distribution.—This species is known only from the type locality in cloud
forest of the Chapare region of the yungas of Bolivia (Fig. 1).

Etymology.—This species is named in honor of Professor Gaston Bejar-
ano B., government official and naturalist, who encouraged and supported
our work in Bolivia.

Remarks.—The stream where C. bejaranoi was collected is about 0.5 m
wide; the stream bed consists of small rocks and gravel. The frogs were
collected from heights of 1-2 m above the stream by night within 25 m of
the road. This species was not observed to be calling.

Lynch and Duellman (1973) distinguished three patterns of peritoneal pig-
mentation among Centrolenella: 1) clear parietal, white visceral (C. fleisch-
manni and pulverata groups, and C. albotunica, C. eurygnatha, and C.
vanzolinii of southeastern Brasil); 2) white parietal, clear visceral (C. pro-
soblepon group); and 3) white parietal, white visceral (C. antioquensis and
C. resplendens).

In the pattern of peritoneal pigmentation, as well as in other characters,
C. bejaranoi fits most readily, though not clearly, in the C. prosoblepon
group. The visceral peritoneum is without pigment; only a small portion of
the parietal peritoneum, immediately ventral to the heart, is white. There
may be intraspecific variation in the extent of the white pigment in centro-
lenid frogs, but such variation is not exhibited in the type series of C.
bejaranoi.

Centrolenella bergeri, new species
Fig. 3
Centrolenella munozorum (partim)—Duellman, 1976, p. 2.

Holotype.—KU 182363, adult male, 24.8 mm SVL, from 58.1 km SW

720 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Villa Tunari (by road), 1,980 m, Departamento Cochabamba, Bolivia
(65°50’'W, 17°11'S), obtained 21 January 1979, by Thomas J. Berger and
David C. Cannatella.

Paratypes.—KU 182364-8, collected 21-22 January 1979, same locality
and collectors as the holotype.

Referred specimens.—KU 162248-50, Peru:Cuzco:Rio Cosnipata, 4 km
SW Santa Isabel, 1,700 m. KU 162251-8, 162259-60 (eggs), Peru: Ayacucho:
Tutumbaro, Rio Piene, 1,840 m.

Diagnosis.—1) Prevomerine teeth absent; 2) bones white in life; 3) parietal
peritoneum clear; visceral peritoneum white; 4) color in life pale yellow-
green with diffuse yellow spots; color in preservative creamy white; 5) web-
bing formula of fingers III(2.25—2*)IV; 6) webbing formula of toes I(1.5—
2~)IIG1.S—2) TIC .25—2.5)1V(2.75—1.5)V; 7) snout round in dorsal view,
subtruncate in lateral view; 8) dorsal skin shagreened; 9) forearm and tarsus
lacking conspicuous dermal folds; 10) humeral spine absent in males; 11)
tympanum concealed beneath skin, strongly oriented dorsolaterally.

Few Andean Centrolenella (Centrolenella munozorum, C. pellucida, and
C. fleischmanni) are creamy white in preservative. Centrolenella bergeri
differs from munozorum in the extent of webbing and relative size of the
fingers (Fig. 4). Centrolenella fleischmanni has a visible tympanum; it is
concealed in C. bergeri. Centrolenella pellucida has more extensive web-
bing on the hands and bears well-developed dermal folds on the limbs and
below the anus (see remarks); C. bergeri lacks such dermal folds.

Description.—Adult males of moderate size for centrolenid frogs; SVL
22.7—-26.5 mm (x = 24.0, n = 13); females 23.2—26.3 mm (x = 24.8, n = 2);
A Mann-Whitney test between SVL’s of males (n = 5) from Rio Santa Is-
abel, Bolivia, and males (n = 7) from Rio Piene, Peru, demonstrates that
the samples are not different in SVL (0.3 > P > 0.2). SVL is correlated
with tibia length (Spearman rank correlation coefficient, r = 0.80; P <
0.001); the mean ratio of tibia length/SVL equals 0.524, n = 12.

Head wider than body; snout short, round in dorsal view, subtruncate in
lateral view; canthus round, indistinct, loreal region barely concave; lips
not flared; nostrils nearly terminal, directed laterally; internarial region flat.
Eyes large, protuberant, directed anterolaterally. Supratympanic fold ab-
sent; tympanum concealed beneath skin. Prevomerine dentigerous process-
es and teeth absent; choanae large, oval; tongue round, barely free poste-
riorly; short vocal slits present in males, extending from angle of jaw almost
to midlateral border of tongue.

Humeral spine absent; conspicuous dermal folds lacking; order of fingers
from shortest to longest 2-1-4-3; fingers slightly webbed (Fig. 4); finger discs
moderately large, rounded; fingers with narrow lateral fringes; subarticular
tubercles small, low; supernumerary tubercles absent; palmar tubercle in-
distinct.

VOLUME 93, NUMBER 3 72M

Hind limbs slender; tibia length 52.4% (mean) of SVL; conspicuous der-
mal folds lacking; inner metatarsal tubercle small, ovoid; outer metatarsal
tubercle absent; subarticular tubercles small, low; supernumerary tubercles
absent; toes about three-fourths webbed (Fig. 5); discs on toes round, small-
er than those on fingers.

Skin on dorsal surfaces of head and body shagreened; skin on venter
granular; skin on other surfaces smooth; anal opening directed posteriorly
at upper level of thighs; anal ornamentation and tubercles absent.

Color in preservative.—Dorsal surfaces creamy white with minute purple
flecks (chromatophores); ventral skin transparent.

Color in life.-—Dorsum pale green with diffuse yellow spots; venter and
concealed surfaces without pigment; digits orange-yellow; parietal perito-
neum clear, pericardial peritoneum clear; visceral peritoneum white, bones
white. Iris metallic yellow-white with a few black flecks.

Distribution.—The species is known in Bolivia only from the type locality
on the Amazonian slopes of Departamento Cochabamba (Fig. 1) and from
two localities in cloud forest of the Cordillera Oriental in southern Peru (see
maps in Duellman, 1976, 1978). The known elevational range of the species
is 1,700—1,980 m. There is a gap of about 700 km between the Bolivian and
Rio Cosnipata (Peru) populations.

Etymology.—This species is named for Thomas J. Berger, who provided
comic relief while securing part of the type series.

Remarks.—Duellman (1976) tentatively referred specimens from southern
Peru to Centrolenella munozorum. He remarked that specimens from the
Rio Cosnipata and Rio Piene have less webbing on the hands and feet and
are larger than C. munozorum from Ecuador, differences of a magnitude
usually seen between species of Centrolenella but perhaps representing cli-
nal variation. Duellman also noted that the Ecuadorian and Peruvian pop-
ulations were identical in coloration and lacked structural differences.

I have compared directly the holotype and most of the paratypes of C.
munozorum with C. bergeri from Peru and Bolivia. In the former species
the fingers are relatively much shorter and more fully webbed than those of
the latter (compare Fig. 4A, this paper, with Fig. 3D, Lynch and Duellman,
1973). The samples from southern Peru and Bolivia exhibit no differences
among themselves, but are substantially different from C. munozorum.
Specimens of C. munozorum reported by Duellman (1976) and Toft and
Duellman (1979) from 200 m in Departamento Huanuco, Peru, are identical
with the Ecuadorian C. munozorum.

Centrolenella bergeri is most similar to C. pellucida, from the Amazonian
versant of Ecuador. The two species agree closely in many characters; C.
pellucida has more webbing on the hands and feet than does C. bergeri, but
the proportions of the hands and feet are quite similar. The species are
identical in coloration. One of the diagnostic features of C. pellucida—

722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Palmar view of hands: A, Centrolenella bergeri, paratype, KU 182366; B, Centro-
lenella pellucida, holotype, KU 143298; C, Centrolenella bejaranoi, paratype, KU 182371.
Line equals 5 mm.

known only from the holotype—is an unscalloped dermal fold on the outer
edge of the hand and forearm, and on the foot and tarsus; a second character
is a transverse dermal fold below the anus (Lynch and Duellman, 1973).
Having examined the holotype of C. pellucida (KU 143298), I believe that
the subanal dermal fold represents a preservation artifact. The dermal folds
along the limbs also may be due partly to preservation.

The holotype of C. pellucida is a gravid female, SVL 22.0 mm. This is
smaller than the smallest male (22.7 mm) and female, also gravid, (23.2 mm)
of C. bergeri.

Centrolenella bergeri obviously belongs to the group of species—C.
fleischmanni, C. munozorum, and C. pellucida—that has white bones in
life, white visceral peritoneum, clear parietal peritoneum, white ground col-
or in preservative, and lacks prevomerine teeth and dentigerous processes,
humeral spines, subanal tubercles, and distinct canthi (Lynch and Duellman,
1973). The southernmost reported locality of C. fleischmanni 1s Quevedo,
Pacific lowlands of Ecuador (Noble, 1924). The lowland species C. muno-
zorum ranges south to the Rio Llullapichis in central Peru (Toft and Duell-
man, 1979). Centrolenella pellucida is known only from the cloud forest of
the Rio Coca drainage of the Amazonian slopes of Ecuador. No members
of the group are known from the cloud forests of southern Ecuador and

VOLUME 93, NUMBER 3 123

Fig. 5. Plantar view of feet: A, Centrolenella bejaranoi, holotype, KU 182369; B, Centro-
lenella bergeri, paratype, KU 182366. Line equals 5 mm.

northern Peru; a gap of about 1,500 km airline separates the closest records
of C. pellucida and C. bergeri.

Life history notes.— Several of the males of the type series were observed
calling from the undersides of leaves 1-3 m above the stream at night. KU
182364 called from the underside of a leaf while sitting about 4 cm from an
egg clutch (KU 182372). The egg clutch consists of 29 embryos at Stage 18
(Gosner, 1960), arranged in a monolayer on the underside of the leaf, not
hanging from the leaf tip. In life, the jelly is clear and the embryos are pale
green (Fig. 2). Duellman (1976) commented on a male calling adjacent to a
clutch of 25 eggs at the Rio Piene in Peru. Centrolenella bergeri and C.
bejaranoi were taken at the same stream in Bolivia. At the Rio Cosnipata
in Peru, C. bergeri was found with C. spiculata, C. truebae, and a specimen
referred to C. ocellata. At the Rio Piene C. bergeri occurred with specimens
referred to C. siren (Duellman, 1976).

724 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

Thomas J. Berger accompanied me in the field in Bolivia; I am grateful
for his companionship. Professor Gaston Bejarano B. of the Departamento
Nacional de Vida Silvestre, Parques Nacionales, Caza y Pesca, of Bolivia
issued collecting and export permits and made our work in Bolivia enjoy-
able. William E. Duellman and John D. Lynch offered support and encour-
agement. Field work was supported by a grant (DEB 76-09986, William E.
Duellman, principal investigator) from the National Science Foundation,
and by a National Science Foundation Graduate Fellowship to the author.

Literature Cited

Cochran, D. M., and C. J. Goin. 1970. Frogs of Colombia.—Bull. U.S. Nat. Mus. 288:i—xii,
1-655.

Duellman, W. E. 1976. Centrolenid frogs from Peru.—Occ. Pap. Mus. Nat. Hist. Univ. Kans.

(52):1-11.

. 1977. Liste de rezenten Amphibien und Reptilien: Hylidae, Centrolenidae, Pseudi-

dae.—Das Tierreich (95):1-225.

1978. New species of leptodactylid frogs of the genus Eleutherodactylus from the

Cosnipata Valley, Peru.—Proc. Biol. Soc. Wash. 91(2):418—430.

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on
identification.—Herpetologica 16(3):183-190.

Lynch, J. D., and W. E. Duellman. 1973. A review of the centrolenid frogs of Ecuador, with
descriptions of new species.—Occ. Pap. Mus. Nat. Hist. Univ. Kans. (16):1—66.
Noble, G. K. 1924. Some neotropical batrachians preserved in the United States National

Museum with a note on the secondary sexual characters of these and other amphibi-
ans.—Proc. Biol. Soc. Wash. 37:65-—72.
Savage, J. M., and W. R. Heyer. 1967. Variation and distribution in the tree-frog genus
Phyllomedusa, in Costa Rica, Central America.—Beitr. Neotrop. Fauna 5(2):111—-131.
Toft, C. A., and W. E. Duellman. 1979. Anurans of the lower Rio Llullapichis, Amazonian
Peru: A preliminary analysis of community structure.—Herpetologica 35(1):71-77.

Museum of Natural History, The University of Kansas, Lawrence 66045.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 725-742

RECORDS OF ANTHURIDS FROM FLORIDA, CENTRAL
AMERICA, AND SOUTH AMERICA (CRUSTACEA:
ISOPODA: ANTHURIDAE)

Brian Kensley

Abstract.—Notes on morphology and new distributional records for
Apanthura magnifica Menzies and Frankenberg, Haliophasma curri Paul
and Menzies, Malacanthura caribbica Paul and Menzies, Skuphonura la-
ticeps Barnard, and Xenanthura brevitelson Barnard are presented. Sku-
phonura itapuca new species, and S. ecuadorensis new species, are de-
scribed.

As part of a series of short contributions on anthurid isopod taxonomy,
several scattered records in the Smithsonian collections from Florida, the
Caribbean, Central and South America are combined prior to a generic
revision of the group.

Family Anthuridae

Apanthura magnifica Menzies and Frankenberg
Figs. 1, 2

Apanthura magnifica Menzies and Frankenberg, 1966:40, fig. 17.—Schultz,
1969:99, fig. 134.—Kruczynski and Myers, 1976:353, figs. 1, 2.—Camp,
Whiting, and Martin, 1977:15.

Previous records.—Off Georgia, 17-137 m; Off Florida, 7-11 m.

Material examined.—Gandy Shell Bar, Florida: 10 July 1966, 2 2; 1 Oct.
1966, 90 2, 43 juv.; 9 Nov. 1966, 27 2, 10 juv.; 14 Dec. 1966, 32 2, 10juv.; 18
Jan. 1967, 36 2, 11 juv.; 12 Apr. 1967, 18 Apr. 1967 6 3 (7.5%) 80 2, +100 juv.;
17 May 1967, 8 ¢ (12.9%) 62 2, +100 juv.; 14 June 1967, 8 o (6.4%) 47
2; 17 Aug. 1967, 66 2, 50 juv.; 20 Sept. 1967, 2 5; St Joseph Bay Florida,
Dey

Remarks.—Kruczynski and Myers (1976) noted an immature (submale)
Stage in A. magnifica, characterized by the possession of a multiarticulate
antennular flagellum, but lacking whorls of aesthetascs, and suggested that
the species is protogynous as is Cyathura. The large sample from Gandy
Shell Bar, Florida, illustrates the low percentage of males in this anthurid
population. This sample also provides evidence of possibly two stages of
immature male. The first is in all respects similar to the mature female,
except for pereopod 1 which is identical with that of a mature male, with

726 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Apanthura magnifica: a, Anterior cephalon in dorsal view; b, Inner view of max-
illiped showing end@jte; c, Mouthparts in situ, maxillipeds removed; d, Pereopod 1, propodus
and dactylus.

its more strongly developed palmar tooth and distal carpal process. The
second submale stage shows both the typical male pereopod | as well as
the multiarticulate flagellum of the antennule lacking aesthetascs. The en-
dopod of pleopod 2 does not carry an appendix masculina. The mature male
possesses both aesthetascs on the antennule as well as an appendix mas-
culina on the second pleopod.

Kruczynski and Myers (1976) state that the basal antennal peduncular
segment is grooved to accommodate the antennule. In fact, it is the second
segment which is grooved.

Both Menzies and Frankenberg (1966), and Kruczynski and Myers (1976)
figure the pleon of A. magnifica as having pleonites 1-3 free, and pleonites
4 and 5 partially fused. Clearing a specimen in lactic acid and chlorozol
black, as well as scanning electron micrographs clearly show pleonites 1—5
to be fused, with sutures indicated laterally by slits, and with slight dorsal
folds linking the lateral slits (Fig. 2a, b).

VOLUME 93, NUMBER 3 Ta,

«laa —— Tap tal oe os Cae, ee ee a aa
Fig. 2. Apanthura magnifica: a, Pleon and telson in dorsal view; b, Pleon in lateral view;
c, Dorsal openings of statocysts at base of telson; d, Pereopod 7.

Haliophasma curri Paul and Menzies
Fig. 31

Haliophasma curri Paul and Menzies, 1971:39, figs. 16, 17.—Poore,
19752532.

Previous records.—Off Venezuela, Cariaco Basin, 11°03’N, 64°37’W,
25) 100%

Material examined.—USNM 136418 Holotype 2 TL 7.0 mm. USNM
136419 Paratype 2 TL 7.0 mm. USNM 173520, 2 TL 7.7 mm, Culebra
Island, Bay of Panama, intertidal.

Remarks.—Poore (1975) mentions that the lack of dorsal pereonal grooves
and pits, and the presence of a 5-segmented maxilliped (as given in the

728 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Malacanthura caribbica: a, Adult 2 in dorsal and lateral view; b, Antennule; c,
Antenna; d, Mandible; e, Maxilla; f, Apex of maxilla; g, Uropod; h, Pleopod 1; i, Haliophasma
curri: maxilliped.

ee

VOLUME 93, NUMBER 3 729

original description) would exclude H. curri from Haliophasma s.s. as re-
defined by him. Examination of the type-material and the Panamanian spec-
imen shows that dorsolateral grooves are present (as they are in all anthu-
rids), and that Paul and Menzies probably mistook a fold in the second
maxillipedal segment for a suture. The maxilliped is 4-segmented and
except for the presence or absence of dorsal pits on pereonites 4—6 (a dif-
ficult feature to see in such small and transparent specimens) H. curri fits
Poore’s definition of Haliophasma.

This second record of the species, from the Pacific side of the Panama Canal,
raises the questions of earlier distributional patterns, and the movement of
species through the canal, but gives little basis for fruitful speculation.

Malacanthura caribbica Paul and Menzies
Figs. 3, 4

Malacanthura caribbica Paul and Menzies, 1971:34, figs, 8, 9.
Malacanthura cumanensis Paul and Menzies, 1971:35, figs. 10, 11.

Description.—®°. Body 17 times longer than wide; integument moderately
indurate. Cephalon half length of pereonite 1, with large eyes. Proportions:
Cle <a) 5 <4 — 5) — 6) = 7, Pereonites) 2 and >) with) middorsal
rectangular pit-like depression on anterior margin; pereonites 4—6 with lon-
gitudinal middorsal slit-like depressions. Pleonites 1-5 fused, segments
faintly indicated ventrolaterally; pleonite 6 free, with middorsal notch in
posterior margin. Telson strongly indurate, narrowly oval, with 2 basal
Statocysts and strong middorsal carina.

Antennular peduncle 4-segmented, 4th segment short; flagellum of 6 ar-
ticles each bearing 2 aesthetascs. Antennal peduncle 5-segmented, 2nd seg-
ment longest and broadest, grooved to accommodate antennule; flagellum
of 4 articles. Mandibular palp 3-segmented, terminal segment with 11 serrate
spines; incisor with 3 cusps; molar reduced, rounded; lacinia with 8 or 9
serrations. Maxilla slender, with 7 distal spines. Maxilliped 5-segmented,
3rd segment with concave outer margin, 4th segment with row of 7 short
serrate spines; scattered serrate spines on all segments as well as short fine
setules. Pereopod 1 unguis one-third length of dactylus; propodus relatively
narrow, palm bearing 4 or 5 slender spines and few setae, plus group of 4
spines distally near dactylar articulation. Pereopod 2 similar to pereopod 1.
Pereopods 5-7 with unguis one-fifth length of dactylus; propodus with short
triangular tooth at posterodistal corner; carpus half length of propodus, not
underriding propodus, with posterodistal triangular spine. Pleopod 1 exopod
operculiform, indurate, with dense plumose setae on distal margin; endopod
half width and three-quarters length of exopod, with 7 distal plumose setae;
basis with 8 retinaculae. Uropod strongly indurate, exopod with outer mar-

730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Malacanthura caribbica: a, Pereopod 1; b, Pereopod 7; c, Maxilliped.

gin sinuate, apically subacute, margins with strong serrations, shorter than
basis; latter with mediodistal corner somewhat produced, 3 times length of
endopod; latter apically acute, margins strongly serrate.

Previous records.—Off Venezuela, 11°03'N, 64°37’W, 95 m.

Material examined.—USNM 136422, Holotype of M. caribbica, 2° TL
17.0 mm. USNM 136423, Paratype of M. caribbica, 2° TL 9.0 mm. USNM
136424, Holotype of M. cumanensis, 2 TL 7.5 mm. USNM 173521, 3 °,
TL 27.1 mm, 25.7 mm, 18.0 mm; off Cape La Vela, Colombia, 42—44 m.

Remarks.—The two large specimens from Colombia prompted re-exam-
ination of Paul and Menzies’ two species of Malacanthura, both coming
from the same locality off Venezuela. As all the above material shows very
similar structure of the uropods and telson, mouthparts, and pereonites, it
was concluded that only one species is involved here. Paul and Menzies
separated M. cumanensis from M. caribbica by the former’s lack of pig-
mentation, by maxillipedal structure, and by the lack of serrations on the
uropodal exopod. With the exception of species of Mesanthura, pigmen-
tation would seem to be an unreliable character for specific separation in
the anthurids. Re-examination of the maxilliped and uropod showed that

VOLUME 93, NUMBER 3 731

what was taken for a sixth segment in the maxilliped of M. caribbica is an
integumental fold in the fifth segment, and that transparent serrations are
present on the uropodal exopod. Perhaps the most important feature com-
mon to all the specimens is the presence on pereonites 2 and 3 of a middorsal
rectangular pit-like depression on the anterior margin, and middorsal slit-
like depressions on pereonites 4—6. As the Colombian specimens are con-
siderably larger than Paul and Menzies’ type-material the above revised
description is given.

Skuphonura Barnard 1925

Diagnosis.—Cephalon wider distally than proximally; anterolateral lobes
extending beyond level of rostrum. Pleonites 1—S fused, segments indicated
ventrolaterally; pleonite 6 free, with middorsal slit in posterior margin. Some
form of midventral tubercle or spine(s) at base of mouthparts. Flagella of
antennae and antennules of single article in male and female; antennular
flagellum in male with single tuft of aesthetascs. Pleopod | operculiform.
Carpus of pereopods 4—7 triangular, underriding propodus. Pereopod 1
showing sexual dimorphism, carpus always distally produced into spinose
process.

Remarks.—The genera Mesanthura and Skuphonura are very similar in
the triangular underriding carpus of the posterior pereopods, in the S-seg-
mented maxilliped, and in the fusion of the pleonites. Two features, how-
ever, immediately separate them. The presence of a persistent pigment pat-
tern in Mesanthura (absent in Skuphonura), and the uniarticulate flagella
of both antennae and antennules in Skuphonura (Mesanthura has flagella
of more than one article).

Skuphonura laticeps Barnard
Fig. 5

Skuphonura laticeps Barnard, 1925:145, fig. 10.—Schultz, 1969:91, Figs.
115q, 119.

Description.—6. Integument not indurate. Cephalon with rounded an-
terolateral lobes extending beyond level of triangular rostrum, bearing dor-
solateral eyes each of 10 ocelli. Blunt rounded lobe at base of mouthparts.
Rropontions; ©2— i> 92 = 3 <4 —5 = 6 = 7. Pereonite 1 with strong
forwardly-directed ventral spinose process. Telson widest at midlength, api-
cally rounded, with broad transparent margin and several lateral and distal
setae. Antennular peduncle 4-segmented, basal segment almost as long as
3 distal segments together; 4th segment short, set obliquely into 3rd seg-
ment; flagellum of single short article bearing cluster of aesthetascs. Anten-
nal peduncle 5-segmented, distal segment equal in length to 3rd and 4th

732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Skuphonura laticeps: a, 3 in dorsal view; b, ¢ Cephalon and pereonite | in lateral
view; c, Pereopod 14; d, Maxilliped; e, Antenna; f, Pereopod 7; g, Antennule 6; h, Pleopod
23; i, Pleopod 1.

VOLUME 93, NUMBER 3 BS

segments together; flagellum article setose. Maxilliped 5-segmented, ter-
minal segment short, obliquely set on penultimate segment, bearing 6 setae;
4th segment twice length of 3rd. Pereopod 1 unguis about one-third length
of dactylus, latter with slight distal bulge at base of unguis; propodal palm
concave, with numerous slender spines distally, membranous convexity
proximally bearing 7 slender spines; carpus triangular, produced distally
into narrowly triangular process. Posterior pereopods with carpus short,
underriding propodus; propodus and dactylus bearing fringed scale-like
spines on posterior margin. Pleopod | exopod operculiform, endopod slight-
ly shorter and one-third width of exopod; basis with 4 retinaculae. Pleopod
2 endopod with straight apically acute appendix masculina, extending well
beyond rami. Uropodal exopod oval, with distinct distal notch, fringed with
setae; endopod almost reaching telsonic apex, distally rounded, bearing nu-
merous setae.

Type-material.—Copenhagen Museum, syntypes 3 6, St. Thomas, West
Indies, 8—40 m.

Remarks.—Barnard (1925) described this species from three males from
St. Thomas in the West Indies, and one female from Tobago. The latter,
however, has not been included in this discussion, as there is some doubt
that it is the same species. The telsonic shape and the dactylus of pereopod
1 differs from the males, while in the two new species described below, the
dactylus is similar in both species. The flagellum of the antennule in this
female has two articles, while in both the new species the flagella are uniar-
ticulate. The female characters are thus omitted from the description of this
species.

Skuphonura itapuca, new species
Figs. 6, 7

Description.—o. Bodysproportionss€ <1 => 2>3 <4 — 5 — 6 7.
Cephalon anteriorly slightly wider than posteriorly; eyes present; lateral
ridge well developed; medioventral toothlike process at base of mouthparts.

_ Pereonites 4, 5, and 6 with faint proximal groove or constriction, becoming
_ obsolete on pereonite 7 and pleon. Pleonites 1—S fused, segments indicated

_ ventrolaterally; pleonite 6 free, posterodorsal margin convex. Telson dis-
_ tally rounded, dorsally with slight proximal median ridge; 2 statocysts pres-
~ ent; ventrally flattened.

Antennular peduncle 3-segmented, basal segment equal in length to 2
distal segments. Antennal peduncle 5-segmented, 2nd segment longest and
widest, grooved to accommodate antennule. Mandibular palp 3-segmented,
terminal segment about half length of middle segment, bearing 3 distal setae;
incisor of 2 blunt cusps; molar of 2 rounded lobes, separated from incisor
by 9-serrated lamina dentata. Maxilla slender, with 6 distal spines. Maxilliped

734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by
W

Fig. 6. Skuphonura itapuca: a, Holotype in dorsal view; b, ¢ Cephalon; c, 2 Base of
mouthparts; d, Pleon in lateral view; e, Antenna; f, Pleopod 2d; g, Pleopod 1; h, Mandible;
i, Maxilla; j, Antennule 3; k, Antennule 9; 1, Maxilliped; m, Uropod.

VOLUME 93, NUMBER 3 135:

Fig. 7. Skuphonura itapuca: a, Pereopod 146; b, Pereopod 12; ¢, Pereopod 2; d, Pereopod 7.

5-segmented, outer margin of 3rd segment concave; distal segment short,
inserted at outer angle of 4th segment. Pereopod 1 unguis one-third length
of dactylus, with small spine at base; palm of propodus excavate, with 3
rounded tubercles on median edge, outer margin sinuous; carpus produced
ventrodistally into narrowly triangular lobe. Pereopod 2 unguis one-quarter
length of dactylus; propodus with short strong spine at ventrodistal corner;

736 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

carpus triangular, produced obliquely beneath propodus. Pereopods 4—7
dactylus with spinose scales; propodus cylindrical, with strong serrate spine
at ventrodistal corner, ventral surface bearing short spinose scales; carpus
triangular, underriding propodus, with strong ventrodistal spine; elongate
setae on all segments. Pleopod 1 exopod operculiform, barely indurate,
twice width and slightly longer than endopod; both rami with distal plumose
setae; basis with 5 retinaculae. Pleopod 2 endopod with stylet articulating
at about midlength of median margin, slightly longer than ramus, distally
broadened, apically acute; both rami with 6 distal plumose setae. Uropodal
exopod slightly longer than basis, with distal notch, margin lined with plu-
mose setae; endopod distally rounded, bearing simple setae.

2. Medioventral process posterior to mouthparts with 2 strong toothlike
spines. Antennular peduncle 4-segmented, segments 2 and 3 bearing elon-
gate ventral setae; segment 4 short. Pereopod | propodal palm with median
tooth and smaller distal tooth, proximal tooth not as developed as in 6;
carpus with ventrodistal triangular extension.

Material examined.—Nictheroy, Rio de Janeiro, Brasil, intertidal, collected
by Dr. Waldo Schmitt in 1925. USNM 173522, Holotype 6 TL 8.9mm. USNM
173523, Allotype 2 TL 9.3 mm. USNM 173524, Paratypes 9 °, 3 juveniles.

Etymology.—The specific name derives from the name of the beach at
which the specimens were collected.

Skuphonura ecuadorensis, new species
Figs. 8, 9

Description.—6 . Integument indurate, brittle. Body proportions: C < 1
>2>3<4=5>6>7. Cephalon broader distally than proximally, with
anterolateral corners lobe-like and rounded, extending well beyond rostrum,
bearing eyes of 6 or 7 small ocelli; lateral margin of cephalon forming a
rounded ridge; small medioventral rounded tubercle posterior to mouth-
parts. Pereonites 4—6 with faint proximal constriction. Pleonites 1—5 fused,
segments indicated ventrolaterally by short slits; pleonite 6 free, with me-
diodorsal slit in posterior margin. Telson lanceolate, ventrally flattened,
dorsally with proximal area raised to rounded median ridge, spreading from
about midlength to broad area sloping to rounded apex.

Antennular peduncle 3-segmented; flagellum of single article bearing clus-
ter of filiform aesthetascs. Antennal peduncle 5-segmented, second segment
longest and broadest, grooved to accommodate antennule; flagellum of sin-
gle setose article. Mandibular palp 3-segmented, terminal segment one-third
length of 2nd, with 3 distal setae; Ist and 2nd segments with elongate setae;
incisor of 2 rounded cusps; molar of 3 rounded teeth; lamina dentata with 10—
11 serrations. Maxilla slender, with 6 distal spines. Maxilliped 5-segmented,
terminal segment situated at outer distal corner of 4th segment, 3rd segment

VOLUME 93, NUMBER 3 137)

Fig. 8. Skuphonura ecuadorensis: a, Holotype in dorsal view; b, Cephalon 6; ¢c, 2 Base
of mouthparts; d, Pleon in lateral view; e, Mandible; f, Pleopod 246; g, Antennule 2; h, Max-
illiped; i, Maxilla; j, Antenna; k, Antennule 6; 1, Pleopod 1.

with concave outer margin. Pereopod 1 unguis one-third length of dactylus,
with small supplementary spine and slight proximal concavity on ventral
margin; propodal palm concave, with small distal and large proximal tooth-
like tubercle; carpus distally narrowed into blunt tubercle. Pereopod 2 un-

738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Skuphonura ecuadorensis: a, Pereopod 1¢; b, Pereopod 12; ce, Pereopod 2; d,
Pereopod 7.

guis one-quarter length of dactylus; propodus with several ventral setae,
strong spine at ventrodistal corner, serrate on one edge; carpus triangular,
with sensory spine distally; merus and ischium bearing elongate setae. Per-
eopods 5-7 dactylus bearing short fringed scales; propodus cylindrical, with
ventrodistal spine, serrated on one margin, and 2 spines serrated on both

VOLUME 93, NUMBER 3 739

margins; carpus short, underriding propodus; all segments with several elon-
gate setae. Pleopod | exopod operculiform; endopod shorter than and slight-
ly less than half width of exopod; both rami bearing distal plumose setae;
basis with 4 retinaculae. Pleopod 2 copulatory stylet on endopod extending
beyond rami, distally acute. Uropodal exopod with distal notch, fringed
with plumose setae, not quite reaching endopod.

2. Medioventral process posterior to mouthparts consisting of 2 tuber-
cles. Antennular peduncle 4-segmented, 4th segment very short; flagellum
of single article bearing 2 aesthetascs and few simple setae. Pereopod | with
2 toothlike tubercles on distal half of propodal palm; triangular carpus pro-
duced distally into toothlike projection.

Material examined.—La Libertad, Ecuador, intertidal. USNM 173525,
Holotype 6 TL 7.2 mm. USNM 173526, Allotype 2 TL 7.2 mm. USNM 173527,
Paratypes 2 TL 7.2 mm, 2 juveniles.

Etymology.—tThe specific name derives from the country of origin of the
species, viz. Ecuador.

Remarks.—Skuphonura itapuca and S. ecuadorensis seem to be closely
related, judging from the many structural similarities, including the midven-
tral cephalon process of the female, the antennules, antennae, mouthparts,
pereopods 2-7, pleopods, and uropods. The main differences lie in the first
pereopods of both the male and female, which show subtle differences in
the size and placing of the toothlike tubercles of the propodal palm and
carpus, the midventral cephalon process in the male, and the outline of the
cephalon seen in dorsal view. S. ecuadorensis is markedly broader distally
and the anterolateral lobes are more produced than in S. itapuca. The re-
lationship between the 3 species of Skuphonura is difficult to assess. It is
possible that a single species was distributed over the Caribbean-Central
American region before the final emergence of the isthmus of Panama. Sub-
sequent geological and hydrological events could have led to the isolation
of 3 populations, giving rise to the present species.

The major differences between the 3 species are summarized in the fol-
lowing table.

S. laticeps S. itapuca S. ecuadorensis
Pereopod 16 Carpus well pro- Carpus well pro- Carpus slightly
duced duced produced
Propodus with 3 short propodal Propodus with 1
low convexity spines large and 1 small
spine
Pereopod 1° — Carpus slightly | Carpus well pro-
produced duced
— Propodus with Propodus with 2
large proxi- distal spines

mal and small
distal spine

740 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘ ® alg ALAC ST f vi 4 =

Fig. 10. Xenanthura brevitelson: a, Mouthparts in situ; b, Antennule showing robust sen-
sory seta; c, Reduced mandibular palp; d, Pereopod 1.

Midventral 6 single, blunt ¢ single, conical <o single, blunt
cephalon pro-
cess g a 2 2 conical 2 1 conical spine,
spines 1 blunt tubercle
Pereonite | Strong medio- Spine lacking Spine lacking

ventral spine

Xenanthura brevitelson Barnard
Fig. 10

Xenanthura brevitelson Barnard, 1925:138, fig. 8.—Nierstrasz, 1941:240.—
Pillai, 1963:263.—Menzies and Frankenberg, 1966:38, fig. 15.—Schultz,
1969:92, fig. 122.—Camp, Whiting, and Martin, 1977:16.

VOLUME 93, NUMBER 3 741

Previous records.—St. Thomas, West Indies, 50-60 m; Off Georgia, 20-
145 m; Off Florida, 8-10 m.

Type-material.—The type-material of X. brevitelson from the Copenha-
gen Museum, collected by T. Mortensen, consists of three vials, each la-
belled ‘Type’. One vial has a pencilled note in K. H. Barnard’s autograph,
a second has an inked label also in Barnard’s autograph. North East of Hans
Lollik (North of St. Thomas) 8 March 1906, Syntypes, 3 d, 1 sub 6, 24
2, 5 juveniles. North East of Hans Lollik (North of St. Thomas) 50-60 m,
8 March 1906, Syntypes, 1 sub ¢,8 2, 6juveniles. St. Thomas, West Indies,
50-60 m, Syntypes 3 ¢ 3 @.

Remarks.—Figure 10b shows what Barnard (1925) mistook for a second
flagellum on the antennule to be a large sensory seta, as was suggested by
Menzies and Frankenberg (1966).

The presence amongst the syntypes of two submales with elongate an-

_ tennules lacking aesthetascs along with fully mature males, suggests that X.

brevitelson is another anthurid which displays protogyny, and that at least
two molts are involved in the change from female to male.

Acknowledgments

My sincere thanks are due to Mr. H. Kaufman for the Panama material,
Mr. J. Vogel for the Apanthura magnifica material from Florida, Dr. Torben
Wolff of the Zoological Museum, Copenhagen, for the loan of the Sku-
phonura and Xenanthura material, and especially to Dr. T. E. Bowman of
the Smithsonian Institution, for reading the manuscript and for his valuable
criticisms. I am grateful to Susann G. Braden and Mary-Jacque Mann of the
Smithsonian Scanning Electron Microscope Laboratory, and Mr. M. Car-

| penter of the Department of Invertebrate Zoology, for the preparation of

the micrographs used here.

Literature Cited

Barnard, K. H. 1925. A revision of the family Anthuridae (Crustacea Isopoda) with remarks
on certain morphological peculiarities —Journal of the Linnaean Society, London (Zo-
ology) 36:109-160.

Camp, D. K., N. H. Whiting, and R. E. Martin. 1977. Nearshore marine ecology at Hutch-
inson Island, Florida: 1971-1974. V. Arthropods.—Florida Marine Research Publica-
tions 25:1-63.

Kruczynski, W. L., and G. J. Myers. 1976. Occurrence of Apanthura magnifica Menzies and
Frankenberg, 1966 (Isopoda Anthuridae), from the west coast of Florida, with a key to
the species of Apanthura Stebbing, 1900.—Proceedings of the Biological Society of
Washington 89:353-360.

Menzies, R. J., and D. Frankenberg. 1966. Handbook on the common marine isopod Crus-
tacea of Georgia.—University of Georgia Press; Athens, Georgia, vili + 93 pp.
Nierstrasz, H. F. 1941. Die Isopoden der Siboga-Expedition IV. Isopoda Genuina III. Gna-
thiidea, Anthuridea, Valvifera, Asellota, Phreatocoidea.—Siboga Expedition monograph

32d:235-306.

742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Paul, A. Z., and R. J. Menzies. 1971. Sub-tidal isopods of the Fosa de Cariaco, Venezuela,
with descriptions of two new genera and twelve new species.—Boletin del Instituto
Oceanografico del Universidade Oriente 10:29—48.

Pillai, N. K. 1963. Observations on the genus Xenanthura (Isopoda, Anthuridae).—Crusta-
ceana 5:263-—270.

Poore, G. C. B. 1975. Australian species of Haliophasma (Crustacea: Isopoda: Anthuridae).—
Records of the Australian Museum 29:503-—533.

Schultz, G. A. 1969. How to know the marine isopod crustaceans.—W. C. Brown, Dubuque,
Iowa, vii + 359 pp.

Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 743-756

ON THE CARAPACE SHAPE OF SOME EUROPEAN
FRESHWATER INTERSTITIAL CANDONINAE
(OSTRACODA)

Dan L. Danielopol

Abstract.—The interstitial Candoninae have a large number of species
with trapezoidal triangular and widely arched carapace shapes. Some of the
interstitial Candoninae display carapaces with extreme morphological fea-
tures, e.g. a very elongate shape with a pointed posterior, a triangular shape
with a large dorsal protuberance.

It is suggested that such species, as Mixtacandona elegans Danielopol &
Cvetkov and Phreatocandona motasi Danielopol, with elongate carapaces
and pointed posterior should have a low fecundity. The Candoninae species
with this extreme type of carapace shape are known also from the sublittoral
of Lake Baikal (‘‘Candona’’ fossiliformis Mazepova) and from the eastern
Paratethys during the Pliocene (Pontoniella schemachensis Mandelstam).
It is hypothesized that the elongate carapace shape with a pointed posterior
is an adaptation less suited for Candoninae living in unpredictable surface
habitats but acceptable for those species inhabiting physically and/or bio-
logically stable habitats which exist over geological epochs.

The Freshwater Interstitial Candoninae of Europe

The subfamily Candoninae (sensu Danielopol, 1976, 1978) contains a high
number of true interstitial species living in freshwater habitats in Europe
(Danielopol, 1977). This subfamily is also one of the most ‘‘species-rich’’
ostracod groups living in European surface waters (Loffler & Danielopol,
1978). The Candoninae are creeping forms with unpigmented carapace and
OWE.

I consider “‘true interstitital’’ ostracods to be species found in interstitial
or in interstitial and cavernous aquatic habitats, which show eye reduction
and/or long antennal aesthetascs ‘‘Y’’ (see Danielopol, 1973). In cases where
data on these characteristics is not available, species recorded exclusively
from the above mentioned habitats are referred to as “‘true interstitial’’
forms (Table 1). Several species are not included in this category because
reliable data about their typical habitat is lacking (see Danielopol, 1978).

The carapace shapes of the interstitial Candoninae can be roughly clas-
sified as follows: a) almost rectangular; b) elongate, slightly triangular; c)
widely arched dorsally; d) ‘“‘highly’’ triangular; e) trapezoidal; f) elongate
with pointed posterior. Table 1 presents data available for the interstitial

744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—The freshwater interstitial Candoninae recorded in Europe (i—interstitial habitat;
k—karstic habitat; a—almost rectangular; b—elongate, slightly triangular; c—widely arched
dorsally; d—‘‘highly’’ triangular; e—trapezoidal; f—elongate with pointed posterior). Data
from Danielopol, 1978; Danielopol & Cvetkov, 1979; Sywula, 1976.

Taxa Habitat Shape Height
Pseudocandona
eremita (Vejdovsky) s.1. hae IS d >1/2
zschokkei (Wolf) i e >1/2
puteana (Klie) d S12
schellenbergi (Klie) 1 e >1/2
triquetra lap Ik d > II
belgica (Klie) 1 e SD,
hertzogi (Klie) 1 e > 1/2
brisiaca (Klie) 1 € 2
insueta (Klie) 1 e <1/2
bilobata i C >1/2
spelaea (Klie) i+ k a 1/2
dichtliae (Brehm) I e S17
szocsi (Farkas) 1 d WP
bilobatoides (LOoffler) i € <I/2
pseudoparallela (Loffler) 1 a 1/2
profundicola (Loffler) 1 a 1/2
tyrolensis (LOffler) 1 a 1/2
altoalpina (LOffler) 1 a 1/2
rouchi Danielopol sb IK e > i/2
delamarei Danielopol i a 1/2
Ps. n. sp. aff. eremita 1 d > 1/2
mira Sywula l e <1/2
Cryptocandona
Dhraeticola (Klie) 1 b <M
leruthi (Klie) 1 b <<a)
kieferi (Klie) 1 b <<a
juvavi (Brehm) 1 b? NYY
matris Sywula i b >1/2
Fabaeformiscandona
latens (Klie) 1 a SZ
wegelini Petkovski 1 a <1/2
Candonopsis
boui Danielopol i b < Il
Phreatocandona
motasi Danielopol 1 f <a
Mixtacandona
laisi (Klie) I e <ah/2
stammeri (Klie) i+k c <7
chappuisi (Klie) 1 e <Il/2
transleithanica (LOffler) i a ea2

VOLUME 93, NUMBER 3 745

Table 1.—Continued.

Taxa Habitat Shape Height
loffleri Danielopol i S >1/2
botosaneanui Danielopol i e <1/2
pietrosanii Danielopol & Cvetkov 1 b <2
elegans Danielopol & Cvetkov 1 f <1/2
Mixtacandona n. sp. i d SZ
Mixtacandona sp. aff. chappuisi 1 e <1/2
pseudocrenulata Schaffer i+ k? e <EIli/72
cottarellii n. sp. (in litt.) 1 e <1/2
jJuberthieae Danielopol 1 e <2.

Nannocandona
stygia Sywula i S <1/2

Candoninae. Similar carapace shapes also exist in the surface-dwelling Can-
doninae (review in Danielopol, 1978).

Hartmann (1973) showed that among the marine creeping interstitial os-
tracods (mainly Cytheracea) the elongate carapace (height less than 12 the
length) is most commonly found. From Table 1 one can see that 41% of the
freshwater interstitial species have carapaces with maximal height of more
than %2 the carapace length. One should notice too that the carapace shapes
of many species are trapezoidal, “‘highly’’ triangular and dorsally widely
arched. Some of the European interstitial Candoninae have carapaces with
extreme morphological features. Mixtacandona elegans Danielopol & Cvet-
kov (Fig. 1A, B) from southeastern Bulgaria has the most dorsoventrally
depressed and elongate carapace of all the Candoninae (Danielopol & Cvet-
kov, 1979). Two other subterranean Candoninae display similar carapace
shapes, 1.e. Mixtacandona ljovuschkini (Rudjakov) from a cave in Trans-
caucasia (Rudjakov, 1963) and Phreatocandona motasi Danielopol (Fig. 1C,
D) from the southern Carpathians in Romania (Danielopol, 1980). Mixta-
candona n.sp. (Fig. 1F) lives in the same habitat and area as M. elegans.
This species has the largest dorsal protuberance occurring in the Candoni-
nae, 1.e. 4 the total height (Danielopol & Cvetkov, 1979). The only epigean
Candoninae I[ could find with similar extreme morphological characteristics
is ““Candona’’ fossiliformis Mazepova (Fig. 1E), a species which lives in
the sublittoral of Lake Baikal between 20 and 50 m depth (Mazepova, 1970;
Loffler, pers. comm.). This species has an elongate carapace with a pointed
posterior. A similar species has been recorded by Mandelstam et al. (1962)
and Vekua (1975) from the Pliocene deposits of the eastern Paratethys in
the southwest of the Soviet Union (1.e. Pontoniella schemaschensis Man-
delstam, Fig. 3B). This species lived in a shallow Caspi-brackish environ-

746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. A,B, Mixtacandona elegans, 3 and ¢ carapaces (Zvetdetz); C, D, Phreatocandona
motasi, 2, (Jiblea); C, left valve; D, Carapace, dorsal view; E, ‘“Candona’”’ fossiliformis (Lake
Baikal); F, Mixtacandona n. sp., °., (Zvetdetz); G, Pseudocandona dorsoconcava, 2? (Lake
Baikal).

ment. The specimen of Pseuwdocandona dorsoconcava (Bronstein) figured
here (Fig. 1G) was found in Lake Baikal at 20 m depth. It has a very
conspicuous dorsal protuberance (‘/, of the total height located in the an-
terior half of the carapace). No other surface Candoninae has been recorded
with such a high dorsal protuberance.

Mazepova (1970) suggested that ‘“Candona’’ fossiliformis from Lake Bai-
kal has phylogenetic affinities with the fossil species belonging to the Can-
dona of the group lobata from the Paratethys and with the Recent ‘‘ Can-
dona’ ljovuschkini from Transcaucasia. Danielopol & Cvetkov (1979)
showed that ‘‘Candona”’ ljovuschkini belongs to the Candona group mixta,
now considered a valid genus, Mixtacandona Klie (Danielopol, 1977/1978).
This species has the 2nd thoracic leg with 2 short and 1 long distal setae,
the clasping palps of the maxilla slightly asymmetric and with digitiform
tips, characteristics that can be found in all the Mixtacandona species.
Mixtacandona ljovuschkini has close affinities with Mixtacandona elegans.
Phreatocandona motasi Danielopol has 3 short setae on the exopodite of
the antenna, a long distal seta on the Ist thoracopod; the 2nd thoracopod
is similar to those of Psewdocandona, having distally 3 long setae. In these

VOLUME 93, NUMBER 3 747

Fig. 2. Epigean (E) and hypogean (Hy) Candoninae: A, B, Pseudocandona pratensis, °,
(E), (Dourdou); C, Pseudocandona aff. pratensis, 3, (E). (Seewinkel, temporary pond nr. 12);
D, Pseudocandona n. sp. aff. eremita, 3, (Hy), (Jiblea); E, F, Mixtacandona aff. chappuisi,
2, 6, (Hy), (Bogdan Voda) (ov—ovocytes, zo—Zenker’s organ, He—hemipenis, te—testes).

peculiarities Phreatocandona differs from other genera of Candoninae.
‘““Candona’’ fossiliformis has the antennae, the clasping palps of the maxilla
and the thoracopods similar to those of Pseudocandona species. These data
suggest that the elongate carapace shape with an extremely pointed poste-
rior, occurring in species found in both ground water (in Europe) and in
Lake Baikal, was produced by parallel evolution in related phylogenetic
lineages.

The Functional Importance of the Carapace Shape
in the Candoninae

Why within the Candoninae does this characteristic carapace shape, very
elongate and pointed, occur only in species which live in subterranean hab-
itats, in Lake Baikal and in the Paratethys? To answer this question one has
to understand, first of all, the functional importance of the carapace. Sec-
ondly one has to find what such different environments as the groundwater
(in southeastern Europe), the eastern Paratethys Sea (e.g. during the Plio-
cene) and the sublittoral of Lake Baikal have in common.

748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Throughout the evolution of the Candoninae the different carapace shapes
developed several times in the different phylogenetic lineages (Danielopol,
1978). Here I showed the example of the elongate carapace with pointed
posterior. There is evidence to show that the almost rectangular, the slightly
triangular, the trapezoidal shaped carapaces allow for the development of
large internal organs: e.g. in the female the oviducts, which store commonly
large numbers of ovocytes; and in the male the seminal vesicles, the
Zenker’s organ with long spikes and the hemipenes, with well developed
internal and external processes. Figure 2 shows such examples. The surface
dwelling Candoninae with more or less rectangular carapaces like Candona
dancaui Danielopol, Candona angulata G. W. Muller, Pseudocandona pra-
tensis (Hartwig), Pseudocandona albicans (Brady), Pseudocandona mar-
chica (Hartwig), display high fecundity (Danielopol, 1980). During the be-
ginning of the reproductive period the females have the ability to store the
ovocytes in the oviducts (Fig. 2A). One can find in some species like Ps.
pratensis up to 20 ovocytes on each side stored in the lateral and the dorsal
parts of the body. In the parthenogenetic species like Ps. albicans (Fig. 3A)
the eggs are laid abruptly without fertilization. In the Candoninae the surface
area and also the volume of the central and posterior parts of the carapace
show marked interspecific differences. McGregor & Kesling (1969) com-
puted the surface area in lateral view of a large number of Candoninae. To
permit a comparison they expressed the data in terms of converted areas of
the carapace using the following formula

CA

CaS (LA)?

x 100

where Ca“ is the converted area, CA is the area of the carapace in lateral
view, L“ is the maximal length of the carapace. Table 2 (part A) shows the
converted areas of several epigean Candoninae (females) living in shallow
water bodies. The second part (B) of the table shows the values for the 4
species with elongate carapace shape and pointed posterior. The latter group
has the lowest values. Kesling (1965) and McGregor & Kesling (1969) point-
ed out that the interspecific differences in the converted areas are due to
differences in the shape of the posterior part of the carapace which holds
the reproductive organs. Table 3 supports this idea. It shows the converted
area of the anterior 4 and of the posterior 24 of the carapace (see also Fig.
3B), calculated by the following:
A P
Ca — TALE x 100 and Ca?’ = CALA x 100

Ca“ and Ca? are the converted areas respectively of the anterior % and the
posterior % of the carapace, A and P are areas of the respectively anterior

VOLUME 93, NUMBER 3 749

A
Nr | eggs-]]
OVeGy teS-|1-side) _O-
{2
10 5 5
‘\ on TN"
ps Q- Y
g Ot Ps O
6 a ee
4 o a C ||
(OF. n 5 ie)
2 ccoarare
, \e WAIT
A OeEC OMI 149slo BIC —20 22 2526 8 30
B Days
2/3 p 113A

— eo

ng

Fig. 3. A, Pseudocandona albicans, 2 , the evolution of the ovocyte number in the oviduct
and the egg laying rhythm (e—the last molt); B, Pontoniella schemachensis, \eft valve (after
Vekua, 1975).

4 and the posterior *%4 carapace in lateral view. L“™ is the maximal length
of the carapace. Whereas the converted area of the anterior part varies
within a narrow range, that of the posterior % varies to a larger extent. All
the epigean Candoninae listed here have almost rectangular, elongate, slight-
ly triangular, ‘“‘highly’’ triangular and trapezoidal carapaces. Most of these
species are widely arched posteriorly. The male copulatory organ as well
as the clasping organs are normally held in the inner carapace space, i.e.
within the space limited by the inner margin of the carapace (Fig. 2B, D,
F).

A comparative study has been made in the following 21 Candoninae
species of the area of the inner carapace space (ICA), excluding the calcar-
eous lamellar area, and of the lateral surface area of the hemipenis (HeA)
(Fig. 4A): 1. Mixtacandona elegans (Hy), 2. M. laisi vindobonensis (Hy),
3. M. pietrosani (Hy), 4. Mixtacandona sp. aff. chappuisi (Hy), 5. M.
loffleri (Hy), 6. M. cottarellii n. sp. (Hy), 7. M. tabacarui (Hy), 8. Can-

750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Converted areas of equivalent carapaces (lateral view) in female Candoninae. Data
compilation from Danielopol, 1978; Danielopol & Cvetkov, 1979; Delorme, 1970; Furtos, 1933;
Hoff, 1944; Klie, 1938; Mazepova, 1970; McGregor & Kesling, 1969; Vekua, 1975. (Ba—Lake
Baikal; Eu—Europe; H—Holarctic realm; NA—North America; P—Palearctic realm; d—ditch-
es; gw—interstitial groundwater; ll—littoral lakes; m—marshes; pp—permanent ponds; rp—
rock pools; se—shallow channel; tp—temporary ponds; tr—temporary running water; s—sub-

littoral; s—shallow lakes).
Converted

Taxa area (%) Distribution
A.
Candona miilleri Hartwig 63.2 Eu—tp
“C.”’ renoensis Gut. & Bens. 48.8 NA—tp
C. weltneri Hartw. 46.6 Eu—ll,tp
C. candida (O.F.M.) 44.4 H—ll,pp,tp,d
“C.”’ neglecta Sars 43.7 P—d,m,ll,pp,tp,tr
“C.”” truncata Furt. 43.6 NA—m, pp,tp
Pseudocandona sucki (Hartw.) 42.4 Eu—tp
C. angulata G. W. Mull. 42.5 Eu—ll,tp
“C.”” distincta Furt. 41.6 NA—m,sc,sl
Fabaeformiscandona hyalina (Br. & Rob.) 40.8 Eu—m,ll,sl
“C.’’ scopulosa Furt. 39.4 NA—m,rp
“C.”” eriensis Furt. 39.3 NA—Il,rp
“C.”’ acuta Hoff. 39.3 NA—tr
F. fabaeformis (Fisch.) 38.3 P—ll,tp
F. acuminata (Fisch.) 37.6 H—Il,tp
F. fragilis (Hartw.) 36.0 Eu—ll,tp
“C.”’ ohioensis Furt. 33.8 NA—II
B.
Phreatocandona motasi Dan. 27.0 Eu—gw
“C.”” fossiliformis Mazep. Do) Ba—s
Mixtacandona elegans Dan. & Cvetk. 22.9 Eu—gw
Pontoniella schemachensis Mand. 20.1 Paratethys

donopsis kingsleii (E), 9. C. boui (Hy), 10. Pseudocandona compressa (E),
11. Ps. simililampadis (Hy), 12. Pseudocandona n. sp. aff. eremita (Hy),
13. Ps. pratensis (E), 14. Ps. rouchi (Hy), 15. Ps. delamarei (Hy), 16. Ps.
brevipes (E), 17. Fabaeformiscandona fabaeformis (E), 18. Candona welt-
neri (E), 19. C. dancaui (E), 20. C. fasciolata (E), 21. C. neglecta (E).
The correlation between the converted areas of the inner carapace space
(Ca'“) and the lateral area of the hemipenis (Ca#*4) have been computed
(Fig. 4A). There is a significant correlation (P < 0.05) but the coefficient of
determination (r?) is only 0.253. This means that only 25% of the variation
in one of the 2 variables is explained by the other variable, therefore there
is not a close relationship. It is suspected that the large hemipenis processes
improve the success of the copulation due to a better sensorial contact with

VOLUME 93, NUMBER 3 751

Table 3.—Converted areas of the anterior (Ca“) and posterior carapace (Ca¥) sides in several
epigean (E) and hypogean (Hy) Candoninae species (see also explanation in Table 2).

Taxa Ca“ CaP Distribution
Pseudocandona
Ps. albicans (Br.), ° Wi 89 H—(B),lI,m,rp,tp,tr
Ps. compressa (Koch), ° 77 93 H—(B),Il,tp
Ps. n. sp. aff. eremita, 3 76 88 Eu—(Hy),gw
Fabaeformiscandona
F. fabaeformis (Fischer), @ 78 118 P—(E),ll,tp
Candonopsis
C. kingsleii Br. & Rob., @ 78 106 H—(E),lIl,sc,sl
Mixtacandona
M. aff. chappuisi, 3 76 87 Eu—(Hy),gw
M. pietrosanii Dan. & Cvetk., ° 73 91 Eu—(Hy),gw
M. tabacarui Dan. & Cvetk., 3 76 86 Eu—(Hy),gw
M. elegans Dan. & Cvetk., 2 78 32 Eu—(Hy),gw
re) 76 372 Eu—(Hy),gw
Phreatocandona
Ph. motasi Dan., 2 84 38 Eu—(Hy),gw
‘““Candona”’
“C.”’ fossiliformis Mazep., @ 83 38 B—s

Pontoniella
P. schemachensis Mand. 76 32 Paratethys

the genital lobes of the female (Danielopol, 1980). I also noticed marked
differences in the development of Zenker’s organ of several Candoninae
belonging to the genera Mixtacandona, Candona, Pseudocandona, Fabae-
formiscandona and Candonopsis, which have high carapace shapes (height
more than 2 the length). These species also have large Zenker organs.
However, I did not find any significant correlation between the width of
Zenker’s organ and the maximum height of the inner carapace space (Fig.
4B).

Relationship between the Carapace Shape and the Environment

Most of the epigean Candoninae listed in Tables 2 and 3 are species with almost
rectangular, elongate, slightly triangular carapace shapes. These species are
widespread in the Holarctic realm. They live commonly in shallow water
bodies which are strongly dependent on the climatic changes of the atmo-
sphere (Klie, 1938; Loffler & Danielopol, 1978; Bronstein, 1947; Delorme,
1970; Furtos, 1933; Hoff, 1942). Such habitats are physically unstable and

752 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Carapace

MIXTACANDONA SP.

Ca ICA PSEUDOCANDONA ROUCH DAN,

ICA
CalfA =[reaye100

f y=0251x+33,59

r =0,503°
n=21

@ HYPOGEAN SP.

© EPIGEAN SP.

Caen fee % 100

60

20 40 60 7;

20 40

CaltA

Fig. 4. A, Regression of area of inner carapace space upon lateral area of the hemipenis,
with their statistical correlation (see also text); B, Relationship between the width of Zenker’s
organ and the maximum height of the inner space. (The numbers refer to the 21 species men-
tioned in the text.)

unpredictable. Some of them are short-lived habitats such as temporary
ponds, ditches, rock pools, and temporary streams. A great success in re-
production and a high fecundity are at a premium for animals living in
unstable water bodies (Southwood, 1976). Comparing the shape of the an-
terior and posterior part of the carapace of Mixtacandona elegans, Phrea-
tocandona motasi, “‘Candona’’ fossiliformis and Pontoniella schemachen-
sis (Table 3, Fig. 1A, B, C, E), one will notice that the anterior 4% is highly
similar to the other Candoninae quoted above whereas the posterior part is
very narrow. These morphological features suggest that the fecundity of
these species is low. Further field and laboratory evidence should confirm
this prediction.

Mixtacandona elegans Dan. & Cvetk. was found in a well from Zvezdets
in southeastern Bulgaria near the border with Turkey (Danielopol & Cvet-
kov, 1979). Phreatocandona motasi was found in a well from Jiblea, near
Calimanesti, in the Olt Valley in the southern Carpathians, in Romania

VOLUME 93, NUMBER 3 753

(Danielopol, 1979a). Whereas the short-term temperature fluctuations of the
atmosphere in Europe cause thermic fluctuations in the shallow surface
water bodies, they have only slight impact on the groundwater. For in-
stance, Kriz (1973) showed that in southern Slovakia the maximum
amplitude of the temperature in a well supplied by a shallow aquifer between
1933 and 1970 was only 0.2°C. Moreover, whereas high temperature or
severe droughts can easily lead to the desiccation of restricted shallow water
bodies above ground, they will affect a groundwater aquifer to a lesser
extent, only by modifying its boundaries. These arguments suggested to me
that the groundwater habitats of Mixtacandona elegans and Phreatocan-
dona motasi are physically more stable compared to the epigean shallow
water bodies. If one considers the geographical position of the localities
where M. elegans and Ph. motasi occur one will notice that they are situ-
ated on or near old land masses which have existed at least since the be-
ginning of the Neogene (for Bulgaria data see Cvetkov, 1975 and Luttig &
Stephens, 1976; for Romania see maps in Decou & Negrea, 1969). This
suggests that the aquifers in the southern Carpathians and in southeastern
Bulgaria could have a very long span (millions of years). Lake Baikal is one
of the oldest lakes; it has existed at least since the beginning of the Neogene,
and according to Kozhov (1963) may have existed since the Oligocene. ‘The
emergence of Baikal as a deep lake with borders approximately of their
present shape is dated by Dumitrashko to the end of the Pliocene... .”’
(Kozhov, 1963:270). The sublittoral of Lake Baikal is also a stable environ-
ment highly similar to the groundwater habitats. Kozhov (1963:194) mentioned:
‘The water of the bottom layer in the sublittoral is subjected to much weak-
er seasonal temperature flucuatuions than the littoral. At a depth of 50 m
the annual amplitude of temperature fluctuations does not exceed 5-6°C.
The influence of turbulence is practically imperceptible. Light intensity is
insufficient for the development of macrophytes... .’’ In the sample
received from Prof. Loffler, collected at 20 m depth in the Baikal, I
found besides ‘‘Candona’’ fossiliformis 8 other endemic species of
Candoninae, 4 endemic Cytherissa species and a remarkable Cytherid.
I could not find any cosmopolitan Siberian ostracod species. This is in
keeping with the data reported by Bronstein (1939, 1947) and suggests
that in the sublittoral of Lake Baikal the ostracod association is a highly
biologically accommodated community where each species has a narrow
niche.

During the Pliocene in the central and eastern Paratethys a diverse en-
demic Candoninae fauna developed (Hanganu & Papaianopol, 1976; Krstic,
1972; Marinescu & Olteanu, 1973/1974; Sokac, 1972; Vekua, 1975, etc.)
which recall those of Lake Baikal. From the data presented by the above
mentioned authors it seems that there are few cosmopolitan Candoninae
dwellers in the Paratethys. This suggests that the Pliocene ostracods from

754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the Paratethys were also species with good competitive possibilities. The
endemic Caspi-brackish fauna prospered for several millions of years. Vek-
ua (1975) showed that Pontoniella schemachensis, a species with elongate
carapace and pointed posterior, lived during the Pontian, Klimmerian and
Kuyalnikian in the eastern Paratethys. That means that the species was
present there for more than 2.5 million years (using the paleomagnetic time
scale proposed by Kochegura & Zubakov, 1978).

Conclusion

From these data I hypothesize that within the group of Candoninae the
elongate carapace shape with a pointed posterior is less suited for wide-
spread species living in unstable surface habitats, but acceptable for species
living in physically and/or biologically stable habitats which exist over geo-
logical epochs.

Acknowledgments

Prof. H. Loffler (Vienna) offered the ostracods from Lake Baikal used in
this paper. Dr. F. Schiemer (Vienna) and Dr. D. Keyser (Hamburg) criti-
cized parts of this paper presented as a poster lecture at the 3rd International
Meiofauna Conference in Hamburg, 1977. Mr. A. Gunatilaka (Vienna), Dr.
H. J. Oertli (Pau) and an unknown colleague reviewed the manuscript. Con-
cerning the Paratethys Candoninae I am much indebted for discussions and/
or material to Dr. R. Benson (Washington D.C.), Dr. E. Hanganu (Bucha-
rest), Dr. M. Krstic (Belgrade), Dr. R. Olteanu (Bucharest) and Dr. M.
Stancheva (Sofia). Dr. H. J. Oertli (Pau), Dr. M. Gramm (Vladivostock),
Dr. I. Neustrueva (Leningrad), Dr. M. Vekua (Tbilisi) and Dr. N. Grekoff
(Paris) helped with bibliographical information.

The C.N.R.N.S. (France) supported my investigations on the biology of
surface and hypogean Candoninae in 1972-1973 at the Laboratoire Souter-
rain, in Moulis.

Addendum

After sending the manuscript to the publisher, Dr. L. Ram (Zoological
Survey of India, Patna) sent me the description of a new Candoninae with
elongate carapace and pointed posterior. The Indian species lives in ground-
water from Bihar.

Literature Cited

Bronstein, Z. S. 1939. On the origin of the ostracod fauna in Lake Baikal.—C. R. Acad. Sc.
USSR 15(4):333-337.

. 1947. Ostracoda presnih vod.—Fauna USSR 31(1):1-334, Izd. Akad. Nauk SSSR,
Moskva.

VOLUME 93, NUMBER 3 TD

Cvetkov, L. 1975. Composition, position zoogéographique et origine des Crustacés thalas-
sophréatiques en Bulgarie.—Hydrobiology (Sofia) 1:35—53 (in Bulgarian, French sum-
mary).

Danielopol, D. L. 1973. Sur la morphologie des aesthetascs chez quelques Ostracodes hypogés

de la sous-famille des Candoninae (Cyprididae, Podocopida).—Ann. Spéléol. 28:233-

245.

1976. The superfamily Cypridacea: Some remarks on the phylogenetical affinities
between the main ostracod Cypridacean groups.—Abh. Verh. Naturwiss. Ver. Hamburg
(NF) 18/19(Suppl.):77-85.

. 1977. On the origin and the diversity of European freshwater interstitia! ostracods.

In Loffler, H. & D. L. Danielopol, eds.—Aspects of Ecology and Zoogeography of

Recent and Fossil Ostracoda: 295-305, Junk Publ., The Hague.

. 1977/1978. Ostracodes hypogés du Sud de la France. 1. Mixtacandona juberthieae n.

sp.—Int. J. Speleol. 9:235-249.

. 1978. Uber Herkunft und Morphologie der siiwasserhypogaischen Candoninae (Crus-

tacea, Ostracoda).—Sitz. Ber. Osterr. Akad. Wiss., Math.-Nat. KI. Abt. I, 187(1-5):1-

162.

. 1980. Three new groundwater Candoninae (Ostracoda) from Romania.—Int. J. Spe-

leol. 10 (in press).

. 1980. Sur la biologie de quelques Ostracodes Candoninae épigés et hypogés d’ Europe.

Bull. Mus. Natn. Hist. nat. Paris. 4th ser. 1, Sect. A, 2 (in press).

, and L. Cvetkov. 1979. Trois nouvelles espéces du genre Mixtacandona (Ostracoda,
Cyprididae, Candoninae).—Hydrobiologia 67(3):249-266.

Decou, V. Gh, and St. Negrea. 1969. Apercu zoogéographique sur la fauna cavernicole ter-
restre de Roumanie.—Act. Zool. Cracoviensia 14(20):471-—54S.

Delorme, L. D. 1970. Freshwater ostracodes of Canada. Part 3. Family Candonidae.—Can.
J. Zool. 48(5): 1099-1127.

Furtos, N. C. 1933. The Ostracoda of Ohio.—Ohio Biol. Survey 5(6):413-524.

Hanganu, E., and I. Papaianopol. 1976. Les subdivisions du Dacien fondées sur les associa-
tions de malacofaune et d’ostracofaune.—Bull. Soc. Belge Géol. 85(1—2):63-88.
Hartmann, G. 1973. Zum gegenwartigen Stand der Erforschung der Ostracoden interstitieller

Systeme.—Ann. Spéléol. 28:417—426.

Hoff, C. C. 1942. The ostracods of Illinois. Their biology and taxonomy .—lIllinois Biol. Mono-
ger. 19(1—2):1-196.

Kesling, R. V. 1965. Anatomy and dimorphism of adult Candona suburbana Hoff. In Kesling,
R. V. et al.—Four Reports of Ostracod Investigations. Nat. Sc. Fund. Project GB-26,
Rep. 1:1—56.

Klie, W. 1938. Krebstiere oder Crustacea III. Ostracoda. Muschelkrebse. Jn E. Dahl, ed.,
Die Tierwelt Deutschlands 37: 1-230.

Kochegura, V. V., and V. A. Zubakov. 1978. Palaeomagnetic time scale of the Ponto-Caspian
Plio-Pleistocene deposits.—Palaeogr. Palaeoclimat. Palaeoecol. 23:151-160.

Kozhov, M. 1963. Lake Baikal and its life. Junk Publ. The Hague. 334 pp.

Kriz, H. 1973. Groundwater in the southern part of the Ceskotrbovska Vrchovina.—Acta Sc.
Nat. Brno 7(11):1—43.

Krstic, N. 1972. Genus Candona (Ostracoda) from Congeria beds of southern Pannonian
basin.—Serbian Acad. Sc. and Arts Monogr. 140(39):1-145.

Loffler, H., and D. L. Danielopol. 1978. Ostracoda. Jn Illies, J., ed., Limnofauna Europaea,
196-208. G. Fischer Verl. Stuttgart.

Luttig, G., and P. Steffens. 1976. Paleogeographic atlas of Turkey from the Oligocene to the
Pleistocene.—Bundesanstalt fiir Geowiss. u. Rohstoffe, Hannover.

Mandelstam, M. I., L. P. Markova, T. R. Rozyeva, and N. E. Stepanaitis. 1962. Ostracody

756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pliotsenovyh i postpliotsenovyh otlozhenii Turkmenistana (Spravochnik)—Izd. A. N.
Turk. SSR., Ashabad (after Vekua, 1975).

Marinescu, F., and R. Olteanu. 1973/1974. Considérations sur les associations des mollusques
et d’ostracodes du Méotien de la partie occidentale du bassin Dacique.—Dari de Seama
ale Sedintelor Inst. Geol. 59: 113-127.

Mazepova, G. F. 1970. A new species of Ostracode from Lake Baikal and its relations to the
transcaucasian subterranean fauna.—Zool. Zhur. 49:1632—1637 (in Russian, English
summary).

McGregor, D. L., and R. V. Kesling. 1969. Copulatory adaptations in ostracods. Part 2.
Adaptation in living ostracods.—Contr. Mus. Paleont. Univ. Michigan 22(17):221-239.

Rudjakov, I. A. 1963. O fauna peshterni zapadnovo Zakavkazya.—Zool. Zhur. 42(1):32—40.

Sokac, A. 1972. Pannonian and Pontian ostracode fauna of Mt. Medvenita.—Palaeont. Ju-
goslavica 11:1—140.

Southwood, T. R. E. 1976. Bionomic strategies and population parameters. Jn R. May, ed.
Theoretical ecology. Principles and applications: 26—48, Blackwell Sci. Publ., Oxford.

Sywula, T. 1976. New species of Ostracoda (Crustacea) from subterranean waters.—Bull.
Acad. Pol. Sc. Ser. Biol. 24(5):271-278.

Vekua, M. L. 1975. The ostracods of the Klimmerian and Kujalnikan deposits of Abkhazia
and their stratigraphic significance.—Metzniereba Tbilisi (in Russian, English summary).
137 pp.

Limnologisches Institut der Osterreichischen Akademie der Wissenschaf-
ten, Berggasse 18, A-1090 Wien.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 757-759

THE SUBSPECIES OF GRASSHOPPER SPARROW
(AMMODRAMUS SAVANNARUM) IN PANAMA
(AVES: EMBERIZINAE)

Storrs L. Olson

Abstract.—A specimen of Ammodramus savannarum pratensis from Al-
mirante, Bocas del Toro, is the first record for Panama and the southernmost
known for this migrant subspecies. A very pale resident population, A. s.
beatriceae subsp. nov., restricted to the llanos of Coclé Province, is de-
scribed. A disjunct population in eastern Panama Province is referable to
A. s. bimaculatus of Middle America.

The Grasshopper Sparrow, Ammodramus savannarum, 1s a secretive,
though widely distributed, grassland finch found in most of North America
and the Greater Antilles (except Cuba), through Middle America as far as
Panama, with two subspecies isolated in Curacao and Bonaire and in Co-
lombia and Ecuador. In Panama, the species is poorly known and there is
little in the literature concerning it. Material in the National Museum of
Natural History, Smithsonian Institution (USNM), indicates that three dis-

_ tinct subspecies may be found in Panama, one of which is a migrant and

two of which are resident. Of the last two, one represents a very distinct
new taxon.

Ammodramus savannarum pratensis (Vieillot)

Fringilla pratensis Vieillot 1817, Nouv. Dict. Hist. Nat., nouv. ed. 25:24.
Type-locality, New York.

Specimens examined.—PANAMA. BOCAS DEL TORO: Almirante. An
unsexed adult taken in November 1963 by personnel of the Gorgas Memorial
Laboratory (USNM 533402).

Remarks.—This specimen is readily separated from any of the resident
populations of Ammodramus savannarum in Middle America by its greater
wing length (63.4 mm vs. 53.2—59.1 mm for A. s. bimaculatus, n = 23). It
is darker and more richly colored than the birds of western North America
(A. s. perpallidus [Coues]), but is inseparable from individuals of A. s.
pratensis, the migratory population breeding in eastern North America. The
present specimen marks a significant range extension, the southernmost
occurrence of the subspecies heretofore being in Belize (British Honduras).
Ridgely (1976:339) mentions a specimen of this species that was captured

758 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and released, also at Almirante, on 4 November 1967. He suggested that
this was a migrant, which seems likely in view of the identity of the above
specimen and the general lack of sufficient habitat in the Almirante area to
sustain a breeding population of this species.

Ammodramus savannarum bimaculatus Swainson

Ammodramus bimaculatus Swainson 1827, Philos. Mag. (n.s.) 1(6):435.
Type locality, Temiscaltipec (=Temascaltepac), Mexico.

Specimens examined.—PANAMA. CHIRIQUI: Francés (4, AMNH).
PANAMA PROVINCE: Rio Pacora, 10 miles W of Chepo (2, USNM, one
of which is juvenile); 10 miles NW of Chepo (1, USNM).

Remarks.—The specimens from the Chepo area have been discussed fully
by Aldrich (1945), who assigned them to the subspecies bimaculatus, which
ranges from Mexico to Chiriqui, Panama. I concur in this and have little to
add to Aldrich’s account. The Chepo birds are inseparable from Costa Rican
specimens in fresh plumage, though I am not prepared to say that these in
turn are the same as Mexican birds, as those Mexican specimens available
to me were in quite worn plumage. The birds from Chepo, which is in the
eastern extreme of the Pacific slope savannas of Panama Province, certainly
represent a breeding population, as evidenced by the juvenile specimen.
Considering their apparent isolation from the nearest known population of
bimaculatus in Chiriqui, it is somewhat curious that these birds have not
differentiated, particularly in light of the distinctiveness of the birds from
the savannas of Coclé, described below.

Ammodramus savannarum beatriceae, new subspecies

Holotype.—USNM 477168, female, taken 10 March 1962 by Alexander
Wetmore at El Coco, Coclé Province, Panama (original number 24894).

Characters.—Palest of all the subspecies of A. savannarum. Equal in size
to A. s. bimaculatus but throat, breast, flanks and undertail coverts pale
pinkish buff, not rich ochraceous yellow. Dorsum much grayer than in Di-
maculatus and median crown stripe very pale, almost white, instead of deep
buff. The subspecies A. s. caucae Chapman, of Colombia and Ecuador, is
also separable from bimaculatus by its paler coloration, but beatriceae is
paler still. In a series of seven specimens of caucae from Yumbo, Depart-
ment of Valle, Colombia (CM), the streaks in the nape are dark, rather
blackish brown, never as distinctly reddish as in beatriceae or bimaculatus.
The single available specimen of A. s. caribaeus (Hartert) from Curagao
(USNM), is also rather pale, but is more ochraceous on the breast and
cheeks, more brownish above, and has a decidedly smaller bill than beatri-
ceae.

VOLUME 93, NUMBER 3 Ue

Range.—Known only from the savannas of southern Coclé Province,
Pacific slope of Panama.

Specimens examined (all USNM).—PANAMA. COCLE: El Coco (2); 4
miles W of Penonomé (2); Penonomé, Puerto Posada (1); Gago, Gaucho (1);
3 miles NW of Anton (4).

Etymology.—Affectionately dedicated to Mrs. Alexander Wetmore, who
accompanied her husband on most of his collecting trips to Panama and
who attended him with unselfish devotion through their 25 years of marriage.

Remarks.—As this is perhaps the most easily distinguished of all the
forms of Ammodramus savannarum, it is indicative of a fair degree of iso-
lation of the grassland birds of the llanos of Coclé. Further evidence of this
is supplied by another endemic subspecies of Emberizinae, Sicalis luteola
eisenmanni, described by Wetmore (1953) from most of the same localities
as A. s. beatriceae. This is the only population of S. luteola occurring
between Guatemala and Colombia.

Acknowledgments

I am grateful to Kenneth C. Parkes, Carnegie Museum of Natural History
(CM), Pittsburgh, for lending specimens of A. s. caucae and for commenting
on the manuscript. This is contribution Number 4 of the Wetmore Papers,
a project supported in part by trust funds from the Smithsonian Institution
for completing unfinished work and study of undescribed material left by
the late Alexander Wetmore.

Literature Cited

Aldrich, J. W. 1945. Notes on Panamanian birds.—Auk 62:111-114.

Ridgely, R. S. 1976. A Guide to the Birds of Panama.—Princeton University Press, Princeton,
N. J. 394 pages.

Wetmore, A. 1953. Further additions to the birds of Panama and Colombia.—Smithsonian
Miscellaneous Collections 122(8): 1-12.

Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 760-770

SPONGICOLOIDES GALAPAGENSIS, A NEW SHRIMP
REPRESENTING THE FIRST RECORD OF THE
GENUS FROM THE PACIFIC OCEAN
(CRUSTACEA: DECAPODA: STENOPODIDEA)

Joseph W. Goy

Abstract.—A new species of stenopodidean shrimp from the Galapagos
Islands is described and figured. It is the first species of the genus Spongi-
coloides Hansen recorded from the Pacific Ocean. The new species, Spon-
gicoloides galapagensis, is closely related to S. profundus and in a few
characters is very similar to the genus Spongiocaris Bruce and Baba, but
is easily distinguished from both by differences of the telson, spination of
the carapace, and branchial formula. A table of some morphological differ-
ences for the known species of Spongicoloides is included.

The four known species of Spongicoloides are from the Atlantic Ocean
from Iceland to the Caribbean (Holthuis, 1946). While examining some un-
identified stenopodid material from the National Museum of Natural His-
tory, Washington, D.C., I found a single example of an undescribed Spon-
gicoloides, collected by the steamer Albatross in deep water off the
Galapagos Islands. In addition to its geographical location, several morpho-
logical characters also distinguish this specimen from the other members of
the genus.

Spongicoloides galapagensis sp. nov.
Figs. 1-4

Material examined.—1 9°, ovigerous, Galapagos Islands, 00°29'S,
89°54'30” W. U.S. Fish Commission Steamer Albatross, 15 April 1888; Sta-
tion 2818; 717 m, white and black sand, large beam trawl; bottom temper-
ature 6.6°C.

Diagnosis.—A fairly large stenopodidean shrimp, with a stout, depressed
body, generally glabrous; telson with 6—7 lateral teeth and 8 smaller teeth
on posterior margin; uropodal exopodite with 13-18 teeth on outer margin,
dorsally | strong and | weak ridge; dactyli of fourth and fifth pereiopods
triunguiculate; third maxilliped with spines; scaphocerite with 10 teeth.

Description.—Holotype (female, USNM 180064). Rostrum (Figs. 1, 2A)
short, compressed, extending almost to distal end of antennular peduncle
basal segment, tapering gradually into slightly upturned acute tip. Dorsal

VOLUME 93, NUMBER 3 761

Fig. 1. Spongicoloides galapagensis, holotype, female. Scale bar in mm.

margin bearing 7 strong spines; ventral margin bearing 4 strong spines. Ros-
tral base triangular with each lateral margin bearing 3 small spines.

Carapace (Figs. 1, 2A) of very thin texture, with distinct cervical groove;
less distinct, shorter postcervical groove. Dorsal midline bearing 5 small
spines anterior to cervical groove, 4 slightly larger spines posterior to post-
cervical groove. Frontal region behind rostrum up to cervical groove bearing
8 small spines. Three small spines present directly behind anterior part of
cervical groove, which in dorsal view appear to be on cervical groove itself.

Small antennal spine, large branchiostegal spine and 4 small pterygostomian

spines with pterygostomial region also bearing numerous minute spinules
somewhat arranged in rows. Branchial region having numerous scattered
spines; branchiostegite bearing large spine close to posterolateral border.
Ventrolateral angle of carapace somewhat rounded while posterolateral an-
gle of branchiostegite obtusely angled, slightly concave.

Abdominal somites glabrous, lacking carinae. Pleura of first 3 rounded,
without setae ventrally. Anterolaterally, second and third somites bearing
articular knobs, while fourth somite with distinct knob. Pleura of fourth and
fifth somites ending in 3 small teeth.

Telson (Fig. 2B) broad, roughly rectangular, slightly constricted at base.

Dorsal surface with 2 longitudinal ridges; left ridge having 7 spines and right
8. Anteriorly, 2 spines and another outside proximal part of left ridge. Lat-

|
|
|

762 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

LL
f He aw =
LEAL, ‘ Sel
( y ies g KS
Kor 4 ~ TSS RASS
Vy Ste SN WSs

SSS a Pr

Fig. 2. Spongicoloides galapagensis, holotype, female: A, Carapace, dorsal view; B, Tel-
son, dorsal view; C, Antennule, dorsal view; D, Antenna, dorsal view; E, Right mandible,
ventral view; F, Same, dorsal view; G, Maxillule; H, Maxilla. Scale bars represent 1.0 mm.

VOLUME 93, NUMBER 3 763

eral borders bearing 6 left and 7 right acute teeth; borders setose along distal
third. Posterior margin broadly rounded, densely setose, with 8 small teeth.

Eyes well developed, of moderate size, cornea globular and devoid of
black pigment. No spinules on ophthalmic peduncle.

Antennular peduncle (Fig. 2C) short, extending to middle of scaphocerite.
Basal segment more than twice length of second segment, third much short-
er than both. Inner margin of basal segment straight, bearing 18 plumose
setae of moderate length. Outer margin slightly expanded proximally bearing
minute stylocerite about midlength, while distal extremity comes to sharp
point with 4 plumose setae. Second segment with 2 small spines on inner
margin; distal segment glabrous. Lateral flagellum slightly stronger than
mesial flagellum, both with some segments bearing short setae.

Antenna (Fig. 2D) with stout basicerite, having strong spine on distal
margin. Carpocerite short, not exceeding second antennular peduncle seg-
ment. Antennal flagellum broken at sixteenth segment. Scaphocerite broad,
quadrangular, dorsally bearing 2 faint longitudinal carinae, with one bearing
small knob about mid-length; outer margin slightly concave bearing 9 small
teeth and larger distal tooth, inner margin convex, fringed with 84 long
plumose setae.

Mandible (Figs. 2E, F) robust, with short fused molar and incisor pro-
cesses. Molar surface with 6 small teeth; incisor bearing 2 stout teeth distally
followed by 11 small teeth proximally. Palp well developed, 3-segmented.
Proximal segment without setae; middle segment setose mesially, on distal
outer margin; distal segment broad, densely setose laterally and distally.

Maxillule (Fig. 2G) bearing simple undivided endopodite with terminal
short seta. Distal endite narrow with numerous fine setae and 11 stout spines
distally; proximal endite also narrow with many fine setae distally.

Maxilla (Fig. 2H) with following setation on coxal and basal endites: 32
on proximal lobe, 16 on distal lobe of coxal; 20 on proximal lobe, 38 on
distal lobe of basal. Endopodite long, slender, with 36 lateral and distal
plumose setae. Scaphognathite well developed; anterior lobe broad, poste-
rior One narrow with 208 plumose setae along margin.

First maxilliped (Fig. 3A) with stout 3-segmented endopodite. Proximal
segment long, narrow proximally, broader distally, bearing 18 long plumose
setae on outer margin, 15 shorter plumose setae mesially, and 4 plumose
setae on inner margin. Middle segment broad, less than half length of basal
segment, with 18 long plumose setae on outer margin and 2 long plumose
setae distally. Distal segment very small, vestigial, with 2 short plumose
setae. Basipodite large, rounded anteriorly, with straight mesial border bear-
ing dense fringe of long setae. Coxopodite represented by somewhat round-
ed lobe bearing 10 short setae. Exopodite well developed bearing 11 long
plumose setae laterally and distally. Large epipod present with broad prox-
imal and distal lobes.

764 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 3. Spongicoloides galapagensis, holotype, female: A, First maxilliped; B, Second max-
illiped; C, Third maxilliped, with dactylus missing; D, E, Chela of first pereiopod; F, G, Chela
of second pereiopod. Scale bars represent 1.0 mm.

Second maxilliped (Fig. 3B) with 7-jointed endopodite. Dactylus suboval,
about twice as long as broad, with densely setose distodorsal margin. Pro-
podus about 1.2 length of dactylus with dense fringe of setae on distodorsal
margin, while carpus short, triangular about half propodal length with nu-
merous long simple setae at distodorsal angle. Merus long, straight, twice
propodal length with dense fringe of long setae on inner border. Ischium

ee

VOLUME 93, NUMBER 3 765

and basis not fused, both short with many setae on inner margins, few on
outer margins. Coxa with 3 short setae on inner margin, 6 setae mesially,
and simple epipod bearing small podobranch laterally. No exopodite pres-
ent.

Third maxilliped (Fig. 3C) with 7-segmented endopodite, but dactylus
missing on specimen. Propodus long, narrow, with 5 spines on inner margin;
numerous long and short simple setae mesially, on outer and inner margins.
Carpus slightly shorter than propodus with 2 long distal setae on inner bor-
der, numerous long and short setae on outer border. Merus longest segment
with 3 long, few short setae on outer margin, few setae on inner margin.
Ischium as long as carpus with few setae, strong spine on inner margin;
small distal spine, some short setae on outer margin. Basis without exo-
podite but with 3 short setae on outer border. Distal end of coxa produced
into rounded point bearing 3 short setae on outer margin; inner margin with
3 short setae, small epipod. Arthrobranch, pleurobranch present.

First pereiopod (Figs. 3D, E) small, slender, glabrous; when extended
reaching slightly beyond scaphocerite. No setiferous organ present. Dac-
tylus less than half propodal length. Fingers slightly compressed with slight-
ly hooked tips. Cutting edges rather indistinct with propodus and dactylus
bearing chitinous ridge along inner margins. Dactylar ridge proximally forms
one large, 2 smaller acute teeth. Both tips of propodus and dactylus round-
ed, heavily chitinized. Fingers and distodorsal extremity of palm with small
tufts of long setae. Carpus longest segment, more than twice propodal
length, with few simple setae on inner and outer margins. Ischium same
size aS propodus; merus slightly longer. Basis and coxa short, unarmed.

Second pereiopod (Figs. 3F, G) similar to first, but longer, stronger. Fin-
gers and distodorsal extremity of palm bearing small tufts of long setae.
Cutting edges provided with 14 small, stout, peg-like teeth separated by
rectangular chitinous lamellae; near tips each edge with small, spike-like
tooth; tips hooked, heavily chitinized. Carpus longest segment with 5 short
simple setae along outer margin. Merus almost equal in length to carpus,
ischium one-third carpal length. Basis and coxa short, unarmed.

Third pereiopod (Figs. 1, 4A) largest, strongest, almost as long as entire
body length. Dactylus with chitinous ridge along cutting edge with large,
rounded, serrate tooth almost at midlength opposing deep notch in serrate
chitinous ridge of propodal cutting edge. Fingers elongate with sharp,
hooked, chitinized, crossing tips, bearing small tufts of long setae. Propodus
longest segment, distoventrally bearing 7 small knobs. Carpus less than half
propodal length, narrowing proximally. Merus almost same length as pro-
podus, with distoventral angle forming sharp spine with smaller mesial spine
directly above. Ischium equal to carpal length, ventrally bearing 4 spines,
distodorsal angle produced into large curved spine with smaller mesial spine
below. Basis and coxa short, stout, unarmed.

Fourth, fifth pereiopods (Figs. 4B, C) long, slender, subequal in length.

766 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 4. Spongicoloides galapagensis, holotype, female: A, Chela of third pereiopod; B,
Propodus and dactylus of fourth pereiopod; C, Propodus and dactylus of fifth pereiopod; D,
First pleopod; E, Second pleopod; F, Right uropod. Scale bars represent 1.0 mm.

Dactylus of fourth triunguiculate with much smaller proximal tooth on ac-
cessory spine. Unguis long, curved, clearly separated from dactylar corpus;
accessory spine similar but shorter. Propodus and carpus undivided, with
propodus about one-third carpal length, bearing 20 movable spines, 2 long
setae ventrally; 13 long dorsal setae. Carpus longest segment with 11 dorsal
setae, 4 ventral setae, movable spine at distoventral angle. Merus almost
same length as carpus with 3 long distal setae dorsally. Ischium same length
as propodus; unarmed. Basis and coxa short, stout. Fifth pereiopod with
similar dactylus as fourth but much smaller unguis and accessory spine.
Propodus half carpal length bearing 16 movable spines, 6 long setae ven-
trally; 13 long dorsal setae. Carpus and merus equal length; merus unarmed
and carpus bearing 14 dorsal setae, 7 ventral setae. Ischium same length as
propodus; unarmed. Basis and coxa short, stout.

VOLUME 93, NUMBER 3 767

First pleopod (Fig. 4D) uniramous, second (Fig. 4E) to fifth biramous, all
lacking appendices. First pleopod smallest, with exopodite slightly longer
than basipodite. Dorsal, ventral margins of basipodite densely covered with
long plumose setae. Exopodite with 28 long plumose marginal setae. Rami
of second pleopod slightly longer than basipodite. Distodorsal margin of
basipodite bearing 5 short plumose setae, ventral margin with numerous
longer plumose setae. Exopodite, endopodite respectively bearing 36, 42
long plumose marginal setae. Third to fifth pleopods generally similar, de-
creasing in size, setation posteriorly.

Uropods (Fig. 4F) well developed, about as long as telson. Basal segment
strong with distodorsal angle produced into large tooth, large mesial tooth.
Exopodite, endopodite ovate with outer margin of left exopodite bearing 13
teeth, right exopodite bearing 18 teeth. Unarmed margin of exopodite with
78 long plumose setae, dorsal surface bearing strong median ridge, much
weaker submedian ridge. Margins of endopodite unarmed, bearing 102 long
plumose setae. Dorsal surface with median ridge with spine located about
mid-length.

There are only 20 large eggs, about 2.0 mm long and 1.3 mm broad.

Branchial formula:

Maxillipeds Pereiopods

I I Il I eS eevee NV
Pleurobranchs —_ — 1 1 l 1 1 l
Arthrobranchs —_- — 1 l l 1 1 —
Podobranchs — a = => SS SS =|
Epipods 1 l —- —- —- — —
Exopods Lo —- —- — — —

Measurements (in mm): Postorbital carapace length, 14.0. Rostral cara-
pace length, 17.0. Total length, approx., 39.0. Length of third pereiopod,
approx., 36.5.

Coloration: The color of the preserved specimen is a pale yellow, the
color of the living animal is unknown.

Type-locality.—Eastern Pacific Ocean, off the Galapagos Islands.

Remarks.—The new species, Spongicoloides galapagensis, follows the
definition of the genus Spongicoloides Hansen given by Holthuis (1946),
only differing in the spinous third maxillipeds, having 2 dorsal ridges on the
uropodal exopodite, and the triunguiculate dactyli of the last 2 pairs of legs.
Holthuis used the branchial formulae as one of the means of distinguishing
the 4 known species of Spongicoloides. In this regard, S. galapagensis
comes close to S. inermis, but the podobranch on the second maxilliped
and the arthrobranch on the fourth pereiopod are not rudimentary in the
new species. Spongicoloides inermis also bears 4 small knobs on the ventral

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VOLUME 93, NUMBER 3 769

margin of the third pereiopod propodus, similar to the 7 knobs on S. gal-
apagensis. The new species has a few characters similar to the genus Spon-
giocaris Bruce and Baba, namely the dactyli of the fourth and fifth pereio-
pods and the dorsal ridges on the uropodal exopodite. This supports Bruce’s
and Baba’s (1973) statement that the genus Spongiocaris seems to be an
intermediate form between Spongicola and Spongicoloides. In most other
respects, the new species is most closely related to Spongicoloides profun-
dus, but it is easily distinguished from members of the genera Spongiocaris
and Spongicoloides by differences of the telson, spination of the carapace,
and branchial formula. The new species differs from the other members of
the genus Spongicoloides in a number of other characters shown in Table 1.

The 4 previously described species of Spongicoloides are known only
from the Atlantic Ocean. There is not much known of the characteristics of
S. evolutus (Bouvier, 1905), based on a single specimen collected off the
west coast of the Sahara. The only characteristics given by Bouvier (1905a,
1908a) and Milne Edwards and Bouvier (1909) are shown in Table 1. Spon-
gicoloides profundus Hansen, 1908 has only been recorded southwest of
Iceland, 60°37’N, 27°52'W (Hansen, 1908); S. inermis (Bouvier, 1905) was
collected off St. Lucia in the Caribbean (Bouvier, 1905b, 1908b; Milne Ed-
wards and Bouvier, 1909); and S. koehleri (Caullery, 1896) has been col-
lected from the Bay of Biscay and the Atlantic Ocean near the Strait of
Gibraltar (Caullery, 1896; Kemp, 1910; Sund, 1920). The new species is the
first record of the genus from the Pacific Ocean. Three specimens of yet
another Western Atlantic species of Spongicoloides were found in the un-
identified stenopodid material of the U.S. National Museum. This species
differs from S. galapagensis and the other members of the genus by differ-
ences in the rostrum, scaphocerite, telson, and uropods and a full descrip-
tion will be presented in a forthcoming publication.

Acknowledgments

I am extremely grateful to Dr. Raymond B. Manning, Curator, Depart-
ment of Invertebrate Zoology, Smithsonian Institution, who gave me the
opportunity to describe this new species and reviewed the manuscript. I
would also like to thank Drs. Fenner A. Chace, Jr. and Brian Kensley, also
of the Smithsonian Institution, for critically reviewing the manuscript.

Literature Cited

Bouvier, E. L. 1905a. Sur les Pénéides et les Sténopides recueillis par les expéditions fran-
¢aises et monégasques dans I’ Atlantique oriental.—C. R. Acad. Sci. Paris 140:980-983.
. 1905b. Sur les Macroures nageurs (abstraction faites des Carides) recueillis par les
expeditions américaines du ‘‘Hassler’’ et du ‘‘Blake’’.—C. R. Acad. Sci. Paris 141:746—
749.

770 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1908a. Sur les relations zoologiques des Crevettes de la tribu Sténopides.—C. R.

Acad. Sci. Paris 146:887-891.

. 1908b. Catalogue des Crustacés de la famille des Stenopides des collections du Mu-

séum d’histoire naturelle —Bull. Mus. Hist. Nat. Paris 14:150.

Bruce, A. J., and K. Baba. 1973. Spongiocaris, a new genus of stenopodidean shrimp from
New Zealand and South African waters, with a description of two new species (Deca-
poda, Natantia, Stenopodidea).—Crustaceana 25(2): 153-170.

Caullery, M. 1896. Crustacés Schizopodes et Décapodes. In R. Koehler, Résultats scienti-
fiques de la Campagne du *“*‘Caudan’’ dans le Golfe de Gascogne, aout-septembre 1895.—
Ann. Univ. Lyon 26:365-419, pls. 13-17.

Hansen, H. J. 1908. Crustacea Malacostraca, 1.—Danish Ingolf Exped. 3(2):1—120, pls. 1-5.

Holthuis, L. B. 1946. Biological results of the Snellius Expedition. XIV. The Stenopodidae,
Nephropsidae, Scyllaridae, and Palinuridae.—Temminckia 7:1—178, pls. 1-11.

Kemp, S. W. 1910. The Decapoda collected by the ‘‘Huxley’’ from the North side of the Bay
of Biscay in August, 1906.—J. Mar. Biol. Ass. U.K. 8:407—420.

Milne-Edwards, A., and E. L. Bouvier. Les Peneides et Stenopides. Reports on the Results
of Dredging, under the Supervision of Alexander Agassiz, in the Gulf of Mexico (1877-
1878), in the Caribbean (1878-1879) and along the Atlantic coast of the United States
(1880), by the U.S. Coast Survey Steamer ““Blake’’.—Mem. Mus. Comp. Zool. Harvard,
27:177-274, pls. 1-9.

Sund, O. 1920. Peneides and Stenopides.—Rep. Sars North Atlantic Deep Sea Exped. 1910,

Zool. 3(2):1—36, pls. 1-2.

Duke University Marine Laboratory, Pivers Island, Beaufort, North Car-
olina 28516.

ADDENDUM

After this manuscript was sent to the printer the following paper was pub-
lished: Baba, K. 1979. A new stenopodidean shrimp (Decapoda, Natantia)
from the Chatham Rise, New Zealand. Pacific Sci. 33(3):311-314. He de-
scribes Spongicoloides novaezelandiae sp. nov., which differs from S.
galapagensis and the other members of the genus in number of gills and
spination of the rostrum.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 771-780

A NEW SPECIES OF ROCK SHRIMP OF THE GENUS
SICYONIA (PENAEOIDEA), WITH A KEY TO THE
WESTERN ATLANTIC SPECIES

Isabel Perez Farfante

Abstract.—Sicyonia olgae, new species, ranges from Dry Tortugas Is.,
Florida, to Suriname. It differs from Sicyonia typica (Boeck, 1864), its closest
western Atlantic relatives, in possessing sublateral carinae on the carapace,
and in lacking posterior pleural sulci on the first three abdominal somites;
also, distinctive are the sharply pointed, mesially directed, distomesial pro-
jection of the petasma in the male, and in the female the pair of long, slender
spines on sternite XI and rounded posterolateral processes of the median
plate of sternite XIII. A key to the western Atlantic species of Sicyonia is
supplemented by synopses of their geographic and depth ranges which in-
clude many extensions of previously known limits.

The shrimp genus Sicyonia was previously known to be represented by
eight species in the western Atlantic, but an examination of the collections
of these shrimps in the National Museum of Natural History, Smithsonian
Institution (USNM), brought to light a ninth species here described as new.

Some of those Sicyonia are present in relatively large quantities in com-
mercial catches of shrimps of the genus Penaeus, and one, S. brevirostris
Stimpson, 1871, is large and abundant enough to sustain a fishery which
rendered about 3,700,000 Ibs (heads-off weight) in 1979.1 Although the col-
lections examined are rather extensive, the number of available specimens
of the new species is limited; thus additional material may demonstrate
variations in some of the characters treated.

Sicyonia olgae has the broadest bathymetric range—from 33 to 622 m—
of any of its western Atlantic congeners. The species of Sicyonia typically
occur in shallow water, but the range of many of them, like that of the new
species, extends across the continental shelf onto the slope, to depths of
a few hundred meters. Such pattern of bathymetric distribution is not un-
common among penaeoidean shrimps; for example, certain species of Me-
tapenaeopsis have been recorded from less than 20 to more than 300 m, and
Parapenaeus longirostris (Lucas, 1846), a species commercially exploited

' Data provided by Roger W. Hutchinson, Natl. Mar. Fish. Serv., Fish. Develp. Div., U.S.
Dept. of Commer., NOAA.

772 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in the Mediterranean and eastern Atlantic, ranges from less than 20 m to at
least 700.

Because two new species of the genus Sicyonia have been discovered
since the appearance of the most recent key to the western Atlantic species
of the genus (Chace, 19727), a modified version of it is presented below. A
study of extensive collections of Sicyonia from that region shows that the
geographic and depth ranges of many species are considerably greater than
was previously realized. Synopses of geographic and bathymetric ranges
are included after the key.

The occurrence of S. brevirostris in the eastern Pacific is cited under
‘‘Geographic and bathymetric ranges. . . .”’ with hesitation. The only record
of its presence outside the western Atlantic is that of Burkenroad (1934)
who reported a single juvenile specimen from off the Pacific coast of south-
ern Mexico (off Tapachula, Chiapas). I have examined this specimen, and
agree with his determination. However, because S. brevirostris has not been
reported again from the region, nor have I found any representatives among
the many hundred specimens of Sicyonia I have examined from the Pacific
coast of America (including large collections from Mexican waters), its oc-
currence there needs to be confirmed.

The terminology employed in describing the external genitalia, and trans-
verse sulci on the pleonic pleura is that proposed by Kubo (1949) and Bur-
kenroad (1934), respectively, the former slightly modified by Pérez Farfante
(1969). The length of the carapace was measured from the orbital margin to
the midposterior margin, and the total length, from the apex of the rostrum
to the posterior end of the telson. The scales accompanying the illustrations
are in millimeters.

Key to the Western Atlantic Species of Sicyonia

1. Antennal spine absent or occasionally exceedingly weak; second
abdominal somite with narrow notch or perpendicular incision in
anterior half of dorsal carina; first pereopod with short distal spine
onbasissand ischiumio 4.002... PG AUR), ae eee 2

— Antennal spine well developed or clearly distinct; second abdominal
somite with dorsal carina entire, not incised; first pereopod with
basiscandiischiumi unarmed iia0, 31). tse eine 3

2. Postrostral carina armed with 3 unequal teeth, anterior one smallest;
first abdominal somite with 1 well marked vertical (posteromedian)

2 A later key by Huff and Cobb (1979) is restricted to the Gulf of Mexico and the Atlantic
coast of Florida.

VOLUME 93, NUMBER 3 TB

sulcus and sometimes with | short, barely distinct more anterior one

(anteromedian) on pleuron ............. S. laevigata Stimpson, 18713
— Postrostral carina armed with 3 subequal teeth; first abdominal so-

mite with 3 vertical sulci on pleuron, posterior one (posterior pleural)

LESSECONSPICUOUS MM Seeks aud ows le oan hen S. parri (Burkenroad, 1934)
3. Postrostral carina bearing 2 or 3 large teeth posterior to level of

Me PALICES PINCH eietes tl Ns Rien alee SOS. ea Vea posal, 2 4
— Postrostral carina bearing | large tooth posterior to level of hepatic

SOM 2 WH ales Be Bes eek Ce re Oe ee ee eet See eee ee ee 6
4. Postrostral carina with 3 large teeth posterior to level of hepatic

SPM apereheey fic, Hh tcseet nty Aut eke tS EY S. brevirostris Stimpson, 1871
— Postrostral carina with 2 large teeth posterior to level of hepatic

SMO Se gh Bea Se Ree Oe Oe ee CER RE eeeT Neem 5

5. Rostrum armed with 1 or 2 teeth (excluding tip) anterior to orbital

margin; first abdominal somite with 3 vertical sulci on pleuron ....
Re ic ee Sh ee io Ma, denials Nig do as S. typica (Boeck, 1864)

— Rostrum armed with 3 or 4 teeth (excluding tip) anterior to orbital

margin; first abdominal somite with 2 vertical sulci on pleuron (pos-
(PST OP ONS EVE] SNe) Reet eee Rr ee eee oe S. olgae, n. sp.

6. First abdominal somite with tooth at anterior end of dorsal carina

bifurcate; fifth abdominal somite without tooth or sharp angle at
posterior end of dorsal carina ............. S. wheeleri Gurney, 1943

— First abdominal somite with tooth at anterior end of dorsal carina

not bifurcate; fifth abdominal somite with tooth or sharp angle at
POSICHIOLR CNnGeOlGdORSalec amas rys.s ctcya 2 4.s.6 sgssceasms dakedlnesienn la ko Says 7

7. First abdominal somite with anteroventral margin of pleuron con-

cave or straight; fourth abdominal somite with posteroventral margin

of pleura angular or with posteriorly oriented spine; fourth and fifth

abdominal somites with long acute spine on either side of midline of
POSteHOdorSalpmManein es. pe ek ee S. dorsalis Kingsley, 1878

— First abdominal somite with anteroventral margin of pleuron slightly

or strongly convex; fourth abdominal somite with posteroventral

margin of pleura rounded; fourth and fifth abdominal somites with

short, obtuse projection on either side of midline of posterodorsal
(TETRA, ahs, ee ae ee er ran Coa eee ee 8

3 The number of rostral teeth which has been widely used to distinguish S. laevigata from
S. parri is omitted here because both possess 3, contrary to previous statements that S.
laevigata bears only 2. In the young and many adults of the latter the third tooth is placed so
far anteriorly that it appears to constitute a part of the tip dentition; the apical dentition consists
of 1 to 3 minute teeth in addition to a ventral one that usually becomes situated farther pos-
teriorly with increasing size of the shrimp.

774 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

8. First four abdominal somites with anteroventral angle of pleura lack-
ing laterodorsally curved spine (small, ventrally directed obtuse
spine occasionally present); last two abdominal somites with well-
defined tooth at posterior end of dorsal carina...................
PRPS Gots cotiocs 5c ete eae el ay ae ee S. stimpsoni Bouvier, 1905
— First four abdominal somites with anteroventral angle of pleura bear-
ing laterodorsally curved, acute spine; last three abdominal somites
with well-defined tooth at posterior end of dorsal carina. .........
sea Nek Alpe o SRN ees arene ain, MET rer liwh Sera haa S. burkenroadi Cobb, 1971

Geographic and Bathymetric Ranges of Western Atlantic
Species of Sicyonia

Sicyonia brevirostris Stimpson, 1871. Off Norfolk, Virginia, through the
Bahamas to southern Cuba, around the Gulf of Mexico to southern Texas,
and from Bahia de Campeche to northeast Yucatan (it has not been recorded
from the east coast of Mexico); a single specimen from off Chiapas (Pacific
coast), Mexico. Shallow water to 350 m.

Sicyonia burkenroadi Cobb, 1971. Cape Lookout, North Carolina, south-
ward through the Gulf of Mexico and the Caribbean to Ilha Itaparica, Bahia,
Brazil. Depth 29 to 585 m.

Sicyonia dorsalis Kingsley, 1878. Cape Hatteras, North Carolina, south-
ward through the Gulf of Mexico, and along the Caribbean coast of Central
and South America to Enseada de Tijucas, Santa Catarina, Brazil (records
from the West Indies are in need of confirmation). Depth 3 to 420 m.

Sicyonia laevigata Stimpson, 1871. Cape Hatteras, North Carolina, to
Biscayne Bay; along west and northwest Florida; Yucatan; and from the
Bahamas, through the Caribbean southward to Anse de Zimbros, Santa
Catarina, Brazil; also along the Pacific coast of Panama. Shallow water to
at least 100 m.

Sicyonia olgae, n. sp. Dry Tortugas Is., Florida, through the Antilles to
Suriname. Depth 33 to 622 m.

Sicyonia parri (Burkenroad, 1934). Beaufort, North Carolina, through the
West Indies and southward to Ponta do Corumbau, Brazil (it is not known
to occur on the continental shelf of the Caribbean). Shallow water to 37 m.

Sicyonia stimpsoni Bouvier, 1905. Cape Hatteras, North Carolina, to the
Straits of Florida; eastern Gulf of Mexico; and from the Bahamas through
the Caribbean to Suriname. Depth 20 to 420 m.

Sicyonia typica (Boeck, 1864). Off Wrightsville Beach, North Carolina,
southward through the Gulf of Mexico and Caribbean (Antilles, Central
America and South America) and along the Atlantic coast of South America
to SE of Ilha de Santa Catarina, Santa Catarina, Brazil. [Record from Beau-

VOLUME 93, NUMBER 3 TS

fort, North Carolina (Hay and Shore, 1918) actually refers to S. brevirostris
(Burkenroad, 1934:92)]. Shallow water to 101 m.

Sicyonia wheeleri Gurney, 1943. Bermudas and Virgin Is. to Sint Eusta-
tius, Lesser Antilles. Depth 2 to 42 m.

Sicyonia olgae, new species
Figures 1-3

Material.—Holotype: 2, USNM 173666, 12 mm carapace length, about
44 mm total length; type-locality: off Paramaribo, Suriname, 06°37’N,
55°36’W, 35 m, 3 September 1958, Oregon stn 2277.

Paratypes: 1 2, USNM 173667, off Dry Tortugas Is., Florida, 33-37 m,
16 August 1933, J. W. Mills. 2 ¢6, USNM 173668, off Punta Las Tunas,
Puerto Rico, 18°31’N, 66°47’W, 70 m, 8 October 1959, Oregon stn 2668. 1
2, USNM 173669, NE Islas Los Testigos, Venezuela, 11°40’N, 62°33’W,
585-622 m, 24 September 1964, Oregon stn 5039. 1 d6 1 2, USNM 103513,
NE of Georgetown, Guyana, 07°40'N, 57°34’W, 55-49 m, 31 August 1958,
Oregon stn 2249.

Description—Body robust (Fig. 1). Rostrum horizontal, surpassing eye,
reaching or slightly overreaching distal margin of first antennular article, its
length 0.3 to 0.4 that of carapace, armed with 3 or 4 teeth decreasing in size
anteriorly, and with tip bifid or trifid. Carapace with closely set fine setae
dorsally and patches of setae laterally. High postrostral carina bearing 3 or
4 teeth (if 4, rostrum with 3, total number of rostral plus carapace teeth not
exceeding 7); 2 behind hepatic spine, anteriormost located slightly behind
orbital margin (if 4 teeth present on carapace, second situated slightly an-
terior to or at level of hepatic spine); first posthepatic tooth located slightly
posterior to midlength of carapace, and posteriormost at about 0.8 carapace
length from orbital margin. Adrostral carina, parallel to ventral rostral mar-
gin, extending from orbital margin aimost to tip of rostrum. Antennal spine
small and buttressed; hepatic spine, projecting from swollen hepatic region,
Straight, directed anteriorly, and longer than antennal. Postocular sulcus
short. Hepatic sulcus almost horizontal, relatively deep; branchiocardiac
carina horizontal, narrow and rather long, extending from about posterior
end of hepatic sulcus to not far from posterior margin of carapace; sublateral
carina, sometimes weak, almost parallel to branchiocardiac carina.

Stylocerite with short spine distally, long, about 0.8 distance between
proximal end of first antennular article and mesial base of distolateral
spine; distolateral spine extending to about midlength of second article;
antennular flagella short, mesial one gradually tapering distally, slightly
shorter than lateral, length of latter equivalent to that of second and third
articles combined.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

776

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VOLUME 93, NUMBER 3 WI

Fig. 2. Sicyonia olgae, paratype ¢ 10 mmcl, off Punta Las Tunas, Puerto Rico: a, Petasma,
dorsal view; b, Ventral view of same; c, Appendix masculina, dorsolateral view.

Scaphocerite reaching almost to distal end of antennular peduncle; length
of antennal flagellum about 2.4 times that of carapace.

Third maxilliped stouter than pereopods, reaching distal end of antennular
peduncle or at most exceeding it by dactyl and distal 0.1 of propodus.

First pereopod reaching between proximal and distal ends of middle half
of carpocerite. Second pereopod extending to distal end of carpocerite or
surpassing it by 0.5 length of dactyl. Third pereopod overreaching anten-
nular peduncle by tip of dactyl or at most by length of propodus, extending
only slightly farther than third maxilliped. Fourth pereopod reaching be-
tween base and midlength of merocerite. Fifth pereopod extending slightly
beyond fourth, at most to basal 0.2 of carpocerite.

Abdomen covered dorsally with fine setae, studded with minute tubercles,
densely distributed on anterior 4 somites, bearing dorsomedian carina from
first through sixth somites; carina on fifth truncate, ending in abrupt angle,
and that on sixth in conspicuous tooth. First somite with anteromedian
pleural sulcus (beginning at notch on anterior margin of tergum) joining
coalescent posterior tergal and posteromedian sulci ventrally, lacking pos-
terior pleural sulcus. Second and third somites bearing anteromedian and
posteromedian pleural sulci and anterior and posterior tergal sulci, lacking
posterior pleural sulcus. Fourth and fifth somites with posterior tergal and
posteromedian pleural sulci coalescent and continuous with anterior tergal

778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Sicyonia olgae, holotype. Thelycum, ventral view.

sulcus. Sixth somite bearing deep, strongly arched posterior pleural sulcus.
Pleura of first 4 somites with anteroventral extremity angular, somites end-
ing in acute tip; fifth and sixth somites rounded anteroventrally. Postero-
ventral margin of first 3 somites rounded, that of fourth broadly obtuse, and
that of fifth and sixth bearing posteriorly directed spine. Telson with short,
triangular terminal portion flanked by minute pair of fixed subterminal
spines.

Petasma (Fig. 2a—b) with distal part of dorsomedian lobule bearing small,
inwardly curved, convex plate. Dorsolateral lobule produced distally in cor-
nified, tear shaped, acutely pointed projection, bearing proximodorsal sub-
hemispherical prominence, and longitudinal ventral groove; dorsolateral lob-
ule forming inconspicuous, rounded proximal process. Ventrolateral lobule
heavily cornified, produced distally in fleshy, tapering, laterally directed
short projection, with tip sometimes curved proximally; latter projection

VOLUME 93, NUMBER 3 779

situated contiguous to and falling distinctly short of cornified projection of
dorsolateral lobule.

Appendix masculina (Fig. 2c), projecting from free distal part of low ridge
on dorsomedian margin of endopod, small, roughly campanulate, with ven-
tral wall produced distally beyond dorsal one.

Thelycum (Fig. 3) with plate of sternite XIV forming sharp, rounded lat-
eral flanges and bearing obliquely disposed, oblong bulges separated by
broad median depression; latter extending to posterior thoracic ridge. Me-
dian plate of sternite XIII triangular in outline anteriorly, tapering into long
slender spine reaching about midlength of coxae of second pereopods; plate
constricted and deeply excavate at level of coxal plates of fourth pereopods,
then continuing posteriorly into short component with rounded posterolater-
al margins (hiding apertures of seminal receptacles) separated by shallow
angular to broadly obtuse median emargination. Posterior thoracic ridge
with anteromedian margin concave, and fused laterally with plates of ster-
nite XIV. Paired long, slender spines projecting anteriorly from posterior
margin of sternite XI, flanking and extending as far as or beyond spine of
median plate of sternite XIII, reaching at least midlength of coxae of second
pair of pereopods.

Sizes: males 9-10 mm carapace length, about 36-38 mm total length;
females 11-14 mm carapace length, 40—44 total length.

Geographic and bathymetric ranges.—This species has been found in
waters off Dry Tortugas Is., Florida, through the Antilles southward to
Suriname at depths between about 33 and 622 m. Three of the four available
samples were obtained at depths of no more than 70 m. Substrates at each
of the four localities were different, consisting of mud, shell, coral, or
‘““specks’’ respectively.

Affinities: Sicyonia olgae is closely allied to the western Atlantic S. typica
(Boeck, 1864). The two possess a highly vaulted carapace and an elevated
postrostral carina armed with 2 teeth posterior to the level of the hepatic
spine.

Sicyonia olgae is distinguished from S. typica by the following: the ar-
mature of the rostrum, consisting of 3 or 4 teeth (not counting tip) instead
of only 1 or 2; the presence of a well defined sublateral carina which is
lacking or occasionally barely distinct in S. typica; the absence of posterior
pleural sulci on the first 3 abdominal somites that are always clearly distinct
in S. typica; the shorter stylocerite, which extends about 0.8 of the distance
between the proximal end of the first article of the antennular peduncle and
the mesial base of the distolateral spine, but in S. typica extends 0.9 the
distance, or almost reaches the base of the spine. In females of S. olgae,
sternite XI is armed with a pair of long, slender spines, that reach at least
midlength of the coxae of the second pair of pereopods, and the postero-
lateral corners of the median plate of sternite XIII are rounded and separated

780 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by a shallow angular or rounded emargination. In contrast, in S. typica the
spines are small and the posterolateral corners of the median plate of sternite
XIII are subrectangular and separated by a deep subrectangular emargina-
tion. Also in males of S. olgae the dorsolateral lobule of the petasma is
produced distally in a sharp, mesially directed projection, whereas in S.
typica this projection is blunt and twisted—turning mesially and then dor-
sally.

Etymology: This species is named in honor of my sister Dr. Olga Lanio,
who through the years followed with utmost interest my shrimp studies.

Acknowledgments

Horton H. Hobbs, Jr., and Fenner A. Chace, Jr., my colleagues at the
Smithsonian Institution, read the manuscript. I am indebted to them for
improvements resulting from their suggestions. Maria M. Diéguez prepared
the illustrations.

Literature Cited

Boeck, A. 1864. Beskrivelse og fremlagde Tegninger af 4 norske Decapoder, undersggte af
Overlaege Danielssen og ham.—Forhandlinger Videnskabs-Selskabet, Christiania
1(1863): 189-190.

Burkenroad, M. D. 1934. Littoral Penaeidea chiefly from the Bingham oceanographic collec-
tion. With a revision of Penaeopsis and descriptions of two new genera and eleven new
American species.—Bull. Bingham Oceanogr. Collect. 4(7), 109 p.

Chace, F. A. 1972. The shrimps of the Smithsonian-Bredin Caribbean expeditions with a
summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia).—
Smith. Contrib. Zool. 98, 179 p.

Hay, W. P., and C. A. Shore. 1918. The decapod crustaceans of Beaufort, N. C., and the
surrounding region.—Bull. U.S. Bur. Fish. 35:369-475.

Huff, J. A., and S. P. Cobb. 1979. Memoirs of the Hourglass cruises: penaeoid and sergestoid
shrimps (Crustacea: Decapoda).—Fla. Dep. Nat. Resour. Mar. Res. Lab. 5S, pt. 4,
102 p.

Kubo, I. 1949. Studies on penaeids of Japanese and its adjacent waters.—J. Tokyo Coll. Fish.
36(1), 467 p.

Lucas, H. 1846. Crustacés. Histoire naturelle des animaux articulés.—Exploration scientifique
de |’ Algérie pendant les années 1840, 1841, 1842, 1:1-88.

Pérez Farfante, I. 1954. Los camarones comerciales de Cuba. Parte 2.—Contrib. Centro

Invest. Pesq. 6, Habana, 31 p.

. 1969. Western Atlantic shrimps of the genus Penaeus.—U.S. Fish Wildl. Serv., Fish.

Bull. 67:461-591.

Stimpson, W. 1871. Notes on North American Crustacea, in the museum of the Smithsonian
Institution, [1].—Ann. Lyc. Natur. Hist., N.Y. 10:92-136.

Systematics Laboratory, National Marine Fisheries Service, National
Museum of Natural History, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 781-796

STENOPUS PYRSONOTUS, A NEW SPECIES OF
STENOPODIDEAN SHRIMP FROM THE INDO-WEST
PACIFIC REGION (CRUSTACEA: DECAPODA)

Joseph W. Goy and Dennis M. Devaney

Abstract.—A new species of shrimp in the genus Stenopus from the Indo-
West Pacific region is described and figured. Specimens have been collected
from the Hawaiian Islands and Mauritius. The new species, Stenopus pyr-
sonotus, is Closely related to S. hispidus but is easily distinguished by mor-
phological differences in the rostrum, telson, and spinulation of the body as
well as by coloration. A Key to the 4 described species from the Indo-West
Pacific is included.

Three of the 5 previously reported species of Stenopus are found in the
Indo-West Pacific region. Stenopus hispidus has a circumtropical distribu-
tion (Holthuis, 1946), S. tenuirostris is known from the Indian Ocean and
Malay Archipelago (Bruce, 1976), and S. zanzibaricus was recently de-
scribed from off the coast of Zanzibar (Bruce, 1976). Collections over sev-
eral years at Mauritius and in the Hawaiian Islands have provided some
interesting specimens of a Stenopus that show a striking color pattern dif-
fering markedly from the other three Indo-West Pacific species. Besides this
distinctive coloration, morphological characters distinguish the specimens
from the other species in the genus. They are described here as representing
a new species. Specimens recorded herein are deposited in the Bernice P.
Bishop Museum (BPBM), Honolulu and the National Museum of Natural
History (USNM), Washington, D.C. All material recorded below is consid-
ered in the type-series.

Stenopus pyrsonotus, sp. nov.
Figs. 1-6

Material examined.—HAWAIIAN ISLANDS: OAHU, off Pokai Bay,
depth 22.9 m, 13 July 1975, G. Fukuda coll., USNM 173917 (Holotype),
USNM 173918 (Allotype), USNM 173919 (Paratype); off Pokai Bay, depth
24.4 m, 29 July 1969, J. E. Randall, S. Swerdloff and D. Chave coll., BPBM
$8577, 1 specimen; off Lahilahi Point, in cave, depth 27.4 m, 11 July 1970,
J. E. Randall coll., BPBM S8287, 1 specimen; %4 mile east of Manana Islet
(Rabbit Island), from bilge tank of sunken boat, depth 68 m, 14 Oct. 1970,
coll. W. D. Madden et al., BPBM S7887, 3 specimens; south of Pokai Bay,
depth 22.9 m, under ledge along reef outcrop, 14 Dec. 1972, D. M. Devaney

782 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

~~
=~

Fig. 1. Stenopus pyrsonotus, holotype, male. Scale in mm.

coll., BPBM S8578, 1 specimen; off Makua, in cave, depth 36.6 m, 19 Aug.
1973, P. Lobel coll., BPBM S8491, S8492, S8493, 3 specimens; Sharks
Cove, north shore, in small crevice in side of lava opening, depth 21.3 m,
22 Aug. 1976, G. M. Ludwig coll., BPBM S8579, | specimen. HAWAITI,
Pada Bay, South Kona, 19°14’N lat., 155°55’W long., depth 32 m, 27 June
1976, G. M. Ludwig coll., BPBM S8580.

MAURITIUS: West coast of La Fouche, Medine, in cave, depth 30 m,
19 Nov. 1973, D. Pelicier, J. E. Randall coll., BPBM S8494, 1 specimen;
West coast, Cathedral Reef, off Flic en Flac, in cave, depth 30.5 m, 20 Nov.
1973, J. E. Randall, D. Pelicier coll., BPBM S8495, 1 specimen.

VOLUME 93, NUMBER 3 783

Diagnosis.—A moderately large stenopodidean shrimp, with slender,
compressed body, generally covered with spinous processes; telson lacking
marginal spine; scaphocerite with distal marginal tooth separated from oth-
ers; with red mid-dorsal abdominal stripe.

Description.—Holotype (male, USNM 173917). Rostrum (Figs. 1, 2A)
long, extending past middle of last antennular peduncle segment, directed
slightly upwards, gradually tapering to apex. Dorsal margin with 5 strong
spines, distal largest, reaching beyond tip. Ventral margin slightly concave
proximally with 3 spines distally, numerous proximal setae. Lateral margins
with 3 right spines, 4 left spines, carinae poorly developed.

Carapace (Fig. 2A) densely covered with spinules placed in more or less
distinct longitudinal rows. Spinules most strongly developed anterodorsally,
smallest over lower and posterior branchiostegite, directed anteriorly. Cer-
vical groove distinct with 5 spines along upper margin. Orbit with small but
distinct supraorbital spine. Inferior orbital angle bearing large tubercle and
large spine. Large antennal spine present, small hepatic spine at lower end
of cervical groove. Ventrolateral angle somewhat rounded, only ‘slightly
produced anteriorly while posterolateral angle of branchiostegite obtusely
angled and slightly concave.

First 3 abdominal somites with similar forwardly directed spinules similar
to those on carapace. First somite with 2 irregular transverse rows of spi-
nules, pleuron ending in | weak and 2 strong spines. Second somite with 3
transverse rows of spinules. Pleura of second to fifth somite truncate, each
ventrolateral margin with 3 strong spines. Third somite (Fig. 2B) broadly
produced with no bare shield-shaped area present. Distolaterally directed
spinules of fourth somite arranged in 3 somewhat parallel rows, depressed
and squamose; those of fifth somite similar but in somewhat less distinct
rows. Sixth somite bearing 2 longitudinal rows of 6 large spinules in mesial
region, a medial row of 8 smaller spinules, scattered proximal transverse
spinules, and additional lateral spines. Each abdominal somite bearing
strong ventral median spine. Ventral surface of sixth somite also densely
covered with spinules. Length of fifth abdominal somite 0.4 times that of
sixth somite which is about 1.5 times longer than its greatest breadth.

Eyes (Fig. 2C) well developed, with peduncle slightly longer than cornea.
Ophthalmic peduncle dorsally with 4 small spinules, 3 extending over cor-
nea; proximodorsal side with 4 small spinules, one extending over cornea.

Telson (Fig. 2D) lance-shaped, with median groove flanked by 2 distinct
longitudinal carinae. Telson length approximately 1.6 times that of sixth
abdominal somite, 2.5 times longer than anterior width of telson. Carinae
provided with 8 left, 6 right strong, posteriorly directed spines, some having
1 or 2 long plumose setae at outer basal side. Strong anterior spine present
laterally at telson base, with 5 pairs of median spines occurring between
each carina and median groove and extending slightly less than half way

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

784

io
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dorsal view; D, Telson; E, Uropods; F, Epistome

male: A, Carapace, dorsal view; B, Third to sixth
and labrum, ventral view. Scale bars represent 1.0 mm.

Stenopus pyrsonotus, holotype,

Fig. 2.
abdominal somites, dorsal view; C, Left eye,

VOLUME 93, NUMBER 3 785

down telson length. Lateral margin lacking spine, provided with 48 long
plumose setae along posterior two thirds. Posterior margin rounded, with
last 2 longitudinal carinal spines overlapping margin.

Uropods (Fig. 2E) well developed with endopodite exceeding telson tip
but narrower and shorter than exopodite. Basal segment stout with single
mesial spine, three outer marginal plumose setae. Exopodite bearing nine
acute teeth and 16 short plumose setae on outer margin. Dorsal surface
bearing distinct median ridge with 2 longitudinal rows of 6, 13 spines outside;
2 longitudinal rows of 4, 9 spines and 7 short plumose setae inside. Outer
proximal margin of endopodite bearing 6 teeth, 5 distal short plumose setae.
Dorsal surface bearing distinct median ridge with scattered spines, denser
proximally, 4 short plumose setae outside; distal row of 6 short plumose
setae inside. Unarmed margins of exopodite, endopodite provided with 36,
52 long plumose setae respectively.

Epistome (Fig. 2F) triangular anteriorly with 2 stout submedian spines
next to small median rounded area. Three pairs of spines laterally; labrum
normally developed. Paragnath (Fig. 3A) bilobed with lobes separated by
median fissure terminating in 2 rounded humps. Thoracic sternites narrow
with 2 submedian spinules on segments 4—6, 8 submedian spinules on seg-
ments 7-8.

Antennular peduncle (Figs. 3B, C) short, extending to middle of scapho-
cerite. Basal, middle segments about same length, distal segment consid-
erably shorter. Basal segment about twice as long as wide with broad rect-
angular process on inner dorsobasal margin; small elongate stylocerite on
outer margin, tapering to sharp point; inner distodorsal angle produced into
prolongated ridge with 3 strong spines, dorsomedially 2 strong spines, 2
small spines and few setae on outer distodorsal angle. Middle segment with
5 spines on ventral margin, | distodorsally, 1 spine on inner proximal mar-
gin, also 3 large spines extending out dorsally. Distal segment about 0.6
times length of basal segment with | spine distodorsally followed by 2 setae
at inner distal margin, smaller medial spine ventrally. Upper flagellum well
developed with proximal part bearing 29 groups of aesthetascs beginning on
tenth article ending on eighteenth. From article 2 to 15, 16 dorsal spines.
Lower flagellum not as strongly developed as upper one.

Antenna (Fig. 3D) with strong basal segment bearing 4 large spines.
Scaphocerite well developed, reaching about 2.5 times beyond tip of ros-
trum. Outer margin strongly concave proximally with small basal spine fol-
lowed by 1 much larger, longer spine and 2 small distal spines; upper part
bearing 24 small teeth followed by considerable non-serrate distance up to
final large tooth. Dorsal surface bearing 2 distinct longitudinal carinae, inner
side of medial carina with row of 33 spinules. Inner margin provided with
69 long plumose setae. Ventral surface with medial longitudinal row of spi-

786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 3. Stenopus pyrsonotus, holotype, male: A, Paragnath; B, Right antennule and anten-
nular peduncle, ventrolateral view; C, Left antennular peduncle, dorsal view; D, Antenna and
scaphocerite, dorsal view; E, Mandible, ventral view; F, Mandible, dorsal view; G, Maxillule.
Scale bars represent 1.0 mm.

nules (not shown). Antennal flagellum well developed, extending beyond tip

of telson.
Mandible (Figs. 3E, F) robust with short, fused molar and incisor pro-

cesses. Molar surface with few minute irregular teeth, incisor bearing 9

VOLUME 93, NUMBER 3 787

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Fig. 4. Stenopus pyrsonotus, holotype, male: A, Maxilla; B, First maxilliped; C, Second
maxilliped; D, Third maxilliped. Scale bars represent 1.0 mm.

small medial teeth. Palp well developed, 3-segmented. Proximal segment
without setae; middle segment about one-third longer than proximal segment
bearing 4 small lateral setae, groups of large distal setae; distal segment
broad, densely covered with setae laterally and distally.

Maxillule (Fig. 3G) with slender undivided endopodite about 6.5 times
longer than broad, bearing 14 plumose setae on lateral border and distally.
Proximal endite moderately broad, somewhat truncated distally with con-

788 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Spicuous seta laterally, 15 simple setae distally, few simple setae on lower
inner border. Distal endite broader, rounded distally, bearing numerous sim-
ple setae, few short setae laterally.

Maxilla (Fig. 4A) with following setation on coxal and basal endites: 27
on proximal lobe, 7 on distal lobe of coxal; 14 on proximal lobe, 23 on distal
lobe of basal. Endopodite long, slender, not exceeding anterior margin of
scaphognathite, with 16 lateral plumose setae, 10 distal plumose setae.
Scaphognathite long, narrow, with 143 plumose setae along margin.

First maxilliped (Fig. 4B) bearing 3-segmented endopodite. Proximal seg-
ment long, 2.5 times as long as broad, with 12 long plumose setae laterally,
15S shorter setae on inner margin. Middle segment about 0.7 times length of
proximal segment, with 12 long plumose lateral setae, 5 minute simple me-
sial setae. Distal segment slender, tapering, slightly shorter than middle
segment, with 2 minute simple terminal setae. Basipodite large, rounded
anteriorly with straight outer broder bearing dense fringe of long setae.
Coxopodite bilobed, with each lobe bearing numerous short setae; proximal
lobe partly covered by 15 short plumose setae. Exopodite well developed,
flagellum arising from incomplete peduncle, bearing 6 short plumose setae
proximomesially, 24 longer plumose setae distally. Large epipod with mod-
erately slender proximal and distal lobes.

Second maxilliped (Fig. 4C) with 7-jointed endopodite. Dactylus suboval,
twice as long as broad, with dense fringe of short setae along distodorsal
margin, 12 short setae along distoventral margin. Propodus about 1.5 times
length of dactylus, densely setose on dorsal margin, ventral margin bearing
3 stout proximal teeth. Carpus short, triangular about 0.5 times length of
propodus, with 4 long simple setae at distodorsal angle, 6 shorter simple
setae mesially and laterally. Merus almost 2.0 times length of dactylus, 2.5
times longer than broad, with straight inner border bearing 4 simple evenly
spaced setae; outer border convex with many long simple setae; outer part
distinctly more compressed and broader than inner part. Ischium and basis
not fused, each with dense mesial setae as for coxa; ventral side of ischio-
basis with distinct projection. Exopodite long, slender, undivided with distal
half bearing 23 long plumose setae. Small epipod present.

Third maxilliped (Fig. 4D) endopodite strongly developed, 7-segmented.
Dactylus slender, tapering, about 6.5 times longer than basal width, with 14
simple setae on outer margin, 19 simple setae mesially and on inner margin.
Propodus same length as dactylus but about 5.0 times longer than broad,
bearing | acute spine, few long and short simple setae on outer margin; 4
spines medially; setiferous organ distally; numerous long simple setae along
inner margin. Carpus slightly longer than propodus, about 4.5 times longer
than broad, with 5 spines, 7 long setae on outer margin; row of 5 spines
medially; 4 small spines, numerous long simple setae on inner margin. Merus
long, slender, 1.5 times longer than carpus, with 9 large spines on ventral

VOLUME 93, NUMBER 3 789

outer margin; row of 8 spines medially; 13 small spines, long setae densely
covering inner margin. Ischium robust, flattened, 0.9 length of merus, with
row of 7 large spines, 17 spaced setae, large submarginal distal spine on
outer margin; mesial row of 9 large spines, 18 short medial setae; 11 large
spines covered by dense fringe of long setae on inner margin. Basis short,
rounded, with 5 long setae on inner margin, laterally bearing well developed
exopodite almost equal to ischium length with 28 plumose setae on distal
half. Coxa short, with small lobe bearing 3 setae on inner margin; laterally
epipod slender, tapering with solid rounded coxal lobe dorsally.

First pereiopod (Figs. 5A, B) small, slender, when stretched reaching just
past scaphocerite, all segments generally glabrous. Dactylus less than half
length of propodus. Fingers slightly compressed, having somewhat hooked
tips. Cutting edges indistinct with only dactylus bearing chitinous ridge
along inner margin. Dactylus covered with numerous short simple setae,
propodus covered with numerous short and long simple setae. Fingers and
distodorsal extremity of palm bearing small tufts of long setae. Distoven-
tral part of carpus and proximoventral part of propodus provided with se-
tiferous organ. Carpus longest segment about 1.3 times length of propodus,
narrowing slightly proximally. Merus about 0.8 times length of carpus, is-
chium about of equal length, both bearing numerous short and long simple
setae. Basis short, bearing 5 ventral setae; coxa stout, unarmed, but bearing
small epipod.

Second pereiopod (Figs. SC, D) similarly built as first, but longer, strong-
er. No setiferous organ present. Finger tips more strongly hooked than first,
cutting edges each with small rounded proximal tooth, dactylar tooth slightly
in advance of one on propodus; also edges provided with 18 small, stout,
peg-like teeth separated by rectangular chitinous lamellae. Fingers and dis-
todorsal extremity of palm bearing small tufts of long setae, surfaces of
dactylus, propodus covered with few simple setae. Carpus same length as
propodus, slightly longer than merus or ischium, bearing 8 dorsal spines,
generally covered with numerous long setae. Merus, ischium of equal length
about 0.7 times length of carpus, with merus bearing 9 curved spines, 16
long setae dorsally; ischium with few long setae. Basis short with 4 ventral
setae; coxa stout with small epipod.

Third pereiopod (Fig. 5E) largest, strongest, longer than entire length of
body, extending beyond scaphocerite by half length of merus and length of
carpus and chela. Palm of chela longest segment with dorsal row of 23 strong
forward curved spines, below this 15 smaller spines dorsomesially, followed
by median groove with 17 similar sized spines. Ventral margin bearing 26
strong forward curved spines, 15 smaller spines ventromesially, less distinct
row of 11 small spines on outer side. Between dorsal, ventral marginal
spines numerous long simple setae. Fingers elongate, with sharp hooked
crossing tips. Dactylus bearing 10 dorsal acute spines, 3 smaller spines dor-

790 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

| |
Kaa

NEC Crete

g

Fig. 5. Stenopus pyrsonotus, holotype, male: A, First pereiopod; B, Chela of first pereio-
pod; C, Second pereiopod; D, Chela of second pereiopod; E, Third pereiopod, mesial view;

F, Fourth pereiopod; G, Fifth pereiopod; H, First pleopod; I, Second pleopod. Scale bars
represent 1.0 mm.

somesially at proximal end, numerous long simple setae. Dactylar cutting
edge bearing 4 small sharply pointed teeth proximally merging into row of
14 stout, peglike teeth separated by rectangular chitinous lamellae. Cutting
edge of propodus with large blunt tooth, more acute tooth proximally, op-
posing 4 small proximal dactylar teeth; also distally bearing 14 peglike teeth
separated by chitinous lamellae. Fingers distally bearing small tufts of long

VOLUME 93, NUMBER 3 791

setae. Carpus about 0.3 times shorter than propodus narrowing gradually
proximally. Dorso-outer margin bearing 19 spines, 18 smaller spines dor-
somesially; ventral margin with 13 spines, less distinct row of 10 small spines
ventromesially. Scattered long setae on dorsal, mesial, outer, and ventral
margins. Merus almost as long as propodus, narrower than carpus, with
dorso-outer row of 19 large spines, 11 smaller spines dorsomesially, 11 large
spines on inner side. Ventral margin with 17 large spines, 7 smaller spines
ventromesially, 8 spines on outer side. Numerous long setae interspaced
between dorsal, ventral marginal spines. Ischium short, about half length of
propodus with 14 spines on inner side, 12 spines on ventral margin, 5 ven-
tromesial spines, few short simple setae. Basis and coxa short, fragile with
1 small spine and 5 spines respectively on ventral margins, few short setae,
small epipod.

Fourth, fifth pereiopods (Figs. 5F, G) long, slender, very similar. Dactylus
of fourth biunguiculate with unguis long, curved, not clearly separated from
dactylar corpus; accessory spine short, straight, about half length of unguis.
Propodus subdivided into 7 segments bearing 25 movable ventral spines, 23
long dorsal setae. Carpus slender, straight, longest segment of pereiopod,
subdivided into 15 smaller segments with 6 small spines on distoventral
margin of distal 7 segments; 29 long, well spaced, unbranched setae. Merus
slender, elongate, with 5 ventral spines, 9 dorsal spines, 5 long setae at
distodorsal extremity. Ischium, basis unarmed except for few long setae.
Coxa ventrally bearing large distal spinule, 2 smaller proximal spinules.
Fifth pereiopod with similar dactylus as fourth, but unguis not as long with
small distal ridge. Propodus subdivided into 5 segments with 20 movable
ventral spines, 12 long dorsal setae. Carpus longer than fourth pereiopod’s,
subdivided into 14 segments with 5 small distoventral spines on distal 5
segments, | proximal dorsal spine, 16 long dorsal setae. Merus with 5 ventral
spines, | proximal and distodorsal spine, 4 long setae at distodorsal extrem-
ity. Ischium, basis unarmed except for few long setae; coxa bearing 3 ventral
spinules.

First pleopod (Fig. 5H) uniramous, second (Fig. 51) to fifth biramous, all
lacking appendices. First pleopod smallest, with exopodite equal in length
to basipodite. Dorsal margin of basipodite with single mid-proximal spine,
2 distal plumose setae; ventral margin with median plumose seta. Exopodite
bearing 28 plumose marginal setae. Rami of second pleopod about twice
length of basipodite. Ventral margin of basipodite bearing 5 spines, 4 simple
setae; | spine ventromesially. Ventral surface of exopodite bearing row of
6 spines, 30 plumose marginal setae; that of endopodite with 8 spines, 36
plumose marginal setae. Third to fifth pleopods generally similar, decreasing
in size, spinulation posteriorly.

Branchial formula:

Te PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Maxillipeds Pereiopods

I Il Il I Te TS Ve ay)
Pleurobranchs — | I l l 1 l l
Arthrobranchs — | Dy 2 yy y Ppt
Podobranchs —- tt — SS SS ee
Epipods l | l 1 1 lL —
Exopods l — eS eee

Measurements: The measurements of the holotype are included in Table
1 and compared with other specimens examined.

Coloration: The carapace, eyestalk, antennular peduncle, antennal scale,
and rostrum are a pinkish-white. The antennal flagella are all completely
Opaque white. The third maxillipeds are a translucent pink. The coxae,
basises, ischia and meri are translucent pink on all the pereiopods, and the
rest of these appendages are opaque white. The abdomen is largely white
except for a red central stripe covering about one third of the dorsal surface
and extending from the first abdominal somite to the sixth somite (often
with a very narrow central white portion) becoming broader by the third
abdominal somite. The telson is white and the uropods are a translucent
pink.

Allotype: (female, USNM 173918). Generally similar to holotype, but dif-
fering in the following respects. The rostrum has 7 dorsal, 2 ventral, 3 left
lateral, and 4 right lateral teeth. The telson’s longitudinal carinae each bear-
ing 7 strong spines. The mouthparts were removed from the right side, but
show no major differences. The dactylar cutting edge of the third pereiopod
has the 4 proximal teeth coalesced into one large sharp tooth. There are no
ventral median spines present on the abdominal somites. The first pleopod
is not shorter relative to the rest and the exopodite is narrow and acuminate,
longer than the basipodite, which lacks the proximal spine on its outer
margin.

Paratypes: The new species is variable in the number of body and appen-
dage spines. For example, our specimens showed the rostrum with from 5—
7 dorsal, 1-7 ventral, and 0-4 lateral spines (Fig. 6). The number of spines
on the proximal part of the outer margin of the scaphocerite in our material
varies from 2 to 4, while the number on the distal part varies from 15 to 30.
The third maxilliped sometimes has a small spine proximal to the setiferous
organ. The first and second pereiopods sometimes have short proximal spi-
nules on their ischia and basises. Some specimens have minute peglike teeth
separated by chitinous lamellae on the cutting edge of the first pereiopod.
The basipodite of the first pleopod has 0-3 spines. We could not find any
correlation between the number of spines and the size or sex of the animals,
but some of these differences probably reflect allometric growth changes
and the normal variation in the species. Some specimens have the carapace
and proximal parts of the walking legs tan or light golden brown.

793

VOLUME 93, NUMBER 3

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794 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Gina C

Fig. 6. Stenopus pyrsonotus, paratypes: A, Rostrum BPBM S8577; B, Rostrum BPBM
S7887; C, Rostrum BPBM S8491. Scale bars represent 1.0 mm.

A large ovigerous specimen (BPBM S8493) has a medial ventral abdom-
inal spine present on each segment except the sixth which bears 2 spines.
A juvenile character of the new species is the absence of pleopods on the
first abdominal segment in specimens with rostral carapace length less than
11 mm or post-orbital carapace length less than 7 mm (BPBM S8287, S8580).

Type locality.—Hawaiian Islands, Oahu, west coast off Pokai Bay.

Habitat.—Most of the specimens were taken in crevices, caves, or under
ledges on well developed coral reef systems. This cryptic behavior is also
exhibited by other members of the genus. In at least 2 observations, the
new species was found with a large green-bordered moray eel, Gymnothorax
flavimarginatus. The specimen from Pada Bay, a juvenile, was one example
seen with an eel, but other large specimens were also said to have been
seen with this eel species (G. M. Ludwig, written communication). These
observations and the fact that these shrimp have long prominent white an-
tennal flagella, may indicate that the new species enters into cleaning sym-
biosis with cooperating fish, much like its congener, Stenopus hispidus.

VOLUME 93, NUMBER 3 795

Etymology.—The specific name is derived from the Greek ‘‘pyrsonotos’’
meaning red-backed; it is given because of the distinctive deep red stripe
on the dorsal surface of the abdomen.

Remarks.—The new species, Stenopus pyrsonotus, follows closely the
definition of the genus Stenopus Latreille given by Holthuis (1946), only
differing in the absence of the lateral spines on the telson. This character
separates it from all other species in the genus. It is most closely related to
S. hispidus, but differs in color, rostrum, telson and the body spines being
less erect.

Key to the Described Indo-West Pacific
Species of Stenopus
1. Lateral marginal spines absent on telson; abdominal somites with

broadtmid=dorsal redksttipe w45 see «ee S. pyrsonotus sp. nov.
— Lateral marginal spines present on telson; abdominal somites oth-
EiNvISesDICIneMuCde tone. ee FOU IRAE TEI) TE ed ee eS, 2

2. Outer margin of scaphocerite serrate up to final (terminal) tooth;
abdomen with oval red patches on lateral sides of second and fifth

S@MMCC Stead cass Why MLTR E tee oer tnesti tian: insta? S. zanzibaricus Bruce, 1976
— Outer margin of scaphocerite with considerable nonserrate space
before final (terminal) tooth; abdomen otherwise pigmented....... 3

3. Rostrum exceeding antennular peduncle in length, with 2—9 ventral
spines; abdomen with red transverse bands on third and fifth somites
5 65 RTE LE I rs ee eee eee ae S. tenuirostris De Man, 1888
— Rostrum not exceeding middle segment of antennular peduncle in
length, without ventral spines; abdomen with red transverse bands
Onethindeand spxtheSomites’. 5-4 eee S. hispidus (Olivier, 1811)

Two species of stenopodidean shrimp have so far been reported from the
Hawaiian Islands, Stenopus hispidus (Rathbun, 1906; McNeill and Ward,
1930; Edmondson, 1946; Tinker, 1965; Hobson and Chave, 1972; Titcomb
et al., 1979) and Spongicola henshawi (Rathbun, 1906). Stenopus hispidus
has also been recorded from Mauritius (White, 1847; Hoffmann, 1874) along
with many Indo-Pacific and Western Atlantic records reported by Holthuis
(1946). The only other stenopodidean shrimp recorded from Mauritius be-
long to the genus Microprosthema. Richters (1880) reported Microprosthe-
ma validum, M. scabricaudatum and M. plumicorne from fle aux Fougquets,
Mauritius. Stenopus tenuirostris is moderately common on western Indian
Ocean reefs according to Bruce (1976). It has also been reported from nu-
merous Indonesian localities (Holthuis, 1946), the Seychelles and Coetivy
Islands (Borradaile, 1910), the Solomon Islands (Yaldwyn, 1968) and New
Caledonia (Monod, 1976). Stenopus zanzibaricus was reported on the basis
of a single specimen taken from Mwemba Island off Zanzibar (Bruce, 1976),
but one of us (JWG) has examined another small male (total length 19.0
mm) collected from Canton Island (USNM 181402).

796 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A specimen of yet another new species of Stenopus was recently collected
off Makua, Oahu. This was an ovigerous female (total length approx. 24
mm) that differs in coloration and numerous morphological characters from
S. hispidus and S. pyrsonotus. Preliminary comparison with other descrip-
tions indicates a closer relationship to the Western Atlantic S. scutellatus,
and a full description will be presented in a forthcoming publication.

Acknowledgments |

We are grateful to Dr. John E. Randall, of the Bernice P. Bishop Museum,
who made it possible to examine several specimens of Stenopus pyrsonotus
he collected and for providing the photograph of the new species taken off
the west coast of Oahu.

Literature Cited

Borradaile, L. A. 1910. Penaeidea, Stenopidea, and Reptantia from the western Indian
Ocean. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the
leadership of Mr. J. Stanley Gardiner, M.A., vol. 2, pt. 10—Trans. Linn. Soc. Lond.
Zool. ser. 2, 13:257-264.

Bruce, A. J. 1976. Studies on Indo-West Pacific Stenopodidea. I. Stenopus zanzibaricus sp.
nov., a new species from East Africa.—Crustaceana 31(1):90-102.

Edmondson, C. H. 1946. Reef and shore fauna of Hawaii.—B. P. Bishop Mus., Spec. Publ.
22:1—381.

Hobson, E. and E. H. Chave. 1972. Hawaiian reef animals.—University Press of Hawaii,
Honolulu, Hawaii, 135 pp.

Hoffmann, C. K. 1874. Crustacés et Echinodermes de Madagascar et de I’ Ile de La Réunion.—
In F. P. L. Pollen and D. C. van Dam, Recherches sur la Faune de Madagascar et des
ses Dépendances 5(2):1—58.

Holthuis, L. B. 1946. Biological results of the Snellius Expedition XIV. The Decapoda Ma-
crura of the Snellius Expedition. I. The Stenopodidae, Nephropsidae, Scyllaridae, and
Palinuridae.—Temminckia, 7:1—178.

McNeill, F. A., and M. Ward. 1930. Carcinological notes. No. I.—Rec. Australian Mus.
12357 —383).

Monod, T. 1976. Sur quelques Natantia (Crust. Déecapodes) de Noumea (Nouvelle Calédon-
ie).—Cahiers du Pacifique 19:7—28.

Rathbun, M. J. 1906. The Brachyura and Macrura of the Hawaiian Islands.—Bull. U.S. Fish.
Comm. 23:837—930.

Richters, F. 1880. Decapoda.—In K. Mobius, Beitrage zur Meeresfauna der Insel Mauritius
und der Seychellen, pp. 139-178.

Tinker, S. W. 1965. Pacific Crustacea.—Charles E. Tuttle, Rutland, Vermont, 134 pp.

Titcomb, M. et al. 1979. Native use of marine invertebrates in old Hawaii.—Pacific Sci.
32(4):325—386.

White, A. 1847. List of species of Crustacea in the collection of the British Museum. 143 pp.,
London.

Yaldwyn, J. C. 1968. Records of, and observations on the coral shrimp genus Stenopus in
Australia, New Zealand and the South-west Pacific.—Australian Zool. 14(3):277-289.

(JWG) Duke University Marine Laboratory, Pivers Island, Beaufort,
North Carolina 28516; (DMD) Bernice P. Bishop Museum, Department of
Zoology, P.O. Box 19000-A, Honolulu, Hawaii 96819.

eee

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 797-829

ONUPHIDAE (POLYCHAETA) FROM BELIZE,
CENTRAL AMERICA, WITH NOTES
ON RELATED TAXA

Kristian Fauchald

Abstract.—Material collected at Carrie Bow Caye and other locations in
Dangriga district, Belize, can be assigned to the following taxa: Americon-
uphis magna (Andrews), Diopatra cuprea (Bosc), Onuphis (Nothria) dan-
grigae, new species, Onuphis (Onuphis) geminata, new species, O. (O.)
pulchra, new species, and O. (O.) virgata, new species. O. (N.) inter-
media Kinberg, O. (N.) veleronis, new name, O. (O.) fragilis Kinberg, and
O. (O.) lineata, new name, are discussed based on examination of the type
materials.

Introduction

Twenty species of onuphid polychaetes are presently known from Gulf
of Mexico and the Caribbean Seas (Table 1). Of these, about half are known
only from waters deeper than 200 m, at least as far as their distribution in
this region is concerned. The family appears poorly represented in shallow
water and intertidal areas here compared to regions at similar latitudes in
the eastern Pacific Ocean (Fauchald, 1968; Hartman, 1968). Especially strik-

_ ing is the dearth of species of Diopatra in shallow water. Members of the

genus Onuphis are frequent in slope and bathyal depths, whereas Diopatra
is more species-rich in the shallows of the warm water regions of the eastern
Pacific Ocean and in the Mediterranean Sea (Fauvel, 1923; Fauchald, 1972).
Paxton (1979) demonstrated, however, that members of the genus Onuphis
are common in shallow water in Australia.

The discrepancy in representation of the two genera between the eastern
Pacific Ocean and the Western Atlantic, could be caused by differences in
collecting efforts; however, the presence of four new species of Onuphis in
very shallow water in Belize, without a corresponding increase in the num-
ber of species of Diopatra, tends to affirm the preliminary impression of a
large-scale difference in the number of shallow water species in the two
genera between the two oceans.

The material was collected as part of the Smithsonian Institution’s Inves-
tigation of Marine Shallow Water Ecosystems (IMSWE-program) coordi-
nated by Dr. Klaus Ruetzler. A series of individual collections were made
by Drs. Meredith L. Jones (stations marked CB-1 through CB-41C), Joan
D. Ferraris (st. JDF-24-1), Brian F. Kensley (K-44 and K-48) and the author
(F-9 through F-33). Most specimens were collected by hand or with shovel,
narcotized using oil of cloves, fixed in approximately 4% formaldehyde in

798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sea water neutralized with hexamine and transferred to 70% ethyl alcohol
for storage. All material is deposited in the collections of the National Mu-
seum of Natural History, Smithsonian Institution, Washington, D.C. Types
of other species were borrowed from Riksmuséet, Stockholm, Sweden and
from the Allan Hancock Foundation, University of Southern California, Los
Angeles.

Station List
Collected by Kristian Fauchald

F-9. 12 April 1979. Carrie Bow Caye, north of the island, sand and small rubble, 30—60 cm
depth.

F-19. 23 October 1979. Carrie Bow Caye, southwest of island, Thalassia and sand, coarse-
screened in 2 mm screen.

F-21. 28 October 1979. Carrie Bow Caye, east of island, about 3 m inside inner edge of
Thalassia-bed, 30-40 cm depth, coarse-screened in 2 mm screen.

F-28. 4 November 1979. Carrie Bow Caye, east of island, about 60 feet from shore, Thalassia
and sand, about 50 cm depth. Coarse-screened in 2 mm screen.

F-33. 8 November 1979. Carrie Bow Caye, north of island, Thalassia, sand and rubble, 60 cm
depth.

Collected by Joan D. Ferraris

JDF-24-1. 30 April 1975. Carrie Bow Caye, lagoon, sand patch N of dock, 30 cm, shovel and
4 inch mesh screen.

Collected by Meredith L. Jones

CB-1. 4 April 1976. Carrie Bow Caye, sparse Thalassia just west of island, | m depth.

CB-2. 5 April 1976. Carrie Bow Caye, northwest of island, general transect from bare sand
area to coral rubble, .5 to 1.5 m depth.
CB-3. 5 April 1976. Carrie Bow Caye, east of laboratory, sand areas mixed with coral

rubble, just inside reef crest, about | m depth.

CB-6. 6 April 1976. Carrie Bow Caye, sieved from sandy areas at top of intertidal around
island, less than 30 cm depth.

CB-7. 6 April 1976. Carrie Bow Caye, associated with old conch shell and sand, on south-
west edge of island, 30 cm depth.

CB-8. 6 April 1976. Carrie Bow Caye, east side of island, associated with coral pieces at
water edge and above.

CB-11. 7 April 1976. Carrie Bow Caye, 100 m north northeast of island, associated with
Acropora curvicornis rubble, stagnated overnight.

CB-14. 9 April 1976. Carrie Bow Caye, back side of reef crest, coral rubble, stagnated
overnight.

CB-16. 11 May 1977. Carrie Bow Caye, sparse Thalassia.

CB-18. 11 May 1977. Carrie Bow Caye, 100 m southwest of island, sieved from coral rubble.

CB-23. 12 May 1977. Carrie Bow Caye, sand immediately behind inner reef crest at far side
of flat, northeast of laboratory.

CB-28. 13 May 1977. Carrie Bow Caye, northwest of island, near inner end of permanent
transect, Thalassia, apparent luxuriant growth, but no compact rhizome masses,
coarse sand, 1—1.5 m depth.

VOLUME 93, NUMBER 3 799

CB-34. 14 May 1977. Carrie Bow Caye, 100 m north of island, 1.5 m depth.

CB-40A. 16 May 1977. Halfway between Colson Point and Salt Creek, Dangriga district, Tha-
lassia, fine sand, almost mud, | m depth.

CB-40B. 16 May 1977. Halfway between Colson Point and Salt Creek, Dangriga district, bare
sand from .5 m depth to shore, much warmer water toward shore, about 15 cm
depth. Fine sand, almost mud.

CB-41C. 17 May 1977. Commerce Bight Pier, Dangriga district, transect over 75—80 m, from
1.5 m depth to shore, at 1.5 m soft brown mud over more compact black mud.

Collected by Brian F. Kensley.

K-44. 8 April 1979. Twin Cayes, rocks and coral rubble, 30 cm depth.
K-48. 10 April 1979. Carrie Bow Caye, coarse calcareous sand in upper spur and groove zone,
no rubble or algae included, 1 m depth.

The terms applied to the morphological features are by and large the same
used by Hartman (1944a) and Fauchald (1968). Some clarification of struc-
tural relations in the parapodia may be useful. In anterior parapodia 2—4
acicula support an acicular lobe which usually is distally truncate or round-
ed. The presetal lobe which is anterior to the acicular lobe, is usually a low
fold that follows the outline of the acicular lobe closely; it is however fre-
quently cut away on the ventral side leaving the acicular lobe exposed (Fig.
Ib). The postsetal lobe usually has a broad, somewhat flattened base and
a digitate or clavate distal part, extending well beyond the acicular lobe
(Fig. 1b and others). The anterior parapodia can be extended or retracted.
In the latter condition a contraction fold (Fig. 3b) may develop across the
parapodium. This fold is normally at the base of either the dorsal or ventral
cirrus or both. The contraction fold, called the low transverse fold, was
treated, undeservedly, as a structural feature by Fauchald (1968); its pres-
ence depends entirely on the state of contraction of the parapodium and
while its presence is interesting it is not of any great taxonomic importance.

The acicula are thick gently tapering rods inside the body; at emergence
they are bent and taper abruptly to fine, needle-like tips.

The setal distribution in the anterior parapodia vary somewhat, but in
general two distinct fascicles of setae can be recognized. Above and in front
of the acicula (i.e. in a superior preacicular position) is found a fascicle of
tapering, limbate setae. Below and slightly inferior to the acicula (i.e. in an
inferior, postacicular position) are found pseudocompound hooded hooks.
A single simple seta may be present directly below the acicula. If compound
spinigers are present, these are always ventral, usually distinctly posterior
to the acicula. The limbate setae shift to postacicular positions by setiger
20 in most species; they remain above the acicula in anteromedian setigers,
but are directly behind the acicula in posterior setigers.

Several different kinds of hooks may be present; in this paper are treated
species with three different kinds. In anterior setigers, usually in 2 to 5

800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

setigers, are found pseudocompound hooded hooks. These hooks may be
distally entire and falcate, or, more usually, bi- or tridentate. Most species
have usually a single sort of hook, either uni- bi- or tridentate, but some
have two sorts of hooks (e.g. both bi- and tridentate).

Another kind of hook may also be present in some anterior setigers. The
large hooks are always tridentate, are usually less distinctly compound and
at least twice as thick-shafted as the pseudocompound ones in the same
setiger. Both kinds of hooks can be present in the same setiger, but the
large hooks are usually present in more setigers than are the pseudocom-
pound ones. 7

A pair of bidentate, hooded subacicular hooks are present in all median
and posterior setigers. Each hook originates above the base of the acicula,
but emerges from the acicular lobe well below the acicula, thus the trajec-
tory of the subacicular hook forms a distinct angle with that of the acicula.

It is possible to have both pseudocompound and large hooks in the same
setiger as mentioned above; likewise, compound spinigers and large hooks
may occur in the same setiger. However, pseudocompound hooks and com-
pound spinigers do not occur in the same setiger. Both compound spinigers
and large hooks usually terminate before the start of the subacicular hooks.
One or several segments without any kinds of hooks may be present be-
tween the last large and the first subacicular hook.

Pectinate setae are present in most median and posterior setigers. The
distal edge may be at right angles to the shaft (transverse) or be at another
angle with the shaft (oblique); the number of teeth vary from about 10 to
more than 25.

The taxonomic treatment in this paper follows traditional lines with one
exception; the inclusion of simple statistical tables demonstrating the pat-
terns of variability within each species is unusual in the polychaete litera-
ture. A couple of interesting and potentially very important features
emerged from this treatment. First, within a population variability is re-
stricted, even over a two-year time span (Onuphis (Onuphis) pulchra at
Carrie Bow Caye); second, in comparing different taxa, the same characters
do not stay invariant for all taxa (cf. the different species of Onuphis (On-
uphis) in the tables below). Two conclusions can be drawn. No single char-
acter can be used to discriminate among three or more taxa even when these
are closely similar. The limited variation exhibited within a taxon from a
single location emphasizes the necessity of extremely careful comparisons
among various populations referred to the same taxon. A simple overlap in
selected character states is insufficient to demonstrate identity between two
populations and a much more exhaustive analysis must be performed in
order to demonstrate such identity.

A character usually avoided in the polychaete literature is the color pat-
tern. These patterns were found to be highly characteristic in the present

VOLUME 93, NUMBER 3 801

material and are described in some detail below; they have been omitted
from the illustrations to avoid clutter. It should be noted that the color
pattern was easily identifiable in alcoholic specimens.

Family Onuphidae Kinberg, 1865
Genus Americonuphis Fauchald, 1973
Americonuphis magna (Andrews, 1891)

Diopatra magna Andrews, 1891a:121, pl. 2, figs. 1-7; 1891b:286-287, pl.
14, figs. 14-20.—Wilson, 1900:351.

Onuphis magna.—Treadwell, 1921:78-81, pl. 7, figs. 1-5, textfigs. 279-
287.—Pearse, 1936:181.—Hartman, 1944a:70; 1944b:21; 1945:24;
1951:51.—Day, 1973:54. ?Monro 1928:89. [Not Monro, 1933:76, nor
Berkeley and Berkeley, 1939:336].

Americonuphis magna.—Fauchald, 1973:22.—Gardiner, 1976:193, fig.
25a—d.

Material examined.—CB-40A (1 specimen).

Remarks.—A. magna can be separated from the closely similar A. reesei
Fauchald (1973) by the branchial structure and maxillary formula. A. magna
has about 12 branchial filaments where the branchiae are best developed,
A. reesei has, maximally, seven branchial filaments. The maxillary formula
for A. magna is 14+1, 8+8, 6+0, 5+9 and 1+1; for A. reesei the formula is
1+1, 10+12, 14+0, 4+9 and 1+1 (this formula was incorrectly reported in
Fauchald, 1973). The specimen reported as Onuphis magna by Berkeley
and Berkeley (1939:336, USNM # 38269) has been re-examined and is here
referred to A. reesei. Monro (1928) did not describe his specimen adequately
so his record must be considered doubtful. There are currently no validated
reports of A. magna from the eastern Pacific Ocean.

Gardiner (1976) pointed out that the generic name Americonuphis was
applied independently by Orensanz (1974) to a different group of species.

~ Paxton (1979:279) rectified the situation by renaming Orensanz’s taxon Aus-
— tralonuphis.

A. magna is known from intertidal and shallow subtidal sand flats from

- North Carolina through the Caribbean Sea and the Gulf of Mexico.

Genus Diopatra Audouin and Milne Edwards, 1833
Diopatra cuprea (Bosc, 1802)

Nereis cuprea Bosc, 1802:143-144 (illustrations published 1830).

Diopatra cuprea.—Pettibone, 1963:250-254, fig. 66a—e (and references
therein).—Gardiner, 1976:185, fig. 23e-1.
Diopatra cuprea cuprea.—Day, 1967:417, fig. 17:12a—d; 1973:54.

Material examined.—CB-1 (11 specimens); CB-2 (2); CB-6 (1); CB-11 (1);
CB-16 (1); CB-28 (6); CB-34 (2); CB-40B (3); CB-41C (5); F-9 (1); F-28 (1).

802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—D. cuprea is usually described as being evenly reddish brown
without distinct dorsal color pattern. Specimens from Belize differ in that
they have scattered brown pigment spots over most of the dorsum and
transverse dark brown bands at setiger 5 or setgers 5 and 6. Most specimens
are small compared to specimens reported from more northerly locations.

D. cuprea 1s very similar to D. ornata Moore (see Hartman, 1968:659 for
a description). The two can be separated by the structure of the anterior
hooks. The proximal tooth of each pseudocompound hook is gently curved
in D. cuprea and nearly parallel to the axis of the hook; it is strongly curved
and nearly at right angles with the axis of the hook in D. ornata (Hartman,
1944a:54). There are also differences in the patterns of papillae on the cer-
atostyles. D. cuprea has papillae of two kinds, organized in 16-18 longitu-
dinal rows, whereas D. ornata has same-sized papillae densely scattered
over the ceratostyles (Hartman, 1944a:54—-55). These characters were re-
examined in the present collections; while somewhat obscure, they appear
absolutely consistent.

D. cuprea has been reported from New England to Brazil in the Western
Atlantic Ocean and along west Africa to the Indian Ocean. In view of the
great similarity between D. cuprea and related species a careful examination
of material from all parts of its range should be undertaken.

Genus Onuphis Audouin and Milne Edwards, 1833
Subgenus Nothria Malmgren, 1867
Onuphis (Nothria) dangrigae, new species
Figs. 1-2, Tables 2-3

Material examined.—CB-1 (1 specimen); CB-3 (9 paratypes, USNM 61225);
CB-6 (4); CB-7 (10); CB-8 (1); CB-11 (3); CB-14 (1); CB-16 (2); CB-18 (1);
CB-23 (1, holotype, USNM 61224); CB-28 (1); K-44 (2); K-48 (2 juveniles);
JDF 24-1 (1).

Description.—The holotype is an incomplete specimen with 66 setigers
that is 28 mm long and 1.5 mm wide with setae. Other, complete specimens
have about 120-130 setigers and are about twice as long as the holotype,
but no wider. All specimens are white and lack color patterns, as preserved.
The anterior part of the body, including the first 5 setigers is cylindrical; in
the rest of the body the dorsal side is strongly flattened and the ventral side
convex. Two anal cirri are present.

The prostomium (Fig. la) is about as wide as long and has a pair of very
small, black eyespots near the base of the frontal antennae. Each frontal
antenna is triangular and slightly pustulate. The ceratophores of all 5 occip-
ital antennae have 4 or 5 rings, of which the distalmost is much longer than
any of the others. The outer lateral antennal styles reach the anterior margin
of the second setiger; the inner lateral and median styles are similar in length

VOLUME 93, NUMBER 3 803

. p
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Fig. 1. Onuphis (Nothria) dangrigae (holotype, USNM 61224): a, Anterior end, dorsal view;
b, Parapodium of setiger 1, anterior view; c, Parapodium of setiger 2, anterior view; d, Para-
podium of setiger 3, anterior view; e, Parapodium of setiger 4, anterior view.

and reach the ninth or tenth setiger. The peristomium is about twice as wide
as the prostomium and carries a pair of peristomial cirri near the anterior
margin. Each cirrus is slender and barely reaches the middle of the prosto-

804 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VULNS
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wo E
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Fig. 2. Onuphis (Nothria) dangrigae (holotype, USNM 61224): a, Parapodium of setiger 6,
anterior view; b, Parapodium of setiger 5, anterior view; c, Compound spiniger, setiger 6; d,
Subacicular hook, setiger 50; e, Pectinate seta, setiger 50; f-o0, Pseudocompound hooks, f-g,
From setiger 1; h-i, From setiger 2; k-l, From setiger 3; m—n, From setiger 4; 0, From setiger 5.

mium. The separation between the two peristomial rings is distinct on the
ventral side, but cannot be seen on the dorsal side.

The parapodia are attached at the anterior edge of each of the 5 first
setigers. The great development of basal parapodial muscles in each of these
setigers makes each segment much wider anteriorly than posteriorly. The
first 3 pairs of parapodia are directed obliquely forward and ventrally. Para-
podium 4 is transitional; from parapodium 5 each parapodium is fully lateral
in position and is situated at about the mid-length of each segment. Further
posteriorly the parapodia take a position at the junction between the flat
dorsum and the curved ventrum and in these segments the dorsal cirri ap-
pear to be attached directly to the dorsum. Each of the first parapodia (Figs.
lb-d) has a distally rounded acicular lobe; the presetal lobe is a low, trans-
verse fold covering the bases of the setae. The postsetal lobe is clavate and

VOLUME 93, NUMBER 3 805

about as long as the parapodium from the base to the tip of the acicular
lobe. The dorsal cirri have distinct cirrophores in the first 3 parapodia.
Thereafter, the cirrophores are indicated only as thickenings near the base
of each cirrus. The dorsal cirri double in length between the first and the
sixth setiger (Figs. Ib—e, Fig. 2a—b) and then decrease slowly in length until
in the last segments (in complete specimens) they are of about the same
length as in the first setigers. The dorsal cirri decrease in width posteriorly
and in far posterior setigers they are slender and nearly filamentous. Ventral
cirri are distinctly cirriform through 4 or 5 setigers and are pad-shaped from
setiger 6. The structure of the ventral cirrus in setiger 5 varies from a short,
tapering cirrus, shorter, but similar in shape to those found in the first 4
setigers, to a glandular pad similar to the ones found in posterior setigers.
In median parapodia the postsetal lobes decrease in size and are recogniz-
able only as short papillae from about setiger 45.

Simple, flattened strap-like branchiae are first present from setigers 18-21.

Limbate and pectinate setae, pseudocompound bi- and tridentate hooks,
compound spinigers and subacicular hooks are present. Limbate setae are
sparsely present in all parapodia in the anterior one-third of the body. All
acicula project beyond the tips of the acicular lobes as slender, slightly bent
needles. The anterior 5 pairs of parapodia have pseudocompound hooks
distributed in the following manner (examined in 5 specimens): In parapo-
dium 1, 4 slender, tridentate hooks (Fig. 2f) with long appendages and one
stout, tridentate hook (Fig. 2g) with short appendage. In parapodium 2, 3
_ Slender, tridentate hooks (Fig. 2h) and 3 stout tridentate hooks (Fig. 21).
The proximal tooth is very small in all slender hooks and 1s difficult to see.
In parapodium 3 the distinction between slender and stout hooks is absent,
but 3 hooks (Fig. 2k) have slightly longer appendages than 2 others (Fig.
21). The proximal tooth is much more reduced in hooks with long append-
ages than in those with short appendages. Parapodium 4 has 3 bidentate
hooks (Fig. 2n) and a single tridentate one (Fig. 2m). In parapodium 5 there
are 3 bidentate hooks (Fig. 20); tridentate hooks are absent. Compound
Spinigers are found in subacicular positions from setiger 6 through the seg-
ment before the start of the subacicular hooks. Each spiniger (Fig. 2c) has
a heterogomph articulation and a narrow, knife-edged blade with fine ser-
rations. Subacicular hooks are first present from setigers 14-16, so the num-
ber of setigers with compound spinigers vary from 8 to 10. Each subacicular
hook (Fig. 2d) is bidentate and has a distinct, short hood. Pectinate setae
are present from setiger 6; each (Fig. 2e) is distally slightly oblique and has
about 12 teeth. One side has a long spur.

The maxillary apparatus is poorly sclerotized. The maxillary formula 1s
1+1, 8+7, 7+0, 7+8 and 1+1 (based on the dissection of two paratypes).
The mandibles are fused over most of their length.

The tubes consist of a thin, pliable inner lining and a loose cover of
unsorted sand grains.

806 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—Other members of the subgenus Nothria with branchiae start-
ing posterior to setiger 10 include O. guadalupensis (Fauchald, 1968:22—24,
pl. 6, figs. e-l), O. rubrescens Augener (1906:139-141, pl. 4, figs. 76-83),
O. stigmatis Treadwell (1922:176—-178, figs. 22-34, see also Hartman,
1944a:89_-91, pl. 11, figs. 240-247) and two species, originally described as
subspecies of the latter, O. cirrata (Hartman, 1944a:92-93, pl. 11, figs. 248-
253) and O. veleronis, new name (for Nothria stigmatis intermedia Hart-
man, 1944a:93-95, pl. 15, figs. 315-324, see below). Table 2 surveys im-
portant differences among these species. The structure of the pseudocom-
pound hooks in anterior setigers is of particular importance; of the species
listed above, only O. guadalupensis and O. dangrigae have both bi- and
tridentate hooks. O. dangrigae has ringed ceratophores, cirriform ventral
cirri on 4 or S setigers, 5 setigers with pseudocompound hooks and lacks
color patterns. O. guadalupensis has smooth ceratophores, cirriform ventral
cirri on 3 setigers and 3 setigers with pseudocompound hooks; in addition,
it has a color pattern of dark dorsal cross-bars on anterior setigers.

The most widely reported species listed in Table 2 is O. stigmatis. In
order to investigate the relationships between O. dangrigae and O. stig-
matis a number of specimens of the latter, collected at the type locality,
were examined (USNM 33713, False Bay, San Juan Island, Washington,
summer, 1937, coll. M. Miller, det. Marian H. Pettibone). The results of the
comparison is given in Table 3. As can be seen, the number of segments
with pseudocompound hooks vary somewhat in O. dangrigae, but averages
4.5, whereas it is invariant at 3 in O. stigmatis. In contrast, the number of
cirriform ventral cirri is invariant at 5 in O. dangrigae and varies somewhat,
but averages 3.5 in O. stigmatis. Both features imply that O. dangrigae has
One more segment involved in the head-formation than does O. stigmatis.
O. stigmatis also retains a distinct color pattern, even after more than 40
years in alcohol; such patterns are absent even in live specimens of O.
dangrigae.

O. dangrigae is known from several localities near Carrie Bow Caye and
Twin Cayes, Belize, in Thalassia beds, coral rubble, sand-and shells in
shallow water.

Etymology.—The specific name is derived from the name of the district
in Belize in which Carrie Bow Caye and Twin Cayes are located, Dangriga
in Carib, or Stann Creek in English.

Onuphis (Nothria) intermedia Kinberg, 1865
Table 2

Onuphis intermedia Kinberg, 1865:560; 1910:40, pl. 14, fig. 9.—Augener,
1931:296—297, fig. 5.

VOLUME 93, NUMBER 3 807

Material examined.—Atlantic Ocean, off the entrance to the harbor at
Rio de Janeiro, 30—40 fms (holotype, Riksmuséet, Stockholm, marked E.E.
191 and type number 461).

Remarks.—The type-material consists of one anterior end of about 50
setigers that is 9 mm long and about | mm wide and three median fragments,
which may belong to another species. The remarks are based on the anterior
end except where noted. The occipital ceratophores have 4 to 5 rings; bran-
chiae are present from about setiger 25 as single filaments and all branchiae
are single on the anterior fragment. The largest midpiece has 2 to 3 branchial
filaments on each segment. Cirriform ventral cirri are present on the first 5
setigers. Tridentate pseudocompound hooks are present in the first 3 setig-
ers; one hook is clearly thicker than the others, but is otherwise similar. A
single compound spiniger is present in each parapodium from setiger 4
through setiger 13. Subacicular hooks are first present from setiger 13. All
specimens are white without color patterns.

Augener (1931:296) also reviewed the type-material of this species, but
his description does not fit the material nor does it fit the original description
by Kinberg (1865) or the illustrations published in Kinberg (1910). Both
description and illustrations by Kinberg fit the type-material very well and
have been augmented above. It is unclear how Augener’s interpretation
arose; it has apparently led to confusion as to the number of taxa present
in Argentinian waters. Orensanz (1974), using Augener’s study as his au-
thority for the description of Kinberg’s species, reported a number of
species of Onuphis from Argentinian waters. The specimens called O. fra-
gilis by Orensanz (1974:94) clearly do not belong to that species (see below);
they resemble O. intermedia, but differ from this species as well in a number
of features. According to Orensanz, his specimens have branchiae first pres-
ent from setigers 10—21 as single filaments; the ventral cirri are cirriform on
3 or 4 setigers and the occipital ceratophores are smooth. Tridentate pseu-
docompound hooks are present in the first 3 setigers and a single, large hook
is continued through one additional setiger. Compound spinigers are present
from setigers 4 or 5 to setigers 13 or 15 and subacicular hooks are first
present from setigers 14-16.

Orensanz’s specimens and O. intermedia both belong to the same group
of Onuphis in that both have branchiae starting behind setigers 10 and have
compound spinigers present in a number of anterior setigers. They are listed
separately in Table 2.

Onuphis (Nothria) veleronis, new name
Table 2

Nothria stigmatis intermedia Hartman, 1944:93-95, pl. 15, figs. 315-324;
1968:686-687, figs. 1-6. [Not Onuphis intermedia Kinberg, 1865].

808 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Material examined.—Northwest Anchorage, San Clemente Island, Cali-
fornia, 20 fms, 12 September 1933 (holotype, AHF Poly 744; 5 paratypes,
AHF Poly 745).

Remarks.—The holotype is an incomplete specimen with about 60 setigers
that is 19 mm long and about .5 mm wide; the largest paratype consists of
85 setigers and is 32 mm long. Ceratophores are smooth; branchiae are first
present from setigers 29-30 and ventral cirri are cirriform on the first 3
setigers. Tridentate pseudocompound hooks are present in 3 setigers and
compound spinigers are present in setigers 4-13. A single large hook is
present from setiger 4 ending between setigers 6 and 8. The first subacicular
hook is in setigers 10-14. A unique feature is the presence of subdistally
expanded acicula from setiger 4 to about setiger 8.

O. veleronis differs clearly from O. stigmatis (see Table 2) in that the
latter has the first branchiae on setigers 18-22, subacicular hooks from se-
tigers 14-16 and a color pattern consisting of dark cross-bars. O. veleronis
lacks a color pattern. O. veleronis differs clearly from O. intermedia Kin-
berg in setal distribution. Thus a new name was needed for this taxon.

Etymology.—The type-material was collected on one of the shakedown
cruises of the VELERO III of the Allan Hancock Foundation, University
of Southern California, hence the specific name.

Subgenus Onuphis Andouin and Milne Edwards, 1833
Onuphis (Onuphis) fragilis Kinberg, 1865
Table 6

Onuphis fragilis Kinberg, 1865:561; 1910:40, pl. 15, fig. 11.—Augener,
1931:298, fig. 6. [Not O. fragilis.-Orensanz, 1974:94—95, pl. 7].

Material examined.—Atlantic Ocean, South of La Plata (holotype, Riks-
muséet, Stockholm, marked E.E. 307 and type number 465).

Remarks.—The type-material fits very well with Kinberg’s description
and illustrations. Occipital ceratophores are smooth; branchiae are first
present from setiger 6 and are bifid in most segments. Ventral cirri are
cirriform in the first 7 setigers. Tridentate pseudocompound hooks are pres-
ent in the first 6 setigers and a single large hook continues to setiger 12.
Subacicular hooks are first present from setiger 25 and compound spinigers
are absent.

Augener (1931:298) redescribed the type-material, but his description does
not fit the material, not does it fit with the original description. For example,
Augener did not find branchiae before setiger 12; branchiae are clearly pres-
ent from setiger 6 in the type-specimen. It is unclear what caused this lapsus;
apparently it led Orensanz (1974) to refer material, discussed above under
O. intermedia, to this species, to which it clearly does not belong.

VOLUME 93, NUMBER 3 809

Yer ue A

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WY £ € E ~
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SHilsillc Wp). re,

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,

Fig. 3. Onuphis (Onuphis) geminata (holotype, USNM 61239): a, Anterior end, dorsal view;
b, Parapodium from setiger 3, anterior view; c, Parapodium from setiger 6, anterior view: d,
Parapodium from setiger 9, anterior view; e, Parapodium from setiger 12, anterior view; f,
Parapodium from setiger 15, anterior view.

810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mm
O3mm
Oo2mm

a
b-g,
h

|
d
oer
ai lh :
1s ae
h

| C
c f g

Fig. 4. Onuphis (Onuphis) geminata (holotype, USNM 61239): a, Parapodium from setiger
50, anterior view; b-f, Pseudocompound hooks; b, From setiger 3, upper hook; c, From setiger
3, median hook; d, From setiger 3, lowermost hook; e, From setiger 6, upper hook; f, From
setiger 6, lowermost hook; g, Pectinate seta, setiger 50; h, Pectinate seta, setiger 3.

Onuphis (Onuphis) geminata, new species
Figs. 3-4, Table 4

Material examined.—CB-40A (holotype, USNM 61239, 2 paratypes,
USNM 61240).

Description.—The holotype is an incomplete specimen that is 32 mm long
for 81 setigers and 1.1 mm wide with parapodia. The anterior end is cylin-
drical and the posterior part of the fragment is dorsally flattened. All color
patterns are dark brown except where specified. The frontal antennae are
yellow; otherwise the prostomium is evenly light rose-colored. Most cera-
tophoral rings have narrow bands; the ceratostyles have speckled, irregular

VOLUME 93, NUMBER 3 811

markings. The peristomium has a band across the anterior edge on the dorsal
side; setiger 1 lacks pigmentation, but from setiger 2 at least through setiger
50, each setiger has a narrow band across the dorsum at the anterior 4 of
each segment. Setiger 3 has the most distinct of these bands, which become
increasingly faint posteriorly; the band is broken up into 2 dorsolateral rows
of patches for about 10 segments posterior to setiger 50. The whole ventrum,
the parapodia and the remainder of the dorsum are evenly light rose- or
flesh-colored and lack distinctive color patterns.

The prostomium (Fig. 3a) is a small, rounded lobe; the frontal antennae
are nearly as long as the prostomium and spindle-shaped. The occipital
antennae are disposed in a semicircle. The outer lateral ones reach setiger
2, with more than % their length in the ceratophore. The inner lateral and
median occipital antennae reach setiger 6. Each ceratophore has 9-10 rings
of which the distalmost one is about as wide as 3 of the other rings com-
bined. The peristomium is about as long as the first setiger and carries a pair
of slender peristomial cirri on its frontal margin.

The first parapodia are no larger than those of the next several setigers
and similar in shape. Each (Fig. 3b) has a flattened, rather wide base. The
acicular lobe is distally rounded; the superior half of the presetal lobe fol-
lows the acicular lobe closely; the inferior half projects beyond the acicular
lobe as a rounded, flattened flap with a distinct excision on the inferior edge.
The dorsal cirri in the first 5 setigers are extremely long and slender and at
setiger 3 (Fig. 3b) are at least twice as long as the whole parapodium from
the base to the tip of the acicular lobe. The postsetal lobes are about as long
as the parapodial bases and are spindle-shaped. The ventral cirri are at-
tached near the base of the parapodia; each tapers evenly to a fine point.
The dorsal cirri rapidly decrease in length posteriorly (Figs. 3b-f, 4a) and
develop distinct swellings near the base. At setiger 9 (Fig. 3d) each dorsal
cirrus is about as long as the postsetal lobe and rather similar in shape.
Ventral cirri are cirriform through the first 9 setigers and are pad-shaped
thereafter. In setigers 8 and 9 the tip of the ventral cirrus is distinctly on
the anterior face of the parapodia and the pads that replace the ventral cirri
are placed on the anterior face of the parapodia (Figs. 3e—f). Postsetal lobes
remain well developed in all parapodia present. A small, distinct cirrus is
present on the superior edge of the parapodium near the base of the dorsal
cirrus in setiger 9.

Limbate setae, pectinate setae, pseudocompound and subacicular hooks
are present. Large hooks and compound spinigers are absent. Limbate setae
are present in all setigers, but are especially numerous in setigers 8-25. Two
different groups of pectinate setae are present. Setigers 2-6 have narrow,
distally transverse setae with about 8 teeth (Fig. 4h). Pectinate setae are
absent in setigers 7 through about setiger 18. From about setiger 18 another
kind is present; each (Fig. 4g) is distally oblique and has about 12 teeth.

812 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pseudocompound hooks are present in the first 7 setigers. Most of the hooks
(Fig. 4b, c, e) are tridentate with 2 distal teeth of about the same size and
the third tooth reduced and closely appended to the second one. In a few
hooks, the lower tooth appears bifid (Fig. 4b). In addition, a single bidentate
hooks (Figs. 4d, f) with subequal teeth is present inferiorly in each fascicle.
Subacicular hooks are first present in setiger 23; each (Fig. 41) is bidentate.

Branchiae are first present from setiger 6 (Fig. 3e) and are bifurcated from
the first. Maximum number of branchial filaments is 8 or 9. The filaments
are arranged as a series of lateral filaments on a main branchial stem in all
setigers (Figs. 3d-f, 4a), but in median and posterior setigers the branchial
stem becomes increasingly short compared to the length of the lateral fila-
ments. In addition the filaments tend to curl up, so the branchiae may appear
superficially as tufted filaments rather than the pectinate arrangement ac-
tually present.

The jaws are well sclerotized; the maxillary formula is 1+1, 7+8, 6+0,
6+7 and 1+1 in one of the paratypes.

Only two of the three specimens could be examined in detail; the third
specimen clearly belongs to the same species in terms of color pattern and
the structure of the branchiae, but the anterior end is regenerating so no
valid counts or measurements could be made.

Tubes are delicate with a thin, flimsy inner lining, sparsely covered with
sand grains.

Remarks.—O. geminata is here compared (Table 4) to other species with
branchiae first present from about setiger 6 and without simple hooks in
anterior setigers.

O. zebra Berkeley and Berkeley (1939:337-—338, figs. 9-10) differs from all
other species in this group by having compound spinigers in anterior setigers
and subacicular hooks are absent; the holotype consists of only 35 setigers
(USNM 32898); hooks may be present in other members of the species.

O. acapulcensis Rioja (1944: 139-143, figs. 1-11), O. cedroensis Fauchald
(1968:3 1-34, pl. 8, figs. a-g), O. multidentata Hartmann-Schroder (1960:24—
26, figs. 50-55), O. nannognathus Chamberlin (1919:270-274, pl. 43, figs.
8-11, pl. 44, figs. 1-5), O. oligobranchiata Orensanz (1974:93—94, pl. 6), O.
pourtalesii (Ehlers, 1879:273, see also Ehlers, 1887:74—75, pl. 19, figs. 6—10,
pl. 20, figs. 1-6), O. proalopus Chamberlin (1919:265—269, pl. 40, figs. 3-8,
pl. 41, figs. 1-10), O. profundi Fauchald (1968:40-41, pl. 10), O. similis
Fauchald (1968:28-29, pl. 4, figs. g-i), O. lineata, new name (for O. striata
Hartmann-Schroder, 1965: 164—167, figs. 135-137, see below) and O. tenuis
Hansen (1882:10, pl. 3, figs. 15-22, see also Orensanz, 1974:87-89, pl. 3) all
have exclusively tridentate pseudocompound hooks; other species in this
group have both bi- and tridentate hooks. Of these, O. dorsalis (Ehlers,
1897:71-74, pl. 5, figs. 108-118, see also Hartmann-Schro6der, 1962: 114-117,
figs. 115-119) and O. heterouncinata (Hartmann-Schroder, 1965:161—164,

VOLUME 93, NUMBER 3 813

figs. 132-134) have subacicular hooks first present on setigers 14 and 12
respectively. O. dorsalis has 3 branchial filaments and O. heterouncinata
a single filament per segment (thus, by definition, a member of the subgenus
Nothria, but with so great similarities to species of the subgenus Onuphis
that it is considered here for the sake of completeness). O. geminata and
O. investigatoris Fauvel (1932:147-148, pl. 6, figs. 1-6, textfig. 21), the two
remaining species in the group, have at least 8 branchial filaments where the
branchiae are best developed. O. geminata has subacicular hooks starting
at setigers 23; the start of the subacicular hooks is not stated in Fauvel
(1932), but judging from his Plate 6, figs. 4 and 5, they must start posterior
to setiger 10 and anterior to setiger 30. It is here assumed that O. geminata
cannot be separated from O. investigatoris on this character. The two
species can be separated on the structure of the occipital antennae. The
inner lateral occipital antennae are at least twice as long as the median one
in O. investigatoris and reach setiger 15; in O. geminata these 3 antennae
are of about the same length and reach setiger 6. There are also minor
differences in the distribution of cirriform ventral cirri, pseudocompound
hooks and in the number of branchial filaments as indicated in Table 4.
Additionally, O. geminata is oculate, O. investigatoris is not.

O. geminata is known from a single locality in Thalassia and fine sand
in a mangrove area about 10 km north of the city of Dangriga, Belize.

Etymology.—The specific name, Latin for twinned, refers to the close
similarity between this and other species of the genus in Belize.

Onuphis (Onuphis) lineata, new name
Table 4

Onuphis striata Hartmann-Schroder, 1965: 164—167, figs. 135-137.

Material examined.—Off Chile (holotype, Hamburg Staatsmuseum,
P- 14293).

Remarks.—The specimen is as described by Hartmann-Schroder (1965).
Pseudocompound hooks are present in the first 7 setigers and postsetal lobes
are distinct in the first 50 setigers. Branchiae are present from setiger 6 at
least through setiger 99; an additional 30 setigers are present in the type,
but are in recent regeneration and lack branchiae.

The name O. striata is preoccupied in the combination O. striata Ushak-
ov (1950), originally described as Onuphis parva striata Ushakov,
(1950:193, fig. 25, see also Ushakov, 1955:235-—236, fig. 74B and 77J). Fau-
chald (1968:37) elevated Ushakov’s subspecies to separate specific status.
Hartmann-Schroder (in litt.) requested that a new name be given her taxon
from Chile.

O. lineata is known from the type-locality only.

Etymology.—The specific name, Latin meaning (in part) striped, was se-

814 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lected to parallel as closely as possible the meaning of the name initially
used for this taxon; both names refer to the presence of numerous narrow
transverse bands of dark brown pigment across the dorsum in anterior se-
tigers.

Onuphis (Onuphis) pulchra, new species
Figs. 5-6, Tables 5, 6

Material examined.—CB-1 (holotype, USNM 61241; 58 paratypes, USNM
61242); CB-16 (45); F-19 (13); F-21 (3); F-28 (7); F-33 (1).

Description.—The holotype is a complete specimen with 240 setigers that
is 70 mm long and 1.3 mm wide including parapodia. The anterior part of
the body, including the S first setigers is cylindrical. Posterior to setiger 6
the body becomes increasingly flattened dorsally and the ventrum convexly
curved, creating a space above the animal in the tube. The complex color
pattern consists of dark brown pigment rings and patches. The prostomium
and peristomial rings lack color patterns. Each of the outer lateral cerato-
styles has a narrow ring near the base; the remaining occipital antennae lack
color patterns. There is a broad band across the posterior half of the first
setiger, darkening distinctly towards the intersegmental groove to the sec-
ond setiger. The anterior half of the first setiger appears translucently white
(due to the presence of a field of epidermal glands). Each setiger back to
about setiger 30 has a transverse band at the posterior edge. These bands
become reduced medially at about setiger 30 and become interrupted into
two rows of dorsolateral transverse patches at about setiger 30. These two
rows are continued posteriorly for about 10 more setigers. Each dark band
is bordered anteriorly by a narrow translucent band of epidermal glands;
these bands disappear at about setiger 30. The ventrum and the rest of the
body are evenly light rose-colored in alcohol and lack distinct color pattern.
The color pattern is as described also in live specimens.

The prostomium (Fig. 6a) is nearly circular and has a pair of short, tri-
angular frontal antennae. The outer lateral occipital antennae barely reach
the posterior edge of the peristomium; in most individuals they are curved
anteroventrally. Each has a spindle-shaped style and a short ceratophore
with 4 rings. The inner lateral occipital antennae reach setiger 4—5; the
median one reaches setigers 3—5 in most specimens (see Table 5); each has

—

Fig. 5. Onuphis (Onuphis) pulchra (holotype, USNM 61241); a, Parapodium from setiger
1, anterior view; b, Parapodium from setiger 2, anterior view; c, Parapodium from setiger 4,
anterior view; d, Parapodium from setiger 5, anterior view; e, Parapodium from setiger 6,
anterior view; f, Parapodium from setiger 7, anterior view; g, Parapodium from setiger 8,
anterior view; h, Parapodium from setiger 12, posterior view.

VOLUME 93, NUMBER 3 815

= |
4 |
fm \
7 = ( |
= \\ i!
Y 3 NA) U1
y \J = AYR \
\ } \ = 7 e
. = i
\ ~ a———
: A
me
\ iby \
| x
\ Ss

.2mm

816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a short ceratophore with 4 to 5 rings of which the distalmost one makes up
about one-half of the length of the ceratophore. The peristomial cirri are
attached at the anterior edge of the peristomial rings; each is spindle-shaped
and reaches just beyond the bases of the occipital antennae. A pair of black
eyespots are present between the bases of the outer and inner lateral occip-
ital antennae.

The first few parapodia (Fig. 5a) are attached laterally. The first projects
anteriorly; each ramus is about as long as the body is wide. The presetal
lobe has a rounded distal edge with a distinct excision on the ventral side.
A contraction fold can be present across the parapodium at the base of the
presetal lobe. The acicular lobe is rounded. The postsetal lobe is about as
long as the base of the parapodium and is spindle-shaped. The dorsal cirrus
is distinctly longer than the postsetal lobe and reaches well beyond all other
parapodial parts and is spindle-shaped. The ventral cirrus is digitate and
reaches the tip of the acicular lobe. The second and third parapodia (Fig.
5b) are similar in shape, except that the parapodial bases are shorter and
relatively wider than that of the first parapodium. In the fourth and fifth
parapodium (Figs. Sc—d) the acicular lobes become increasingly wider and
distally more truncate than that of the first one. The presetal lobes are
flattened to follow the outline of the acicular lobe closely. The dorsal cirri
are longer compared to the other parapodial parts and the ventral cirri be-
come shorter and are placed on the anterior face of each parapodium. The
ventral cirri are cirriform through setigers 9-10 (Figs. Se—g, Table 5). Dorsal
cirri are present in all setigers to the posterior end; they become increasingly
slender posteriorly but are of about the same length as those in the anterior
setigers. Postsetal lobes (Fig. 5h) can be recognized in the first 50 setigers
as a distinct digitiform lobe on the distal end of a low, rounded fold that
closely follows the outline of the acicular lobe.

Branchiae are first present from setiger 6 (Table 5); the first 10 setigers
have a single branchial filament; thereafter, the numbers increase to 3 or 4;
the branchiae again become single filaments at about setigers 70-100 (ap-
proximately '/, the length of the specimen) and the last '/; of the length of
each specimen lacks branchiae.

Limbate and pectinate setae, two kinds of anterior hooks and subacicular
hooks are present. Limbate setae are most common in the first 50 setigers.
Pseudocompound tridentate hooks (Fig. 6b—h) are present in the first 6 se-
tigers. Each has a blunt hood and the teeth are curved and slender, decreas-
ing more or less evenly in size from the distal to the proximal tooth. Large
tridentate hooks (Fig. 6k—n) are present from setiger 4 to the last segment
before the start of the subacicular hooks. Each large hook is at least twice
as heavy as the pseudocompound hooks of the same segment; the median
tooth is larger than the distal one in all large hooks. The median tooth
becomes increasingly curved and the distal tooth more erect posteriorly

VOLUME 93, NUMBER 3 817

5mm
b-n .02mm

Fig. 6. Onuphis (Onuphis) pulchra (holotype, USNM 61241): a, Anterior end, dorsal view,
b-h, Pseudocompound hooks; b, From setiger 1; c, From setiger 2; d-e, From setiger 3; f,
From setiger 4; g, From setiger 5; h, From setiger 6; i, Pectinate seta, from setiger 3; k—n,
Large hooks; k, From setiger 4; 1, From setiger 5; m, From setiger 6; n, From setiger 12.

818 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Fig. 6n). Bidentate subacicular hooks are present from setigers 16—20 (Ta-
ble 5); usually 2 hooks are present in a segment. Pectinate setae (Fig. 61)
are present from the second setiger; each is slightly oblique distally and has
about 12 teeth.

The maxillary formula is 1+1,6—7+8, 8+0, 5-6+8 and 1+1 as observed
by dissection of four paratypes. The whole jaw-apparatus is poorly sclero-
tized and soft.

The tubes have a tough, translucent inner lining and are densely and
evenly covered with shell-fragments and coral debris.

Remarks.—The relations between O. pulchra, O. virgata (see below) and
related taxa can best be clarified by discussing them jointly. Table 6 surveys
species of Onuphis with branchiae first present from about setiger 6 and
with large tridentate hooks in some anterior setigers, in addition to the
pseudocompound hooks present in all species of the genus.

O. fragilis Kinberg (1865:561, see also Kinberg, 1910:40, pl. 15, fig. 11
and above) has smooth occipital ceratophores; all other species have dis-
tinctly ringed ceratophores. Three species, O. jenneri Gardiner (1976: 192—
193, fig. 240-t), O. microcephala Hartman (1944a:78-80, pl. 3, figs. 67-75,
pl. 18, fig. 339, see also Gardiner, 1976:191-192, fig. 24k—n) and O. pig-
mentata Fauchald (1968:38—39, pl. 9, figs. f-n) have pad-shaped ventral cirri
from setiger 3; the other species have cirriform ventral cirri through at least
setiger 4. O. taeniata Paxton (1979:284—288, figs. 43-55) and O. vermilli-
onensis Fauchald (1968:41—43, pl. 11) have large hooks through setiger 10,
the remaining three species, O. setosa Kinberg (1865:560, see also Kinberg,
1910:40, pl. 14, fig. 10 and Orensanz, 1974:89-93, pls. 4-5) and the two new
species have large hooks at least through setiger 13.

It is difficult to define O. setosa precisely in relation to the two new
species. It differs from O. pulchra in that it has fewer segments with cirri-
form ventral cirri and fewer segments with pseudocompound hooks. Or-
ensanz (1974:89-93) indicated the range of variation of these features he
found in O. setosa, but one cannot determine from Orensanz’s description
how this variability is distributed. The two species differ clearly in color
patterns in that O. setosa has dark pigment bands on both pro- and peris-
tomium according to Orensanz (1974:89, pl. 4, fig. 1); all specimens of O.
pulchra lack pigment on both pro- and peristomium. O. setosa differs from
O. virgata in that it has 4 to 8 setigers with cirriform ventral cirri and 4 to
5 setigers with pseudocompound hooks; O. virgata has 11-13 setigers with
cirriform ventral cirri and 7 setigers with pseudocompound hooks. O. pul-
chra has pseudocompound hooks in 6 setigers and cirriform ventral cirri in
9 to 10 setigers. O. virgata has color patches and bands on the pro- and
peristomium; O. pulchra lacks such color patterns. The numerical differents
appear minor, but are statistically different (chi squared, at the .01 level,

VOLUME 93, NUMBER 3 819

even if the sample of O. virgata is too small for strict application of any
Statistical tests).

O. pulchra is very common in Thalassia-flats in shallow water near Carrie
Bow Caye, Belize.

Etymology.—The specific name, Latin meaning pretty or beautiful, refers
to the striking color pattern of the anterior end.

Onuphis (Onuphis) virgata, new species
Fig. 7, Tables 6, 7

Material examined.—CB-40A (1 specimen); CB-40B (holotype, USNM
61248, 3 paratypes, USNM 61249).

Description.—The holotype is an incomplete specimen with 149 setigers
that is 57 mm long and .7 mm wide with parapodia. The anterior end, in-
cluding the 5 first setigers, is cylindrical, median and posterior parts of the
body are dorsally flattened and the parapodia become increasingly dorso-
lateral in position. A distinct color pattern of dark brown bands and patches
is present on the anterior end of the body. A narrow band is at the base of
all ceratostyles and a large dark patch is on the prostomium in front of the
occipital antennae. The peristomium has a wide band across the dorsum
and the first setiger is evenly light brown with a darker band across the
posterior half on the dorsal side only. The next dozen or so segments have
dark bands across both dorsum and ventrum. The pigment bands on the
ventrum rapidly fade posteriorly and are absent after setiger 20. The dorsal
pigment bands break up into dorsolateral patches which continue to about
setiger 50. Each of the first 5 or 6 pairs of parapodia has a dark patch near
the distal end of the acicular lobe on the posterior face. The rest of the body
is pale pink-colored.

The prostomium (Fig. 7f) is frontally rounded and has the occipital an-
tennae attached in a semicircle; the frontal antennae are narrowly triangular.
The outer lateral occipital antennae barely reach the posterior edge of the
peristomial rings. The inner lateral and median occipital antennae reach
about the middle of setiger 3; each ceratophore has about 4 rings of which
the distalmost is longer than the 3 others combined. The peristomial rings
are about half as long as the first setiger; the anteriormost carriers a pair of
short, slender peristomial cirri on the frontal edge.

The first parapodia are directed forward and ventrally; they are of about
the same size as those of the second and third setiger. Each (Fig. 7a) has
a wide, antero-posteriorly flattened base; there may be a contraction fold
across the anterior face of each parapodium at the level of the bases of the
dorsal and ventral cirri (not shown in the illustration). The presetal lobe is
obliquely rounded and is distinctly excised on the ventral side. The acicular

820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sy

—STY
sae
4 Vee an ri!
Tt ale <
/ = ‘ 2,
iy vf, | | | YW ="
j Wp jy st
Y/ j Z '
Ly
/
a “J

mM n

Fig. 7. Onuphis (Onuphis) virgata (holotype, USNM 61248): a, Parapodium from setiger 3,
anterior view; b, Parapodium from setiger 6, anterior view; c, Parapodium from setiger 9,
anterior view; d, Parapodium from setiger 15, anterior view; e, Parapodium from setiger 18,
anterior view; f, Anterior end, dorsal view; g—k, Pseudocompound hooks, g, From setiger 3;
h, From setiger 6; i-k, From setiger 7 of a paratype; I-n, Large hooks, 1, From setiger 12; m,
From setiger 6; n, From setiger 7 of a paratype.

VOLUME 93, NUMBER 3 821

lobe is evenly rounded. The postsetal lobe is spindle-shaped and is about
as long as the base of the parapodium. The dorsal cirrus is digitate; it is
shorter than the postsetal lobe. The ventral cirri remain cirriform through
about 12 setigers (see Tables 6 and 7); they decrease in size from the first
setiger (Fig. 7a—e) and move on to the anterior face of each parapodium
from about setiger 9. The postsetal lobes remain distinct in all segments in
the fragments available (of which the holotype is the longest), but decrease
rapidly in size at about setigers 20-25; they are short, digitate extensions
from the middle of a wide, rounded lobe in all posterior setigers. The dorsal
cirri become somewhat more slender in posterior setigers, but remain about
the same length.

Branchiae are first present from setigers 6—7 (Table 7); the first branchia
is a single filament, more posteriorly the branchial filaments increase to
about 5. The number of filaments decrease in more posterior segments; in
all specimens examined the branchiae have at least 2 filaments, even on the
last setigers present.

Limbate and pectinate setae, pseudocompound, large and subacicular
hooks are present. Limbate setae are present in most setigers, but are dens-
est in anterior setigers; pectinate setae are present from about the third
setiger; each has a transverse distal margin with about 8 teeth. Pseudocom-
pound hooks (Figs. 7g—k, Table 7) are present in the first 7 setigers. Each
is slender and has 3 slender teeth; the hoods are short and distally blunt.
The teeth decrease rather evenly in size from the distalmost one. Large
hooks (Figs. 7l—n) are present from setiger 4 through about setiger 21; each
is at least twice as thick as the pseudocompound hooks of the same segment,
and tridentate; the distal tooth is short and slender and is erect in at least
the last 10 segments. Bidentate subacicular hooks are present from about
setiger 22. There are no segments with both large and subacicular hooks.
Compound spinigers are absent.

The maxillary formula is 1+1, 7+9, 7+0, 5+9 and 1+1 as examined in
one paratype. The jaw-apparatus is well sclerotized. The teeth on left max-
illa TV are unusually long and slender.

The tubes have a thin inner lining and are covered with fine sand-grains.

The relationships between O. virgata and similar species have been dis-
cussed above.

O. virgata is known from an area about 10 km North of the city of Dan-
griga, Belize, in Thalassia and fine sand in mangroves.

Etymology.—tThe specific name, Latin meaning (in part) colored stripes,
refers to the striking color pattern of this species.

Acknowledgments

I would like to thank Drs. Meredith L. Jones and Brian F. Kensley of the
Smithsonian Institution and Joan D. Ferraris, Mount Desert Island Biolog-

822 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ical Laboratory for permission to study material collected in Belize. Drs.
Lars Orrhage and Jerry D. Kudenov arranged loans of type material in their
care, without which this study would have beem impossible. Funding for
collecting trips to Belize were provided in part by a grant from the Exxon
Foundation to the Smithsonian Institution. I am very grateful to Dr. Mer-
edith L. Jones for a careful and critical review of the manuscript.

The paper is Contribution No. 62 from the IMSWE Program.

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a

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Hartmann-Schroder, G. 1960. Zur Polychaeten-Fauna von Peru. Part !.—Beitr. Neotrop.
Fauna 2:1—44.

. 1962. Die Polychaeten des Eulitorals. Jn Hartmann, G. and G. Hartmann-Schroder:

Zur Kenntnis der Eulitorals der chilenischen Pazifikktste und der argentinischen Ktiste

Siidpatagoniens unter besonderer Berucksichtigung der Polychaeten und Ostracoden.—

Mitt. Hamburg Zool. Inst. u. Mus. Erg. B. zu 60:57—167.

. 1965. Die Polychaeten des Sublitorals. Jn Hartmann, G. and G. Hartmann-Schroder:

Zur Kenntnis des Sublitorals der chilenischen KUuste unter besonderer Berucksichtigung

der Polychaeten und Ostracoden (Mit Bemerkungen uber den Einfluss sauerstoffarmer

Stromungen auf die Besiedlung von marinen Sedimenten).—Mitt. Hamburg Zool. Inst.

u. Mus. Erg. B. zu 62:59-305.

Kinberg, J. G. H. 1865. Annulata nova.—Ofv. Vet. Akad. Stockholm Forh. 21:559-574.

1910. Zoologi 3. Annulata.—Kongliga Svenska Fregatten Eugenies Resa omkring
jorden under befal af C. A. Virgin aren 1851-1853. Vetenskapliga Iagttagelse pa Konung
Oscar den Forstes befallning utgifna delen. Almquist 0. Wicksells, Stockholm. 78 pp.

Kiseleva, M. J. 1968. Policheti sublitorali tsentralino-Amerikanskikh Morei. Otvyid Eunice-
morpha.—Issled. Tstentralino-Amerikanskikh Morei. Akad. Nauk. Biol. St. Sevastopol
2:75-98.

McIntosh, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. Challenger
during the years 1873-76.—Challenger Reports 12, 554 pp.

Monro, C. C. A. 1928. On the Polychaeta collected by Dr. Th. Mortensen off the coast of

Panama.—Vidensk. Medd. dansk naturh. Foren. 85:75—103.

. 1933. The Polychaeta Errantia collected by Dr. C. Crossland at Colon in the Panama

region and the Galapagos Islands during the expedition of the S.Y. St. George.—Proc.

Zool. Soc. London 1933 (1):1—96.

Orensanz, J. M. 1974. Los anelidos poliquetos de la provincia biogeografica Argentina, V:
Onuphidae.—Physis, Argentina Sec. A 33(86):75—122.

Paxton, H. 1979. Taxonomy and aspects of the life history of Australian beachworms (Poly-
chaeta: Onuphidae).—Australian Jour. Mar. Freshwater Res. 30:265—294.

Pearse, A. S. 1936. Estuarine animals at Beaufort, North Carolina. Jour. Elisha Mitchell Sci.
Soc. 52:174-222.

824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pettibone, M. H. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae
through Trochochaetidae.—Bull. U.S. Natl. Mus. 227(1), 356 pp.

Rioja, E. 1944. Estudios anelidologicos XI. Notas sobre algunas especies de poliquetos de las
costas mexicanas del Pacifico.—Anales Inst. Biol. Univ. México 15:139-145.

Rullier, F. 1974. Quelques annélides polychetes de Cuba recueillies dans les €ponges.—Trav.
Must. Hist. Nat. Gregore Antipa 14:9-77.

Suarez, A. M., and R. Fraga. 1978. Poliquetos bentosicos cubanos I.: Lista de poliquetos
errantes.—Univ. Habana Cuba, Ciencias Ser. 8: Invest. Marinas 33. 60 pp.

Treadwell, A. L. 1921. Leodicidae of the West Indian region.—Pub. Carnegie Inst. Washing-

ton 293. 131 pp.

. 1922. Polychaetous annelids collected at Friday Harbor, State of Washington in Feb-

ruary and March, 1920.—Publ. Carnegie Inst. Washington 312:171-181.

. 1934. New polychaetous annelids.—Smithsonian Misc. Coll. 91(8). 9 pp.

. 1939. Polychaetous annelids of Porto Rico and vicinity.—Scientific Survey of Porto

Rico and the Virgin Islands, New York Acad. Sci. 16(2):151-319.

Ushakov, P. V. 1950. Mnogoshchetinkovye chervei (Polychaeta) Okhotskoyo moreii.—Issled.
Dalynevost. Morei SSSR 2:140-234.

—. 1955. Mnogshchetinkovye chervei dal’nevostochnykh morei SSSR.—Akad. Nauk,
USSR. Keys to the Fauna of the USSR 56, 433 pp. (translated to English 1965).

Wilson, H. V. 1900. Marine biology at Beaufort.—Amer. Nat. N.Y. 34:339-360.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

Table 1.—Onuphidae (Polychaeta) previously reported from the Gulf of Mexico and Carib-
bean Sea. Some species (e.g. Diopatra cuprea) have been reported frequently from the U.S.
Gulf coast; no attempts were made to include those records in the table.

s Local depth
Name Record distribution
Americonuphis magna (Andrews, 1891) Treadwell, 1921 Intertidal-shallow
Hartman, 1951 subtidal
| Suarez and Fraga, 1978
Diopatra cuprea (Bosc, 1802) Augener, 1906 Intertidal to about
Monro, 1933 200 m

Hartman, 1951
Fauchald, 1977
Suarez and Fraga, 1978

Hyalinoecia bilineata Baird, 1870 Rullier, 1974 Shallow subtidal

H. branchiata Treadwell, 1934 Treadwell, 1934 Deeper than 200 m
Treadwell, 1939
Suarez and Fraga, 1978

' HH. juvenalis Moore, 1911 Hartman, 1944b Shallow subtidal
Kiseleva, 1968
Suarez and Fraga, 1978

VOLUME 93, NUMBER 3

Table 1.—Continued.

Name

H. tubicola (O. F. Miller, 1776)

H. varians Baird, 1870

Onuphis (Nothria) conchylega Sars, 1835

O. (N.) opalina Verrill, 1873

O. (N.) rubrescens Augener, 1906

O. (N.) sombreriana (McIntosh, 1885)

Onuphis (Onuphis) eremita
Audouin and Milne Edwards, 1833

O. (O.) eremita oculata Hartman, 1951

O. (O.) pourtalesii (Ehlers, 1879)

O. (O.) vermillionensis Fauchald, 1968
Paradiopatra fragosa Ehlers, 1887

P. glutinatrix Ehlers, 1887

Paronuphis gracilis Ehlers, 1887

Rhamphobrachium agassizii Ehlers, 1887

Record

Augener, 1906
Hartman, 1942
Hartman, 1951
Suarez and Fraga,

Baird, 1870
Ehlers, 1887
Kiseleva, 1968
Suarez and Fraga,

Augener, 1906
Kiseleva, 1968
Suarez and Fraga,

Augener, 1906
Kiseleva, 1968
Suarez and Fraga,

Augener, 1906
Suarez and Fraga,

McIntosh, 1885
Suarez and Fraga,

Kiseleva, 1968
Rullier, 1974
Suarez and Fraga,

Hartman, 1951

Ehlers, 1879
Ehlers, 1887
Augener, 1906
Hartman, 1938
Suarez and Fraga,

Fauchald, 1977

Ehlers, 1887
Hartman, 1938

Ehlers, 1887
Augener, 1906
Hartman, 1938

Ehlers, 1887
Hartman, 1938

Ehlers, 1887
Hartman, 1938
Treadwell, 1939
Kiseleva, 1968
Suarez and Fraga,

1978

1978

1978

1978

1978

1978

1978

1978

1978

825

Local depth
distribution

Deeper than 200 m

Shallow subtidal

Deeper than 200 m

Deeper than 200 m

Deeper than 200 m

Deeper than 200 m

Shallow subtidal

Shallow subtidal
Deeper than 200 m

Shallow subtidal
Deeper than 200 m

Deeper than 200 m

Deeper than 200 m

Deeper than 200 m

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 5.—Summary statistics on numerical features, Onuphis (O.) pulchra.

Number
Range Mean Stand. dev. of observ.
Occipital antennae
length of inner lateral 4-7 4.87 58 38
length of median 3-6 4.26 79 38
number of rings 5 invanant
Branchiae
start 5-7 5.98 .20 104
max. # filaments 2-5 3.37 si 38
Cirriform ventral cirri 9-10 YA 56 104
Pseudocompound hooks to setiger # 6 invanant
Simple hooks
Start 4 invanant
end 15-19 16.16 eS 38
Subacicular hooks from setiger # 16-20 17.34 97 104
Table 6.— —
Table 7.—Summary statistics on variable features, Onuphis (O.) virgata.
Number
Range Mean Stand. dev. of observ.
Occipital antennae
length of inner lateral 3-5 3.60 .89 B)
length of median 3-4 3.40 5) 5
number of rings 4-5 4.20 45 5
Branchiae
Start 6-7 6.20 45 5)
max. # filaments 3-5 4.40 .89 5
Cirriform ventral cirri I SI13} 12.20 .84 5
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Simple hooks
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PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 830-844

THE INDO-PACIFIC PIPEFISH GENUS
UROCAMPUS (SYNGNATHIDAE)

C. E. Dawson

Abstract.—The genus Urocampus Gunther is rediagnosed and compared
with other genera with confluent superior trunk and tail ridges. Recognized
species (U. nanus Gunther, U. carinirostris Castelnau) are redescribed and
illustrated and notes are provided on variation, distribution, and pouch lar-
vae. The littlhe known Urocampus southwelli Duncker is provisionally re-
ferred to the genus Siokunichthys Herald.

This report, a continuation of revisionary studies on pipefishes, treats the
genus Urocampus Gunther which is in part characterized by the presence
of confluent superior trunk and tail ridges. This feature is shared with a wide
variety of temperate-tropical genera and some of these have been reviewed
recently by Dawson (1976, 1977a, 1977b, 1979) and Dawson and Allen
(1978). The two recognized species of Urocampus (nanus, carinirostris) are
poorly known and existing descriptions are largely inaccurate or incomplete.
Present study material is limited but the type material has been examined
and males and females of each species have been illustrated.

Measurements are in millimeters (mm); proportional data are referred to
standard length (SL) or head length (HL); total length (TL) measurements
are given for pouch larvae; color descriptions are from specimens preserved
in alcohol; depths are in meters (m); other methods follow Dawson (1977b).
Abbreviations for repositories of materials examined follow: AMS—Aus-
tralian Museum, Sydney; BMNH—British Museum (Natural History);
CAS—California Academy of Sciences; CAS-SU—former Stanford Uni-
versity specimens housed as CAS; FMNH—Field Museum of Natural His-
tory; GCRL—Gulf Coast Research Laboratory Museum; MNHN—Muséum
National d’ Histoire Naturelle, Paris; NMV—National Museum of Victoria,
Melbourne; QM—Queensland Museum, Brisbane; QVM—Queen Victoria
Museum, Launceston, Tasmania; SAM—South Australian Museum, Ade-
laide; UMMZ—University of Michigan Museum of Zoology; USNM—Na-
tional Museum of Natural History, Smithsonian Institution; WAM—West-
ern Australian Museum, Perth; YCMP—Yokosuka City Museum.

Urocampus Gunther

Urocampus Gunther, 1870:179 (orig. descr.; type-species U. nanus Gunther
1870, by monotypy).

VOLUME 93, NUMBER 3 831

Diagnosis.—Superior trunk and tail ridges confluent, superior ridges not
arched dorsad below dorsal-fin base, inferior trunk and tail ridges discon-
tinuous near anal ring, lateral trunk ridge confluent with inferior tail ridge.
Median dorsal snout ridge distinct, entire, usually elevated in adult males,
not confluent with supraorbital ridges, usually fails to reach interorbital;
supraorbital ridges elevated slightly with interorbital depressed between;
supraopercular ridges absent; median dorsal head ridges low; median lon-
gitudinal opercular ridge essentially straight, usually crosses half or more
of opercle in subadults and adults and margined with fine striae above and
below; pectoral-fin base not protruding strongly laterad, without prominent
ridges. Principal body ridges distinct, not elevated strongly, the margins
entire, indented but not deeply notched between rings; scutella small, with-
out longitudinal keels or other ornamentation; dorsum of body flat to some-
what concave anteriad, becoming convex toward dorsal-fin insertion, angled
clearly upward about dorsal-fin base, becoming gradually flattened caudad;
venter of trunk V-shaped, usually with a median keel-like ridge in adults;
venter of tail essentially flat; posterior tail rings usually shortened progres-
sively toward caudal-fin base. Without spines or prominent denticulations;
subadults and adults usually with simple or branched dermal flaps on head
and/or body, flaps essentially round in section. Without odontoid processes
(Dawson and Fritzsche, 1975) in jaws. Brood pouch under tail, pouch plates
absent, pouch closure the semi-type of Herald (1959). Head length ca. 10-
13 in SL; snout length-ca. 2.1-3.6 in HL; rings 7-12 + 49-59; dorsal-fin
rays 13-17; dorsal-fin origin on 5th—9th tail ring, its base distinctly elevated,
the membranes closely bound to fin rays; total subdorsal rings 2.5—4.0;
pectoral-fin rays 7-10; anal-fin rays typically 2; caudal fin minute in sub-
adults and adults, typically with 10 rays.

Comparisons.—Among some 22 genera of pipefishes with confluent su-
perior trunk and tail ridges, subadults and adults of only Urocampus and
Siokunichthys Herald share the combination of confluent-lateral trunk and
inferior tail ridges, presence of pectoral fins and dorsal-fin origin on the 2nd—
9th tail ring. Urocampus is characterized by the presence of a well-devel-
oped anal fin, dermal flaps on most subadults and adults and a small to
rudimentary caudal fin. The anal fin and dermal flaps are lacking in Siok-
unichthys and the caudal fin is relatively long and well developed. Further-
more, the head and body ridges are distinct and rather prominent in Uro-
campus, whereas the few persistent ridges in most Siokunichthys are low
or vestigial and difficult to see even under x60 magnification.

Remarks.—Since Gunther’s (1870) original description, the anal fin has
been variously reported as present or absent in Urocampus. Although some-
times concealed within the anterior portion of the brood pouch in mature
males, the anal fin is present in all examined subadults and adults. This fin
is not visible in early pouch larvae but it is developed in specimens as small

832 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

as 14mm SL. Dermal flaps occasionally are absent or lost from some spec-
imens but they are present in most subadults and adults and are usually best
developed in mature males. The caudal fin is very small and accurate counts
of fin rays are difficult. Although atypical counts are not uncommon, the
count is 10 in pouch larvae (of carinirostris) and most subadults and adults.

Duncker (1910) described Urocampus southwelli from two adult fish (40—
45 mm TL) from Ceylon and later (1915) tentatively referred this species to
the synonymy of U. carinirostris Castelnau. The type material is no longer
available, but Duncker’s (1910) description and figures show that southwelli
(without anal fin) is distinct from carinirostris and it is here provisionally
referred to Siokunichthys.

The two recognized species of Urocampus appear to frequent shallow
coastal marine and estuarine habitats and are commonly associated with
algal covered sedimentary bottoms.

Urocampus nanus Gunther
Figs. 1-2

Urocampus nanus Gunther, 1870:179 (orig. descr.; Manchuria).
Urocampus rikuzenius Jordan and Snyder, 1901:10, pl. 7 (orig. descr.; Mat-
sushima Bay, Japan).

Diagnosis.—Snout long, its length averages 2.2 in HL, trunk rings 10-12
(usually 10-11), dorsal-fin rays usually 16.

Description.—Dorsal-fin rays 15-17 (x = 15.9), pectoral-fin rays 7-9
(8.2), rings 10-12 + 53-56 = 63-68 (66.2), total subdorsal rings 3.25—4.0
(3.5), dorsal-fin origin on 7th—9th (7.9) tail ring; see Tables 1—4 for additional
counts. Proportional data based on 7 specimens 84.0—127.5 (105.9) mm SL
follow: HL in SL 10.5-12.4 (11.5), snout length 2.4—4.0 (3.2), length of
dorsal-fin base in HL 1.4—1.8 (1.6), anal ring depth in HL 4.0-6.1 (5.2),
trunk depth in HL 2.1-3.3 (2.8), pectoral-fin length in HL 5.7-6.9 (6.4),
length of pectoral-fin base in pectoral-fin length 1.5—2.1 (1.8).

Median dorsal snout ridge (Figs. 1-2) low to slightly concave in young
and females, somewhat elevated in mature males but protrudes little above
horizontal through dorsal margin of eye. Supraorbital ridges continue for-
ward to terminate bilaterally above nares and near posterior end of median
dorsal snout ridge; lateral profile of head not clearly depressed behind the
eye; pectoral-fin base with two faint ridges.

Dermal flaps often distally bifurcate on dorsum of predorsal rings and
profusely branched on median ventral trunk ridge, elsewhere mostly simple.
Head with an enlarged barbel-like flap bilaterally near posterior end of man-
dible and single short flaps bilaterally above middle of opercle; pectoral-fin
base with a minute flap; most trunk rings with a long flap near midline of
dorsum, a short flap on or just above each lateral ridge and a long flap on

VOLUME 93, NUMBER 3 833

Fig. 1. Urocampus nanus: Lateral and dorsal aspects of head and anterior trunk rings,
together with section of body illustrating ridge configuration, dorsal and anal fins, and brood
pouch. Top: YCMP 3522 (84.0 mm SL, juvenile or female). Remainder: GCRL 15701 (99.5 mm
SL, brooding male).

the median ventral ridge; predorsal tail rings with single median or paired
bilateral flaps on the dorsum and single minute flaps just above each inferior
ridge; short single or paired bilateral flaps present on dorsum to about the
9th postdorsal ring, the tail elsewhere devoid of flaps (description from
MEMIP3522):

Coloration.—Ground color tan to dark brown, the dorsum of body often
lighter than sides and venter; head and body occasionally plain but most
often irregularly marked with pale mottled areas and scattered pale spots
(Fig. 2); median ventral trunk ridge and associated dermal flaps mainly dark

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Upper pair.—Urocampus nanus. Top: YCMP 1223 (127.5 mm SL, female). Bottom:
GCRL 15701 (105.5 mm SL, brooding male). Lower pair.—Urocampus carinirostris GCRL
15507 (top, 72.0 mm SL, brooding male; bottom, 67.5 mm SL, female).

brown; large, more or less regularly spaced, pale spots on superior and
inferior ridges of postdorsal tail rings impart a grossly banded appearance
to some specimens. Dorsal and pectoral fins hyaline or the fin rays are finely
peppered with brown microchromatophores, caudal fin mainly brownish.

VOLUME 93, NUMBER 3 835

Table 1.—Frequency distributions of trunk and tail rings in species of Urocampus.

Trunk rings Tail rings
Species Le i
locale Tw ®» i0 i RM ~ 0 Sh ~2D SB GF 95.55 Ss Sh oO
U. nanus O29 el l Sree Quem
U. carinirostris
New Guinea l 1 1 1
Australia
Queensland Ona greet] ewD
New South Wales 3 33 31 1 ib 2b I Way eg a!
Victoria See One” l DATA MAS) te 32 8
S. Australia 1 1
Tasmania 1 1
W. Australia 3 5 hea | 4 1 1

Comparisons.—Urocampus nanus has somewhat higher counts of trunk
rings and dorsal-fin rays than its only known congener (usually 10-11 and
16 against 8-9 and 14 in carinirostris). The snout length in HL ratio is lower
in U. nanus (averages 2.2 against 3.1 in carinirostris), and the snout depth
in snout length and pectoral-fin length in HL ratios are both higher than
those of U. carinirostris (respectively average 3.2 and 6.4 against 1.5 and
4.9). The median dorsal snout ridge of U. nanus is usually not clearly par-
alleled posteriad by anterior continuations of the supraorbital ridges (parallel
for some distance in most carinirostris) and the median dorsal snout ridge
of mature males is lower than that of comparable U. carinirostris (Figs. 1-
3). The enlarged barbel-like mandibular flaps present in many U. nanus are

>

Table 2.—Frequency distributions of total rings and dorsal-fin rays in species of Urocampus.

Total rings Dorsal-fin rays
Species cD es Ee a OE eS 0 ee 1
locale SS oa COL ma Olen 62 2 PE63 6465 66 67-68 [1 CY athe wel. a
U. nanus l 302 OD anil «1
U. carinirostris
New Guinea 1 | 2
Australia
Queensland 10 2 13
New South Wales Serle. 22 ae Ii. 7, 4 oe? | ak al
Victoria ASOmm Onn lane | 2

Tasmania l

4
S. Australia 1 |
l
W. Australia ll 4 | l 6

836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Frequency distributions of pectoral-fin rays and subdorsal rings in species of
Urocampus.

Pectoral-fin rays Subdorsal rings
Species Ca Ee Pee Gem
locale 7 8 9 10 2.50 DS 3.00 3.25 3.50 3.75 4.00
U. nanus l 21 9 6 3 7 l
U. carinirostris
New Guinea 4 | l
Australia
Queensland 1 9 9 2 6 4
New South Wales 6 74 39 D 3 11 Di 24 IS ]
Victoria l 15 7 l 5) 8 2
S. Australia 2 1
Tasmania y) |
W. Australia 2 3 l 4 py) |

replaced by small and rather inconspicuous flaps in U. carinirostris and,
when well developed, dermal flaps are generally more abundant and most
profusely branched in the latter species.

Types.—Duncker (1915) indicated that the type material of U. nanus con-
sisted of two uncataloged females in the BMNH collections but Gunther’s
(1870) description was based on a single 96 mm female specimen. Duncker
evidently examined two fish (96-111 mm SL) now cataloged as BMNH
1863.1.22.2 and labeled ‘‘Manchuria, Adams.’’ The larger specimen is from
an unknown locality and was added to the jar containing the holotype after
1870 (A. C. Wheeler, pers. comm.). Giinther (1870) counted 11 + 50 rings

Table 4.—Frequencies of dorsal-fin origin on Sth through 9th tail rings in species of Uro-
campus.

Tail rings
Species
locale 5.00 5.25 5.50 5.75 6.00 6.25 6.50 6.75 7.00 7.25 7.50 7.75 8.00 8.25 8.50 8.75 9.00
U. nanus AED) 35 ae l
U. carinirostris
New Guinea l |
Australia
Queensland et? | ANS! GL
New South Wales Der Qh Ss) Se a7 whe On ivalSa Otmann: D
Victoria 1 » Ll BE She ade eRe Ore Fy)
S. Australia |
Tasmania |

W. Australia l 1 | ite l 1

VOLUME 93, NUMBER 3 837

and 15 dorsal-fin rays in the holotype, indicated a total of 3 subdorsal rings
and stated that there was no anal fin. I count 10 trunk rings in the holotype
(the anus and 2-rayed anal fin share the 11th), 53 tail rings, 16 dorsal-fin rays
and 3.5 subdorsal rings. Although not mentioned in the original description,
the holotype has long mandibular flaps, long flaps near the dorsal midline
of most trunk rings, some flaps on the median ventral trunk ridge and there
are paired bilateral flaps on the dorsum of the Ist, 4th and 7th tail rings as
well as a single flap on the left side of the dorsum of the 6th.

Jordan and Snyder (1901) reported 59 tail rings and 5 subdorsal rings in
the holotype of U. rikuzenius, whereas I find 56 tail rings and 3.75 subdorsal
rings in this specimen (CAS-SU 6520).

Remarks.—Jordan et al. (1913) implied that U. rikuzenius was conspecific
with U. nanus and Duncker (1915) tentatively referred rikuzenius to the
synonymy of the latter species. These early statements have been largely
ignored or overlooked and most recent authors have incorrectly employed
the specific name rikuzenius.

Among the study material, dermal flaps are abundant and well developed
on the holotype and an 84 mm juvenile or female, in some specimens only
the mandibular flaps persist, and flaps are absent from two mature males
(95.5-105.5 mm SL). Well-developed flaps were noted in smaller fish (45.5—
75.2 mm SL) by Takai and Mizokami (1961).

The brood pouch extends below 9-11 rings in three males examined (95.5—
105.5 mm SL). The largest specimen has eggs in about four crowded trans-
verse rows and in 1-2 layers; there are about 17 eggs in the outer left row
through 6 of the 9 pouch rings. Takai and Mizokami (1961) reported two
layers of two rows of pouch eggs in a 62.4 mm male, noted that the ovoid
pouch eggs were about 0.5 x 0.7 mm in diameter, and counted 69 ovarian
eggs in each of two females (45.5—75.2 mm SL).

Examined materials include few data on depth of collections or habitat
but U. nanus is evidently most common in protected shallow inshore en-
vironments. The species was reported as a year-round resident of the Zos-
tera zone in the Amakusa Islands by Kikuchi (1968, 1970) and additional
notes on habits and behavior were provided by Takai and Mizokami (1961).
The holotype was reportedly collected in ‘‘Manchuria’’ and, without addi-
tional evidence, this was interpreted to be “‘in Yellow Sea off coast of
northeast China’’ by Lindberg and Legeza (1965).

Although the type locality is uncertain and the geographic range is pres-
ently undefined, U. nanus has been reported from Pusan and Masan, Korea
(Mori, 1952) and from Sado Island and Matsushima to Kochi Prefecture,
Japan (Kamohara, 1964). The southernmost record is evidently represented
by 4 specimens (FMNH 83875) reportedly collected in the Haneji River,
Okinawa (ca. 26°35'’N, 128°05’E).

Material examined.—17 specimens, 84-133 mm SL, including holotype.

838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Holotype. —BMNH 1863.1.22.2 (96.0 mm SL, female), Manchuria, A.
Adams.

Other material.—KOREA, Fusan: UMMZ 205279 (1, 101.5). JAPAN,
Kyoto Pref.: GCRL 15701 (2, 99.5—105.5). Kanegawa Pref.: UMMZ 205271
(3, 111.5-133), UMMZ 205274 (1, 130), YCMP 1223 (1, 127.5), YCMP 3522
(1, 84). Kagoshima Pref.: CAS-SU 6520 (117.5, holotype of U. rikuzenius).
Okinawa: FMNH 83875 (4, 99-112). Loc. uncertain: USNM 70773 (1, 103).
LOC. UNKNOWN: BMNH 1863.1.22.2 (1, 111).

Urocampus carinirostris Castelnau
Figs. 2—4

Urocampus carinirostris Castelnau, 1872:200 [orig. descr.; Melbourne (Aus-
tralia)].

Urocampus coelorhynchus Gunther, 1873:103 [orig. descr.; Sydney (Aus-
tralia)].

Urocampus Gintheri Duncker, 1909:242, figs. 1-2 (orig. descr.; Sharks Bay,
W. Austr.).

Urocampus carinorostris. Coleman, 1933:87 (Misspelling).

Urocampus guentheri. Whitley, 1948:14 (emended spelling).

Stigmatophora boops (not of Castelnau). Bertin and Estéve, 1950:50 (mis-
identification).

Diagnosis.—Snout short, its length averages 3.1 in HL; trunk rings 7-10,
usually (94%) 8-9; dorsal-fin rays usually (92%) 14.

Description.—Dorsal-fin rays 13-15 (x = 14.0), pectoral-fin rays 7-10
(8.3), rings 7-10 + 49-59 = 58-68 (62.0), total subdorsal rings 2.5—4.0 (3.4),
dorsal-fin origin on Sth—8th (6.6) tail ring; see Tables 1-4 for additional
counts. Proportional data based on 32 specimens 56.0-95.0 (66.5) mm SL
follow: HL in SL 9.7-13.1 (10.8), snout length in HL 2.8-3.6 (3.1), snout
depth in snout length 1.3-1.8 (1.5), length of dorsal-fin base in HL 1.2-1.7
(1.6), anal ring depth in HL 2.3-4.6 (3.5), trunk depth in HL (3 fish) 1.9-
2.4, pectoral-fin length in HL 3.9-6.4 (4.9), length of pectoral-fin base in
pectoral-fin length 1.4—2.2 (1.7).

Median dorsal snout ridge (Figs. 2—3) more or less linear to slightly con-
cave in females, not elevated to or above horizontal through dorsal margin
of eye; ridge clearly convex and usually elevated to or above dorsal margin
of eye in mature males. Supraorbital ridges continued anteriad on each side
of median snout ridge to near vertical from nares; lateral profile of head
depressed behind the eye but somewhat elevated over posterior part of
head.

Dermal flaps simple or irregularly branched distally. In well-preserved
males, head flaps may include long branched flaps over eye and on subor-
bital and short simple or branched flaps below angle of gape, on ventral

VOLUME 93, NUMBER 3 839

Fig. 3. Urocampus carinirostris: Lateral and dorsal aspects of head and anterior trunk
rings, together with section of body illustrating ridge configuration and dorsal and anal fins.
Top: 64.5 mm SL, male. Remainder: 66.5 mm SL, female (GCRL 14799).

midline of snout, on median dorsal snout ridge, on lower half of opercle and
opercular membrane, as well as on the posterior part of the supraorbital
ridge and on the frontal ridge. In females, head flaps are usually reduced to
short simple flaps on the supraorbital and frontal ridges. In males, each
trunk ring may have a pair of long bilateral flaps on the dorsum, a long flap

840 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Partially straightened brood-pouch larva of Urocampus carinirostris (ca. 11.5 mm
10).

on each lateral ridge and one on the median ventral ridge; tail flaps are
usually confined to the anterior 20-25 rings and each ring may bear paired
flaps on the dorsum and single long flaps on the inferior ridges which, on
anterior 5-8 rings, may be accompanied by a row of 4—7 minute simple
flaps. Flaps are usually absent from the median ventral trunk ridge in fe-
males and those on the dorsum and sides of body are typically shorter and
less obvious than those of males.

Coloration.—Ground color tan to dark brown; head mainly mottled with
pale, often with a dark brown stripe on ventral midline; sides and dorsum
of body plain or mottled, occasionally with indications of narrow pale bars
(ca. 1-2 rings wide) crossing dorsum of trunk and anterior part of tail; venter
often with dark brown stripe on midline of trunk but elsewhere plain or
mottled. Dorsal-fin rays usually with 2—S brown spots, the membrane hya-
line; pectoral fin shaded lightly with brown; caudal fin brownish, often mar-
gined with pale.

Comparisons.—See under U. nanus.

Types.—Castelnau (1872) failed to designate type material but his descrip-
tion indicates that both males and females were examined, that the largest
specimen was “‘three and a-half inches’’ (TL) and that the species was rather
common. Examination of pipefishes in the Paris collection shows that the
10 fish in MNHN A.1435, listed as ‘“‘paratypes’’ of Stigmatophora boops

VOLUME 93, NUMBER 3 841

Castelnau by Bertin and Estéve (1950), are actually Urocampus cariniros-
tris. These specimens, including males and females (52-72 mm SL) labeled
‘‘Australia, Yarra River, Castelnau,’’ are here considered to be the pre-
sumptive syntypes of U. carinirostris. Both carinirostris and boops were
treated by Castelnau (1872) and subsequent errors in labeling or cataloging
appear certain. The holotype of U. Giintheri Duncker, deposited in the
Hamburg Museum, has been destroyed.

Remarks.—McCulloch (1909) incorrectly reported the presence of 4 lat-
eral ridges on the body (trunk) of U. carinirostris and implied that a pair of
these were confluent with the inferior tail margin. Evidently misled by
McCulloch’s description, Munro (1958) reported a “‘double’’ lateral trunk
ridge in this species and Scott (1961) employed this nonexistent “‘double”’
ridge as a differentiating character in his key to the Tasmanian Syngnathi-
dae.

Examined Australian collections include brooding males taken in Western
Australia (Feb., Oct.), Victoria (Nov.), New South Wales (Feb., Apr.,
Sept., Oct., Dec.) and Queensland (July). Scott (1971) noted brooding males
collected in Tasmania during April.

The brood pouch extends below 8—11 (usually 9-10) anterior tail rings in
34 examined males 45—95 mm SL. The free margins of the pouch membranes
are usually edged with narrow, laterally directed, folds and the margins meet
or, more commonly, fail to meet on the ventral midline of the egg-filled
pouch. Pouch eggs are usually deposited in a single layer of 1—4 transverse
rows and are often absent from the posteriormost pouch rings. The smallest
examined brooding male (48 mm SL) contained only 2 eggs in the 8-ring
pouch, a 57.5 mm fish had 2 rows of 19 eggs through 8 of 10 pouch rings,
and there were 4 rows of 17 eggs through 9 of 10 rings in a 76 mm specimen.
Rather large (ca. 10-14 mm TL) larvae (Fig. 4) are often found coiled within
the open membranous compartments lining the sides and dorsum of the
brood pouch. The anal fin and pectoral-fin rays are not evident at x60
magnification in 11-12 mm pouch larvae but they are distinct in a 14 mm
specimen and the dorsal and caudal fins are well developed in 10 mm larvae.
The 14 mm larva had the head and body peppered with brown microchro-
matophores and about 6 narrow brown bands encircled the tail behind the
dorsal fin.

There were 7—9 trunk rings in 57 adult males and the count was 8 in 82%,
whereas the range was 8-10 in 50 adult females and the modal value was 9
(88%). Other meristic data (Table 1) suggest clinal variation wherein tail ring
frequencies are highest in samples from Tasmania and Victoria and lowest
in material from New Guinea, Queensland and Western Australia. Addi-
tional study is required for confirmation of this apparent trend in U. cari-
nirostris but similar geographic variation is indicated (author’s unpublished
data) for other Australian pipefishes.

Although dredge collections among Zostera sp. were noted by Scott

842 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(1965), the majority of examined material was seined in depths to ca. 3 m
and all collections appear to be from the lower reaches of rivers, estuaries
or other protected inshore habitats. A number of samples indicate that col-
lections were from areas of algae or Zostera. This is a small pipefish which
probably seldom attains 100 mm SL. The species is presently known only
from Australia and New Guinea. Examined Australian specimens were col-
lected along southern coasts from Narang, Queensland (ca. 24°53’S,
151°48’E) to Crawley Bay, Swan River, Western Australia (ca. 32°01’S,
115°48’E).

Material examined.—151 specimens (excluding pouch larvae), 35.5—95.0
mm SL, including 10 presumptive syntypes.

Presumptive syntypes.—MNHN A.1435 (10, 52.0-72.0 mm SL), Yarra
River, Victoria, Australia, F. de Castelnau.

Other material—NEW GUINEA, Tobriand Is.: AMS [I.107095-010 (1,
59.5), USNM 215314 (1, 67). AUSTRALIA, Queensland: QM 1.8729 (1, 51),
QM 1.8733 (1, 64.5), QM 1.13379 (4, 59.5-65.5), QM I.-6705 (7, 51-63.5).
New South Wales: AMS 1.16475-017 (1, 63.5), AMS 1.19488-001 (1, 71),
BMNH 1873.4.3.74-5 (2, 84-95, syntypes of Urocampus coelorhynchus),
BMNH 1873.4.3.210 (1, 88), BMNH 1890.2.26.199 (1, 58), GCRL 14799 (5,
62.5-71), GCRL 15506 (1, 71.5), GCRL 15507 (7, 64-72), GCRL 16268 (1,
ca. 55.5), GCRL 16357 (4, ca. 51-69), GCRL 16367 (2, 54.5-57), GCRL
16371 (2, ca. 35.5—47.5), GCRL 16373 (2, 36-63.5), GCRL 16378 (8, 38-76),
GCRL 16448 (35, 47—58.5), USNM 215308 (7, 55-64), USNM 215309 (1, 67),
USNM 215310 (20, 37-68). Victoria: NMV A.551 (8, 50.5—91), NMV A.555
(1, 79), QM 1.16657 (2, 56.5-65). South Australia: CAS 36433 (1, 68.5), SAM
F.3441 (1, 65.5). Tasmania: GCRL 14766 (1, 93), QVM 1968.5.33 (1, 84).
Western Australia: AMS IA.7445-8 (4, 49.5—65.5), AMS I.15724-007 (1, 64),
NMV A.696 (2, 63.5-73.5), WAM P.25701-001 (1, 55), WAM P.26473-001
(1, 50), WAM P.26475-001 (1, 50).

Acknowledgments

For permission to examine materials in their care and for other courtesies
and assistance, I thank G. R. Allen and J. B. Hutchins (WAM), R. M.
Bailey and Ellie Koon (UMMZ), M. L. Bauchot (MNHN), W. N. Esch-
meyer and Pearl Sonoda (CAS), C. J. M. Glover (SAM), M. F. Gomon
(NMV), R. H. Green and E. O. G. Scott (QVM), R. K. Johnson (FMNH),
E. A. Lachner and Susan Karnella (USNM), R. J. McKay (QM), J. R.
Paxton and D. F. Hoese (AMS), and A. C. Wheeler (BMNH). Special ac-
knowledgment is due K. Meguro (The Crown Prince’s Household, Japan)
and M. Hayashi (YCMP) for their cooperation in arranging the loan of im-
portant material. Valuable gifts of specimens were received from I. Naka-
mura (Fish. Res. Sta., Kyoto Univ.), J. R. Paxton and D. F. Hoese (AMS),

VOLUME 93, NUMBER 3 843

E. O. G. Scott (QVM) and from Alison and Rudie Kuiter (Sydney, Austr.).
Dr. T. Abe (Univ. Tokyo) provided locality data for the Haneji River. Draw-
ings are by Mrs. Nancy Gordon.

Literature Cited

Bertin, L., and R. N. Estéve. 1950. Catalogue des types de poissons du Muséum National
d’Histoire Naturelle. 6° partie. Haplomes, Hétéromes, Catostéomes.—Imprimerie Na-
tionale, Paris, 1-60.

Castelnau, F. de. 1872. Contribution to the ichthyology of Australia.—Proc. Zool. Acclim.
Soc. Victoria 1:29-242.

Coleman, E. 1933. Gleanings from Marlo—the pipefish.—Victorian Naturalist 50(4):86—-88,
figs.

Dawson, C. E. 1976. Review of the Indo-Pacific pipefish genus Choeroichthys (Pisces: Syng-

nathidae), with descriptions of two new species.—Proc. Biol. Soc. Wash. 89:39—66, figs.

1-9.

. 1977a. Review of the Indo-Pacific pipefish genus Lissocampus (Syngnathidae).—Proc.

Biol. Soc. Wash. 89:599-620, figs. 1-7.

. 1977b. Synopsis of syngnathine pipefishes usually referred to the genus Ichthyocam-

pus Kaup, with description of new genera and species.—Bull. Mar. Sci. 27:595—65S0,

figs. 1-19.

1979. The Indo-Pacific pipefish genera Notiocampus gen. nov. and Nannocampus

Gunther.—Proc. Biol. Soc. Wash. 93:483-—492, figs. 1—S.

, and G. R. Allen. 1978. Synopsis of the “‘finless’’ pipefish genera (Penetopteryx,

Apterygocampus and Enchelyocampus, gen. nov.).—Rec. West. Austr. Mus. 6:391-

411, figs. 1-7.

, and R. A. Fritzsche. 1975. Odontoid processes in pipefish jaws.—Nature 257:390, fig.

Duncker, G. 1909. Pisces, I. Teil: Syngnathidae. Pages 233-250, figs. 1-20. In W. Michaelson

and R. Hartmeyer, eds.—Die Fauna Stidwest-Australiens, 2. Jena.

. 1910. On some syngnathids (“‘pipe fish’’) from Ceylon.—Spolia Zeylanica 7(25):25—

34, pl.

. 1915. Revision der Syngnathidae.—Mitteil. Naturh. Mus. Hamburg 32:9-120, pl.

Gunther, A. 1870. Catalogue of fishes in the British Museum. Vol. 8. 549 pp. London.

1873. Erster ichthyologischer beitrag nach Exemplaren aus dem Museum Godef-
froy.—J. Mus. Godeffroy 1(2):1—128, 83 pls. ig

Herald, E. S. 1959. From pipefish to seahorse—a study of phylogenetic relationships.—Proc.
Calif. Acad. Sci., 4th ser., 29:465—473, figs. 1-3.

Jordan, D. S., and J. O. Snyder. 1901. A review of the Hypostomidae and lophobranchiate

fishes of Japan.—Proc. U.S. Nat. Mus. 24:1-20, pls. 1-12.

, 8. Tanaka, and J. O. Snyder. 1913. A catalogue of the fishes of Japan.—J. Coll. Sci.,

Tokyo Imp. Univ. 33:1—497, figs. 1-396.

Kamohara, T. 1964. Revised catalogue of fishes of Kochi Prefecture, Japan.—Rep. Usa Mar.
Biol. Sta. 11:1-99, figs. 1-63.

Kikuchi, T. 1968. Faunal list of the Zostera marina belt in Tomioka Bay, Amakusa, Kyushu.—

Publ. Amakusa Mar. Biol. Lab. 1:163—192.

. 1970. Fauna and flora of the sea around the Amakusa Marine Biological Laboratory.

Part VIII. Fishes.—Amakusa Mar. Biol. Lab., 52 pp.

Lindberg, G. U., and M. I. Legeza. 1965. Ryby Yaponskogo morya i sopredel’nykh chastei
Okhotskogo 1 Zheltogo morei. (Fishes of the Sea of Japan and adjacent parts of the Sea
of Okhotsk and Yellow Sea.)—Acad. Sci. USSR 2:1-391, figs. 1-324 (1969 English
trans., Israel Program Sci. Transl.).

844 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

McCulloch, A. R. 1909. Studies in Australian fishes. No. 2.—Rec. Austr. Mus. 7(4):315-321,
fig. 18, pls. 90-91.

Mori, T. 1952. Check list of the fishes of Koreaa—Mem. Hyogo Univ. Agr., Biol. Ser. 1:1-
228.

Munro, I. S. R. 1958. Family Syngnathidae. Handbook Austr. Fishes. 21:85—88.—/n Austr.
Fish. Newsletter 17(3):17—20, figs. 587-615.

Scott, E.O.G. 1961. Observations on some Tasmanian fishes: part X.—Pap. Proc. Roy. Soc.
Tasmania 95:49-65, figs. 1-3.

1965. Observations of some Tasmanian fishes: part XIII.—Pap. Proc. Roy. Soc.

Tasmania 99:53-65, fig.

. 1971. Observations on some Tasmanian fishes—part XVIII.—Pap. Proc. Roy. Soc.

Tasmania 105:119-143.

Takai, T., and A. Mizokami. 1961. Some notes on the morphology and ecology of the pipefish,
Urocampus rikuzenius (Jordan and Snyder).—J. Shimonoseki Coll. Fish. 11:79-84, figs.
1-3.

Whitley, G. 1948. A list of the fishes of Western Australia.—Fish. Bull., W. Aust. Fish. Dept.
2:1-35.

Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi
39564.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 845-848

NEW FRESHWATER SHRIMP RECORDS FOR TOBAGO,
WEST INDIES, WITH A SUMMARY OF RECORDS
FOR THE LESSER ANTILLES
(Atyidae and Palaemonidae)

Caw Hart auir-

Abstract.—Collections made on Tobago, W.I., in April 1978 increased
the known freshwater shrimp fauna of that island to 10 species—Atya in-
nocous, A. scabra, Jonga serrei, Potimirim glabra, Xiphocaris elongata,
Macrobrachium acanthurus, M. carcinus, M. crenulatum, M. faustinum,
and Palaemon (P.) pandaliformis. Before that time only 2 freshwater
shrimp species were known from Tobago.

In April, 1978, several staff members! from the Chesapeake Biological
Laboratory of the University of Maryland carried out a 2-week fisheries
survey of near-shore and freshwater habitats on Tobago, West Indies. M.
L. Wiley was primarily responsible both for collecting the decapods and
sending them to Austin B. Williams of the National Marine Fisheries Service
Systematics Laboratory, Washington, D.C. Subsequently, Dr. Williams
made the freshwater shrimps available to me.

I thank Drs. Wiley and Williams; Mr. Hardy, who initiated the survey;
and Mr. H. E. Wood, Senior Fisheries Officer, Ministry of Agriculture,
Land and Fisheries Division, Port-of-Spain, Trinidad, who gave permission
for the survey to be made.

Collections containing palaemonid and atyid shrimps were made at 6 lo-
calities (Fig. 1) on Tobago; night and daytime collections were made at
Station C; and rotenone was used to aid in collecting at Station B.

The station localities are as follows:

A.—Bloody Bay River, lower coastal lagoon just above beach. 6 April 1978.
11°18’N; 60°38'W.

B.—Bloody Bay River at barrier beach pool (collected with rotenone). 15
April 1978. 11°18'N; 60°38’W.

C.—Doctor’s River, in small dammed pond near Speyside (collections made
in daylight and at night). 9 April 1978. 11°18’N; 60°32’W.

D.—Merchiston River, below Windward Highway down to barrier pool. 9
ANT NOV Se tl alGwN "60232 We

‘J. D. Hardy, Jr., leader; L. Lubbers III; F. D. Martin; D. Shelton; and M. L. Wiley.

846 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Tobago

~

Fig. 1. Tobago, W. I., showing locations of collecting stations.

E.—Roxborough River, about 1 mile N of windward Highway. 9 April 1978.
11°14’N; 60°35’W.

F.—Goldsborough of Great River, just upstream of bridge at Windward
Highway. 7 April 1978. 11°12’N; 60°38’ W.

Some 200 specimens of caridean shrimps, representing 8 species in 6
genera, were collected from the 6 localities. Data concerning the species are
given in Table 1.

When Chace and Hobbs (1969) summarized the available information on
freshwater and terrestrial decapods of the West Indies, they mentioned
Macrobrachium faustinum as the only freshwater shrimp having been re-
corded from Tobago. Later, a record of Atya scabra was discovered by
Hobbs (Ortman, 1895), and this, together with the 8 species collected by the
Chesapeake Biological Laboratory team, bring the island total of known
caridean shrimp species to 10 (Table 2).

In discussing the Antillean distribution of the monotypic genera Jonga
and Micratya, Chace and Hobbs (1969:20) surmised that those species
reached the Greater Antilles from the Central American-Mexican region and
spread southeastward. They based their reasoning on the observation that
Jonga and Micratya were apparently absent from the lower islands of the
Lesser Antillean chain and from South America. The new information does

847

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o =: I =
> 2k 2K OK OK ok KOK *
9 —— —— ——
¢ — —— —
roa 2K OK OK OK ae = *
ak = ae G
Be =? i C
ar 2k 2k OK Ok a *
ia —- a C
oa =o a 174
a =e aan 14
<- oe = 9
ro nF a ¢
i. = a IT
me = == 6l
ial ar as C
9 ak it ¢
I aay el ¢
d a d @)
suoleis

(SIAO)

(e101) 6

(B10) @
((e101) 4
2

XS

smm4sofijppund (*q) uowavjvg

SNANYJUDID WNIYIDAGOAIDPV

WHNIDINUIAD WNIYIDAGOAIDY

SNUIDADI WINIYIDAGOAIDG
snoJouul vay

DIDsUO]a S1IADIOYALY

DAGD]S U1dUNO”d

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soisedsg

"8/61 [Lidy ul ‘[' AA ‘O8eqoO] UO poaxda]Joo sduiys Uvapled JayeMYSOIJ JO SaIdeds g ay} UO RIVGQ—'| I1qGR]

. wc —— + - — ——EE —

848 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Summary of freshwater shrimp records for the Lesser Antilles.

St. Martin

Barbuda

Montserrat

e + _ 4 JI

Guadeloupe| @ ) |} eo U|®@ (a)
; | | | | |
@

Dominica

Martinique |

St. Lucia | @ ® @
St. Vincent} @ | @ | |
Barbados @ | @ ® @ @
@
@

Grenada

Tobago wil Ww IL ee | See | Sar
Trinidad e@ @

Ju + +

Bonaire
+

w
@
@
Curacao e@ ® |
@
Ww

4 uL + {| +

Aruba

New Tobago records

@ Chace & Hobbs LJ Léeveque A Oriman

not dispute this conclusion as it relates to Micratya, but the occurrence of
Jonga on Tobago leads one to believe that it may have entered the Antillean
chain from either Central or South America. The presence of Jonga on an
island so close to South America suggests that it may also be present, but
overlooked, in the mainland waters of the continent.

Reports of the behavior of another shrimp (H. E. Wood, personal com-
munication) found in areas away from the coastline, indicate that Macro-
brachium heterochirus is, in all probability, also to be found on Tobago.

Literature Cited

Chace, Fenner A., Jr., and Horton H. Hobbs, Jr. 1969. The freshwater and terrestrial decapod
crustaceans of the West Indies with special reference to Dominica.—United States Na-
tional Museum Bulletin 292: 1-258.

Lévéque, Christian. 1974. Crevettes d’eau douce de la Guadeloupe (Atyidae et Palaemoni-
dae).—Cahiers O.R.S.T.O.M., serie Hydrobiologie 8(1):41-49.

Ortman, A. E. 1895. A study of the systematic and geographical distribution of the decapod
family Atyidae Kingsley.—Proceedings of the Academy of Natural Sciences of Phila-
delphia 46(for 1894):397-416.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Instituion, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(3), 1980, pp. 849-874

TWO NEW PHOXOCEPHALID GENERA, FUEGIPHOXUS
AND PHOXORGIA, FROM MAGELLANIC SOUTH
AMERICA (AMPHIPODA: CRUSTACEA)

J. L. and C. M. Barnard

Abstract.—Parharpinia fuegiensis Schellenberg, 1931, from Magellanic
South America, is made the type-species of Fuegiphoxus a new genus gen-
erally in the Brolginae, a dominantly Australian subfamily. Fuegiphoxus
appears to be a plesiomorph of the Australian Elpeddo. Two new Magellanic
species are added to Fuegiphoxus along with the Antarctic Pontharpinia
uncinata Chevreux, 1912. Phoxorgia is described for Parharpinia sinuata
K. H. Barnard, 1931, and appears to be plesiomorphic both to the Australia
Parharpinia and the predominantly North American Metharpinia.

Two more new genera of Phoxocephalidae are now extracted from the
polytypic Paraphoxus Sars, a genus which should be reduced to 2 species
(see Barnard and Drummond, 1976, 1978 and J. L. Barnard, 1979, 1980).

We thank Dr. Charlotte Holmquist of the Swedish Museum of Natural
History for loaning us Schellenberg’s (1931) material for the second time in
25 years. We hope this time will be the last and that the species are now
adequately analyzed. We thank Elizabeth B. Harrison, Janice Clark, Irene
F. Jewett and Roland H. Brown for their considerable help. Dr. Wim Vader
of Troms®@ offered valuable advice.

Antecedents for styles and methods of description are found in Barnard
and Drummond (1978).

Key to Illustrations

Capital letters refer to appendages and parts of the amphipod; lower case
letters to left of capitals refer to specimens cited in legends and Voucher
Material; lower case letters to right of capitals or within parts refer to con-
ditions as listed below: A, antenna; B, buccal complex, lateral view; D,
dactyl of a pereopod; E, epimeron; F, accessory flagellum; G, gnathopod;
H, head; I, inner plate or ramus; L, lacinia mobilis; M, mandible; N, molar;
O, outer plate or ramus; P, pereopod; R, uropod; S, maxilliped; T, telson;
U, upper lip; W, pleon; X, maxilla; Y, palp of mandible. b, broken; d, dorsal;
1, left; m, medial; p, apex of peduncle; r, right; s, setae removed.

Fuegiphoxus, new genus

Diagnosis.—Eyes present. Flagella of antennae 1—2 unreduced in female,
article 2 of antenna | shortened to ordinary, ventral setae confined apically.

850 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Article 1 of antenna 2 not ensiform, article 3 with 2 setules, facial spines on
article 4 in one main row or 2 rows, article 5 ordinary in size. Right man-
dibular incisor with 3 teeth, molar not triturative, small, pillow-shaped, bear-
ing 2 short and one elongate spines, not bearing patch of pubescence, palpar
hump small. Palp of maxilla | biarticulate, inner plate with 4 setae. Setation
of maxilla 2 ordinary. Inner plate of maxilliped ordinary, apex of palp not
strongly protuberant, dactyl elongate, apical nail distinct, medium. Gnatho-
pods small but dissimilar, gnathopod 2 weakly to moderately enlarged, ar-
ticle 5 of gnathopod | of ordinary length but on gnathopod 2 very short and
almost cryptic, without eusirid attachment, palms oblique, hands of gnatho-
pods I—2 respectively ovatorectangular and slightly broadened, poorly se-
tose anteriorly. Article 2 of pereopod 5 of broad form, articles 4—5 of per-
eopods 5-6 narrow to medium, article 2 not setose posteriorly, pereopod 7
ordinary, article 2 naked ventrally, article 3 ordinary, dactyl normal. Pe-
duncle of uropod | normally elongate, without apical spike, without dis-
placed apical spine, only peduncular apices of uropods 1—2 with faint comb,
inner ramus of uropod | with one row of marginal spines, no rami contin-
uously spinose to apex, inner ramus of uropod 2 ordinary, uropod 3 ordi-
nary, bearing article 2 of outer ramus, carrying 2 medium to long apical
setae. Telson ordinary, each lobe with 2 apical spines plus setules on each
lobe, without special dorsal and lateral spines or setae. Epimera 1-2 bearing
Sparse short posterior setae, without midfacial setae above ventral facial
ridge, epimeron 3 ordinary, bearing 3 or more long setae. Urosomite | with-
out large lateral armament, bearing or lacking one or more midventral cres-
cents or bundles of setae. Urosomite 3 without dorsal hook or process.

Description.—Rostrum fully developed, unconstricted. Pubescence on
article | of antenna | in male present; calceoli on male primary flagellum of
antenna | present, calceoli on article 5 of male antenna 2 present, flagellum
in male with calceoli. Prebuccal parts ordinary. Right lacinia mobilis bifid,
subflabellate, article 1 of mandibular palp short to slightly elongate, palp
thin, apex of article 3 oblique, article 2 without outer setae. Lower lip
bearing cones. Outer plate of maxilla 1 with 10-11 spines, one spine espe-
cially thickened. Inner plates of maxillipeds with one main spine, ordinarily
setose. Coxae 2—4 without special anterodorsal humps. Posterior spines on
article 6 of pereopods 3-4 thick and stiff, midapical spine or seta absent;
article 2 of pereopod 7 without facial setae. Peduncle of uropod 1 with
dorsolateral spines confined apically (possibly widely spread on wncinata),
medial spines widely spread. Peduncle of uropod 2 with only one medial
spine confined apically, peduncle of uropod 3 lacking extra subapical setae
or spines. Telson with ordinary pair of midlateral or dorsal setules on each
side.

Etymology.—From *‘Tierra del Fuego,’’ modified for euphonic reasons
and from **Phoxocephalus,’’ the type-genus of the family. Masculine.

b)

VOLUME 93, NUMBER 3 851

Type-species.—Parharpinia fuegiensis Schellenberg, 1931.

Composition.—Pontharpinia uncinata Chevreux, 1912; Fuegiphoxus in-
utilus, new species; F. abjectus, new species.

Remarks.—Fuegiphoxus differs from Paraphoxus Sars in the: (1) weak
division of the spine rows on article 4 of antenna 2; (2) proximal position of
the dorsal notch on article 4 of antenna 2; (3) elongation of the third spine
on the mandibular molar; (4) presence of 4 (not 2) setae on the inner plate
of maxilla 1; (5) presence of a thick apical spine on the inner plate of the
maxilliped; (6) elongate article 5 of gnathopod 1; (7) significantly enlarged
gnathopod 2; (8) cryptic article 5 of gnathopod 2; (9) well developed facial
setae on articles 4-5 of pereopods 3—4; (10) apical digitation of article 6 on
pereopod 7; and (11) lowered flexibility of the apical nail on the rami of
uropods 1-2.

Fuegiphoxus differs from Wildus Barnard and Drummond in the: (1) clear
division of spines into rows on article 4 of antenna 2; (2) slightly longer
article 5 of antenna 2; (3) diversity of spine size on the mandibular molar;
(4) presence of 4 setae on the inner plate of maxilla 1 and the presence of
11 spines on the outer plate (not 9); (5) presence of the thick spine on the
inner plate of the maxilliped; (6) deeper cleft on the inner plates of the
maxillipeds; (7) incompleteness of the cryptic condition on the wrist of
gnathopod 2; (8) strong facial setation on articles 4—5 of pereopods 3-4; (9)
retention of apical digits on article 6 of pereopod 7; (10) presence of ventral
setae on epimeron 3; and (11) deeply immersed apical nails on the rami of
uropods 1-2. In addition the type-species of Fuegiphoxus has more baso-
facial setae on the peduncle of uropod 1 (also satisfactory in inutilus but
unknown in uwncinatus) and the nondisplacement of the apicomedial spine
on the peduncle of uropod 1 (unknown in wncinatus and see W. waipiro, an
exception to the Wildus pattern). Generally, Fuegiphoxus has a more prox-
imal dorsal notch on article 4 of antenna 2 and larger outer plates of the
maxillipeds. :

Fuegiphoxus resembles Eyakia J. L. Barnard in the elongation of one
spine on the mandibular molar but differs from Eyakia in the: (1) short thick
article 2 on antenna 1; (2) additional proximal spine(s) in the formula on
article 4 of antenna 2 (but unknown in uwncinata); (3) absence of pubescence
on the molars; (4) presence of only one (not 2) main spine on the inner plate
of the maxilliped; (5) distinctly enlarged gnathopod 2; (6) untapered article
2 of pereopod 5; (7) poorly setose epimeron 3; and (8) more immersed apical
nails on the rami of uropods 1-2.

Elpeddo Barnard and Drummond (1978) in Australia may be an apomorph
of Fuegiphoxus because that monotypic genus has a peculiar male antenna
1 like the appendage of Fuegiphoxus abjectus. Elpeddo differs from Fue-
giphoxus in the loss of 2 of the 4 setae on the inner plate of maxilla 1, the
loss of the main spine on the inner plates of the maxilliped, has developed

852 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

WAH
SS Gh

S

Fig. 1. Fuegiphoxus fuegiensis: f = lectotype female ‘‘f’’ 6.44 mm, g = male ‘‘g’’ 7.83 mm.

VOLUME 93, NUMBER 3 853

a displaced spine on the apicomedial margin of the peduncle on uropod 1,
has giant calceoli on article 5 of antenna 2 and bears the neotenic or juvenile
form of uropod 3 in which article 2 on the outer ramus is elongate.

Key to the Species of Fuegiphoxus

1. Epimeron 3 with long thin blunt posterior teeth ........ F. uncinatus
Epimeron 3 rounded or broadly quadrate posteriorly ............. 2
2a. Epimera 1-2 with small posteroventral tooth, spine formula on ar-
ticle:4 ofsantenna 2 = 024-4-1 9 fan ctalosjen his Saoer (Lok F. inutilus
2b. Epimera 1-2 rounded posteroventrally, spine formula on article 4
Ofgantennar® = 0-6- hasan dacatelioaiss 2 Rie sole tes! F. fuegiensis
2c. Epimera 1-2 subquadrate posteroventrally, spine formula on article
ATOleantenna 2a=*323-3- lt jotioney 8 ota sinks eoees deka. | F. abjectus

Fuegiphoxus fuegiensis (Schellenberg)
Figs. 1—3 (part)

Parharpinia fuegiensis Schellenberg, 1931:78—-80, fig. 40.—Stephensen,
1949:5-6.
Paraphoxus fuegiensis.—J. L. Barnard, 1960:271, pl. 42.

Description of lectotype female ‘‘f.’’—Head about 20 percent of total
body length, greatest width about 75 percent of length, rostrum unconstrict-
ed, broad, short, reaching middle of article 2 on antenna 1; eyes large, clear
of pigment, ommatidia ordinary; article 1 of peduncle on antenna | almost
1.5 times as long as wide, about twice as wide as article 2, ventral margin
with about 12 setules (not all illustrated), produced dorsal apex with 3 set-
ules, article 2 about 0.55 times as long as article 1, with apicoventral cycle
of 7-8 setae, primary flagellum with 10 articles, about 0.8 times as long as
peduncle, bearing one short aesthetasc on each of articles 4-9, accessory
flagellum short, with 7 articles. Spine formula of article 4 on antenna 2 =
3-3-2, dorsal margin with notch bearing 4 setae, ventral margin with 5-6
groups of 1—4 long to medium setae, one ventrodistal long spine, article 5
about 0.7 times as long as article 4, facial spine formula = 1, dorsal margin
bearing one set of small setae, ventral margin with 3 sets of one seta each,
3 ventrodistal long to medium spines, one of these set subdistally; flagellum
about 0.95 times as long as articles 4-5 of peduncle combined, with 8 arti-
cles.

Mandibles with weak palpar hump, right incisor with 3 teeth, left incisor
with 3 humps in 2 branches, right lacinia mobilis bifid, distal branch much
shorter than proximal, flabellate, broad, subbifid, proximal branch simple,
blunt, with marginal denticles, left lacinia mobilis with 4 teeth plus one
accessory tooth, middle teeth shortened, right rakers 8 plus 2 rudimentaries,

854 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

left rakers 5 plus one rudimentary, molars composed of bulbous protrusions,
each molar with one long serrate and 2 short spines plus granulated callus,
without plume, palp article 1 slightly elongate, article 2 with one medium
inner apical seta and 2 other shorter inner setae, article 3 about 1.1 times
as long as article 2, oblique apex with 7 and 6 spine-setae, basofacial for-
mula = 1-1 and 0-1. Each outer lobe of lower lip with cone. Inner plate of
maxilla 1 large, thin, bearing one long apical pluseta, one shorter apico-
medial seta, 2 apicolateral much shorter setae, left outer plate with 10
spines, right outer plate with 11 spines, one thick on each side, palp article
2 with one apical spine, one apicolateral, 3 medial spines, and 4 submarginal
setae. Inner plate of maxilla 2 slightly shorter than outer, outer scarcely
broader than inner, outer with 5S apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with one large thick apical spine, 2 apicofacial
setae, 3 medial setae, outer plate with 8 medial spines, one apicolateral seta
and cusp, palp article 2 without apicolateral seta, article 2 with one apico-
lateral seta, medial margin of article 2 weakly setose, article 3 with 4 facial
setae, one lateral seta, nail of article 4 medium, with 2 accessory setules.

Coxa | not expanded apically, anterior margin weakly convex, main ven-
tral setae of coxae 1-4 = 10-11-11-10, posteriormost seta of coxae 1—4
scarcely shortened, anterior and posterior margins of coxa 4 almost parallel,
posterior margin convex, posterodorsal corner rounded, posterodorsal mar-
gin V-shaped, width-length ratio of coxa 4 = 6:7. Gnathopods with elongate
hands, gnathopod 2 larger than gnathopod 1 and with subcryptic wrist, width
ratios on articles 5-6 of gnathopods 1-2 = 30:38 and 32:50, length ratios =
66:88 and 50:88, palmar humps large, palms strongly oblique, article 5 of
gnathopod 1 elongate, ovate, posterior margin rounded-flat, article 5 of
gnathopod 2 triangular, posterior margin rounded-angular.

Pereopods 3—4 similar to each other, facial setae formula on article 4 =
6 and 5, parallel to apex, on article 5 = 5 and 5, main spine of article 5
extending to M. 90-100 on article 6, article 5 with no proximoposterior
spines, spine formula of article 6 = 3 + 2 and 4 + 2 plus no middistal seta,
spines especially long, medial spines tightly grouped (and inserted from
lateral side, not truly medial), acclivity on inner margin of dactyls of per-
eopods 3-4 obsolescent, emergent setule short, almost fully immersed, mid-
facial pluseta ordinary but highly anteriad. Coxae 5-7 posteroventral setule
formula = 3-3-4, gills of these coxae large. Articles 4-5 of pereopods 5-6
narrow, facial spine rows sparse, facial ridge formula of article 2 on per-
eopods 5-7 = 0-1-1, article 2 of pereopod 5 scarcely tapering distally. Width
ratios of articles 2, 4, 5, 6 of pereopod 5 = 41:23:18:10, of pereopod 6 =
64:22:15:9, of pereopod 7 = 80:16:13:7, length ratios of pereopod 5 =
69:26:36:37, of pereopod 6 = 88:48:53:55, of pereopod 7 = 100:20:25:30,
article 2 of pereopod 7 much broader than in female ‘“‘d’’ of Barnard (1960)
exceeding middle of article 4, posterior margin with 7 small serrations, one

VOLUME 93, NUMBER 3 855

Fig. 2. Fuegiphoxus fuegiensis: d = female ‘‘d’’ 6.0 mm, f = lectotype female “‘f’ 6.44
mm, g = male *‘g’”’ 7.83 mm.

856 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

for

Fig. 3. Upper, Fuegiphoxus fuegiensis: f = lectotype female ‘‘f’ 6.44 mm, g = male “‘g”’
7.83 mm. Lower, Fuegiphoxus inutilus, holotype female “‘p’’ 7.80 mm.

VOLUME 93, NUMBER 3 857

elongate posteroventral setule, medial apex of article 6 finely to coarsely
combed, bearing 7-8 digital processes.

Posteroventral corner of eipmeron | rounded-quadrate, posterior margin
almost straight, with 4 setules, anteroventral margin with 9 medium setae,
posteroventral margin with 3 long setae. Posteroventral corner of epimeron
2 rounded, posterior margin weakly convex, with 3 setules, facial setae =
6, no lateral ridge. Posteroventral corner of epimeron 3 rounded, posterior
margin straight, with 2 setule notches, ventral margin with 2 setae in pos-
terior half.

Urosomite | with 2 groups of 2 ventral setae, no lateral setae, articulation
line complete, urosomites unprotuberant dorsally. Rami of uropods 1-2 with
fused, immersed apical nails, outer ramus of uropod 1 with 1-2 dorsal
spines, inner with one, rami of uropod 2 naked, peduncle of uropod 2 with
6 basofacial to ventral setae and 2-3 apicolateral spines, narrowly confined
medially, with 4 marginal setae and spines, apicalmost weakly enlarged, but
not displaced, plus accessory slit. Peduncle of uropod 2 with 11 dorsal
spines, medially with one large apical spine, apicolateral corners of pedun-
cles on uropods 1-2 with faint comb. Peduncle of uropod 3 with 6-7 ventral
spines, dorsally with one long lateral spine, one small medial spine and
setule, rami submasculine, inner extending to M. 95 on article 1 of outer
ramus, apex with 2 setae, medial and lateral margins with 3 and one setae,
article 2 of outer ramus short, 0.17, bearing 2 medium to long setae, medial
margin of article 1 with 3 setae, lateral margin with 2 acclivities, spine
formula = 1-1-2, setal formula = 0. Telson long, length-width ratio = 6:5,
almost fully cleft, each apex of medium width, subtruncate, acclivity shal-
low, bearing short lateral setule, spine next medial longer than setule or
with lateral and medial spines separated by setule, midlateral setules di-
verse, largest setule small.

Description of male ‘‘g.’’—Article | of antenna | with medial pubescence,
article 2 with 9 ventral setae, primary flagellum with 14-articles, one cal-
ceolus each on articles 2-10, aesthetasc each on articles 2—11+, 3 aesthe-
tascs on article 1. Facial spine formula on article 4 of antenna 2 = 3-3-3,
article 5 with 3 dorsal sets of male setae and one small calceolus, ventro-
distal apex with 2 thin short spines, calceolar formula of elongate flagellum =
IEP 46.79 lil 13s | na Orokensae 17):

Right mandibular rakers = 7 plus 6 rudimenataries, left = 7 plus 4 rudi-
mentaries. Basofacial setal formula on article 3 of mandibular palp = 2-3
(opposite and offset) and 2 inner setae; left palp formula = 2-2.

Coxa 4 slightly broadened and posterodorsal corner very broadly round-
ed. Facial and setal spine formulas of pereopods 3-4 on article 4 = 5 + 5,
on article 5 = 5 + 6, on article 6 = 4 + 2 and 5 + 2, main spine on article
5 reaching only M. 75 on article 6. Article 2 of pereopod 7 narrower than

858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in female, article 5 with special form of figure 3gP7, but lacking special male
posterior spines found in birubiins.

Epimera 1-2 broadened, posterior margin of epimeron 3 bulbous, setal
formulas: epimeron | anteroventral = 11, posteroventral = 6 weakly facial,
epimeron 2 facial = 12-13, occasional pair vertical, epimeron 3 posterior =
5 setules, facial = 0, ventral = 4. Spine formulas of uropods: uropod 1 pe-
duncle apicolateral = 3, basofacial = 10, uropod 2 peduncle dorsal = 12,
dorsal spines on outer ramus of uropod 1 = 1-2, of uropod 2 = 0, inner
ramus of uropod | = 1, of uropod 2 = 0, ventral spines on peduncle of
uropod 3 = 9, spine formula on article 1 of outer ramus = 1-1-1-1-1-1, setal
formula = 1-1-1-1-1-1. Telson slightly broadened, distal spines greatly short-
ened, each lobe with short basodorsal row of denticles.

Juvenile “‘j.’’—Recognizable as member of this species but epimeron 3
lacking ventral setae, rami of uropod 1 lacking dorsal spines, eye very small,
inner ramus of uropod 3 reaching M. 55 on article 1 of outer ramus, only
seta present fully apical.

Lectotype.—Female ‘‘f’’ 6.44 mm, Swedish Museum of Natural History
6632.

Type-locality.—Hope Harbor, 30 April 1896, 6-10 fd (fathoms).

Material.—Type-locality, female “‘d’’ 6.0 mm (old length not remea-
sured), male ‘‘g’’ 7.83 mm and one other specimen. Swedish Museum of
Natural ee 3637, South Georgia, Boiler Bay (Kochtopfbucht), 54°22’
S, 36°28’W, stones and algae, juvenile ‘‘j’’ 2.96 mm and one adult; 3633,
Puerto Madryn, 2—5 fms, 9 November aes (1); 3634, Puerto Condor, 50
fms, 26 February 1896 (2 small); 3635, Punta Arenas, “‘Ebbestrand,’’ 1 De-
cember 1895 (2).

Remarks.—As the type-species of the genus, fuegiensis forms the model;
and comparisons can therefore be made through the key or in remarks of
following species.

Distribution.—Magellanic-Fuegian Archipelago, 0-91 m; South Georgia,
0-311 m; Tristan da Cunha, surface (corrected summary from Barnard,
1960).

Fuegiphoxus inutilus, new species
Figs. 3, 4 (part)

Parharpinia fuegiensis Schellenberg, 1931:78 (in part, see text here).
Paraphoxus fuegiensis.—J. L. Barnard, 1960:pl. 42, figs. S, T (not most of
Schellenberg, 1931).

66

Description of female holotype ‘‘p.’’—Head about 18 percent of total
body length, greatest width about 80 percent of length, rostrum unconstrict-
ed, broad, short, reaching middle of article 2 on antenna 1. Eyes medium,
clear of pigment, ommatidia ordinary. Article 1 of peduncle on antenna |

VOLUME 93, NUMBER 3 859

i

WW

I
1
. 1
N 1
‘ 1
\ 1
\ '
\ '
\
\ .
1 .
‘ N
‘

N

Fig. 4. Fuegiphoxus inutilus: p = holotype female ‘‘p’’ 7.80 mm; s = young male ‘‘s”’
6.51 mm.

860 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

about 1.3 times as long as wide, about 2.3 times as wide as article 2, ventral
margin with about 4 setules, produced dorsal apex with 3 setules, article 2
about 0.47 times as long as article 1, with apicoventral cycle of 9 setae,
primary flagellum with 10 articles, about 0.9 times as long as peduncle,
bearing one each short aesthetasc on articles 1-9, accessory flagellum short,
with 6 articles. Antenna 2 not ensiform, spine formula of article 4 = 4-4-1,
dorsal margin with 2 notches bearing 2 setae and 3 setae and one spine,
ventral margin with 7 groups of 2—3 long to medium setae, one ventrodistal
long spine, article 5 about 0.72 times as long as article 4, facial spine for-
mula = 3-2, dorsal margin bearing one set of small setae, ventral margin
with 5 sets of one each long setae plus setules, 3 ventrodistal long to medium
spines counted in facial formula above; flagellum about as long as articles
4-5 of peduncle combined, with 9 articles.

Epistome unproduced. Mandibles with weak palpar hump, right incisor
with 3 teeth, left incisor with 3 humps in 2 branches, right lacinia mobilis
bifid (trifid), distal branch much shorter than proximal, broad, subbifid, dis-
tal subbifidation denticulate, proximal branch simple, pointed, with marginal
denticles, left lacinia mobilis with 4 deep teeth, middle teeth not shortened,
right rakers 6 plus one rudimentary, left rakers 7 plus 1-2 rudimentaries,
molars composed of bulbous protrusions, each molar with one weakly ser-
rate medium spine and 2 short spines, molars without plumes, palp article
1 greatly elongate, article 2 with 2 long to medium inner apical setae and 2
other shorter inner setae, article 3 about 1.2 times as long as article 2,
oblique apex with 7 and 6 spine-setae, basofacial formula = 1-1. Each outer
lobe of lower lip with one cone. Inner plate of maxilla | large, broad, bearing
one long apical pluseta, one similar apicomedial seta, 2 apicolateral much
shorter setae, outer plate with 11 spines, palp article 2 with one thin apical
spine, I—2 apicolateral, 5 medial and 4 submarginal setae. Inner plate of
maxilla 2 shorter than outer, outer not broader than inner, outer with 5
lateral setae, inner with one medial seta. Inner plate of maxilliped with one
large thick apical spine, 2 apicofacial setae, 4 medial setae, outer plate with
11 medial spines, 2 apicolateral setae and cusp, palp article 1 without api-
colateral seta, article 2 with one group of 2 apicolateral setae, medial margin
of article 2 moderately setose, article 3 with 7 facial setae, 2 lateral setae,
nail of article 4 short, with 2 accessory setules.

Coxa | scarcely expanded apically, anterior margin convex, main ventral
setae of coxae 1-4 = 13-13-14-17, posteriormost seta of coxae 1-3 slightly
shortened, anterior and posterior margins of coxa 4 parallel, posterior mar-
gin convex, posterodorsal corner rounded, posterodorsal margin short,
V-shaped, width-length ratio = 7:8. Gnathopods with elongate hands,
gnathopod 2 larger than gnathopod 1 and with subcryptic wrist, width ratios
on articles 5-6 of gnathopods 1-2 = 30:40 and 33:50, length ratios = 65:87
and 50:90, palmar humps ordinary, palms strongly oblique, article 5 of

VOLUME 93, NUMBER 3 861

gnathopod 1 elongate, ovate, posterior margin rounded-flat, article 5 of
gnathopod 2 triangular, posterior margin rounded, angular.

Pereopods 3—4 similar to each other, facial setae formula on article 4 =
5 and 5S, parallel to apex, on article 5 = 5 and 6, main spine of article 5
extending to M. 80-70 on article 6, article 5 with no proximoposterior spines,
spine formula of article 6 = 4 + 5 and 4 + 5 plus no middistal seta, spines
especially long, medial spines inserted from lateral side; acclivity on inner
margin of dactyls of pereopods 3—4 obsolescent, emergent setule fully im-
mersed, midfacial pluseta ordinary but highly anteriad. Coxae 5-7 postero-
ventral setule formula = 2-4-1. Articles 4—5 of pereopods 5-6 narrow, facial
spine rows sparse, facial ridge formula of article 2 on pereopods 5-7 = 0-1-
1; article 2 of pereopod 5 tapering apically. Width ratios of articles 2, 4, 5,
6 of pereopod 5 = 46:24:21:8, of pereopod 6 = 64:20:15:9, of pereopod 7 =
83:15:13:8, length ratios of pereopod 5 = 65:27:30:31, of pereopod 6 =
88:42:49:44, of pereopod 7 = 100:18:23:29, article 2 of pereopod 7 exceeding
middle of article 4, posterior margin with 9-10 small serrations, one scarcely
elongate posteroventral setule, medial apex of article 6 coarsely combed,
bearing 7-8 digital processes.

Posteroventral corner of epimeron 1 weakly toothed, posterior margin
convex, setulose, corner with setule, anteroventral margin with 8-9 long to
medium setae, posteroventral margin with 3 long setae. Posteroventral cor-
ner of epimeron 2 with small sharp tooth guarded by setule sinus, posterior
margin convex, setulose, facial setae = 11, none set vertically. Posteroven-
tral corner of epimeron 3 rounded, protuberant, with setule sinus, posterior
margin oblique, weakly concave, setulose, ventral margin with 3 setae near
middle.

Urosomite 1 with lateral setule at base of uropod 1, no ventral setae,
articulation line complete, urosomites scarcely protuberant dorsally. Rami
of uropods 1-2 with fused and immersed apical nails, outer ramus of uropod
1 with 3 dorsal spines, inner with 2, outer ramus of uropod 2 with one dorsal
spine, inner with no dorsomedial spines, peduncle of uropod 1 with 5 ba-
sofacial setae and 3 apicolateral narrowly confined spines, medially with S—
6 marginal spines, apicalmost weakly enlarged but not displaced, no acces-
sory slit. Peduncle of uropod 2 with 10 dorsal spines, medially with one
small apical spine, apicolateral corners of peduncles on uropods 1-2 with
obsolescent comb. Peduncle of uropod 3 with 6 ventral spines, dorsally with
2 long lateral spines, one short hooked medial spine and 2 setules or spinule,
rami submasculine, inner extending to M. 100 on article | of outer ramus,
apex with [unknown] setae, medial and lateral margins with 2 and 2 setae,
article 2 of outer ramus short, 0.15, bearing 2 ?long setae [broken], medial
margin of article 1 with 5 setae, lateral margin with 3 or | acclivities, spine
formula = 1-1-1-1 or 1-2, setal formula = 0-0-0-1 or 0-1 (variable on 2 sides).
Telson long, length-width ratio = 13:11, almost fully cleft, each apex nar-

862 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

row, rounded, lateral acclivity broad, shallow, bearing short lateral setule,
spine next medial longer than setule or with lateral and medial spines sep-
arated by setule (variable), midlateral setules diverse, largest of small size.

Female ‘‘s.’’-—Eyes about as large as in fuegiensis.
Male ‘‘r.’’—Too young to differentiate from female.
Juvenile “‘t.’’—Spine formula on article 4 of antenna 2 = 0-3-3-0, on ar-

ticle 5 = 3 apicofacial spines. Formula on article 6 of pereopods 3—4 = 2
+ 3 + 0. Epimera 1-2 with posteroventral tooth, epimeron 3 like adult but
with only one ventral seta. Inner ramus of uropod | lacking spine, both rami
of uropod 2 lacking spines.

Holotype.—Female ‘‘p’’ 7.80 mm, Swedish Museum of Natural History
3638.

Type-locality.—South Georgia, mouth of Cumberland Bay, 54°11’S,
36°18’W, 252-310 m, N:o 34 5/6 1902, gray clay with small (few) (‘weni-
gen’’) stones, bottom temperature + 1.45° [C?] [translated from label written
in German, see Schellenberg, 1931:78 as Parharpinia fuegiensis quoted as
250-310 m and “‘einigen Steinen’’].

Material.—Swedish Museum of Natural History 3636, South Georgia,
outer half of May-Bay, 75 m, 54°17’S, 36°28’W, N:o 22, 14/5 1902, clay, also
some algae, bottom temperature +1.5° [C?] [translated from label written
in German, see Schellenberg, 1931:78 as Parharpinia fuegiensis quoted as
‘‘Grytviken’’], female “‘r’’ 6.21 mm, young male ‘“‘s’’ 6.51 mm, juvenile
tC) Sef Ik tating

Illustrations: Parts not illustrated generally like those of P. fuegiensis
herein and in J. L. Barnard (1960:pl. 42); uropod 3 of female **s’’ added to
illustrations here to better show apex broken in the holotype.

Remarks.—The juvenile ‘“‘t’’ is needed to confirm the validity of this
species because the material was originally determined as fuegiensis by
Schellenberg (1931) and again by Barnard (1960). This species differs from
F. fuegiensis, the type-species, in the following characters: (1) Eyes of the
female are only medium in size; (2) article 5 of antenna 2 has 3 apicofacial
thick spines instead of one; (3) article 6 of pereopods 3-4 has more spines;
(4) the epimera are broader, epimera 1-2 have a posteroventral tooth, and
epimeron 3 is protrusive; (5) urosomite | lacks ventral setae; (6) the pleo-
some is larger and the urosome smaller; (7) outer ramus of uropod 3 bears
a dorsal spine; (8) uropod 3 is shorter.

Distribution.—South Georgia, 75—250 m (confirmed minimum range).

Fuegiphoxus abjectus, new species
Figs. 5, 6

Description of holotype ‘‘h’’ male.—Head about 18 percent of total body
length, greatest width about 75 percent of length, rostrum unconstricted,

VOLUME 93, NUMBER 3 863

Fig. 5. Fuegiphoxus abjectus: holotype male “‘h”’ 10.55 mm.

broad, of ordinary length, tapering evenly, reaching apex of article 2 on
antenna 1. Eyes small or absent or diffuse. Article 1 of peduncle on antenna
1 about 1.4 times as long as wide, about 1.3 times as wide as article 2,
ventral margin with about 6 setules, weakly produced dorsal apex with 2-
3 setules, article 2 about 0.55 times as long as article 1, with ventral row of
9 setae (broken), primary flagellum with 16 articles, about 0.8 times as long
as peduncle, bearing one each tiny aesthetasc on articles 7-15, basal third
of flagellum swollen, no medial fuzz on antenna 1, accessory flagellum with
? articles [broken]. Antenna 2 not ensiform; spine formula of article 4 =
3-3-3-1, dorsal margin with 2 notches bearing 6 and 4 (distal to proximal)
setae and one spine (proximal), ventral margin with 8 groups of 1-3 long to
short setae, one ventrodistal long spine, article 5 about 0.75 times as long
as article 4, facial spine formula = 1, dorsal margin bearing one set of small

864 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distal setae, ventral margin with 6 sets of one long seta each, 3 ventrodistal
short to medium spines, one of these strongly facial, flagellum [broken after
article 10]. Epistome not produced. Mandibles with weak palpar hump, right
incisor with 4 teeth!, left incisor with mainly 3 humps in 2 branches, right
lacinia mobilis bifid, distal branch much shorter than proximal, flabellate,
denticulate, proximal branch simple, pointed, left lacinia mobilis with 4
teeth, middle teeth slightly shortened, right rakers 9 plus 3 rudimentaries,
left rakers 10 plus 2 rudimentaries, molars composed of bulbous protrusions,
each with one very long serrate and 2 medium spines, molar without plume,
palp thin but very elongate, article 1 short, article 2 with 1 long to medium
inner apical seta and 2 other short inner setae, article 3 about 1.3 times as
long as article 2, oblique apex with 9 thin spine-setae, basofacial formula =
0-2. Each outer lobe of lower lip with one cone. Inner plate of maxilla 1
especially large, thin apically, broad basally, bearing one medium subapical
pluseta, one longer similar facial seta, 2 apicolateral much shorter setae,
outer plate with 11 spines, palp article 2 with 2 apical, 4 medial spines and
5 submarginal setae. Plates of maxilla 2 extending subequally, outer scarcely
broader than inner, outer with 4 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with one large thick apical spine, 3 apicofacial
setae, 4 medial setae, outer plate with 11 medial spines, 2 apicolateral setae
and 2 cusps, palp article 1 with one apicolateral seta, article 2 with one
group of 3 apicolateral setae, medial margin of article 2 moderately setose,
article 3 with 4 facial setae, 2 lateral setae, nail of article 4 medium, with 2
accessory Setules.

Coxa | not expanded apically, anterior margin weakly convex, main ven-
tral setae of coxae 1-4 = 17-12-12-14, only posteriormost seta of coxa 1
strongly shortened, anterior and posterior margins of coxa 4 slightly diver-
gent, posterior margin convex, posterodorsal corner rounded, posterodorsal
margin ordinary, concave, width-length ratio = 5:6. Gnathopods generally
ordinary, gnathopod 2 scarcely longer than gnathopod 1, width ratios on
articles 5—6 of gnathopods 1-2 = 27:32 and 27:34, length ratios = 65:61 and
61:62, palmar humps ordinary, palms strongly oblique, article 5 of gnatho-
pod 1 elongate, ovate, posterior margin rounded-flat long, article 5 of
gnathopod 2 elongate, ovate, posterior margin rounded.

Pereopod 4 slightly stouter than pereopod 3 especially on article 4, facial
setae formula on article 4 = 8 and 6, parallel to apex, on article 5 = 9 and
8, main spine of article 5 extending to M. 100-90 on article 6, article 5 with
no proximoposterior spines, spine formula of article 6 = 6 + 5 and 7 + 6
but no middistal seta, medial members = clump of thin apical setae, spines
especially long, acclivity on inner margin of dactyls of pereopods 3—4 rep-
resented by slit, emergent setule short, midfacial pluseta anteriad, short.
Coxae 5-7 posteroventral seta formula = 6-5-1. Articles 4—5 of pereopods
5-6 narrow, facial spine rows sparse, facial ridge formula of article 2 on

VOLUME 93, NUMBER 3 865

—— ig

c=. é-
EZ
GEE

Y/))

Fig. 6. Fuegiphoxus abjectus: holotype male ‘‘h’’ 10.55 mm; dotted spine on R3 shown for

same member on right side of animal.

866 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pereopods 5-7 = 0-1-1. Width ratios of articles 2, 4, 5, 6 of pereopod 5 =
40:25:25:15, of pereopod 6 = 72:29:23:14, of pereopod 7 = 90:23:20:12,
length ratios of pereopod 5 = 78:32:42:41, of pereopod 6 = 95:48:60:58, of
pereopod 7 = 100:26:28:39, article 2 of pereopod 7 exceeding middle of
article 4, posterior margin with 7 medium serrations and several long pos-
terodorsal setae (very unusual), setae of articles 3-4 very thick and stiff,
medial apex of article 6 scarcely combed, bearing 9 digital processes.

Posteroventral corner of epimeron 1 quadrate, posterior margin straight,
with setule, anteroventral margin with 10 medium setae, posteroventral face
with horizontal row of 7 long setae. Posteroventral corner of epimeron 2
rounded-quadrate and weakly protuberant, posterior margin weakly convex,
with 2 setules, facial setae = 15, occasionally middle pairs set vertically.
Posteroventral corner of epimeron 3 rounded, overall protuberant but pos-
terior margin weakly convex, with 2 setule notches, ventral margin with 4
setae mainly in posterior half.

Urosomite | with lateral setule at base of uropod 1, brush of setae ventral
to uropod 1 and brush of midventral setae, articulation line complete, uro-
somites unprotuberant dorsally. Rami of uropods 1-2 with articulate but
tightly fixed apical nails, outer ramus of uropod 1 with 3 dorsal spines, inner
with 2, outer ramus of uropod 2 with one dorsal spine, inner with one
dorsomedial spine, peduncle of uropod 1 with 7 basofacial and ventral
cluster of 5 setae, one apicolateral spine, medially with 6 marginal setae and
spines, apicalmost enlarged but not displaced. Peduncle of uropod 2 with
7-8 dorsal spines, basalmost thin and elongate, medially with one medium
apical spine, apicolateral corners of peduncles on uropods 1-2 with very
weak comb. Peduncle of uropod 3 with 6 ventral spines, dorsally with one
lateral spine, one medial spine and 2 setules, rami submasculine, inner ex-
tending to M. 100+ on article | of outer ramus, apex with 4 setae, medial
and lateral margins with 2 and one setae, article 2 of outer ramus short,
0.16, bearing 2 short to medium setae, apicomedial margin of article 1 with
3 setae, lateral margin with 1-2 acclivities, spine formula = 0-2-5 or 0-4-4,
setal formula = 0-0-1 or 1-0-1. Telson long, length-width ratio = 33:28, al-
most fully cleft, each apex of medium width, rounded, lateral acclivity
broad, shallow, with short lateral and long medial spines separated by short
setule, midlateral setules diverse, largest of small size, denticles absent.

Notes.—Male unusual: antenna 2 not elongate, uropod 3 not fully devel-
oped, denticles on telson absent; therefore male possibly youthful.

Holotype.—Swedish Museum of Natural History 3631, male “*h’’ 10.55
mm. Unique.

Type-locality.—Bahia Inutil, 23 January 1896, 20-30 fd.

Remarks.—This specimen was formerly called Parharpinia fuegiensis by
Schellenberg, then Paraphoxus fuegiensis by J. L. Barnard (1960) and was

VOLUME 93, NUMBER 3 867

considered to be a female but now is thought to be a male of character
similar to that of Elpeddo kaikai Barnard and Drummond (1978:119).

The remarkable antenna | of this unique male links Fuegiphoxus with the
Australian Elpeddo as noted in remarks after the description of Fuegiphoxus
above. In fact, abjectus has more strong characters of difference from its
sympatriots fuegiensis and inutilus in South America than from Elpeddo
but the characters noted earlier in this paper seem to be more important to
distinguish genera than to invoke distinctions simply by counting the number
of unevaluated differences among taxa.

Fuegiphoxus abjectus differs from both F. inutilus and F. fuegiensis in:
(1) the formula, spacing and thickness of the spines on article 4 of antenna
2; (2) the odd male antenna 1; (3) the short article 1 of the mandibular palp;
(4) the scarcely enlarged gnathopod 2; (5) the odd setal clump on article 6
of pereopods 3-4; (6) the shape of epimeron 3; and (7) the setal brush on
the apex of urosomite 1.

Fuegiphoxus? uncinatus (Chevreux)

Pontharpinia uncinata Chevreux, 1912:4; 1913:100—104, figs. 10-12.
Paraphoxus uncinatus.—J. L. Barnard, 1960:283.

This species is provisionally assigned to Fuegiphoxus but many charac-
ters need confirmation, for example: spine formulas on antenna 2, laciniae
mobiles, mandibular palp, maxillipedal spine counts and dactyl formations,
setal formulas of epimera 1—2 and urosomite 1, spine formulas on apices of
rami on uropods 1-2, and medial margins of peduncles.

For the moment, the species is distinguished from others in its genus by
the long thin posteroventral tooth of epimeron 3.

Distribution.—Antarctica, Port Lockroy, Chenal de Roosen, 60—70 m.

Phoxorgia, new genus

Diagnosis.—Eyes present. Flagella of antennae 1—2 unreduced in female,
article 2 of antenna | shortened, ventral setae confined apically; article 1 of
antenna 2 not ensiform, article 3 with 3 lateral setae, facial spines on article
4 in 2 or more rows, article 5 ordinary in size. Right mandibular incisor with
3 teeth, molar not triturative, small, pillow shaped, bearing 4 or more
splayed spines, bearing pubescence, palpar hump moderate. Palp of maxilla
1 biarticulate, inner plate with 4 setae; setation of maxilla 2 ordinary. Inner
plate of maxilliped ordinary, apex of palp weakly protuberant, dactyl elon-
gate, apical nail weakly distinct, medium.

Gnathopods ordinary, small, similar, wrists elongate, unlobed, without
eusirid attachment, palms transverse, hands bell-shaped, poorly setose an-

868 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

teriorly. Article 2 of pereopod 5 of broad form, scarcely tapering apically,
articles 4-5 of pereopods 5-6 narrow, article 2 not setose posteriorly, per-
eopod 7 ordinary, article 2 naked ventrally, article 3 ordinary, dactyl normal.

Peduncle of uropod | normally elongate, without apical spike, with dis-
placed apicomedial spine, uropods 1—2 without comb, inner ramus of uropod
1 with one row of marginal spines, inner ramus of uropod 2 more or less
continuously spinose to apex (with one accessory nail), apical nails well
developed and that on inner ramus of uropod | flexibly articulate, inner
ramus of uropod 2 normal (not fused). Uropod 3 ordinary, bearing article
2 of outer ramus, carrying 2 short apical setae. Telson ordinary, each lobe
with 2 apical spines and attendant setule, without other special seta. Epi-
mera 1-2 bearing sparse short posterior setules, without midfacial setae
above ventral facial ridge, epimeron 3 ordinary and bearing 3 or more long
setae in adult. Urosomite | without large lateral armament, bearing one or
more large bundles of ventral setae, urosomite 3 without hook or special
process.

Description.—Rostrum fully developed but weakly constricted. Pubes-
cence and calceoli on male antennae [unknown]. Prebuccal parts ordinary.
Right lacinia mobilis bifid or multitoothed, article 1 of mandibular palp short,
palp thin, apex of article 3 oblique, article 2 without outer setae. Lower lip
bearing cones. Outer plate of maxilla 1 with 11 spines, one spine especially
thickened. Inner plates of maxillipeds with 3 main spines, ordinarily setose.
Coxae 2—4 without special anterodorsal humps. Posterior spines on article
6 of pereopods 3-4 thick and stiff, midapical seta present; article 2 of per-
eopod 7 without facial setae. Peduncle of uropod | with dorsolateral spines
widely spread, medial spines widely spread; peduncle of uropod 2 with only
one medial spine confined apically; peduncle of uropod 3 lacking extra sub-
apical setae or spines. Telson with one or more apical spines plus one dis-
junct subapical spine on each lobe plus attendant setules.

Etymology.—From ‘‘Phoxocephalus,’’ the type-genus of the family and
‘‘orgyia,’’ Greek for fathom, modified so as to simplify. Feminine.

Type-species.—Parharpinia sinuata K. H. Barnard, 1932.

Composition.—Unique.

Relationship.—This genus is very close to Parharpinia Stebbing from
Australia (see Barnard and Drummond, 1978). It resembles that genus in
broad generalities of uropod 1, especially the diverse pattern of spination,
and in the presence of 3 main spines on the inner plate of the maxiiliped.
There are many other similarities such as extreme antennal spination, slight
taper to article 2 of pereopod 5 (strong in Parharpinia, barely discernible
in Phoxorgia), presence of at least remnants of accessory apical spination
on uropods 1|-2, extra telsonic spination (dorsal in Parharpinia, marginal in
Phoxorgia) and generalities of head, mandible, and uropod 3.

Phoxorgia differs from Parharpinia in the apical shift of setae on article

VOLUME 93, NUMBER 3 869

2 of antenna | (characteristic of juveniles in Parharpinia), the lack of true
dorsal spines on the telson, the absence of ventral setae on article 2 of
pereopod 7, and the poor posterior setation on epimera 1-2. These may
ultimately not be adequate to segregate Phoxorgia but for the moment can
be correlated with geography.

Phoxorgia appears to be a very good plesiomorph both to Parharpinia
and Metharpinia, the latter an American genus already revised by J. L.
Barnard (1980). Phoxorgia is plesiomorphic to both genera because of the
absence of dorsal telsonic spines, plesiomorphic to Parharpinia in addition
because of almost normal pereopod 5, and poor epimeral and pereopod 7
setation. It is plesiomorphic to Metharpinia which differs from Phoxorgia
in the strongly reduced rostrum, widely spread ventral setation on article
2 of antenna 1, reduction of main spination on inner plate of maxilliped
(from 3 down to 2 or 1), and the loss of true apical nails, only the accessories
remaining. The connection to Microphoxus through Metharpinia can be
determined in Barnard (1980).

Phoxorgia shares with Fuegiphoxus the odd position of setae on article
2 of antenna | but otherwise differs in numerous characters from Fuegi-
phoxus, as follows: (1) stronger spination on article 4 of antenna 2; (2)
stronger and more even spination on mandibular molars; (3) presence of 3
(not 1) spines on inner plate of maxilliped; (4) displaced spine of uropod 1;
and (5) well developed apical and accessory apical spines on the rami of
uropods 1-2. Fuegiphoxus therefore appears to be apomorphic to Phoxorgia
and is actually in the Brolginae.

Phoxorgia sinuata is very close to Foxiphalus but differs from an unde-
scribed species (here called species S which will be described in a later
paper) in the following ways: (1) slightly constricted and highly flattened
head; (2) shortened article 2 of antenna 1 with the setae shifted apicad; (3)
presence of more than 2 facial setae on article 3 of antenna 2; (4) flabellate
and subbifid distal branch of the right lacinia mobilis; (5) presence of more
than one apical spine on the inner plate of the maxillipeds; and (6) the
presence of an accessory apical nail on the inner ramus of uropod 1.

Phoxorgia sinuata (K. H. Barnard)
Fig. 7

Parharpinia villosa, Schellenberg, 1931:75—78, fig. 39 (not Haswell, 1879);
NSS 2527

Parharpinia sinuata K. H. Barnard, 1932:103-104, fig. 52.

Paraphoxus sinuatus, J. L. Barnard, 1958:147-148; 1960:278-282, pl. 45.

Description of female ‘‘v.’’—Head very flat, about 22 percent of total
body length, greatest width about 72 percent of length, rostrum slightly
constricted near eyes, then continuing broad but tapering, short, reaching

870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Phoxorgia sinuata: v = female ‘‘v’’ 10.5 mm; w = female “‘w”’ 18.1 mm; y = male
“‘y’? 10.3 mm.

apex of article 2 on antenna |. Eyes medium, clear of pigment, ommatidia
ordinary. Article 1 of peduncle on antenna | about 1.4 times as long as wide,
about twice as wide as article 2, ventral margin with about 14+ setules,
weakly produced dorsal apex with 2 setules, article 2 about 0.5 times as
long as article 1, with apicoventral cycle of 9 setae and 8+ lateral setae at
apex, primary flagellum with 15 articles, about 1.2 times as long as peduncle,
bearing one short aesthetasc each on articles 8-12, accessory flagellum with
11 articles. Antenna 2 not ensiform though article 1 very large, article 3 with
3 lateral setae, spine formula of article 4 = 1-3-6-5-1 or 1-3-5-5-1, dorsal

VOLUME 93, NUMBER 3 871

margin with weak notch bearing 3 setae, and 2 spines, ventral margin with
5 groups of 3—5 long to medium setae, one ventrodistal long spine, article
5 about 0.8 times as long as article 4, facial spine formula = 3-4, dorsal
margin bearing 2 sets of small setae, ventral margin with 4 sets of 1-2 long
to medium setae, 3 ventrodistal long to medium spines, one of these strongly
facial; flagellum about 1.5 times as long as articles 4-5 of peduncle com-
bined, with 16 articles.

Epistome bulbous anteriorly, fused with upper lip. Mandibles with me-
dium palpar hump, right incisor with 3 teeth, left incisor with 3 humps in 2
branches, right lacinia mobilis bifid, distal branch much shorter than prox-
imal, distal branch broad, subbifid, proximal branch simple, blunt, left la-
cinia mobilis with 4 teeth, right rakers 10 plus one rudimentary, left rakers
9+, molars composed of elongate bulbous protrusions, right molar with 6
long spines, 6 short teeth, plus one ragged spine weakly disjunct, left molar
with 6 long spines, plus 4 short teeth plus ragged spine weakly disjunct,
each molar with plume, palp article | short, article 2 with one medium inner
apical seta and 3 other shorter inner setae and one outer seta, article 3 about
1.1 times as long as article 2, oblique apex with 9 spine-setae, basofacial
formula = 1-2. Each outer lobe of lower lip with one cone. Inner plate of
maxilla 1 ordinary, bearing one long apical pluseta, one similar apicomedial
seta, 2 apicolateral much shorter setae, outer plate with 11 spines, palp
article 2 with 2 apical setal-spines, 2 medial setal spines, one apicolateral
and 4 medial marginal setae. Inner plates of maxilla 2 scarcely shorter than
outer, outer not broader than inner, outer with 6 apicolateral setae, inner
with 2 medial setae. Inner plate of maxilliped with 3 large thick apical spines,
2 apicofacial setae, 6 medial setae, outer plate with 11 medial spines, 6
apicolateral setae, palp article 1 without apicolateral seta, article 2 with one
apicolateral seta, medial margin of article 2 moderately to weakly setose,
article 3 with 10 slightly strewn facial setae, 3 lateral setae in clump, nail of
article 4 mostly fused but short and internally visible, with 2 accessory
setules.

Coxa 1 expanded apically, anterior margin weakly convex or sinuous,
main ventral setae of coxae 1-4 = 7-6-6-4, posteriormost seta of coxae 1-3
very short, anterior and posterior margins of coxa 4 weakly divergent, pos-
terior margin almost straight, posterodorsal corner rounded, posterodorsal
margin, ordinary, concave, width-length ratio of coxa 4 = 14:17. Gnatho-
pods generally ordinary, width ratios on articles 5—6 of gnathopods 1-2 =
22:33 and 25:34, length ratios = 66:60 and 60:58, palmar humps ordinary,
palms weakly oblique, article 5 of gnathopod | elongate, ovate, posterior
margin flat, long, article 5 of gnathopod 2 ovate, posterior margin rounded,
short.

Pereopods 3-4 similar to each other, facial setae formula on article 4 =
4 and 3, parallel to apex, on article 5 = 4 and 5, main spine of article 5

872 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

extending to M. 100 on article 6, article 5 with 3 and 2 proximoposterior
spines, spine formula of article 6 = 4 + 5 and 4 + 5 plus strong middistal
seta (like Foxiphalus, not Fuegiphoxus), some spines long, acclivity on
inner margin of dactyls of pereopods 3—4 weak, subsharp, produced as
tooth, emergent setule almost fully immersed, midfacial pluseta ordinary
but highly anteriad. Coxae 5-7 posteroventral seta formula = 3-3-7. Articles
4-5 of pereopods 5-6 narrow, facial spine rows sparse, facial ridge formula
of article 2 on pereopods 5-7 = 0-1-1. Width ratios of articles 2, 4, 5, 6 of
pereopod 5 = 48:30:26:15, of pereopod 6 = 78:35:28:15, of pereopod 7 =
96:23:22:12, length ratios of pereopod 5 = 83:36:41:52, of pereopod 6 =
97:67:59:81, of pereopod 7 = 100:30:32:43. Article 2 of pereopod 7 not
reaching middle of article 4, posterior margin with 10 small serrations, me-
dial apex of article 6 finely combed, bearing 6 digital processes.

Posteroventral corner of epimeron 1 quadrate, posterior margin straight,
with 2 setules, corner with long seta, anteroventral margin with 6 short to
medium setae, ventral margin with one long seta. Epimera 1-2 with lateral
ridge. Posteroventral corner of epimeron 2 rounded-quadrate, posterior mar-
gin straight, with 2 setules, facial setae = 6, posteriormost pair set vertically
and positioned above tangent of others, posteroventral corner of epimeron
3 broadly rounded, then with 4 crowded setae at ventral end of straight
posterior margin, above with one setule notch, ventral margin with 4 short
setal spines widely spread.

Urosomite 1 with lateral spinule at base of uropod | and midventral setal
brush, articulation line almost complete, urosomites unprotuberant dorsally.
Rami of uropods 1-2 with articulate but tightly fixed apical nails, except
inner ramus of uropod | with flexible nail, plus accessory nail, spines on
outer rami small and numerous and extending almost to apex, outer ramus
of uropod | with 11 dorsal spines, inner with 3 large spines, outer ramus of
uropod 2 with 6 dorsal spines, inner with 2 large dorsomedial spines, pe-
duncle of uropod 1 with 3 widely spread basofacial spinules, 9 packed api-
colateral spines, medially with 4 thin marginal spines and apical displaced
enlarged spine, peduncle of uropod 2 with 8 dorsal spines, basalmost short,
medially with one medium apical spine, apicolateral corners of peduncles
on uropods 1-2 without comb. Peduncle of uropod 3 with 8 ventral spines,
dorsally with one lateral spine, one medial spine, rami submasculine, inner
extending to M. 75 on article 1 of outer ramus, apex with 2 setae, medial
and lateral margins with 6 and one setae, article 2 of outer ramus ordinary,
0.23, bearing 2 short to medium setae, medial margin of article 1 with 3
setae, lateral margin with 4 acclivities, spine formula = 2-2-2-2-2, setal for-
mula = 0-0-0-0-1. Telson ordinary, length-width ratio = almost 1:1, almost
fully cleft, each apex wide, rounded, but incised apically, lateral acclivity
narrow, weak, bearing short lateral spine, setule next medial of length equal

VOLUME 93, NUMBER 3 873

to spine, medial spine in terminal incision, midlateral setules diverse, larger
of medium size.

Male ‘‘y”’ (not fully terminal).—Eyes not enlarged. Article 1 of antenna
1 with 12 ventral setules, article 2 with 8 anteroventral setae and 5 apico-
lateral setae, primary flagellum with 17 articles, accessory with 11, calceoli
absent, aesthetascs weakly developed, pubescence on antennae absent. Fa-
cial spine formula on article 4 of antenna 2 = 1-3-5-5-1, on article 5 = 3-4,
with no dorsal sets of male setae and calceoli, ventrodistal apex with 3 thick
spines; flagellum weakly proliferate, with 18 articles. Basofacial setal for-
mula of article 3 on mandibular palp = 1-3 or 1-2. Article 2 of only pereopod
7 slightly narrower than in female. Epimera 1-3 not broadened, posterior
margin of epimeron 3 not shortened, setal formulas: epimeron | anteroven-
tral = 5, facioventral = 3, then large gap to posterior corner = 1, epimeron
2 facial = 7 (2 posterior = vertical), epimeron 3 posterior = 4, ventral = 3.
Spine formulas of uropods: uropod | peduncle apicolateral = 9, basofacial =
2, uropod 2 peduncle dorsal = 9, dorsal spines on outer ramus of uropod
1 = 12, of uropod 2 = 6, inner ramus of uropod | = 4 + 1 accessory nail,
of uropod 2 = 2, ventral spines on peduncle of uropod 3 = 7, inner ramus
extending to M. 90 on article | of outer ramus, well setose, spine formula
on article | of outer ramus = 1-2-2-2-2, setal formula = 0-0-0-0-1; telson like
female, distal spines not shortened. Apical spine guarded by 2 setules (fig-
ured), no denticle rows.

Female ‘‘w’’ giant.—See illustration of antenna 2 noting flagellum slightly
proliferate as in males. Epimera 1-3 illustrated, noting epimeron | postero-
ventral corner slightly produced into 2 cusps. Armament formulas, uropod
1 basofacial peduncle = 4 in long string, peduncle apicolateral = 15 (cov-
ering almost half of margin), outer ramus = 17, inner ramus = 5 + one ac-
cessory apical nail; uropod 2 peduncle dorsal = 14, outer ramus = 11, inner
ramus = 3; uropod 3 peduncle ventral = 9, article | outer ramus spines =
1-2-2-2-2-2, setae = 0-0-0-0-0-1, medial margin of article 1 on outer ramus
on both margins of inner ramus strongly setose, article 2 on outer ramus =
0.14. Telson illustrated.

Juvenile ‘‘k.’’—Ventrodistal setae of article 2 on antenna | = 4. Setae on
article 3 of antenna 2 = one long, one short only, spine formula on article
4 = 1-3-4-3, on article 5 = 2 only. Right lacinia mobilis similar to adult,
right molar with 7 main spines and one disjunct, basal setal formula on palp
article 3 = 0-0. Spine formula on inner plate of maxilliped = 3 (like adult).
Epimeron | with 3 anteroventral setae, no others. Epimeron 2 with 4 facial
setae, posterior pair of facials slightly disjunct and obliquely set. Epimeron
3 with one ventral and 2 posterior setae. Spine formulas: uropod | peduncle
apicolateral = 3, outer ramus = 4, inner ramus = 2, on latter accessory nail
rudimentary, only socket truly present; uropod 3 dorsal peduncle = 3, outer

874 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ramus = 2, inner ramus = 1; uropod 3 outer ramus article 1 spines = 1-2-
2, setae = 0-0-0, article 2 about 0.45!!, inner ramus extending to M. 55 on
article 1 of outer ramus, with only one apical seta.

Material.—Swedish Museum of Natural History 2722, Bahia Inutil, 20-
30 Fd., 23/1/96 (11 specimens, including male ‘‘y’’ 10.3 mm and female ‘‘v”’
10.5 mm); 3640, W. Feuerland, 54°43’S, 64°08’ W, 6/1/02, 36 m, pebbles and
gravel, senile female “‘w’’ 18.1 mm; 3631, juvenile ‘‘k’’ 5.18 mm; 2728,
Bahia Inutil, 11-5 fms, 23/1/96 (1); 3639, Punta Arenas, 7-8 fms, 4/12/95
(Guvenile); 2741, Isla Nueva, 30 fms, 7/2/96 (15); 591, Valparaiso, 6—8 fms,
several but others in sample not this genus. Smithsonian Institution: Alba-
tross 2776, Straits of Magellan, 52°41'00"S, 69°55'30"W, 21 fms, 18 January
1888, water surface temperature 51 F (1).

Distribution.—Valparaiso, Chile through Magellanic Archipelago to
South Georgia, 4-159 m.

Literature Cited

Barnard, J. L. 1958. Revisionary notes on the Phoxocephalidae (Amphipoda), with a key to
the genera.—Pacific Science 12:146-151.

. 1960. The amphipod family Phoxocephalidae in the Eastern Pacific Ocean, with anal-

yses of other species and notes for a revision of the Family.—Allan Hancock Pacific

Expeditions 18:175—368, pls. 1-75, 1 chart.

. 1979. Revision of American species of the marine amphipod genus Paraphoxus (Gam-

maridea: Phoxocephalidae).—Proceedings of the Biological Society of Washington

92:368-379.

, and M. M. Drummond. 1976. Clarification of five genera of Phoxocephalidae (Marine

Amphipoda).—Proceedings of the Biological Society of Washington 88:515—547, figs.

1-4.

, and . 1978. Gammaridean amphipoda of Australia, part IIJ.—Smithsonian Con-

tributions to Zoology 245:1-—555, figs. 1-269.

Barnard, K. H. 1932. Amphipoda.—Discovery Reports 5:1—326, plate 1, figs. 1-174.

Chevreux, E. 1912. Diagnoses d’amphipodes nouveaux.—Deuxieme Expédition dans

l Antarctique, dirigée par le Dr. Charcot, 1908—1910.—Bulletin du Muséum d Histoire

Naturelle, Paris 18:208-218.

. 1913. Amphipodes.—Deuxieme Expédition Antarctique Francaise (1908-1910) com-

mandée par le Dr. Jean Charcot. Sciences Naturelles: Documents Scientifiques: 79-186,

figs. 1-62.

Haswell, W. A. 1879. On Australian Amphipoda.—Proceedings of the Linnean Society of
New South Wales 4:245—279, pls. 7-12.

Schellenberg, A. 1931. Gammariden und Caprelliden des Magellangebietes, Sidgeorgiens und
der Westantarktis.—Further Zoological Results of the Swedish Antarctic Expedition
1901-1903, 2(6): 1-290, plate 1, figs. 1-136.

—. 1935. Amphipoden von Chile und Juan Fernandez, gesammelt von Prof. W.
Goetsch.—Zoologische Jahrbtcher, Systematik, 67:225-—234, 3 text figures.

Stephensen, K. 1949. The Amphipoda of Tristan da Cunha.—Results of the Norwegian Sci-
entific Expedition to Tristan da Cunha 1937-1938, 19:1—61, 23 text figures.

Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, DC 20560, USA.

SS eee

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CONTENTS

Two new species of Lizzia (Hydrozoa: Anthomedusae) from the eastern tropical Pacific
: Lourdes Segura
Oratosquilla megalops, a new stomatopod crustacean from Taiwan
Raymond B. Manning
Reappraisal of the gastropod genus Varicopeza Griindel (Cerithiidae: Prosobranchia)
Richard S. Houbrick
Pseudochirella squalida Grice & Hulsemann, 1967, from continental slope waters off
Delaware (Copepoda: Calanoida) Frank D. Ferrari
Larval development under laboratory conditions of the tropical spider crab Mithrax
(Mithraculus) coryphe (Herbst, 1801) (Brachyura: Mayidae)
Liberta E. Scotto and Robert H. Gore
A new species of Pontophilus (Crustacea: Natantia: Crangonidae) from the Gulf of
Mexico and the western Atlantic Mike Dardeau
Four new species of Renocila (Isopoda: Cymothoidae), the first reported from the
New World Ernest H. Williams, Jr., and Lucy Bunkley Williams
A new Strobilops (Mollusca: Pulmonata: Strobilopsidae) from Baja California Sur,
Mexico Walter B. Miller and Carl C. Christensen
A giant extinct insectivore from Cuba (Mammalia: Insectivora: Solenodontidae)
Gary S. Morgan, Clayton E. Ray, and Oscar Arredondo
New Amphipoda from the southern ocean, with partial revisions of the Acanthonoto-

zomatidae and Paramphithoidae Les Watling and Heather Holman
The calls and taxonomic positions of Hyla giesleri and Ololygon opalina (Amphibia:
Anura: Hylidae) W. Ronald Heyer

Two new lamippid copepods parasitic on gorgonians from Hawaii and the Bahamas

Mark J. Grygier
New and little-known Neotropical Kinnaridae (Homoptera: Fulgoroidea) R.G. Fennah
A siphonostome copepod associated with a vestimentiferan from the Galapagos Rift

and the East Pacific Rise Arthur G. Humes and Masahiro Dojiri
A new species of Sclerobregma (Polychaeta: Scalibregmatidae) from off the southeastern
United States Rodney D. Bertelsen and Donald P. Weston

Two new species of Centrolenella from Bolivia (Anura: Centrolenidae)
David C. Cannatella
Records of anthurids from Florida, Central America, and South America (Crustacea:
Isopoda: Anthuridae) Brian Kensley
On the carapace shape of some European freshwater interstitital Candoninae (Ostra-
coda) Dan L. Danielopol
The subspecies of grasshopper sparrow (Ammodramus savannarum) in Panama (Aves:
Emberizinae) Storrs L. Olson
Spongicoloides galapagensis, a new shrimp representing the first record of the genus
from the Pacific Ocean (Crustacea: Decapoda: Stenopodidea) Joseph W. Goy
A new species of rock shrimp of the genus Sicyonia (Penaeoidea), with a key to the
western Atlantic species Isabel Pérez Farfante
Stenopus pyrsonotus, a new species of stenopodidean shrimp from the Indo-West Pacific
region (Crustacea: Decapoda) Joseph W. Goy and Dennis M. Devaney
Onuphidae (Polychaeta) from Belize, Central America, with notes on related taxa

Christian Fauchald

The Indo-Pacific pipefish genus Urocampus (Syngnathidae) C. E. Dawson
New freshwater shrimp records for Tobago, West Indies, with a summary of records
for the Lesser Antilles (Atyidae and Palaemonidae) C. W. Hart, Jr.

Two New Phoxocephalid Genera, Fuegiphoxus and Phoxorgia, from Magellanic South
America (Amphipoda: Crustacea) J. L. and C. M. Barnard

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2s)
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849

Proceedings
of the
BIOLOGICAL SOCIETY
of
WASHINGTON

Volume 93 16 January 1981 Number 4

THE BIOLOGICAL SOCIETY OF WASHINGTON

1980-1981
- Officers

President: Richard Banks Secretary: Michael A. Bogan
Vice President: Raymond B. Manning Treasurer: David L. Pawson

Elected Council
Stephen D. Cairns Leslie W. Knapp

Anne C. Cohen Storrs L. Olson
Stanwyn G. Shetler

PROCEEDINGS
Editor: C. W. Hart, Jr.

Associate Editors

Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman
Plants: David B. Lellinger Vertebrates: Richard Banks

Insects: Robert D. Gordon

Membership in the Society is open to anyone who wishes to join. There are no prerequisites.
Annual dues of $10.00 include subscription to the Proceedings of the Biological Society of
Washington. Correspondence concerning membership should be addressed to the Treasurer,
Biological Society of Washington, National Museum of Natural History, Smithsonian Insti-
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The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly
in February, May, August and November.

Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological
Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash-
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Second class postage paid at Washington, D.C., and additional mailing office.

4
:

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 875-886

A NEW SPECIES OF PIKEBLENNY (PISCES:
CHAENOPSIDAE: CHAENOPSITS) FROM THE
WESTERN ATLANTIC

Philip A. Hastings and Robert L. Shipp

Abstract.—Chaenopsis roseola is described from the middle shelf region
of the northeastern Gulf of Mexico. It is distinguished from its closest known
congener in the western Atlantic, C. stephensi, on the basis of pigmentation
pattern, morphometrics, and palatine tooth pattern. The relationship of the
forms is discussed, but the status of a specimen from off Yucatan, previ-
ously assigned to C. stephensi, is unresolved. The habitat of C. roseola, as
observed from a research submersible, consists of ‘‘windrows’’ of shell
rubble.

The blennioid genus Chaenopsis (Pisces: Chaenopsidae) has been divided
into two species groups based on body length, number of fin-ray elements,
and number of body blotches or bands (Bohlke, 1957b). In the western
Atlantic Chaenopsis ocellata, C. resh, and C. limbaughi compose the long-
bodied, high-count (or ocellata) group while the short-bodied (or coheni)
group has been represented by only C. stephensi Robins and Randall (1965).

Recent baseline studies of fishes of the continental shelf of the north-
eastern Gulf of Mexico using semi-balloon trawls with fine mesh (9.5 mm)
liners and Capetown dredges with inner baskets of 6.4 mm mesh have been
especially productive in capturing previously unknown or poorly known
small, cryptic fish species. Among these is a new species of short-bodied
Chaenopsis which is herein described.

Methods and Materials

Methods of taking measurements follow Hubbs and Lagler (1964) except
for eye diameter for which we measured the pigmented eye as described by
Bohlke (1957a). All measurements from snout include the upper lip. Inter-
orbit equals least bony interorbital width. Pectoral fin length equals length
of the longest ray. MP index equals 10x distance between mandibular pores
3 and 2 divided by the distance between mandibular pores | and 2 (Robins
and Randall, 1965). All measurements were made with dial calipers except
snout length, eye diameter, interorbit, upper jaw, distance between man-
dibular pores, caudal peduncle depth, and caudal peduncle length which
were measured with an ocular micrometer on a Wild MS® stereoscope. Fin-
ray counts of median fins were made from X-rays as these counts were

876 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —

Fig. 1. Holotype of Chaenopsis roseola, USNM 221167. Anterior dorsal fin is depressed.

difficult to take directly from specimens. Head pore terminology follows
Johnson and Greenfield (1976). Abbreviations of institutions cited are as
follows: ANSP, Academy of Natural Sciences of Philadelphia; FMNH,
Field Museum of Natural History, Chicago; FSBC, Florida Department of
Natural Resources; GCRL, Gulf Coast Research Laboratory Museum;
LACM, Los Angeles County Museum; UAIC, University of Alabama Ich-
thyological Collection; UF, Florida State Museum, University of Florida;
UMML, University of Miami, Rosenstiel School of Marine and Atmospher-
ic Sciences; USAIC, University of South Alabama Ichthyological Collec-
tion; USNM, United States National Museum of Natural History.

Chaenopsis roseola, new species
Flecked pikeblenny
Figs. 1-3

Chaenopsis ocellatus (in part). Springer and Woodburn 1960, p. 77. USNM
134923, two specimens.

Holotype. —USNM 221167 (originally USAIC 03661), 42.2 mm SL, male.
30°07’N, 86°45’W, northeastern Gulf of Mexico, about 35 km SSW of Ft.
Walton Beach, Florida, 19 March 1977, 55 m. Collected with a semi-balloon
trawl from a bottom of coarse shell rubble.

Paratypes.—USNM 221168 (3 specimens, 35.1—-36.2 mm SL), collected
with the holotype. GCRL 16893 (1, 42.7), 30°10’N, 86°50’W, about 35 km
SSW of Ft. Walton Beach, FL, 22 May 1976, 53 m. GCRL 16894 (1, 28.4),
29°55'48"N, 86°06'36"W, about 40 km SW of Panama City Beach, FL, 6
Sept. 1977, 37 m. ANSP 143748 (1, 36.3) 30°09’30’N, 86°50’30’W, about 35
km SSW of Ft. Walton Beach, FL, 30 Aug. 1976, 55 m. ANSP 143749 (1,
31.5), 29°50’N, 86°06.5'W, about 30 km SW of Panama City Beach, FL, 20
July 1975, 41 m. UF 27444 (1, 41.0), 29°48’00"N, 86°03'30’W, about 40 km
SW of Panama City Beach, FL, 4 June 1974, 40 m. UF 27445 (1, 29.9),
28°19’00"N, 84°21'00"W, about 60 km SSE of Apalachicola, FL (Florida
Middle Grounds), 18 June 1974, 50 m. LACM 38701-1 (1, 34.8), 29°55'48’N,
86°06'36"”W, about 40 km SW of Panama City Beach, FL, 6 Sept. 1977, 37

VOLUME 93, NUMBER 4 877

Table 1.—Frequency of counts for western Atlantic short-bodied Chaenopsis. All counts
(except pectoral rays) for C. roseola include the holotype, seven paratypes, and two non-type
specimens (USNM 134923). Pectoral fin ray counts include only the type material in which
accurate counts could be made. * = holotype.

Dorsal fin
Spines Rays Total elements
17 18 26 27 28 29 30 44 45 46 47
C. roseola 6 (SaaS ieee) 4 Sc7
C. stephensi (LACM 20157) 1 1 1
C. sp. (UMML 28601) 1 1 1
Anal fin rays Pectoral fin rays Vertebrae
29 30 31 12 13 14 48 49 50 51 52
C. roseola 4* 6 Sm) (Shaye 1 he
C. stephensi (LACM 20157) 1 iy 1
C. sp. (UMML 28601) 1 2 1

m. FMNH 83918 (1, 30.5), 29°55'48"N, 86°06'36”’W, about 40 km SW of
Panama City Beach, FL, 6 Sept. 1977, 37 m. UAIC 5948.01 (1, 28.6),
29°55'48"N, 86°06'36”’W, about 40 km SW of Panama City Beach, FL, 6
Sept. 1977, 37 m. USAIC 06271 (1, 30.0), 29°55'48’”N, 86°06'36”"W, about 40
km SW of Panama City Beach, FL, 6 Sept. 1977, 37 m.

Diagnosis.—A short-bodied species of Chaenopsis with relatively few
vertebrae (48-49), few dorsal fin elements (X VII-XVIII, 26—28; 44—45 total)
and few anal fin elements (II, 29-30). Eight black blotches present along the
side, first through sixth typically inverted triangles, seventh and eighth hor-
izontally elongate blotches. Flecks of rusty or pink pigment scattered over
entire body with two or three prominent (though variable in shape) flecks
on cheek. Dorsal fin low in both sexes. Males with a black blotch on dorsal
fin membranes between spines I and IV. Palatine teeth in one row, those in
anterior section of row moderate in size (none noticeably enlarged), those
in posterior section small.

Description.—Vertebral and fin-ray counts are given in Table 1. Sixteen
precaudal and 32-33 caudal vertebrae. Dorsal fin low in both sexes (Fig. 2),
composed of XVII-XVIII spines and 26-28 unbranched rays (44-45 total
elements). Anal fin with two closely spaced spines and 29-30 unbranched
rays. Pectoral fin rounded, composed of 12-14 unbranched rays. Pelvic fin
I, 3; first and second rays elongate, third short and inconspicuous (about as
long as pelvic spine). Body proportions are given in Table 2.

Snout bluntly V-shaped when viewed from above, i.e., lateral edges con-
verge from the posterior nostril forward (Fig. 3). Forehead sloping when
viewed from side (Figs. 1, 3). Lower jaw projecting slightly, visible from

878 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2.0mm

Fig. 2. Anterior dorsal fins of a) male (USNM 221167, holotype) and b) female (USNM
221168, paratype) of Chaenopsis roseola.

above. Dewlap on lower jaw extending to the anterior edge of orbit. Anterior
nostril tubular, slightly shorter in length than maximum width of bony in-
terorbit. Posterior nostril with a raised rim. Tongue long, slender, coming
to a rounded point. Tip of tongue extends past vomerine tooth patch which
is anterior to origin of palatine tooth row.

Outer tooth row of upper jaw bluntly U-shaped, composed of canines
anteriorly and laterally. Teeth largest at corner of snout and decreasing in
size posteriorly (ANSP 143748 with 5 canines across the left side of the
front of the snout, followed by 19 teeth in the lateral series). Two to 4
irregular rows of fine pointed teeth behind the outer row on anterior part of
mouth, extending in a wedge from mid-line outward and backward to the
fourth tooth of the lateral series. Palatine teeth 17-18, in a single row orig-
inating near the thirteenth or fourteenth tooth of lateral series; anterior teeth
in row moderate in size and pointed (7 in ANSP 143748) followed by a series
of small teeth (10 in ANSP 143748). A few minute teeth on the vomer (3 in
ANSP 143748). Outer tooth row of lower jaw bluntly U-shaped, composed
of canines anteriorly (4 across the left of the front in ANSP 143748), with
lateral series of teeth composed of canines anteriorly grading to low rounded
teeth posteriorly (ANSP 143748 with 5 lateral canines, followed by 12 close-
set moderate, pointed teeth, followed by 12 smaller close-set teeth, those
in posterior part of latter section low and rounded). Teeth behind outer row
similar to those in upper jaw. Two to 4 irregular rows of low pointed teeth
in a wedge extending from middle of jaw back to fifth lateral canine.

Head pores are illustrated in Fig. 3: Nasal, 1 pair; anterofrontal, | pair;
infraorbital, 5 pairs; supraorbital, 3 pairs; commissural, 1 median; supratem-
poral, | median + 1 pair; posttemporal, 4 pairs; preopercular, 5 pairs; man-
dibular, 4 pairs.

Color description.—Holotype, USNM 221167, 42.2 mm SL, male (Fig.
1); notes taken shortly after preservation in formalin and transfer to 45%
isopropyl alcohol. Background color straw. Body with 8 black blotches
along flank, extending ventrad from midline. First located over middle of

VOLUME 93, NUMBER 4 879

Tp pele
AF cP SO D
N bods
: Q aa bE
NA Vs \
5 Se °
O. \ , '
tS Se g
si ee ent ati
IFO Vaan
MD4 POP

Fig. 3. Semi-diagrammatic drawing of the head-pore pattern of €haenopsis roseola (UF
27444). Pores are enlarged to illustrate their positions. Pores within a series are connected by
a dashed line. N = nostrils, NA = nasal, AF = anterofrontal, CP = commissural, SO = su-
praorbital, IFO = infraorbital, MD4 = fourth mandibular, POP = preopercular, PT = post-
temporal, ST = supratemporal, D = dorsal fin origin.

belly, last on caudal peduncle. On left side, blotches 1—3 and S—6 inverted
triangles, 4 more rectangular, 7-8 horizontally elongate; blotches 1 and 2
each with 2 pink flecks within triangles. On right side, 1—6 inverted triangles,
7-8 horizontally elongate; blotches 1, 2, and 4 with pink flecks within tri-
angles. Blotches 1-7 each with 2 rows of melanophore clusters (forming
saddles) extending upward and across dorsum. A saddle also present an-
terior to the pectoral fin bases and concentrations of melanophores present

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 93, NUMBER 4 881

under the opercular flaps at the bases of this saddle. Faint rows of melano-
phores across dorsum between each of the 8 saddles. Clusters of melanophores
scattered along flank. Small, irregularly shaped pink flecks scattered along
dorsum. Eleven pairs of pink dots present on either side of body along anal fin
base. Pink blotch present on upper one-third of right pectoral fin base, not
present on left side. No pigment on belly. Melanophores scattered on head,
most dense behind eye extending posteriorly to edge of preopercle. Melano-
phores present on interorbit, preorbit, suborbit, upper and lower lips, isthmus,
lower branchiostegal membrane (few on upper membrane), chest, pectoral
fin base, and operculum. Pink flecks present on operculum, preoperculum,
snout, and upper lip (including fold above upper lip). Three pink dots present
on anterior edge of lower lip. Iris pink. Cheek with 2 prominent pink hori-
zontally elongate blotches; larger blotch posterior to eye (at level of pupil),
smaller one below anterior edge of larger blotch, above maxillary. On right
side of head this smaller blotch is narrower than on left side. Dorsal fin
membrane between spines I-IV with a concentration of melanophores (Fig.
2) which form an ill-defined but prominent blotch (darkest between II and
IV). Scattered melanophores present on most spines and rays of dorsal fin.
Anal fin with a faint concentration of melanophores on membrane between
spines and first ray. Scattered melanophores present on most rays. Scattered
melanophores present on caudal fin, concentrated at center of base of fin.
Pectoral fin unpigmented. Pelvic fin with few melanophores on bases of
rays.

Paratype, USNM 221168, 36.0 mm SL, female. Color of freshly preserved
specimen similar to holotype except generally less intensely pigmented.
Lateral blotches similar on both sides; 1-6 inverted triangles, 7-8 horizon-
tally elongate. Blotches 2, 3, 4, 6, and 7 with pink flecks within blotches.
Pink blotch present on upper pectoral fin base of both sides. Eleven pairs
of pink dots present along body at anal fin base. Left cheek with upper pink
blotch broken into 2 smaller blotches. Lower blotch also broken into 2
separate blotches. Right cheek with blotch behind eye broken into 3 spots
with no lower blotch present. Melanophores scattered over head although
not as densely as in holotype. Lower jaw with melanophores broken into
4 bands with unpigmented areas between them. No black blotch on anterior
dorsal fin.

Basic color pattern similar in all other specimens examined although in-
tensity varies greatly. Rarely an individual has one of the black lateral
blotches ill-defined or broken, giving the appearance of 2 proximal blotches.
Live specimens show rust-colored instead of pink-colored flecks. These
flecks turn pink upon fixation in formalin and fade entirely after a few
months of preservation in isopropynol. Prominent, though variously shaped,
rusty or pink flecks present on the cheek of all specimens.

882 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Peritoneum of those specimens examined internally with scattered me-
lanophores, flecks of pink and two black blotches (one on either side) in the
shape of inverted triangles directly beneath and corresponding to the first
lateral exterior blotches. One male (ANSP 143748) with melanophores cov-
ering belly and concentrated in a semi-circle in front of the genital area; no
pigment on belly of other specimens. Black pigment at the corner of the
mouth inside the lips of ANSP 143748.

Comparisons.—Of the western Atlantic species, C. roseola is most sim-
ilar to C. stephensi with which it shares a short body and low meristic
counts relative to other Atlantic species of Chaenopsis (Robins and Randall,
1965). Chaenopsis roseola differs from C. stephensi in a number of char-
acters including pigment pattern, palatine tooth pattern, and some morpho-
metric characters.

The color pattern of C. roseola (in preservative) differs from that of C.
stephensi in that 8 instead of 6 lateral blotches are present. Robins and
Randall (1965) indicate that 5 blotches are present along the side of the
holotype of C. stephensi, but we count 6 faint blotches along the side.
Unfortunately, the life colors of C. stephensi are unknown.

The palatine tooth row of C. roseola is composed of teeth of essentially
two sizes, the anterior portion of the row being of moderately sized teeth
with the posterior portion of small teeth. The palatine tooth row of C.
stephensi is composed of 16 or 17 (some are broken) teeth which are irreg-
ular in size with large teeth interspersed throughout the row of otherwise
small teeth.

Several morphometric differences exist between C. roseola and C. ste-
phensi although these may not always be diagnostic when used alone, a
situation common in chaenopsids (Stephens, 1963) including the genus
Chaenopsis (Robins and Randall, 1965). Excluding characters which ap-
parently vary allometrically (see below), C. roseola differs from C. stephensi
in having a shorter predorsal length, a larger eye, and a deeper caudal peduncle
(Table 2). Since only a single specimen of C. stephensi is known (a second
is doubtful), establishment of morphometric variation in that species is pres-
ently impossible; reliable comparison of such characters must await the
collection of more material of C. stephensi.

Chaenopsis roseola differs from the Arrowsmith Bank specimen (UMML
28601) identified as C. stephensi by Robins (1971) in number of vertebrae,
number of dorsal fin rays, number of anal fin rays (Table 1), and number of
blotches along the side (7 in UMML 28601) as well as several non-allometric
morphometric characters including head length, head depth, predorsal
length, caudal peduncle depth, and caudal peduncle length (Table 2).

Chaenopsis roseola differs from the other known western Atlantic Chaen-
opsis species principally in having a shorter body and fewer vertebrae, fin-

VOLUME 93, NUMBER 4 883

ray elements, and lateral body markings. Chaenopsis roseola as well as C.
stephensi belong to the short-bodied species group as defined by Bohlke
(1957b) and thus are allied with C. coheni and C. deltarrhis of the eastern
Pacific.

Range and habitat.—The type-material of Chaenopsis roseola is from the
northeastern Gulf of Mexico, from the head of the De Soto Canyon eastward
and southward to the Florida Middle Grounds. Additional Gulf of Mexico
specimens have been collected as far south as off the Tampa Bay area (see
below). Two specimens referable to C. roseola have been collected from
the Atlantic Ocean off North Carolina by D. J. Stewart.

Recently, the area adjacent to and just east of the northern rim of the De
Soto Canyon has been trawled and dredged extensively; the samples taken
were marked by abundant shell and rubble. This area, from which the ma-
jority of the specimens of C. roseola were captured, was examined during
a 3-hour dive by the research submersible DIAPHUS during June 1978.
Observations made during this dive by one of us (RLS), recorded on audio-
tape for later transcription, revealed a bottom with ‘‘windrows’’ of rubble
and coarse shell hash. These were of about 1-3 m width, with intervening
areas of silica sand, of about the same width. Numerous observations of
small fishes, thought to be the pirate blenny, Emblemaria piratula, and
Chaenopsis roseola (recorded .as Chaenopsis sp.) were made. The fish were
observed to dart to and from rubble and shell fragments, and to retreat
backward into the cover when approached closely by the submersible. On
several occasions the submersible was placed at rest on the bottom for more
prolonged observation. Although slight protrusion of the head region of a
chaenopsid was occasionally detected, this was never close enough to obtain
a diagnostic photographic or videotape record. In addition, the pearly ra-
zorfish, Hemipteronotus novacula, was frequently noted projecting from
burrows. The sand perch, Diplectrum formosum, was also frequently noted
in this area. ‘

Extensive dredge and trawl operations along the northeastern Gulf shelf
show that these shell rubble areas form mosaics north and east of De Soto
Canyon. Several previously unknown or poorly known species have been
collected from this habitat with C. roseola. These include Emblemaria pir-
atula (Chaenopsidae), Gobulus myersi, Palatogobius paradoxus (Gobi-
idae), an undescribed species of Gillelus (Dactyloscopidae), and two un-
described ophichthid eels (Ophichthidae).

Collection depths of C. roseola range from 33 to 64 m, but suitable habitat
is present beyond this depth range. Chaenopsis roseola probably inhabits
the entire lower shelf region of the northeastern Gulf where the preferred
Shell rubble patches exist, as well as the eastern continental shelf of the
southeastern United States.

884 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology.—The name roseola is from the Latin roseus, rosy colored.
This name is selected for the pink or rust colored flecks (in living adults)
reminiscent of roseola or measles.

Discussion.—Stephens (1963) attributed the wide variation seen in mor-
phometric characters in chaenopsids to four factors: 1) growth, 2) individual
variation, 3) sexual dimorphism, 4) error in measurement. A fifth factor,
shrinkage of specimens in preservative, may also account for some variation
in measurements. The holotype of C. stephensi has apparently undergone
some shrinkage since its description, as our measure of its standard length,
44.8 mm, is less than that given by Robins and Randall (1965), 45.8 mm.
This further emphasizes the need of more material of C. stephensi before
an adequate comparison with C. roseola can be made.

At least two morphometric characters vary allometrically in C. roseola
(Table 2). Larger individuals have proportionately shorter pelvic fin rays
and, at least among males, larger individuals tend to have proportionately
longer upper jaws. Bohlke (1957b) discusses the allometry of the upper jaw
in Chaenopsis and points out that the two species groups differ in the rate
of change of the length of the upper jaw relative to the head length. In the
short-bodied coheni group, the maxillary is longer at all stages of growth
than in the long-bodied ocellata group. However, Stephens (1963) points
out that the relative jaw elongation in the coheni group may be due to the
increase in body elongation of the ocellata group. While in the male spec-
imens of C. roseola the relative length of the upper jaw increases with
growth, the opposite appears to be true for the four female specimens (al-
though a good size range of females is lacking). Additional specimens are
needed to determine the growth characteristics of the upper jaw in C. ro-
seola.

Sexual dimorphism is subtle in C. roseola. Both sexes have low dorsal
fins (Fig. 2) but males are distinguishable by a black blotch (which females
lack) on the anterior dorsal fin. In C. ocellata the anterior dorsal fin mark
(also present only in males) is used for display when defending territories
(Robins et al., 1959) and may serve a similar function in C. roseola. Robins
and Randall (1965) describe the holotype of C. stephensi (presumably a
female) as having a dusky area on the anterior part of the spinous dorsal
fin. Females of C. roseola have no such pigment on the dorsal fin.

In the holotype of C. stephensi the second mandibular pore is closer to
the first than to the third. In ten of the specimens of C. roseola the second
pore is closer to the third than the first (MP index less than 10, Table 2).
However, in three specimens the second is closer to the first (MP index greater
than 10). This variation calls into question the use of this character for the
delineation of related groups within the genus Chaenopsis (Bohlke, 1957b).

Chaenopsis stephensi was described from a single specimen apparently
taken from Cubagua Island, Venezuela, at Allan Hancock station A28-39 at

VOLUME 93, NUMBER 4 885

a depth of 2 fathoms (Robins and Randall, 1965). However, in their remarks
on the species, these authors implied some doubt as to the locality data:
‘*Even if a locality error is involved, stephensi is unidentifiable with any
Pacific species.’’ This comment also reflected their assessment that although
close to the Pacific C. coheni, their species was in fact distinct. Subse-
quently, Robins (1971) stated, in reference to this matter, that ‘‘Since C.
stephensi belonged to a species group otherwise known from the Pacific
shore of Central America and since the Allan Hancock collections encom-
passed both coasts, there was reason to doubt the origin of the holotype,”’
but reported on a second specimen (UMML 28601) from Arrowsmith Bank
off Yucatan, Mexico which verified the provenance of C. stephensi.

The taxonomic status of the Arrowsmith Bank specimen reported by Ro-
bins (1971) is unresolved. In some characters (predorsal length, eye size) it
resembles C. roseola. In other characters (caudal peduncle depth) it more
closely resembles C. stephensi. However, in many characters it is unique
(e.g., number of vertebrae, dorsal rays, and anal rays, head length, head
depth, upper jaw length, caudal peduncle depth). More material of this form
is needed to resolve its status. Additional material from Venezuela is also
needed to elucidate relationships within this distinctive sub-group of Chaen-
opsis in the western Atlantic.

Additional material examined.—Chaenopsis roseola: USNM 134923 (2,
29-33). 28°45’00"N, 85°02’00”W, off Cedar Keys, FL, 15 March 1885, about
64 m. FSBC 6567 (1, 22.2), 27°37'N, 83°58'W, off Tampa Bay, FL, 20 Nov.
1966, 55 m. FSBC 6889 (1, 34.6), 27°37'N, 83°58’ W, off Tampa Bay, FL, 2 Aug.
1967, 55 m. Uncatalogued (1, 31.3), R(}V EASTWARD cruise ES5-77, sta. 11,
34°34.2'N, 75°13.4’W, off North Carolina, 3 Aug. 1977, 33 m. Uncatalogued
(1, 26.5), R/V EASTWARD cruise ES5-77, sta. 19, off North Carolina.
Chaenopsis stephensi: LACM 20157, holotype, Cubagua Island, Venezuela,
10°49'25"N, 64°16'00"W, 15 April 1939. 2 fathoms (3.7 m). Chaenopsis sp.
(stephensi of Robins, 1971): UMML 28601, Arrowsmith Bank, off Yucatan,
Mexico, 21°05’N, 86°31’W, 20 Aug. 1970, 275 m.

Acknowledgments

We thank C. R. Robins and W. F. Smith-Vaniz for their assistance in the
early part of this study and R. K. Johnson for his comments on the manu-
script. S. A. Bortone, T. S. Hopkins, C. R. Robins, R. K. Johnson, and D.
J. Stewart provided specimens for examination. R. J. Lavenberg is espe-
cially thanked for the loan of the holotype of C. stephensi. W. R. Courtenay,
C. E. Dawson, E. A. Lachner, and M. M. Leiby allowed use of X-ray
equipment. Linda Lutz drew Fig. 2. Tom Smoyer provided photographic
assistance. This is contribution number 175 of the Harbor Branch Foun-
dation, Inc.

886 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Bohlke, J. E. 1957a. The Bahama species of emblemanid blennies.—Proc. Acad. Nat. Sci.

Philadelphia 109:25—57.

. 1957b. A review of the blenny genus Chaenopsis and the description of a related new

genus from the Bahamas.—Proc. Acad. Nat. Sci. Philadelphia 109:81-103.

Hubbs, C. L., and K. F. Lagler. 1964. Fishes of the Great Lakes region.—Univ. Michigan
Press, Ann Arbor, 213 pp.

Johnson, R. K., and D. W. Greenfield. 1976. A new chaenopsid fish, Emblemaria hyltoni,
from Isla Roatan, Honduras.—Fieldiana: Zoology 70(2): 13-28.

Robins, C. R. 1971. Comments on Chaenopsis stephensi and Chaenopsis resh, two Caribbean

blennioid fishes.—Carib. J. Sci. 11(3—4):179-180.

, C. Phillips, and F. Phillips. 1959. Some aspects of the behavior of the blennioid fish

Chaenopsis ocellata Poey.—Zoologica 44(2):77-88.

Robins, C. R., and J. E. Randall. 1965. Three new western Atlantic fishes of the blennioid
genus Chaenopsis, with notes on the related Lucayablennius zingaro.—Proc. Acad.
Nat. Sci. Philadelphia 117(6):213-234.

Springer, V. G., and W. D. Woodburn. 1960. An ecological study of the fishes of the Tampa
Bay area.—Florida State Bd. Conserv., Prof. Papers 1:1—104.

Stephens, J. S., Jr. 1963. A revised classification of the blennioid fishes of the American
family Chaenopsidae.—Univ. California Publ. Zool. 68:1—165.

(PAH) Harbor Branch Foundation, Inc., RR 1, Box 196, Ft. Pierce, Flor-
ida 33450 [present address: Dept. of Ecology and Evolutionary Biology,

University of Arizona, Tucson, Arizona 85721]; (RLS) Dept. of Biology,
University of South Alabama, Mobile, Alabama 36688.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 887-913

THE RELATIONSHIPS OF THE SOUTH AMERICAN
PYGMY CHARACOID FISHES OF THE GENUS
ELACHOCHARAX, WITH A REDESCRIPTION
OF ELACHOCHARAX JUNKI
(TELEOSTEI: CHARACIDAE)

Stanley H. Weitzman and J. Géry

Abstract.—Elachocharax junki (Géry), previously Known only from the
holotype, is redescribed on the basis of four additional specimens from the
Rio Madeira basin and one from a tributary of the Rio Negro, Amazonas,
Brazil. This species, originally described as Geisleria junki and placed in a
monotypic subfamily, Geisleriinae, is here considered to be a species of
Elachocharax of the characid subfamily Characidiinae. Elachocharax con-
sists of three species, E. pulcher, E. geryi, and E. junki. Their relationships
are discussed and a key is provided for their identification. The relationships
of Elachocharax to the other genera of the Characidiinae is treated. The
Geisleriinae and Elachocharacinae of Géry (1971) are rejected and placed
as synonyms of the Characidiinae.

The first known pygmy characoid related to Characidium Reinhardt and
Jobertina Boulenger but having a very long dorsal-fin base with 17 to 20
rather than 10 to 14 dorsal-fin rays was Elachocharax pulcher Myers (1927)
from the central region of the Rio Orinoco in Venezuela. In recent years,
further collecting in the Amazon basin has resulted in the description of
additional nominal species of Elachocharax. Elachocharax georgiae Géry
(1965) was shown by Weitzman and Kanazawa (1978) to be a synonym of
E. pulcher but two additional valid species have been described. Géry (1971)
established a new genus and species, Geisleria junki (here placed in Elach-
ocharax), from the Rio Novo in an area about 45 km east of Porto Velho
in the basin of the Rio Madeira, Brazil. Weitzman and Kanazawa (1978)
described a new species, Elachocharax geryi, from the middle of the Am-
azon basin near Manaus, Brazil, and from Cano Muco, an upper tributary
of the Rio Orinoco in Colombia. They further reported the extension of the
known range of Elachocharax pulcher from the mid Rio Orinoco basin to
an area covering much of the central Amazon basin.

Geéry (1971) assigned Geisleria junki to a new subfamily, Geisleriinae,
based on its apparent possession of two rows of premaxillary teeth, pre-
sumed absence of ectopterygoid teeth, the presence of what appeared to be
maxillary teeth, a supposed absence of ‘‘suborbital’’ bones, and absence of

888 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a cranial fontanel. At that time Gery recognized two new, but related groups
at the subfamily level, the Geisleriinae and the Elachocharacinae, the latter
based on Elachocharax of Myers (1927). He included these taxa along with
the Characidiinae in a family, the Characidiidae. This family, here tenta-
tively recognized as a subfamily of the Characidae, includes Characidium,
Jobertina, Klausewitzia Géry (1965), Elachocharax, and Ammocryptochar-
ax Weitzman and Kanazawa (1976). Géry (1971) correctly pointed out that
the phylogenetic relationships of this family group to other characoids are
uncertain, a fact agreed to by Weitzman and Kanazawa (1978) who briefly
reviewed the history of the placement of these fishes by ichthyologists. Géry
further noted that the species of the Elachocharacinae and the Geisleriinae
form a group of small fishes related to Characidium but with a long-based
dorsal fin and apparently living in very slow-moving waters. Species of
Characidium, Jobertina, and Ammocryptocharax are ordinarily found in
streams of moderate to rapid flow whereas Elachocharax, Klausewitzia,
and the nominal Geis/eria occur in quiet waters of large river basins. Weitz-
man and Kanazawa (1976) tentatively rejected the Elachocharacinae and
Geisleriinae as valid taxa at the subfamily level and adopted a hypothesis
that all the genera noted above belong to a subfamily, the Characidiinae,
whose characoid relationships are uncertain.

The examination of a specimen from the Rio Negro basin, here considered
a female of the nominal Geisleria junki, of four specimens of this species
from the Rio Madeira basin, and a reexamination of the holotype of G. junki
has allowed a new interpretation to be placed on the phylogenetic relation-
ships among the species Elachocharax pulcher, Elachocharax junki, and
Elachocharax geryi.

A key is presented below to the species of Elachocharax. Elachocharax
junki is redescribed and the relationships among the three species are dis-
cussed. Finally, the possible relationships of Elachocharax with other
species and genera of the Characidiinae is reviewed.

Elachocharax Myers

Elachocharax Myers, 1927:114, original description, type by monotypy
Elachocharax pulcher Myers (1927).—Géry, 1977:122, brief discussion.—
Weitzman and Kanazawa, 1978:160, redefinition and discussion of rela-
tionships.

Elacocharax Weitzman and Kanazawa, 1976:328, misspelling for Elacho-
charax.

Geisleria Géry, 1971:154, original description, type by monotypy Geisleria
junki Géry (1971).—Géry, 1977:122, brief discussion.

Weitzman and Kanazawa (1978) redefined the genus and discussed its
presumed phylogenetic relationships. Inclusion herein of the species E. jun-

VOLUME 93, NUMBER 4 889

ki modifies that description. A discussion of the relationships of the genus
will be found below, after the redescription of E. junki.

Diagnosis.—The following two correlated characters will distinguish
Elachocharax from all other known genera in the Characidiinae. Total num-
ber of dorsal-fin rays 17-22. Anterior origin of the anal fin either ventral to
or in advance of the posterior termination of the dorsal-fin base.

Description.—Other characters useful in defining Elachocharax are as
follows. Premaxillary teeth in single series, partly tricuspid, bicuspid, and
unicuspid, or only unicuspid. Maxillary and mesopterygoid teeth absent.
Dentary teeth in two rows. Outer row of partly tricuspid, bicuspid, and
unicuspid, or only unicuspid teeth. Inner row unicuspid. Ectopterygoid
teeth present, unicuspid, and nearly uniserial. Mouth terminal or snout
slightly in advance of lower jaw. Adipose fin present or absent. Scale rows
between anterior dorsal-fin origin and pelvic-fin origin 8. Predorsal scales
8-10. Perforated lateral-line scales few, 5—9. Scales in a lateral series 26—
33. Scale rows around caudal peduncle, 14 (Weitzman and Kanazawa, 1978,
erroneously reported 12 scale rows around caudal peduncle for E. pulcher).
Vertebrae 29-32, including those of Weberian apparatus and terminal cen-
trum. Branchiostegal rays 4-5. Body relatively short, greatest depth at dor-
sal-fin origin 23-30% of standard length. First and second infraorbital bones
with a well developed laterosensory canal. Third, fourth, and fifth infraor-
bital bones apparently absent.

Key to the Species of Elachocharax

1. Premaxillary teeth tricuspid anteriorly followed by a few bicuspid
and/or unicuspid teeth. Dentary with tricuspid teeth anteriorly fol-
lowed by none, one, or a very few bicuspid teeth and then several
unicuspid teeth lateroposteriorly. Adipose fin present. Branchioste-

WE AUMRDY Sie oe eee ot onc epee ane os oils eho ou tee fon So Ree Elachocharax pulcher
2. Premaxillary and dentary teeth unicuspid. Adipose fin absent. Bran-
CIOS CS TALE AY SA erase ooo Eh sors i SAS RN RS apache eas ie syed: sede ye 3

3. Caudal fin without obvious vertical dark bars. Teeth of ectopterygoid
8—13. Inner row dentary teeth 19-20. Upper limb gill rakers 5-6.
Anterior anal-fin origin in advance of posterior termination of dorsal-
fin base. Snout length about 5.4-6.1% of standard length. Caudal
peduncle length about 19.0—22.0% of standard length ............

er eae Se ETN Oh LEN, tie ee EN geese Elachocharax junki

4. Caudal fin with 13-14 narrow vertical dark bars. Teeth of ectopter-
ygoid 4—5. Inner row dentary teeth 13-16. Upper limb gill rakers 3.
Anal-fin origin ventral to or slightly posterior to posterior termina-
tion of dorsal-fin base. Snout length about 6.0-8.0% of standard
length. Caudal peduncle length about 16.0—19.0% of standard length

Pre APS a SIS as. i Neti Nt Ae ei ea an Elachocharax geryi

890 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A description and synonymies of E. pulcher and a description of E. geryi
appear in Weitzman and Kanazawa (1978). Some additional localities for
Elachocharax pulcher are as follows: FMNH 87812, (2), 13.8-14.6 mm SL,
Peru, Rio Nanay, Mishana, about 33 km west of Iquitos (73°30’ W, 3°50’S),
5 November 1974, Spieler and Poncho; MPM 24,697, (6), 12.1-14.7 mm SL,
same locality as FMNH 87812; AMNH 382339, (2) 21.9-22.1 mm SL, Bolivia,
Dept. Beni, Arroyo Grande, 4 km west of Guayaramerin, new road to Ri-
beralta (65°27'W, 10°50’S), 23 October 1964, R. M. Bailey and W. Villaroel.

Elachocharax junki (Géry)
Figs. 1-6, Table 1

Geisleria junki Gery, 1971:154, original description, Brazil, Rondonia, Rio
Novo, tributary to Rio Jamari about 45 km east of Porto Velho, Rio
Madeira basin.—Gery, 1977:122, listed, brief discussion.—Weitzman and
Kanazawa, 1978:161, discussion of relationships.

Material examined.—Holotype, collection of J. Gery, 22.4 mm SL, Bra-
zil, locality as listed above in synonymy, collected 18 October 1967 by R.
Geisler and W. Junk; MZUSP 14008, (1) 22.3 mm SL, Brazil, Amazonas,
igarapé Cobrinha, left bank of Rio Cuieiras, approx. 20 km from its mouth
(60°30'W, 2°50’S), 28 January 1977, H. Britski and N. Menezes; collection
of J. Géry, (4) 21.5—23.1 mm SL, Brazil, Amazonas, 44 km east of Humaita
along rodovia Trans-Amazonica, Rio Madeira basin (62°35'W, 7°35'S), 22
August 1976, H. R. Axelrod. Figure | is a map of localities.

Description.—Body relatively short in comparison to most members of
Characidiinae; greatest depth at dorsal-fin origin. Profile of head and back
from snout tip to dorsal-fin origin gently convex with a slight concavity at
nape and dorsal to area of eyes. (Note holotype, Fig. 2, preserved with
branchial apparatus and gill covers expanded and head extended dorsally,
giving a greater concavity at nape than is present in specimens with bran-
chial apparatus, gill cover, and head not in extended position, Figs. 3, 4.)
Dorsal-fin origin in advance of pelvic-fin origin. Anal-fin origin in advance
of posterior termination of dorsal-fin base. Profile of ventral surface of head
convex. Belly slightly convex from region of isthmus to pelvic-fin origin.
Profile of body between pelvic-fin origin and anal-fin origin gently convex
to nearly straight. Caudal peduncle relatively short and deep. Both dorsal
and ventral profiles of caudal peduncle slightly concave.

Head moderately long. Mouth nearly terminal or snout slightly in advance
of lower jaw tip. Snout relatively blunt. Eye large, horizontal diameter ex-
ceeding snout length.

Teeth on dentary in 2 rows (Figs. 5, 6). Outer (anterior and lateral) row
with 14 (13-17, x = 15.5) 16 conic teeth. Inner row with 19 (19-20, « =
19.5) 20 conic teeth. Premaxillary (Fig. 5) with a single row of 11 (12-15,

VOLUME 93, NUMBER 4 891

em
\ D.
S 7 N
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4 { Bh
a) Agr
PQ 5 (Zs
, - ae SS wa G2..." Ay
S AL er f
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XRT

Fig. 1. Localities reported for Elachocharax junki. 1. Type locality Rio Novo, tributary to
Rio Jamari, Rio Madeira basin, Brazil. 2. 44 km east of Humaita along rodovia Trans-Ama-
zonica, Rio Madeira basin. 3. Igarapé Cobrinha, Rio Cuieiras, Rio Negro basin.

= 13) 14 conic teeth. Maxillary toothless. Ectopterygoid with 8 (10-13,
= 11.3) 12 conic teeth in nearly a single row.

Branchiostegal rays 4, two slender anterior rays attached to anterior part
of ceratohyal followed by a broad ray attached to posterior part of cerato-
hyal and another broad ray attached to epihyal. Gill rakers 6 (S—6, « = 5.8)
5 on upper limb and 6 (6-7, * = 6.8) 6 on lower limb. Frontal-parietal fo-
ramen greatly reduced, bounded by posterior borders of parietals and su-
'praoccipital. A narrow antorbital bone present dorsal and posterolateral to
posterior dorsal portion of first infraorbital. First (anterior) infraorbital of
moderate size and well ossified in stained face bones of right side of spec-
imen from Rio Cuieiras. First infraorbital with an anterior process extending
dorsal and anterior to anterior dorsal slender process of maxillary bone.
Second infraorbital bone primarily a narrow laterosensory tube but some
additional bone present. Both first and second circumorbitals contain later-
osensory canals. Other infraorbital bones apparently absent.

Scales cycloid, usually 4—5 radii on exposed field. Lateral line with 7 (7—

1

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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896 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Elachocharax junki (Géry), MZUSP 14008, SL 22.3 mm, igarapé Cobrinha, Rio
Cuieiras, Amazonas, Brazil. Lateral view of premaxillary and maxillary and dentary bones of
right side. Anterior is to the right. Outer dentary tooth row lies in a deep groove. Their
replacement teeth develop in that groove. The inner row dentary teeth are seen posterior to
the ridge behind the groove.

VOLUME 93, NUMBER 4 897

Fig. 6. Elachocharax junki (Géry), MZUSP 14008, SL 22.3 mm, igarapé Cobrinha, Rio
Cuieiras, Amazonas, Brazil. Medial view of right dentary bone. Twenty inner row teeth are
visible on the dentary ridge. Nine replacement teeth can be seen posterior to the inner row
teeth. These replacement teeth point inwards toward the oral cavity. Anterior is to the left.

8, x = 7.8) 8 scales. Scales in a lateral series 31 (30-33, « = 31.5) 31. Scale
rows between dorsal and pelvic fins 8 in all specimens. Scale rows around
caudal peduncle 14 in all specimens. Predorsal scales 10 (9-10, * = 9.3) 9.
Area on belly anterior to pectoral fin with scales.

Dorsal-fin rays 111,16 (411,17 in two, i1,18 in one, and 11,19 in one) 11,18,
last ray not split to its base. Adipose fin absent. Anal-fin rays 1i,6 in all
specimens, last ray split to its base. Pectoral fin elongate reaching to or
somewhat beyond pelvic-fin origin. Pectoral-fin rays total 8 in all specimens,
most rays undivided, sometimes middle two or three divided. Pelvic fin 1,7
in all specimens. Pelvic fin relatively short, reaching anus, or near to it.
Caudal fin forked with principal ray count 10/9 in all specimens.

Total number of vertebrae including Weberian apparatus and terminal
centrum 32 in all specimens except one from near Humaita which has 33.

Color in alcohol.—Background color pale yellowish brown, with dark
brown chromatophores relatively dense except in central areas of pale
scales. Definite dark vertical bars not readily apparent in some specimens
(Figs. 2, 4) although in areas where scales have dark centers as well as dark
borders, bars may appear in life as well as in preserved specimens (Fig. 3).
If this is so, about 16 to 18 dark vertical bars appear present. Géry (1971:159)

898 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

remarked that other than scales being bordered in dark pigment, no really
dark markings were present on body of holotype. Body pigment in preserved
specimens of all three species of Elachocharax seems quite variable; see
Weitzman and Kanazawa (1978: Figs. 2, 3, 4, 9, 10) for illustration of this
variation in E. pulcher and E. geryi. E. pulcher especially appears to have
a dark as well as a light colored phase that may be changed in life; compare
Figs. 2 and 3 in Weitzman and Kanazawa (1978). E. junki may have a similar
ability to change color pattern in life by selective expansion and contraction
of dark chromatophores. All specimens from 44 km east of Humaita have
color pattern illustrated in Fig. 3. Detailed examination of Fig. 2 of holotype
of E. junki reveals traces of color pattern of parallel vertical rows of dark
scales clearly shown in Fig. 3; the same is true for the specimen in Fig. 4.
This color pattern of parallel vertical rows of darkly pigmented scales in E.
junki is most like that of E. geryi; compare with Figs. 9 and 10 in Weitzman
and Kanazawa (1978:174) and see discussion below.

Top of head pale brown, darker from dorso-posterior border of eye to
near nape. Snout pale brown dorsally; a prominent dark stripe extends from
snout tip and oral border of premaxillary and anterior part of maxillary to
anterior border of eye. Anterior tip of lower jaw dark brown. Dark brown
chromatophores ventral to eye organized into one or two short vertical bars.
Opercle with scattered dark chromatophores apparently somewhat orga-
nized according to borders of opercular, preopercular, subopercular, and
interopercular bones (Fig. 4). Ventral surface of head pale yellow with small,
scattered brown chromatophores.

Caudal fin without vertical bars; interradial membranes with dark chro-
matophores producing a dusky appearance to fin which in life may be quite
dark when chromatophores expanded. Dorsal fin rather evenly dusky with
distal parts of interradial membranes quite dark, especially anteriorly, in
some specimens (Fig. 2). As in other species of Elachocharax, a dark hor-
izontal band present along length of dorsal fin near its base (sometimes
indistinct as in Fig. 3); see especially Fig. 4. Anal fin with interradial mem-
branes usually very dark; anal fin probably black in life when dark chro-
matophores expanded. Pelvic fins with pigment distributed as in anal fin.
Pectoral fin dusky, with pigment distributed as in pelvic and anal fins but
not as dense as in these fins. Specimens from 44 km east of Humaita appear
to have pale pelvic fins.

Relationships

The three species of Elachocharax are related by the following analysis.
Table 2 shows relative primitive and derived states of 12 characters and Fig.
7 is a diagram showing the relationships of the three species and the distri-
butions of the synapomorphies and autapomorphies discussed below. In the

VOLUME 93, NUMBER 4

899

Table 2.—Characters used in synapomorphy and autapomorphy diagram (Fig. 7).

Character

Plesiomorphic
(relatively primitive)
state

Apomorphic
(relatively derived)
state

. Total number of dorsal-fin
rays

11 to 14

17 to 22

2. Position of anal-fin origin Anal-fin origin Anal-fin origin below,
with respect to dorsal-fin posterior to dorsal- very slightly posterior,
base termination fin base or in advance of

termination of dorsal-
fin base

3. Branchiostegal-ray number 5 4

4. Adipose fin Present Absent

5. Premaxillary and dentary Tricuspid Unicuspid
tooth cusps

6. Number of vertical body 9 to 12 narrow bars 8 wide bars
bars

7. Number of vertical body 9 to 12 16 to 17
bars

8. Position of anal-fin origin Anal-fin origin below Anal-fin origin anterior
with respect to dorsal-fin posterior termination to posterior
base termination of dorsal-fin base termination of dorsal-

fin base

9. Number of teeth in inner 19 to 24 13 to 16
dentary tooth row

10. Number of ectopterygoid 7 to 24 4to5
teeth

11. Number of caudal-fin bars 0 to 5 13 to 14

12. Number of epibranchial gill 5 to 6 3

rakers

discussion of the polarity of the characters below and in Table 2 and Fig.
7, each character is represented by the same numeral.

1. The number of dorsal-fin rays in Elachocharax is 17 to 22. In all other
known species of the other genera of the Characidiinae the number of dorsal-
fin rays is 11 to 14. By far the greater number of new world characoids have
a total of 11 dorsal-fin rays. Thus by outgroup comparison the high number
of dorsal-fin rays in the species of Elachocharax should be a derived shared
character.

2. The reasons for the position of the anal-fin origin, ventral to the pos-
terior termination of the dorsal-fin base, in Elachocharax being considered

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 93, NUMBER 4 901

a uniquely derived character is not as clear as the reasons that the increased
number of dorsal-fin rays is a singularly derived condition. The reasons for
the acceptance of this character as advanced are as follows. Most characoids
including the other Characidiinae have the dorsal fin in advance of the anal
fin. It is only in specialized groups such as the Gasteropelecidae, some of
the glandulocaudin characids, certain genera such as Charax, Iguanodectes,
Agoniates, and Paragoniates, that the dorsal fin is over or posterior to the
origin of the anal fin. The members of each of these characoid groups share
their own unique set of synapomorphies and therefore appear unrelated to
the Characidiinae. Thus it is assumed here that the similarity among these
groups of the position of the anal fin and the dorsal fin is of independent
origin. Two further comments should demonstrate the verity of this state-
ment. In most characoid species or groups with a posteriorly placed dorsal
fin such as Xenagoniates bondi Myers, the gasteropelecids, and some of
the glandulocaudins, the relative position of the dorsal and anal fins has
been derived by an increase in the number of anal-fin rays and an anteriorly
advanced migration of the origin of the anal fin. In Elachocharax the relative
position of the dorsal fin and anal fin is correlated with an increase in the
number of dorsal-fin rays and a posterior extension of the posterior termi-
nation of the dorsal fin over the anal fin, which does not have an increased
number of rays. The relative positions of the dorsal fin in Elachocharax on
the one hand and such characoids as Xenagoniates bondi, the species of
the gasteropelecids and of the glandulocaudins on the other, are not the
same thing. Characters 1 and 2 discussed here are correlated synapomor-
phies for the species of Elachocharax.

The other characters mentioned in the generic description above are not
unique to Elachocharax and only form a combination which will separate
the species of Elachocharax from other species of the Characidiinae. We
cannot at present offer evidence, pro or con, that the other characters men-
tioned in the generic description above for Elachocharax are independently

<_—

Fig. 7. Synapomorphy and autapomorphy diagram for species of Elachocharax. Solid black
squares indicate a derived condition, empty squares indicate a plesiomorphic condition and
half-black squares indicate the same derived condition as the black squares but indicate an
independent origin of that condition. The outgroup line to the left in the diagram labeled ‘‘other
Characidiinae’’ represents species of all other genera of the Characidiinae. The use of half-
black and empty squares along this left hand line indicates that both plesiomorphic and apo-
morphic (derived) conditions of the character in question occur in various of these species.
When the squares are empty along the outgroup line it means that the derived condition of the
character in question is not known outside of Elachocharax. Only a broken line is shown
joining Elachocharax to the ‘‘other Characidiinae’’ and no characters are shown as synapo-
morphies for the Characidiinae because this problem is only partly treated in this paper.

902 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

derived. Many of the characters separating Elachocharax are reductive or
perhaps labile and therefore may have been repetitively evolved, or altered
in similar fashion independently. Such characters appear to be of low value
in phylogenetic evolutionary studies, especially at generic and higher taxo-
nomic levels. :

The following three derived characters can be used to unite E. junki and
E. geryi, and exclude E. pulcher.

3. Branchiostegal rays. Five branchiostegal rays are present in E. pulcher.
According to Weitzman (1962) five branchiostegal rays appear to be primi-
tive for the Characidae and independent repetitive reductions in branchio-
stegal rays appear common in characoid fishes. In some groups of chara-
coids, branchiostegal ray counts appear invariate in species and genera, for
example, the Lebiasinidae (Weitzman, 1964). On the other hand Weitzman
and Kanazawa (1976:330) find that the number of branchiostegal rays can
be variable within genera of the Characidiinae and that the number can
occasionally vary from one side to the other side of the head of a single
specimen. We have not found this kind of variation in Elachocharax, but
our sample size of E. junki and E. geryi is small. In any case the occurrence
of four branchiostegal rays in E. junki and E. geryi is a shared derived state
within Elachocharax and is probably a derived homology and therefore a
synapomorphy for these two fishes at this taxonomic level.

4. Adipose fin absent. Again this character is one that is repeatedly and
independently lost in a wide variety of characoid groups, frequently disap-
pearing in small or miniature species (Myers, 1958; Weitzman, 1962) and
may even be found to be present or absent in individuals of one species of
the Characidiinae (Weitzman and Kanazawa, 1977). Within each species of
Elachocharax the presence or absence of the adipose fin appears stable and
therefore within this genus the absence of the fin may be considered a
derived state shared by E. junki and E. geryi.

5. Premaxillary and dentary teeth unicuspid. As in the previous two char-
acters, the presence of lateral cusps is quite variable among characoids. The
use of the number of tooth cusps in relating the member species and genera
of subfamilies, tribes, and genera in characoids has a long history (see Ei-
genmann, 1912, 1915, 1917; Myers and Bohlke, 1956; Weitzman, 1964). Such
practice may be valid in certain groups of characoids, but it now appears
that using this character at traditional generic and family-group levels should
be done with caution. For example, Myers and Bohlke (1956) proposed the
tribe Xenurobryconini for a group of pygmy characoids apparently related
to glandulocaudin characids. One of the characters used to unite the two
included genera and separate the Xenurobryconini from the remainder of
the glandulocaudins was the possession of unicuspid teeth in the former and
tricuspid teeth in the latter. Weitzman and Fink (in prep.) have described
a new species of Xenurobrycon which in nearly all its characters except the

VOLUME 93, NUMBER 4 903

number of tooth cusps and shape of the enlarged caudal-fin scale is ex-
tremely similar to the previously known species, Xenurobrycon macropus
Myers and P. de Miranda Ribeiro. The new species bears tricuspid and
biscuspid teeth as well as unicuspid teeth. In this instance, at least, the
nature of the cusps appears useful as a species difference only, rather than
at generic or tribal levels. Fewer tooth cusps is a reductive character prob-
ably subject to much independent evolution so that the assumption of its
homology among characoid taxa and its use as a synapmorphy must be
treated with great caution in any given instance. In the genera Characidium,
Jobertina, Ammocryptocharax and Klausewitzia there are species which
have tricuspid teeth in the jaws and others which have only unicuspid teeth.
If these genera, as they are currently constituted, are monophyletic groups,
it would appear that unicuspid teeth evolved at least four separate times in
the Characidiinae, given the assumption that the presence of multicuspid
teeth is primitive for the Characidiinae based on an outgroup comparison
with other characoids. The presence of species with tricuspid or only uni-
cuspid teeth in Elachocharax would make at least a fifth independent oc-
currence of this derived character in the Characidiinae.

Although we are unable to accept the hypothesis that the loss of tooth
cusps is homologous at the currently recognized generic levels in the Char-
acidiinae, we incline to believe that it may be a synapomorphy at the level
of relationships in the case under discussion. We favor this position because
the three characters discussed immediately above represent congruent syn-
apomorphies and thus ‘‘reinforce’’ one another. No synapomorphies were
found to indicate either that E. pulcher and E. geryi are more closely related
to each other than each is to E. junki, or that E. pulcher and E. junki are
more closely related to each other than each is to E. geryi.

The following series of characters appear to be autapomorphies, derived
features allowing separation and identification of the species of Elacho-
charax but providing no information about their relationships.

6. Eight wide vertical bars on body sides. The case for the derived nature
of this character is as follows. In most if not all species of the Characidiinae,
relatively narrow vertical body bars or large mid-side dark marks are pres-
ent, usually 9 to 11 or rarely 12 in number. The wide nature and capability
of these bars to expand so that the whole body is black or dark brown in
E. pulcher would seem, by outgroup comparison with the remainder of the
Characidiinae, to be a derived state.

7. Sides of body with 16 or 17 narrow vertical body bars. This character,
present in E. junki, appears to be an increase in the number of body bars
in comparison with any other known species within the Characidiinae and
is therefore derived. E. geryi, with 11 to 12 vertical body bars, is plesio-
morphic in this respect.

The above analyses of vertical body bars is not without problems. The

904 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

number of vertical body bars in E. pulcher, 8-9, overlaps that, 9-11, found
in the other characidiin genera. However, the width is apomorphic. In E.
geryi the number of vertical body bars is slightly higher, 11-12 versus 9-11,
than those found in most other characidiin species, and again overlaps the
plesiomorphic number. However, the bars in E. geryi and E. junki remain
narrow, a plesiomorphic state.

Two features are found in E. geryi which are not present in E. pulcher
and E. junki and at least most of the other species of the Characidiinae.

8. Anal-fin origin anterior to posterior termination of dorsal-fin base. This
character is a further derived state of the character discussed under 2 above.
The character is autapomorphic for E. junki.

9. Inner dentary tooth row with 13 to 16 teeth. Elachocharax junki and
E. pulcher respectively have 19-20 and 20-24 inner row dentary teeth in
adults. Weitzman and Kanazawa (1978) reported a single exceptional spec-
imen of E. pulcher with 14 inner row dentary teeth. Most other species of
Characidiinae have more than 16 teeth in the inner dentary row of adults
but some pygmy species such as Klausewitzia aphanes have 16 or fewer.
It is to be expected that a reduction in tooth row number would evolve
independently in separately derived pygmy species.

10. Four to five conic ectopterygoid teeth in a single row. E. pulcher has
7-10 and E. junki has 8-13 ectopterygoid teeth. Most other species of the
Characidiinae have more than eight ectopterygoid teeth and sometimes two
Or more rows with as many as 24 teeth. Reductions do occur, apparently
independently, in other genera. For example, Klausewitzia aphanes has 6—
8 in a single row and Ammocryptocharax elegans varies greatly, having 4—
22 in a single row.

Finally E. geryi bears a couple of autapomorphic characters which are
not found anywhere else in the Characidiinae as far as is known.

11. Caudal fin with 13 to 14 narrow vertical dark bars. There are no more
than four or five vertical caudal-fin bars in any other known member of the
Characidiinae, and usually there are none. Caudal-fin bars, especially ver-
tical ones, are not found in most characoids.

12. Three upper limb gill rakers. There are five or six upper limb gill
rakers in the other species of Elachocharax and in the other members of
the Characidiinae known to us. There are usually more than six upper limb
rakers in other characoids. Therefore a reduction to three may be taken as
a derived character for E. geryi.

There are a few other differences among the three species of Elacho-
charax but polarity of derived versus plesiomorphic states of these char-
acters was difficult to hypothesize by outgroup comparison. For example,
peduncle length was 14.4 to 18.0, « = 16.5, n = 36 in E. pulcher; 16.0 to

19.2, « = 17.7, n = 8 in E. geryi; and 19.2 to 21.9, « = 20.3, n = 6in E.
Junki. Elachocharax junki has more lateral series scales, 30-33, « = 31.3,

VOLUME 93, NUMBER 4 905

than either E. pulcher, 26-30, = 28, n = 36, or E. geryi, 27-29, = 27.8,
n = 8. There also appear to be differences in vertebral numbers, 29-31, « =
30.4, n = 62 for E. pulcher; 31-32, « = 31.5, n = 8 for E. geryi; and 32,
yo — 325m — 2 TOR 2. JUNK.

Additional characters were used by Géry (1971) to distinguish E. junki at
generic (Geisleria) and subfamily (Geisleriinae) levels from the Characidi-
inae and from the Elachocharacinae. As noted in the introduction, of these
four taxa the only one recognized here is the Characidiinae. The reasons
for the rejection of the two subfamilies are based on the following analysis.

Géry (1971:162, table, and 163, key) in discussing Geisleria indicates that
E. junki has two rows of premaxillary teeth, a ““suspected’’ presence of
maxillary teeth, a ““suspected’’ absence of ectopterygoid teeth, a stated
absence of ‘‘suborbital’’ bones, and no cranial fontanel. These characters,
as presented by Géry, are in error because no alizarin preparation was avail-
able. The primary character of those above used by Géry to separate Geis-
leria from Elachocharax was the presence of two rows of premaxillary
teeth. Géry’s placement was an attempt to be consistent with a classification
of other characoids as established by Eigenmann (1915, 1917) wherein the
Cheirodontinae were separated from the Tetragonopterinae on the basis of
one versus two rows of teeth on the premaxillary bone. Fink and Weitzman
(1974) have reviewed the problems associated with basing a classification
of characoids at the family-subfamily level on this character. They suggest
the character may not be useful at some taxonomic levels in some characoid
groups due to independent appearance. However, in the instance of Geis-
leria versus Elachocharax the question is moot because the holotype of
Geisleria does not have a second inner row of premaxillary teeth. The
appearance of a second row of teeth is due to thick, very papillose flesh
immediately external to the oral valve and just posterior to the prominent
row of premaxillary teeth which are firmly ankylosed to the premaxillary
bone. Buried in this papillose flesh are some well developed replacement
teeth which are not firmly bound to the premaxillary. They are smaller than
the anterior teeth and point posteriorly and ventrally in the oral cavity.
When the premaxillary teeth of the holotype were first reexamined, two
rows of teeth seemed apparent. In part this was due to the well-developed
state of the replacement teeth and in part to the outward-thrust position of
the premaxillary of the holotype which was preserved with its mouth open
and gill apparatus expanded. However, application of an extremely fine jet
of air to the region to shrink the flesh somewhat away from the second tooth
row revealed that some of the ‘‘teeth’’ deformed and shrank away as pap-
illous flesh. The remaining teeth did not deform but did move with the
receding flesh, indicating that they are not firmly attached to the premaxil-
lary bone. This movement is typical of even well developed replacement
teeth in characoids. Resoaking the fish in alcohol brought the papillous flesh

906 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and replacement premaxillary teeth back into normal position. Examination
of the cleared and stained jaws, orbital bones, palatine and pterygoid arches,
opercle, and branchiostegal rays of the right side of the head in the specimen
of E. junki from the Rio Cuieiras revealed 9 premaxillary replacement teeth
posterior to a single row of 14 prominent teeth firmly ankylosed to the
premaxillary bone. In Fig. 5 the replacement teeth are posterior to the vis-
ible row of teeth and cannot be seen. One tooth has been shed (4th from
the right) and was being replaced by a tooth that is not as yet firmly anky-
losed to the jaw margin. There is only one row of premaxillary teeth in all
the specimens of E. junki examined. Weitzman and Kanazawa (1978) report
only one row of premaxillary teeth in E. pulcher and E. geryi.

Similar investigation of the maxillary bone revealed no maxillary teeth in
the holotype of E. junki and examination of the cleared and stained jaws
showed no maxillary teeth. There were no teeth on the left maxillary bone
in any of the new specimens. There are no maxillary teeth in E. pulcher or
E. geryi.

There are 8 conic ectopterygoid teeth in nearly a single row on both sides
of the holotype of E. junki and 11-13 ectopterygoid teeth in the other spec-
imens examined. Weitzman and Kanazawa (1978:168, 175) report a single
row of 7-10 unicuspid ectopterygoid teeth in E. pulcher and 4-5 in E. geryi.

Gery (1971:154) stated that the circumorbital series was very reduced or
perhaps even totally absent in E. junki. On pages 162 and 163 he states that
the suborbitals are absent and uses this as one of the characters to distin-
guish his Geisleriinae and Elachocharacinae. The suborbitals are usually
considered to be circumorbitals two, three, and sometimes the remainder
of the posterior orbital series exclusive of the supraorbital and antorbital.
Tube-bearing circumorbitals, more properly infraorbitals (see Weitzman,
1962:28 and footnote 7), one and two are present in the holotype of E. junki
and are moderately well ossified on the right side. They are much less
ossified on the left side where they may have been damaged by a greater
exposure to acid formalin. These two bones are well ossified and each bears
a canal along its entire length in the stained side of the specimen of E. junki
from the Rio Cuieiras. Infraorbital three and the subsequent infraorbitals
four, five, and six appear absent or at least not ossified in the holotype. The
bone, if present, did not take up alizarin stain in the specimen from Rio
Cuieiras. In all specimens the skin in the area of infraorbital three is thicker
and tougher than the skin of other parts of the orbit. At least the specimen
of E. junki from the Rio Cuieiras was exposed to formalin for about eight
months and may therefore have had the bone of this normally thin bony
plate dissolved away. No bony canal bone was found in the region of the
infraorbitals three to five. In summary, a slender, tubeless antorbital and
tube-bearing infraorbitals one and two are present. The ‘‘great suborbital’’
of Eigenmann (1912) (infraorbital three) and all the remaining more posterior

VOLUME 93, NUMBER 4 907

orbitals are either greatly reduced in ossification or absent. The same con-
dition occurs in cleared and stained specimens of E. geryi and E. pulcher
examined for this report.

Another reductive character used by Géry (1971:163) to separate Geisleria
and Elachocharax at the subfamily level was the absence of a cranial fon-
tanel in the former and its presence in the latter. The fontanel is not actually
absent in E. junki but occurs as a very small opening where the posterior
medial borders of the parietals of each side meet the supraocciptial and
surround the anterior portion of the supraoccipital fossa. In fishes with this
anatomical arrangement a very slender needle can be passed between the
parietals and supraoccipital into the brain cavity. In the Characidiinae the
posterior cranial fontanel (that part posterior to the epiphyseal bar) is all
that remains. Usually this is represented by a separation of the parietal
bones from each other along the midline. The resulting fontanel is contin-
uous with and part of the median fossa along the dorsal surface of the
supraoccipital spine. Apparently in many species of Characidium the pa-
rietals are completely separated but in some species (and in some species
of Jobertina) they contact each other anteriorly, restricting the fontanel to
a more posterior postion and reducing it in relative size. Near closure of the
fontanel is not restricted to pygmy members of the Characidiinae. There is
an unidentified species of robust, large (to at least 64 mm SL) relatively high
altitude Characidium from the Rio Uruyen on the south slopes of Mt. Au-
yantepuy, State of Bolivar, Venezuela (USNM 219847), which has the pa-
rietals completely jointed along their medial borders as in E. junki, leaving
only the area immediately bordered by the posterior part of the parietals
and the supraoccipital as a fontanel. Elachocharax pulcher and E. geryi
have reduced fontanels, the parietals usually being closely jointed anteriorly
and not posteriorly. The larger specimens of E. pulcher examined by Weitz-
man and Kanazawa (1978) usually had the fontanel more restricted than the
smaller specimens and sometimes it was almost as restricted in E. pulcher
as in E. junki. The extent of the restriction of the fontanel thus seems to
vary somewhat with the size of the specimens examined in the instance of
Elachocharax and a small fontanel is subject to repetitive independent ap-
pearance in the Characidiinae.

Géry (1971:163) separated his Elachocharacinae and Geisleriinae on the
one hand from his Characidiinae on the other by the former two having a
very small size, a long dorsal fin of at least a total of 17 rays and poorly
developed pectoral fins. He also noted that the species of these genera
apparently live a relatively ‘‘secondary’’ specialized existence in a quiet
water habitat whereas Characidiinae live in stream habitats requiring an
active existence to resist current. The habitat differentiation is true when
compared to the larger species of Characidium. However, some other pyg-
my species such as Jobertina eleotrioides Géry and Klausewitzia aphanes

908 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

appear to live in similar, “‘quiet’’ habitats. One of us (Weitzman) has re-
covered specimens (USNM 221046) of miniature Jobertina 13.0 to 14.5 mm
standard length from a mass of algae collected by Bruce Collette in a very
slow moving stream, Rio Marituba, Para, Brazil. Meinken (1969) erroneous-
ly identified Klausewitzia aphanes as Characidium voladorita Schultz while
describing its swimming habits, indicating that it is a quiet water fish, and
Weitzman and Kanazawa (1977) report that aquarium specimens of K.
aphanes appear adapted to quiet waters. The habitat reported for K.
aphanes by Weitzman and Kanazawa is a slow moving blackwater stream.

These two fishes, J. eleotrioides and K. aphanes, also have pectoral fins
somewhat elongate in form and with a reduced number of fin rays, like that
of the three species of Elachocharax recognized here. There is usually a
total of 11 fin rays in the pectoral fin of species of Characidium. Some
species will have one or two less. This is the same count found in most
‘‘tetragonopterin’’ characids and may therefore be primitive for the Char-
acidiinae. In Characidium the number of unbranched anterior rays varies
from one to four or rarely five. Most *‘tetragonopterin’’ characids only have
on unbranched anterior pectoral fin ray. The propensity to have two or more
in the Characidiinae appears to be a synapomorphy for the Characidiinae.
Elachocharax pulcher and E. junki usually have eight pectoral-fin rays and
usually all the rays are undivided, but occasionally one, two or three of the
middle rays may be divided, especially in E. junki. We found one specimen
of E. pulcher with seven undivided rays on one side and nine undivided
rays on the other. Weitzman and Kanazawa (1978:170) report six to nine
rays in this species with very few specimens with any branched rays. Weitz-
man and Kanazawa (1978:177) report six to eight unbranched rays in E.
geryi. Two specimens of a Jobertina, apparently related to Jobertina eleo-
trioides and 13.0 mm in standard length from Rio Marituba, Para, Brazil, had
eight pectoral-fin rays, the anterior four undivided, the posterior four divid-
ed. Two paratypes of Jobertina eleotrioides had nine pectoral-fin rays, the
anterior four undivided, the posterior five divided. Klausewitzia aphanes
has a total of eight, rarely nine, unbranched pectoral-fin rays. Furthermore,
there is a series of 10 or more undescribed species of miniature to small
members of the Characidiinae, mostly from the Amazon basin, which have
10 or fewer pectoral-fin rays. As the genera are currently defined, these
species ‘‘fit’’ Characidium or Jobertina. It appears that a reduced number
of pectoral-fin rays and a trend for all of them to be undivided is common
in small to miniature Characidiinae regardless of their apparent phylogenetic
origin. This character should not be used to define genera and subfamilies
within this group, at least not unless further investigation substantiates a
hypothesis that all miniature Characidiinae belong to a single monophyletic
group.

Thus, of all reasons that Géry (1971) separated the Elachocharacinae and

VOLUME 93, NUMBER 4 909

the Geisleriinae from the Characidiinae, the only characters that we now
can find to serve as synapomorphies for their members are the two char-
acters used above and in Fig. 7 to differentiate Elachocharax as here defined
from the other genera of the Characidiinae.

Géry (1971:163) separated his Elachocharacinae and Geisleriinae on the
basis of the following characters: teeth triscupid, uniserial in the premaxil-
lary, pterygoid teeth present, maxillary teeth apparently absent, infraorbit-
als reduced and with a small posterior cranial fontanel for Elachocharacinae,
and teeth conic, biserial in the premaxillary, pterygoid apparently absent,
maxillary teeth apparently present, infraorbitals absent and no cranial fon-
tanel for the Geisleriinae. Of all these differences or apparent differences
only that of the tooth cusps remains and all the species included by these
two nominal subfamilies have uniserial premaxillary teeth, pterygoid teeth
present, maxillary teeth absent, infraorbitals 1 and 2 present but the posterior
infraorbitals reduced or absent, and with a small to very small cranial fon-
tanel.

As described above, the differences in tooth cusps (the reduction to conic
teeth) has been found to be a synapomorphy uniting E. junki and E. geryi.
Elachocharax pulcher has the less derived state of tricuspid cusped teeth.
This is correlated with two other synapomorphies present in E. junki and
E. geryi, a loss of the adipose fin and a branchiostegal ray number reduced
from five to four. To recognize these two species as belonging to a subfamily
or a genus separate from E. pulcher on the basis of these three synapo-
morphies seems to us poor systematic practice, especially in view of the
fact that these characters could be independent derivations since they are
apparently independently derived in other sections of the Characidiinae.
Elachocharax is then a pygmy genus of the Characidiinae with three known
species, of which E. pulcher is the least derived and of which E. junki and
E. geryi apparently are related and more derived; E. geryi appears the most
derived. Table 2 and Fig. 7 give a summary of the characters and suggested
relationships.

There is not enough information available to attempt a productive analysis
of the relationships of Elachocharax to the other genera of the Characidi-
inae. There is a basic color pattern shared by the species here assigned to
Elachocharax which does not appear in quite the same form in many other
species of the Characidiinae. The pattern of a stripe of dark pigment along
the length of the dorsal fin, near its base or at least nearer than half the
length of the fin rays toward the base, is shared with Klausewitzia aphanes,
another miniature species (up to 16.5 mm SL) in the Characidiinae, and in
Characidium surumnense Steindachner, Characidium boavistae Steindach-
ner, and a few other species of Characidium. Klausewitzia ritae Gery
(1965), also of small size ( at least 25.0 mm SL), is stated to have two very
narrow bands on the dorsal fin. Both species of Klausewitzia have a much

910 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

shorter dorsal fin and have maxillary teeth. Both of these characters are
plesiomorphic to the state found in Elachocharax. The latter character,
the presence of maxillary teeth, is a presumably plesiomorphic character
otherwise found only in the two species of the otherwise specialized genus
Ammocryptocharax. Klausewitzia ritae is probably less specialized than K.
aphanes, being of larger size and having tricuspid as well as unicuspid teeth
(teeth are all unicuspid in K. aphanes), in always having an adipose fin
(present or absent in K. aphanes), and in having a complete lateral line
(much reduced in length in K. aphanes). Klausewitzia ritae needs further
study in order to outline more clearly its possible reductive apomorphies in
orbital bones and other characters, but of the known species of relatively
short-bodied small or pygmy Characidiinae this species appears to be the
most primitive.

Some species of Jobertina have an increase in the number of dorsal-fin
rays, to 14, and some, Jobertina interruptua Pellegrin and Jobertina theag-
eri Travassos, have the longitudinal dorsal-fin stripe as found in Elacho-
charax. Jobertina and Elachocharax (along with Klausewitizia aphanes)
share a reductive character, a short lateral line of 10 or fewer perforated
lateral-line scales. As with most reductive characters, this one is hard to
evaluate because of its repetitive independent appearance in miniature fish-
es. The genus Jobertina, for example, separated from Characidium on the
basis of a lateral line of few perforated scales and a tendency to slightly
increase the number of dorsal-fin rays in some species, may be polyphyletic
and its members may have arisen more than once from a Characidium-like
ancestor. The relationships of Elachocharax are currently indefinable but
may be with certain of the pygmy species of Klausewitzia or Jobertina.

Acknowledgments

Heraldo A. Britski and Naério A. Menezes, Museu de Zoologia da Un-
versidade de Sao Paulo (MZUSP) aided the senior author’s search of their
collections and loaned specimens for study. Herbert Axelrod, Rolf Geisler
and Wolfgang Junk provided specimens through the junior author. Paulo
Vanzolini (MZUSP) allowed the senior author to survey the collections in
his charge, and the Fundacgao de Amparo a Pesquisa do Estado Sao Paulo
supported the collection of specimens used here by the Expedicao Perma-
nente da Amazonia under Vanzolini’s direction. Financial support for ex-
amining the collections at the MZUSP was provided by the Smithsonian
Institution Amazonian Ecosystems Research Program directed by Clifford
Evans and by a grant from the National Geographic Society. Assistance in
sorting and examining specimens in the collections at MZUSP was provided
by Marilyn Weitzman, William L. Fink and Sara V. Fink. The latter three
persons as well as Richard Vari, James E. Bohlke and Antonio Machado

VOLUME 93, NUMBER 4 911

read the manuscript and offered useful suggestions. Sara V. Fink prepared
Figs. 5 and 6, and Marilyn Weitzman prepared Figs. | and 7. Robert H.
Kanazawa provided technical assistance. A. W. Crompton, Karel F. Liem,
and William L. Fink provided space and facilities for research on this project
while the senior author was a Visiting Alexander Agassiz Curator at the
Museum of Comparative Zoology, Harvard University. Naércio Menezes
translated the summary into Portuguese.

Summary

Based on the characters analyzed here, Elachocharax is a genus of the
Characidiinae of uncertain relationships. Its species have a relatively short
body and a long-based dorsal fin (17—22 rays) with its posterior termination
over or posterior to the anal-fin origin. All the known species are miniature,
25 mm or less in standard length. The genus may be most closely related to
Klausewitzia or to certain members of Jobertina. Elachocharax consists of
three species further characterized by reductions in the orbital bones and
other features of the lateral sensory system such as the lateral-line scales
of the body. Elachocharax pulcher is apparently the least derived and has
tricuspid, bicuspid, and unicuspid teeth, usually five branchiostegal rays,
and an adipose fin; E. geryi and E. junki appear more advanced or derived
with regard to these characters and have unicuspid teeth, four branchioste-
gal rays, and no adipose fin. Elachocharax junki is the most specialized
with regard to the origin of the anal fin which is noticeably in advance of
the posterior termination of the dorsal-fin base. This species also has the
largest number of body bars, about 17, presumably an advanced condition.
Elachocharax pulcher may be advanced in a different direction in having
a decrease in the number of vertical body bars and a widening of those bars
when compared to species in other genera of the Characidiinae. Elacho-
charax geryi may be the least advanced in some of its color pattern, having
11 to 12 vertical body bars, but the most advanced in having many narrow,
vertical, caudal-fin bars.

The nominal subfamilies Elachocharacinae and Geisleriinae are rejected,
the type species of the latter being considered a species of Elachocharax
and the Elachocharacinae being recognizable as a member of the Characi-
diinae at the generic level.

Resumo

O exame do holotipo e de cinco novos exemplares da espécie nominal
Geisleria junki, juntamente com uma reconsideracgao sobre as espécies
Elachocharax pulcher e Elachocharax geryi, permitiu uma reavalicao das
subfamilias nominais Geisleriinae e Elachocharacinae, bem como do género
nominal Geisleria, da subordem Characoidei. Ficou evidenciado que alguns

912 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dos caracteres usados por Géry (1971) para separar Geisleria de Elacho-
charax e Geisleriinae de Elachocharacinae nao apresentam diferengas
nestes ‘‘taxa’’ nominais. Todas as espécies anteriormente incluidas nestes
‘‘taxa’’ tem uma unica série de dentes premaxilares, dentes no ectopteri-
goide, maxilar sem dentes, ossos orbitais com configuracao idéntica ou mui-
to semelhante e fontanelas cranianas reduzidas mas distintas. As trés es-
pécies aqui consideradas como pertencendo a Elachocharax, E. pulcher,
E. junki e E. geryi, tem em comum os seguintes caracteres que as separam
de outros géneros de Characidiinae reconhecidos neste trabalho: nadadeira
dorsal alongada, com 17 a 22 raios, sua base terminando acima da origem
da nadadeira anal, ou ultrapassando-a um pouco. Nos outros Characidiinae
ha 11 a 14 raios na nadadeira dorsal e a origem da nadadeira anal é€ muito
posterior a nadadeira dorsal.

Um carater usado por Géry (1971) para separar Geisleria e Geisleriinae
de Elachocharax de Elachocharacinae—a presenga de dentes apenas uni-
cuspides nos primeiros e tricuspides nos segundos—tem sido utilizado para
separar familias, tribos e géneros em outros grupos de Characoidei. Weitz-
man and S. V. Fink (no prelo) demonstraram, por exemplo, que em Xen-
urobryconini 0 carater e util apenas para separar as espécies de um género,
nao tendo portanto utilidade ao nivel de tribo. Em Characidiinae, dentes
unicuspides podem ter aparecido independentemente em espécies de cinco
géneros: Characidium, Jobertina, Ammocryptocharax, Klausewitzia, e
agora Elachocharax.

Originalmente Geisleria foi em parte definido com base na auséncia de
uma nadadeira adiposa e em caracteres que consideramos no presente tra-
balho como nao pertinentes. Além de nao possuir nadadeira adiposa, Geis-
leria pode agora ser definido com base na existéncia de dentes unicuspides
e€ presenca de quatro ao invés de cinco raios branquiostégios. Mostramos
que estes caracteres apomorficos surgiram independentemente em outros
géeneros de Characidiinae. Parece-nos desnecessario continuar a reconhecer
Geisleria e Elachocharax como géneros distintos com base na preseng¢a de
caracteres de reducao que podem ter surgido independentemente.

As subfamilias nominais. Elachocharacinae e Geisleriinae e o género
Geisleria sao rejeitados, a espécie-tipo deste ultimo é€ considerada uma
especie de Elachocharax e Elachocharacinae considerado como membro de
Characidiinae ao nivel de género.

Literature Cited

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1917. The American Characidae.—Mem. Mus. Comp. Zool. Harvard College

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Fink, W. L., and S. H. Weitzman. 1974. The so-called cheirodontin fishes of Central America
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Géry, J. 1965. Poissons characoides sud-Americains du Senckenberg-Museum, II. Characidae

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Aquarien- und Terrarien-Zeitschrift. (DATZ), Jahrgang 22, (4):100-102.

Myers, G. S. 1927. Description of new South American fresh-water fishes collected by Dr.

Carl Ternetz.—Bull. Mus. Comp. Zool., Harvard College 68(3): 107-135.

1958. Trends in the evolution of Teleostean fishes.—Stanford Ichthyological Bull.

7(3):27-30.

, and J. E. Bohlke. 1956. The Xenurobryconini, a group of minute South American

characid fishes with teeth outside the mouth.—Stanford Ichthyological Bull. 7(2):6—-12.

Weitzman, S. H. 1962. The osteology of Brycon meeki, a generalized characid fish, with an

osteological definition of the family.—Stanford Ichthyological Bull. 8(1):1-77.

. 1964. Osteology and relationships of South American characid fishes of subfamilies

Lebiasininae and Erythrininae with special reference to subtribe Nannostomina.—Proc.

U.S. Nat. Mus. 116(3499):127-170.

, and S. V. Fink. In prep. A revision of the glandulocaudin Xenurobryconini with a

description of a new species of Xenurobrycon from Ecuador and Peru. (Teleostei:

Characidae).—Smithsonian Contributions to Zoology.

, and R. H. Kanazawa. 1976. Ammocryptocharax elegans, a new genus and species of

riffle-inhabiting characoid fish (Teleostei:Characidae) from South America.—Proc. Biol.

Soc. Wash. 89(26):325-346.

. 1977. A new species of pygmy characoid fish from the Rio Negro and Rio Amazonas,

South America (Teleostei:Characidae).—Proc. Biol. Soc. Wash. 90(1): 149-160.

. 1978. The South American fish genus Elachocharax Myers with a description of a

new species (Teleostei:Characidae).—Proc. Biol. Soc. Wash. 91(1):158-183.

(SHW) Department of Vertebrate Zoology, National Museum of Natural
History, Washington, D.C. 20560; (JG) Argentonesse, Castels, F-24220,
Saint-Cyprien, France.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 914-919

NEW RECORDS OF STEREOMASTIS SCULPTA
PACIFICA (FAXON) (DECAPODA: POLYCHELIDAE)
IN THE EASTERN PACIFIC OCEAN

Mary K. Wicksten

Abstract.—The range of Stereomastis sculpta pacifica is extended south
from Peru to Chile. Adults are reported for the first time from southern
California. The larvae, taken in midwater trawls, reach a length of 87 mm.
All but two of the adults measured over 70 mm. The large size of the larvae
relative to the adults suggests that much of the life cycle is spent in midwater
zones.

The deep-sea lobster Stereomastis sculpta pacifica is one of the most
striking decapods occurring at bathyal depths in the eastern Pacific Ocean.
The bright scarlet larvae have inflated, globose carapaces unlike those of
other midwater decapods in the area. The adults are recognizable by their
long chelipeds, flattened, hirsute carapace, and eyes without pigment.

In 1978, Paul Gregory and other biologists of the California Department
of Fish and Game brought two specimens of these lobsters to the Allan
Hancock Foundation for identification. Examination of the animals sug-
gested that other polychelids in local collections might belong to the same
species. I found more specimens in the midwater collections of the Allan
Hancock Foundation and at Scripps Institution of Oceanography. Adults
were taken in sablefish traps, otter trawls, and beam trawls. Larvae were
collected in Isaacs-Kidd midwater trawls.

New records of S. sculpta pacifica extend its range south from Peru to
Chile, and north from off San Clemente Island to off Point Conception,
California. This paper presents new records, discusses the synonymy of the
species, and comments on its life history.

Stereomastis sculpta pacifica (Faxon)
Fig. 1

Polycheles sculptus pacificus Faxon, 1893:196—-197; 1895:122—-123, pl. C, fig.
le ta:

Eryonicus caecus?—Faxon, 1893:197-198; 1895:110-111, pl. B, fig. 2; pl.
We) Vee APA E

Eryoneicus Agassizi Bouvier, 1915:2.

Stereomastis sculpta pacifica.—de Man, 1916:5, 8.—Firth and Pequegnat,
ISA NG) IZ

VOLUME 93, NUMBER 4

jp ===}
lcm

Fig. 1. Stereomastis sculpta pacifica. Upper: adult female, total length 102.5 mm. Off Baja
California, Mexico (31°59.6’N, 118°47.4’W), 1,100 m, boat Calafia. Lower: larva, total length
87.0 mm. 34.3 mi. from Morro Hermosa, Baja California, 3,720—3,785 m, Velero IV sta. 11522.

916 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Eryoniscus agassizi.—Schmitt, 1921:105-107, pl. 15, figs. 1-2.

Polycheles sculptus.—del Solar, 1972:11.

Not Pentacheles Agassizi.m—A. Milne-Edwards, 1880:65—66 (=Polycheles
typhlops Heller).

Previous records. —U.S.A.: 2.9 miles, 72°S of east point, Northwest Har-
bor, San Clemente Island, California (approximately 33°0’N, 118°40’W),
1,207-1,300 m, 9-foot Tanner beam trawl, 9 April 1904, Albatross station
4405. MEXICO: Near Las Tres Marias Islands (21°15’0"N, 106°23’0’W),
1,248 m, gray sand, black specks, Globigerina ooze, 18 April 1891, Alba-
tross sta. 3424. Off Acapulco (16°33’0"N, 99°52’30"W), 1,218 m, brown sand,
black specks, 11 April 1891, Albatross sta. 3419. PANAMA: Gulf of Panama
(7°21'0"N, 79°35’0"W), 943 m, dark green mud, 10 March 1891, Albatross
sta. 3394. Gulf of Panama (7°21'0"N, 79°2’0"”W), 3,382 m, 8 March 1891,
Albatross sta. 3383. Gulf of Panama (7°15'0’N, 79°36’0"W), 1,883 m, green
mud, 10 March 1891, Albatross sta. 3393. Off Mariato Point (7°6'15’N,
80°34'0”W), 1,283 m, green mud, 23 February 1891, tow net, Albatross sta.
3353. Gulf of Panama (7°6'0"N, 79°48’0"W), 0-738 m, tow net, 9 March 1891,
Albatross sta. 3388. Gulf of Panama (7°5'30"N, 79°40'0”W), 2,345 m, hard
bottom, 10 March 1891, Albatross sta. 3392. COLOMBIA: Off Malpelo
Island (3°56’0’N, 81°40'15”W), 1,410 m, 5S March 1891, Albatross sta. 3377.
Cocos Island—Malpelo Island (2°34’0"N, 82°29’0"W), 2,217 m, surface tow
net, 4 March 1891, Albatross sta. 3375. PERU: North of Callao (10°45’S,
78°36'W), 870 m, 1970-1972, SNP-/ transect 7105. Off southern Peru
(18°23’S, 71°13’W), 1,100 m, 1970-1972, SNP-/ transect 7201.

Note: Faxon, 1895, as Eryonicus caecus? and Polycheles sculptus paci-
ficus; Schmitt, 1921, as Eryoniscus agassizi; del Solar, 1972, as Polycheles
sculptus.

The range given by Firth and Pequegnat (1971), ‘‘off Panama and the
Galapagos Islands, 934 to 2,323 m,”’’ is incorrect.

Material examined.—U.S.A.: SW of Point Conception, California
(34°30’N, 120°35’W), 923-1,108 m, sablefish trap, 10 May 1979, boat Arista,
1 adult. 11 miles, 235°T to W end, Santa Catalina Island (33°26'0'N,
118°51'30"W to 33°21'42”W), 1,302 m, 23 June 1964, Velero IV sta. 9852, 1
larva. 13.2 miles, 188°T from W end, Santa Catalina Island (33°20'25’N,
118°47'20"W to 33°10'15"N, 118°31'15”W), 1,228-1,278 m, 25 July 1964, Ve-
lero IV sta. 9876, 1 larva. 24.5 miles from SE point, San Nicolas Island
(33°8'20"N, 119°12'35"W to 32°50’15”W, 118°53'35”W, 1,643 m, 14 May 1964,
Velero IV sta. 9661, 1 larva. 26.5 miles, 120°T from Aero Light, San Nicolas
Island (32°52'30"N, 118°59’20"W to 33°05'0"N, 119°11'0”W), 1,588—-1,735 m,
26 February 1969, Velero IV sta. 12726, 1 larva. CHILE: Off Arica
(18°40.5’S, 70°36.0'W), 768-968 m, 25-foot otter trawl, 7 May 1972, Thomas
Washington sta. MV72-II-27, 1 adult. Off Arica (18°42’S, 70°37’), 1,097-

VOLUME 93, NUMBER 4 917

1,152 m, 7 May 1972, 25-foot otter trawl, Thomas Washington sta. MV72-
II-26, 1 adult. Off Valparaiso (30°46’S, 81°31’W), 3,000 m, 4 March 1969,
Piquero trawl 2, 1 larva. Also an additional 32 adults from 20 stations; Costa
Rica, Mexico, and southern California, 750—1,875 m; and 86 larvae from 63
stations; Costa Rica, Mexico, and southern California.

Total lengths of adults.—41.3—102.5 mm.

Total lengths of larvae.—16.9-87.0 mm.

Size distribution of larvae.—10—20 mm (n = 9); 20-30 mm (n = 7); 30-40
mm (n = 35); 40-50 mm (n = 10); 50-60 mm (n = 9); 60 mm or greater (n =
19). Undetermined (n = 2).

Depth distribution of larvae.—1,846 m or less (n = 39); 1,846-2,748 m
(n = 15); 2,748-3,692 m (n = 28); 3,692 m or greater (n = 3); not recorded
(G74):

Remarks.—Larval stages of polychelid lobsters have been called species
of Eryoneicus or Eryoniscus. Bouvier (1915, 1917) noted that the eastern
Pacific species, which he named Eryoneicus Agassizi, was different in its
spination from Eryonicus caecus, taken by the Challenger expedition off
the Canary Islands (Bate, 1888). Although he saw the similarity between S.
sculpta pacifica and his Eryoneicus Agassizi, he considered the two to be-
long to distinct genera. Balss (1925) finally established that the large, open-
water animals actually were larval polychelids. The generic names Ery-
oneicus and Eryonicus have been placed on the Official Index of Rejected
Names and the generic name Stereomastis validated under the plenary pow-
ers (ICZN Opinion 702, 1964). Faxon (1895) considered the genus Stereo-
mastis Bate 1888 as being a synonym of Polycheles Heller 1862. De Man
(1916), however, thought that they were distinct based on their epipodites
and the spination of the carapace.

Studying the life history of an animal living at bathyal depths is difficult.
The eggs of the female taken off Baja California (Fig. 1) measure approxi-
mately 1 mm in diameter. The smallest larval stages have not been collected
by the Isaacs-Kidd midwater trawl, which has a liner mesh of 3 mm. The
size distribution of the larvae suggests that there may be at least five larger
larval stages before settling.

The larvae were collected with myctophid fishes, cephalopods, sergestid
shrimps, and other nektonic animals. Like these animals, larval polychelids
may migrate vertically. Remains of a bony fish were found in the oral field
of one larva.

All but two of the adults measured over 70 mm in total length. The other
two measured 41.3 and 47.6 mm each. Nineteen of the larvae were 60 mm
or greater in length. The small difference between most of the adults and
the largest larvae suggests that maturity is reached soon after settling, per-
haps after only one or two molts. The two very small adults indicate that
not all the lobsters settle at the same size.

918 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Little is known of the habits of adult S. sculpta pacifica. Faxon (1895)
reported them from soft bottoms. Four specimens were caught on or inside
traps for sablefish (Anoplopoma fimbria [Pallas]) baited with chopped fish.
The lobsters may have been trying to scavenge on the fish. Wenner (1979)
found foraminiferans and parts of crustaceans, fish and polychaetes in the
stomachs of S. sculpta sculpta Smith, the Atlantic subspecies.

Wenner and Boesch (1979) found that Stereomastis sculpta sculpta oc-
curred at 575—2,130 m in the western Atlantic Ocean. Most individuals were
taken at 600-—1,200 m, at 3.4-10.5°C. This depth range is similar to that of
S. sculpta pacifica. At 1,300—2,400 m in the Atlantic, they reported Ster-
eomastis nana (Smith) to be more abundant than S. sculpta. In the eastern
Pacific, S. nana has been reported off Colombia and the Galapagos Islands
(Faxon, 1895). It also has been taken off Chile (32°51'S, 72°8’W, 2,580 m,
beam trawl, 19 December 1965, Anton Bruun Cruise 12, one specimen,
Scripps Institution of Oceanography, unpubl. data). At depths greater than
2,400 m in the eastern Pacific, the lobsters Willemoesia pacifica Sund, Wil-
lemoesia challengeri Sund, and Willemoesia inornata Faxon have been col-
lected.

The life history of S. sculpta pacifica is unusual for a decapod in having
such large open-water larvae relative to the adults. Perhaps it is not sur-
prising that earlier writers did not immediately recognize tie midwater forms
as larvae. Much of the lifespan may be spent in midwater zones. The catches
of nektonic shrimp, fishes, cephalopods, and other invertebrates in the
trawls with the larval polychelids suggest that food may be more abundant
in the midwater zones than on the bottom. Spending a long time in the open
water may aid these lobsters in avoiding competition for food and/or pre-
dation on the bottom.

Literature Cited

Balss, H. 1925. Macrura der Deutschen Tiefsee-Expedition. 1. Palinura, Astacura und Tha-
lassinidea.—Deutsche Tiefsee-Expedition 1898-1899 20: 189-216.

Bate, C. S. 1888. Report on the Crustacea Macrura collected by the H.M.S. Challenger during
the year 1873-76.—Rep. Voy. Challenger, Zool. 24:1-942.

Bouvier, M. E.-L. 1915. Observations nouvelles sur le genre Eryoneicus.—Bull. Inst. Ocean-

ogr. Monaco 309:9 pp.

. 1917. Résultats des campagnes scientifiques accomplies sur son yacht par Albert Ier.

Fasc. L. Crustacés decapodes (Macroures marcheurs) provenant des campagnes des

yachts Hirondelle et Princess-Alice (1885-1915). 140 pp.

del Solar, E. M. 1972. Addenda al catalogo de Crustaceos del Peru.—Instituto del Mar del
Peru Informe no. 38, 21 pp.

de Man, J. G. 1916. The Decapoda of the Siboga Expedition. Part III. Families Eryonidae,
Palinuridae, Syllaridae, and Nephropsidae.—Siboga Expeditie 39a’, 122 pp.

Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America
to the Galapagos, to the west coast of Mexico, and in the Gulf of California . . . by the

VOLUME 93, NUMBER 4 919

U.S. Fish Commission steamer ‘‘Albatross’’ during 1891 . . . VI. Preliminary descrip-
tions of new species of Crustacea.—Bull. Mus. Comp. Zool., Harvard Univ. 24(7):149-
220.

. 1895. Reports on exploration off the west coast of Mexico, Central and South Amer-

ica, and off the Galapagos Islands . . . by the U.S. Fish Commission steamer ‘‘ Alba-

tross’’ during 1891 . . . XV. The stalk-eyed Crustacea.—Mem. Mus. Comp. Zool., Har-

vard Univ. 18:1—292.

Firth, R. W., Jr., and W. E. Pequegnat. 1971. Deep-sea lobsters of the families Polychelidae
and Nephorpsidae (Crustacea, Decapoda) in the Gulf of Mexico and Caribbean Sea.—
Texas A&M Univ. Dept. Oceanogr. Ref. 71-11T, 106 pp.

Heller, C. 1862. Beitrage zur naheren Kenntnis der Macrouren.—Sitzungsber. Kais. Akad.
Wiss. Wien 45(1):389-426.

International Commission on Zoological Nomenclature. 1964. Stereomastis Bate, 1888 (Crus-
tacea, Decapoda): Validated under the plenary powers.—Bull. Zool. Nomen. 21(2):111-
U2,

Milne-Edwards, A. 1880. Reports on the results of dredging by the U.S. Coast Survey steamer
‘Blake.’ VIII. Etudes preliminaires sur les Crustacés.—Bull. Mus. Comp. Zool., Har-
vard Univ. 8(1):1-68.

Schmitt, W. L. 1921. The marine decapod Crustacea of California.—Univ. Calif. Publ. Zool.
23: 1-470.

Wenner, E. L. 1979. Some aspects of the biology of deep-sea lobsters of the family Poly-

chelidae (Crustacea, Decapoda) from the western North Atlantic. U.S. Nat. Mar. Fish.

Serv. Fish. Bull. 77(2):435—444.

and D. F. Boesch. 1979. Distribution patterns of epibenthic decapod Crustacea along

the shelf-slope coenocline, middle Atlantic bight, U.S.A.—Bull. Biol. Soc. Wash. 3:106—

1335

Allan Hancock Foundation, University Park, Los Angeles, California
90007.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 920-927

THE IDENTITY OF THE FROG EUPSOPHUS
VANZOLINIT FROM RAMADILLAS, NAHUELBUTA
RANGE, SOUTHERN CHILE

J. R. Formas

Abstract.—The frog Eupsophus vanzolinii is shown to be a species of the
genus Alsodes. The species is redescribed on the basis of 38 fixed speci-
mens, and natural history notes on the species are presented.

Donoso-Barros (1974) described a new species of leptodactylid frog, Eup-
sophus vanzolinii, on the basis of specimens from Ramadillas (73°15’S,
37°15'W), at 100 m elevation on the western slope of the Nahuelbuta Range,
Arauco Province, Chile. Little comment has been made concerning the sys-
tematic status of this species, although Lynch (1978) tranferred E. vanzolinii
to the genus Alsodes on the basis of specimens collected by W. E. Duellman
in Cabreria (73°7'S, 50’W), Parque Nacional Nahuelbuta, 1,030 m elevation,
Malleco Province, Chile. The air line distance between these points is 65 km.

I analyzed the frogs collected by Duellman in Cabreria, which Lynch
(1978) considered Alsodes vanzolinii, and the specimens found by Donoso-
Barros in Ramadillas, and concluded that the frogs of the two places are
different species.

The frogs from Cabreria were previously identified as Eupsophus coppin-
geri (=Alsodes monticola) (Iturra and Veloso, 1975), and as E. vanzolinii
(Duellman and Veloso, 1978). Grandison (1961) used the name Eupsophus
coppingeri (=Alsodes monticola) for the following frogs from Cabreria: Chi-
cago Natural History Museum 44210-44214, 44216-44219, Cabreria, 800 m,
Nahuelbuta, Angol; 40136, Nahuelbuta, Angol. According to my colleague
Alberto Veloso, the frogs of Cabreria are a new species which is now being
described by him.

As a result of the study of the specimens collected by Donoso-Barros in
Ramadillas I conclude that Eupsophus vanzolinii Donoso-Barros, 1974,
must be transferred to the genus Alsodes because the males have spiny
excrecences on the thumb and chest, the distal edge of the xiphisternum is
notched, the nasal bones are large, the cultriform process is long and the
anterior end extends between the palatines.

Redescription of Alsodes vanzolinii
Fig. 1

Diagnosis.—Alsodes vanzolinii is a moderate sized species that can be
distinguished from it congeners by the following combination of characters:

VOLUME 93, NUMBER 4 921

:
ytd
te
Pe
*
Se
“ek

a - 2
wh ‘
a.
: |
om
¥ |

Naat

$52
cs

Fig. 1. Holotype of Alsodes vanzolinii (Donoso-Barros), MUZUC 12063, female.

1) rudiment of web between the fifth and fourth toes; 2) thinner limbs; 3)
notched tongue covered by minute papillae; 4) yellow triangle on the head.

Description (based on 38 fixed specimens).—Head slightly wider than
long. Snout rounded and sometimes truncate in lateral and dorsal profile;
canthus rostralis well defined; loreal region flat, sloping abruptly to lip; lips
not flared. Nostrils lateral, closer to tip of snout than to orbit; length of eye
greater than the distance between eye and nostril; interorbital distance great-
er than internarial distance. Typanum absent. Well developed glandular fold
from behind eye to insertion of arm. Large cordiform tongue, notched be-
hind, covered by minute transparent papillae, posterior one-third free. Rel-
atively small, round choanae; dentigerous process of prevomers lying me-
dian to and slightly below choanae, separated medially, transverse or
slightly oblique, each process bearing 4—6 teeth. Forelimbs thin; first finger
equal in length to second; third finger much longer than fourth; digital length
in decreasing order 3-4-2-1. Palmar webbing absent (Fig. 2a); tips of fingers
slightly expanded. Large elliptical and prominent inner palmar tubercle;
outer palmar tubercle ovoid and smaller than inner; subarticular tubercles
moderate sized, conical, and simple; supernumerary palmar tubercles pres-
ent. Hind limbs slender, tibiotarsal articulation reaching to middle of eye.
Toes long (Fig. 2b), slender and fringed; third and fifth toes equal in length;
toes in decreasing order of length 4-(3,5)-2-1. Outer metatarsal tubercle
ovoid and flattened, inner metatarsal tubercle small and conical. Subartic-

327 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.5 cm

05 cM

Fig. 2. Hand (a), foot (b), and pectoral girdle (c) of Alsodes vanzolinii.

VOLUME 93, NUMBER 4 923

Table 1.—Measurements (mm) and proportions of Alsodes vanzolinii.

15 males 13 females
Character Range x +SD Range 52 se SD)
Snout-vent length (SVL) 3614— 52 Oe Ee. AD — 54: Ali illite sses
Tibia length/SVL .50-.59 5 Sec= 02 .49_-.55 soll se 01
Foot length/SVL .48-.61 .54 + .03 .50-.56 Si Ol
Head length/SVL .35—.42 .38 + .01 .33-.41 .36 + .02
Head width/SVL .38-.43 .40 + .01 31 =.42 39 + .01
Interorbital distance/head width .22—.28 .24 + .02 .22—.28 525923) 04
Eye-nostril/head length .22—.32 .26 + .02 .24—.31 Die OZ

ular tubercles large and conical; minute supernumerary tubercles present.
Tarsal fold present and reduced. Rudiment of web between fifth and fourth
toes. Anal opening oriented transversely, at dorsal level of thighs, and in-
conspicuously ornamented.

Skin smooth, minute tubercles on flanks, dorsum and head; posteroven-
tral areas of thighs with tuberculate skin. Dorsum brown with irregular
whitish spots; head with a yellow triangle. Venter usually whitish, but some
specimens show a dark background with minute irregular spots; juveniles
have black mottled belly with irregular white spots. Throat whitish, loreal
region dark. Dark brown glandular fold behind the eyes. Some specimens
have a white vertebral line that reaches to the middle of the back. Arms,
shanks, thighs and tarsi barred dark brown.

Variation in proportions is summarized in Table 1.

Osteology

Pectoral girdle.—The pectoral girdle is arciferal (Fig. 2c). Clavicles
arched and not in contact medially, tips extending anteriorly to a line be-
tween the anterior edges of the scapulae. Clavicles in contact with the pars
acromialis but not overlying it. Scapula proximally bicapitate. Coracoids
dilated at their distal and proximal ends and slightly wider than the clavicles.
Epicoracoidal cartilages free. Procoracoids fused anteromedially and later-
ally to the clavicle. The large omosternum has no endochondral ossification
and the manubrial portion is well defined. Sternum well developed; the
proximal and medial regions show ossification. Metasternum and xiphister-
num well differentiated. Distal edge of xiphisternum notched.

Cranial osteology.—The nasal bones are large but not in median contact
and are separated from the frontoparietals (Fig. 3a). Nasals have long max-
illary processes which do not contact the maxillae; margins of nasals rest
on the anterolateral margin of the sphenethmoid. Frontoparietal fontanelle
narrow at posterior end and wide at anterior part. Anteriorly, the fronto-

924 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.5 cm

Fig. 3. Skull of Alsodes vanzolinii: dorsal (a) and ventral (b) view.

VOLUME 93, NUMBER 4 925

parietals broadly rest on the posterolateral margin of the sphenethmoid;
posteriorly they rest on the otoccipitals. Maxillary arch complete. Premax-
illae narrow, the alary processes of moderate length and directed dorsally.
The 14-15 premaxillary teeth moderately long; pars palatina narrow, but
the palatine process moderate sized. Pars facialis of the maxilla relatively
deep and restricted to the snout. Maxilla bears 42-44 teeth in a row ex-
tending to a point just posterior to the maxillo-pterygoid junction. Quadra-
tojugal of moderate size and broadly articulating with the maxilla. Otoccip-
ital poorly ossified with low epiotic eminences. Occipital condyles (Fig. 3b)
not stalked and narrowly separated. Columela present. Zygomatic ramus of
squamosal of moderate length; otic ramus short, broad at distal end, and
not in contact with the crista parotica. Parasphenoid cruciform. Cultriform
process long, gradually narrowing anteriorly, the anterior end extending
slightly between the palatines. Parasphenoid alae oriented at right angle to
the cultriform process. Anterior ramus of pterygoid rests on the palatal shelf
of the maxilla, but does not reach the palatines. Palatines broad and arched,
extending from the maxillae to the sphenethmoid. Prevomers small and
broadly separated.

Natural History

Donoso-Barros (1974) considered the holotype of Alsodes vanzolinii a
male; however, my examination shows that this specimen (MUZUC 12063,
Donoso-Barros’s field number 4938) has yellow ovarian eggs. All the females
examined that have been collected during January and February have mod-
erate sized eggs (x = 2.33 + 0.15 mm diameter); 134 immature ovarian eggs
were counted in one female (MUZUC 12216). Adult males found during
these months have nuptial spines on the chest and on the dorsal surface of
the thumb and second finger.

The type locality (Ramadillas) is highly disturbed by human intervention,
there being only small patches of the primitive Nothofagus forest. In this
place we also collected Bufo rubropunctatus, Rhinoderma darwinii, and
Telmatobufo venustus.

Comparisons

Alsodes vanzolinii shows a rudiment of web between the fifth and fourth
toes. This characteristic is also present in Alsodes nodosus (Cei, 1962) and
A. monticola (Formas, personal observation). Webbing is absent in A. il-
lotus (Gallardo, 1962). Alsodes gargola (Gallardo, 1970), A. laevis and A.
montanus (Cei, 1962) share well developed interdigital webbing. Alsodes
vanzolinii and A. nodosus have minute transparent papillae on the tongue,
but this character is absent in A. monticola; the tongue of the latter species
does not have papillae anteriorly.

926 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Alsodes nodosus has 65.8 mm snout-vent length (25 adults) differing in
this respect from A. vanzolinii (54.4 mm, 28 adults). On the other hand,
Alsodes nodosus have robust limbs, which are slender in A. vanzolinii.
Finally, A. vanzolinii has a yellow triangle on the snout, which is absent in
Alsodes nodosus.

Specimens Examined

Abbreviations.—University of Kansas Museum of Natural History (KU),
Museo de Zoologia, Universidad de Concepcion (MUZUC), Instituto de
Zoologia, Universidad Austral de Chile (IZUA).

Alsodes montanus: IZUA 823-24; Estero Covarrubias, Provincia Santia-
go, 2,400 m.

Alsodes monticola: IZUA 1149-1154; Cordillera Pelada, Provincia Val-
divia, 1,020 m.

Alsodes nodosus: IZUA 747-761, 767-776; Aguas Ciaras, Provincia
Aconcagua, 150 m.

Alsodes vanzolinii: MUZUC 12063-70, 12209-18, 12221, 12223 -25,
12227-28, 1223144; Ramadillas, Provincia Arauco, 100 m.

Alsodes sp.: KU 162206-09, 162212, 162221-—22, 162229, 162232; Cabrer-
ias, Parque Nacional Nahuelbuta, 1,030 m.

Acknowledgments

I am grateful to Tomas Cekalovich (University of Concepcion) and Wil-
liam Duellman (University of Kansas) for making specimens available for
me for study, and to the latter for providing field notes on the frogs. Sonia
Lacrampe typed the manuscript. This work was supported by Proyecto de
Investigacion S-79-3, Direccion de Investigacion, Universidad Austral de
Chile.

Literature Cited

Cei, J. M. 1962. Batracios de Chile.—Ed. Univ. Chile, Santiago, i + i—cviii-128.
Donoso-Barros, R. 1974. Nuevos reptiles y anfibios de Chile.—Bol. Soc. Biol. Concepcion
48:217-229.

Duellman, W. E., and A. Veloso. 1978. Phylogeny of Pleurodema (Anura: Leptodactylidae):
A biogeographic model.—Occas. Papers Mus. Nat. Hist. Univ. Kansas 64:1—46.
Gallardo, J. M. 1962. Los géneros Eupsophus y Batrachyla (Anura, Leptodactylidae) en la
Argentina y la verdadera identidad de Paludicola illota Barbour.—Rev. Mus. Argentino

Cs. Nats. Cienc. Zool. 8:113-122.

. 1970. A proposito de los Telmatobinae (Anura, Leptodactylidae) patagonicos.—Neo-

tropica 16:73-85.

Grandison, A. 1961. Chilean species of the genus Eupsophus (Anura: Leptodactylidae).—
Bull. British Mus. (Nat. Hist.) 8:111-149.

VOLUME 93, NUMBER 4 S27

Iturra, P., and A. Veloso. 1975. Citotaxonomia en especies del grupo Eupsophus roseus
(Amphibia, Leptodactylidae). Descripcion del cariotipo de Eupsophus coppingeri Gun-
ther.—XVIII Reunion An. Soc. Biol. Chile: 25—26 (abstract).

Lynch, J. D. 1978. A re-assessment of the telmatobiine leptodactylid frog from Patagonia.—
Occas. Papers Mus. Nat. Hist. Univ. Kansas 72:1-57.

Instituto de Zoologia, Universidad Austral de Chile, Casilla 567, Valdivia,
Chile.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 928-942

THE STATUS OF THE CENTRAL AMERICAN
LEPTODACTYLID FROGS ELEUTHERODACTYLUS
MELANOSTICTUS (COPE) AND ELEUTHERODACTYLUS
PLATYRHYNCHUS (GUNTHER)

Jay M. Savage and James E. DeWeese

Abstract.—Eleutherodactylus melanostictus, a species unique to the ge-
nus in lower Central America in having dark transverse bars on the anterior,
dorsal, and posterior surfaces of the thighs, is redefined. E. platyrhynchus
is conspecific with melanostictus. In external and skeletal morphology the
species seems allied to the wnistrigatus-cruentus series. Jaw musculature
(dfsq + e) allies this species to the fitzingeri-rugulosus series, whereas all
known representatives of the wnistrigatus-cruentus stock have a very dis-
tinctive set of jaw muscles (DFSQ,AT + s). Karyologically the species has
2N = 22, N.F. = 36 and resembles several members of the fitzingeri-ru-
gulosus series, especially E. berkenbuschii of eastern Mexico. Known mem-
bers of the wnistrigatus-cruentus series have 2N = 26, 32, 34; N.F. = 32,
36, 46. These differences make E. melanostictus the sole representative of
a monotypic species group.

Among the numerous species of rain frogs, genus Eleutherodactylus,
found in the cordilleras of Costa Rica and western Panama, is a series of
populations distinctive from all others in having transverse dark bars on the
anterior, dorsal, and posterior surface of the thighs. In life the contrasting
light interspaces on the posterior thighs range from cream through yellow-
green to orange, salmon, magenta, and scarlet to make these animals among
the most conspicuous forms in the genus. Several names have been applied
to the populations—including Hylodes brocchi Boulenger, 1882 (Gunther,
1900), now known to be restricted to Guatemala (Savage, 1975); Lithodytes
melanostictus Cope, 1875; and Hylodes platyrhynchus Gunther, 1900
(Dunn, 1937; Taylor, 1952), each based on Costa Rican frogs.

Prior to 1960 these frogs were known from a few individuals from scat-
tered localities. While the problem of the status of the various samples has
been recognized for some years, it is only now that sufficient material has
been accumulated to make a review of the populations possible.

Comparison of Populations

Frogs of the melanostictus-platyrhynchus population system are found
along the slopes of the cordilleras of Costa Rica and western Panama, from

VOLUME 93, NUMBER 4 929

12

87 86
10 =

KILOMETERS

Fig. 1. Distribution of Eleutherodactylus melanostictus in Costa Rica and western Panama.
Arabic numerals and letters indicate population samples discussed in text. The dotted line
indicates the 1,500 m contour. a

Monteverde de Puntarenas on the north to the slopes of Volcan Chiriqui to
the south, between 1,150 and 2,483 m. Although these creatures are ex-
tremely similar to one another in most features of morphology, seemingly
significant differences between montane (1,900-—2,483 m) and slope (1,150-
1,920 m) populations in coloration that correlated with minor morphological
distinctions led us to regard them as separate species (Savage, 1976). As
additional collections were made the supposed differences became less clear
and we decided to undertake a sample by sample comparison to verify or
refute our tentative conclusion.

The available material of this system has been grouped into a series of 11
populations (Fig. 1). Slope populations are designated by Arabic numerals,

930 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

montane populations by letters. The original comparisons (and greatest re-
corded differences) were made between the slope population (3) on the pass
between volcanes Barba and Irazu of the Cordillera Central of Costa Rica
(La Palma—La Hondura pass) and the montane population (NT) on the Cor-
dillera Talamanca along the Carreterra Interamericana in the same country.
These differences are summarized below:

Population 3

. Snout obtuse, sloping in profile

2. Disk on finger II expanded, palmate; larg-

er than tympanum

. Subarticular tubercles projecting

. Toe disks expanded, palmate, II-IV emar-
ginate; largest at least as large as tym-
panum

. Throat and venter usually light, with scat-
tered punctations

. Light interspaces on posterior thigh sur-
faces yellow, yellow-green, purplish, or
orange in life

. Groin uniform or with two obscure light

Population NT

. Snout rounded in profile
. Disk on finger II barely expanded, round;

smaller or equal to tympanum

. Subarticular tubercles flattened
. Toe disks barely expanded, II-IV not

emarginate; largest smaller than tympa-
num

. Throat and venter dark, heavily pig-

mented

. Light interspaces on posterior thigh sur-

faces scarlet in life

. Groin with well-developed dark bars

spots

It seemed likely that if these differences were maintained with some con-
sistency geographically and/or altitudinally, that two species were involved.
For this reason we carefully compared all other populations for these fea-
tures as follows from northwest to southeast.

1) Cordillera de Tilaran (1,500—1,580 m): snout round to obtuse, sloping;
disk on finger II expanded; subarticular tubercles projecting; toe disks ex-
panded; throat and venter light to dark; light thigh interspaces yellow to
orange; groin not barred (N = 14).

P) Volcan Poas (2,100 m): snout rounded; disk on finger II small, round;
subarticular tubercles flattened; toe disks small; throat and venter dark;
thigh color not known; groin barred (N = 13).

2) East slope of Volcan Poas (1,200-—1,590): snout round to obtuse, sloping
or rounded; disk on finger II expanded; subarticular tubercles projecting;
toe disks expanded; throat and venter light; light thigh interspaces yellow
to magenta; groin not barred (N = 24).

B) Volcan Barba (1,828—1,920 m): snout round; disk on finger II small,
round; subarticular tubercles intermediate; toe disks expanded; throat and
venter dark; thigh color not known; groin barred (N = 5).

3) La Palma—La Hondura Pass (1,150—1,500 m): see summary above (N =
25).

IT) Volcanes Irazu and Turrialba (2,000—2,374 m): snout rounded; disk on
finger II small; subarticular tubercles flattened; toe disks small; throat and
venter dark; light thigh interspaces scarlet; groin uniform (N = 7).

VOLUME 93, NUMBER 4 931

4) South slope of Volcan Irazu (1,600 m): snout obtuse, sloping; disk on
finger II expanded; subarticular tubercles projecting; toe disks expanded;
throat and venter light; light thigh interspaces yellow, groin uniform (N = 2).

5) Tapanti (1,280—1,320 m): snout obtuse, sloping; disk on finger II ex-
panded; subarticular tubercles projecting; toe disks expanded; venter inter-
mediate in dark pigmentation; thigh color not Known; groin not barred (N =
2).

NT) Northern Cordillera de Talamanca (1,900—2,482 m): see summary
above (N = 7).

ST) Southern Cordillera de Talamanca (2,135—2,160 m): snout obtuse,
sloping; disk on finger II rounded; subarticular tubercles flattened; toe disks
expanded; venter intermediate in pigmentation; light thigh interspaces yel-
low; groin not barred (N = 2).

6) Slope of Cerro Pando (1,200 m): snout rounded; disk on finger II small,
palmate; subarticular tubercles projecting; toe disks expanded; venter in-
termediate in pigmentation; thigh interspace color unknown; groin with a
single bar (N = 1).

These comparisons indicate that montane populations from Volcan Poas
(P), volcanes Irazu and Turrialba (IT), and the northern Talamanca region
(NT) are essentially similar. Populations from the Cordillera de Tilaran (1)
and the passes (2, 3) and slopes (4) of the Cordillera Central and northern
Talamanca range (5) are almost identical to one another. Some overlap in
snout profile and ventral coloration occurs between the three montane sam-
ples and the five populations from lower elevations. Nevertheless, it is pos-
sible to divide the material into two altitudinal groups based on these eight
samples, a montane group from 2,000 m and above and a slope sample from
1,150 to 1,600 m, which differ principally in disk size, nature of the subar-
ticular tubercles and presumably in posterior thigh color. Unfortunately
these differences break down in the remaining population samples, one (B)
from an intermediate elevation (1,828—1,920 m) on Volcan Barba and the
others (ST, 6) from the southern portion of the Cordillera de Talamanca
axis in Panama.

The Barba sample consists of three adult males (CRE 6463, 7094, 7130A)
and a juvenile (CRE 7130B). In general coloration they agree with other
montane examples in having dark venters and barred groins. Morphologi-
cally they further resemble montane members of the complex in having the
disk on finger II round and small and the subarticular tubercles somewhat
flattened, but not as much as in other montane populations. The toe disks,
however, are somewhat larger than in other montane individuals and ap-
proach the size typical of slope populations.

The southern Talamanca sample consists of two examples, the holotype
of Lithodytes melanostictus Cope and a single adult female from the vicinity
of Cerro Punta, Provincia de Chiriqui, Panama (2,160 m). The female type

932 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(USNM 30608) of melanostictus is now in very poor condition, so most
characteristics cannot be determined. The Cerro Punta example agrees with
montane samples in all characteristics except that the snout is obtuse in
profile, the venter is not heavily pega, and the light thigh interspaces
were yellow in life.

A single female (KU 114851) from the lower northern slope of Cerro
Pando (1,200 m) also has a mixture of features. The finger disk size resem-
bles those in montane samples, although larger. The shape of the finger and
toe disks and the size of the latter approach those in other slope populations.
The ventral coloration is intermediate.

Several possible interpretations may be made based on the available ma-
terials: 1) the Cordillera de Tilaran—Cordillera Central populations and the
Cordillera de Talamanca samples represent two allopatric races or species;
2) the montane and slope populations represent related but distinct species
differing only in the size of disks; or 3) only a single species is involved.

Alternative 1 does not seem valid, since the differences between montane
and slope samples from each cordillera is greater than between the cordil-
leras and no feature will consistently separate available material into two
geographic groups. Alternative 2 also seems contraindicated, since both
montane and slope populations overlap to some extent in all characteristics
except disk size and the Barba sample, from an intermediate elevation, is
intermediate between the two groups in this feature.

Alternative 3 seems to be the only tenable conclusion based on this anal-
ysis. While montane and slope populations may be separated most of the
time on the basis of disk size, subarticular tubercle profile, and coloration,
we see no need to formally recognize altitudinal races within the complex.
Many of the apparent differences may represent ecotypic influences asso-
ciated with temperature related responses during development. Theoreti-
cally, at least, it is easier to conceive of the similarities among the isolated
high elevation populations as the result of such influences rather than the
result of fragmentation of a formerly continuous population. This idea im-
plies that each montane isolate is more closely related to the adjacent slope
population than to other montane members of the system. The single rec-
ognizable species including the eleven populations compared above is:

Eleutherodactylus melanostictus (Cope, 1875)
Fig. 2

Lithodytes melanostictus Cope, 1875:109, pl. 23, fig. 10.

Hylodes melanostictus.—Brocchi, 1881:56.—Giunther, 1900:236.

Hylodes platyrhynchus Gunther, 1900:230, pl. 67, fig. a (holotype: BM 1905-
7-18,1/1947.2.15.81; Costa Rica).

VOLUME 93, NUMBER 4 933

Fig. 2. Eleutherodactylus melanostictus: upper, male, CRE 7048-1, from 0.5 km W Bajo
La Hondura, Provincia de San Jose, Costa Rica; lower, female, CRE 3895, from Monteverde,
Provincia de Puntarenas, Costa Rica. Photographs courtesy of James L. Vial and James E.
DeWeese.

934 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hylodes brocchi.—Gunther (in part), 1900:236, pl. 68, figs. a—b.
Eleutherodactylus melanostictus.—Dunn, 1937:163.—Taylor, 1952:728.
Eleutherodactylus platyrhynchus.—Taylor, 1952:757, fig. 34.

Holotype.—USNM 30608, adult female, 50 mm in standard length; from
Costa Rica, Provincia de Limon, Canton de Talamanca, Cerro Utyum, 2,135 m.

Diagnosis.—A moderate sized Eleutherodactylus (males to 43, females
to 57 mm in standard length), immediately distinguished from all other
species in lower Central America by having distinct dark thigh bars that
continue down onto the posterior thigh surface. E. melanostictus belongs
to a cluster of species placed in the cruentus group by Savage (1976) that
have large finger and toe disks and a granulate venter and that lack both
webbing and a tarsal fold.

The only two species within the group with which melanostictus might
be confused, E. cruentus and E. cerasinus, have numerous accessory pal-
mar tubercles on the hand and are much smaller in size (cruentus males to
25, females to 38 mm; cerasinus males to 23, females to 35 mm). The pos-
terior thigh surface of cruentus is usually uniform dark brown to black,
often with some clear yellow spots; the posterior thigh surface of cerasinus
is almost uniform dull red in life (brown in preservative) with a very few
small light spots in some examples.

Description.—Head outline from above subovoid to subelliptical; snout
profile rounded to obtuse (sloping) in profile. Canthus rostralis sharp. Loreal
area obtuse, slightly concave in section. Choanae small, ovoid, smaller than
vomerine tooth patches; vomerine tooth patches transverse, posterior but
internal to choanae, very narrowly separated on midline. Paired vocal slits
and single internal vocal sac in males. Surfaces of head mostly smooth, with
a single large supraorbital tubercle near posterior edge of upper eyelid, a
series of weak superciliary tubercles along margin of upper eyelid and sev-
eral smaller tubercles elsewhere on eyelid. Tympanum round in males, oval
in females; internal, indistinct in both sexes, vertical diameter about 4%
length of orbit. A distinct supratympanic fold. Dorsum and upper limb sur-
faces relatively smooth with scattered tubercles. Finger I shorter than II;
disk on finger I rounded. Disks well developed on fingers III-IV, at least 2
times width of digit, larger than tympanum, emarginate; disk on finger II
slightly to definitely expanded, usually palmate. Whitish nuptial pad on
thumb of adult males. Subarticular tubercles of fingers ovoid, flattened to
slightly projecting, globular in profile; no supernumerary tubercles; thenar
tubercle large, elongate; palmar tubercle large cordate, no accessory palmar
tubercles. A distinct calcar. Toe disks smaller than finger disks, largest
about equal to disk on finger II; disks on toes II-IV truncate to palmate,
emarginate; disks on toes I and V very slightly expanded. No webbing.
Subarticular tubercles under toes ovoid, slightly projecting and globular in

VOLUME 93, NUMBER 4 935

profile; no supernumerary tubercles; no plantar tubercles; inner metatarsal
tubercle well developed, elongate, outer indistinct very small, round; no
tarsal fold, but a very weak series of small outer tarsal tubercles may be
present. A distinct small gland in groin; venter granulate.

Coloration.—Dorsal ground color pale tan to dark brown, in life brown,
chestnut olive, olive-green or green; head uniform above or with distinct
light (pink in life) enamel area on anterior surface of snout; dark brown to
black interocular bar in most examples, sometimes bordered anteriorly by
an obscure light area; usually a distinct dark canthal blotch extending from
eye to nostril and involving upper loreal zone; a distinct black supratym-
panic stripe from eye to behind tympanum; tympanum covered by a dark
spot; four distinct to obscure dark lip bars, the first continuous with the
canthal blotch; two bars separated by a light area lie below eye. Iris golden
in life, with a greenish cast. Five principal dorsal color patterns: i) essen-
tially uniform with a few dark blotches and/or short lines on suprascapular
and flank areas, sometimes with a definite W-shaped suprascapular dark
mark; ii) with a broad mid-dorsal stripe, as broad or broad as interorbital
area, contrasting in color with flanks and usually demarcated by an irregular
narrow dark boundary; in life the center stripe is chestnut to reddish-brown;
iii) with a narrow distinctly light (scarlet to yellow in life) mid-dorsal stripe,
about half width of interorbital space, bordered by a black outline; stripe
begins at interorbital or suprascapular region and continues to anus; iv) very
narrow tan mid-dorsal stripe along median raphe, continuing as a narrow
light longitudinal stripe on posterior thigh surface; and v) with distinct to
obscure dark blotches and/or chevron-shaped marks dorsally. Flanks
marked with a series of dark elongate oblique dark blotches, outlined by
black, to give impression of a tiger pattern. Dark areas often flecked with
metallic green, magenta or gold in life. Upper surfaces of forelimbs uniform
to strongly barred with dark. Dorsal, anterior and posterior surfaces of thigh
and calf marked with distinct dark bars; primary dark bars and light inter-
spaces about same widths, although interspaces often split by a very narrow
dark stripe; light interspaces on upper and anterior surfaces of thigh and
calf suffused by darker pigment; on posterior thigh surface interspaces clear
light and usually very distinct in most examples, although sometimes suf-
fused with darker pigment; in life light areas on posterior surface are bright
yellow, yellow-green, orange, salmon-pink, magenta, or scarlet. Throat and
venter and undersides of limbs range from dirty white (in life and preser-
vation) through intermediate stages to being heavily marked with dark pig-
ment; in some examples dark pigment forms distinct dark spots. Groin area
usually uniform like flank, sometimes with two obscure light spots or with
elongate dark blotches similar to those on the flank in the tiger pattern
phase. In life, light areas of groin and adjacent surfaces of venter and hind-
limbs may be suffused with orange, pink, or red.

936 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—The centromeric indices (C.I.) and percent genome values (G) for the 11 pairs of
chromosomes in the karyotype of Eleutherodactylus melanostictus.

Chromosome number (Call G
1 13)7/ 16.9
2 1.47 13.4
3 4.07 12
4 2.04 11.9
5 1.15 9.8
6 1.26 8.8
7 1.28 7.0
8 7.00 6.6
9 7.00 5.1

10 7.00 4.6
11 7.00 BA)

Measurements.—In this section the notation gives the mean followed by
the range in parentheses. Standard lengths (distance from snout to vent) are
given in millimeters, other measurements as percentages of standard length.
Values are based on samples of 10 adult males and 10 adult females ran-
domly selected from montane and slope populations. The smallest juvenile
available is 12 mm in standard length. Standard length, adult males 38.2
(35.0-42.6), adult females 43.8 (35.4—-56.3); head length, males 38.0 (36.2—
40.0), females 37.7 (34.0-39.5); head width, males 41.3 (38.4-45.6), females
40.9 (39.0-42.4); snout length, males 16.6 (15.0-18.3), females 16.9 (14.5-
18.2); loreal length, males 11.1 (10.3-12.4), females 11.6 (9.0—13.0); length
of orbit, males 16.2 (14.5-—18.0), females 14.6 (12.8-17.2); height of tympa-
num, males 6.5 (S.0-7.9), females 6.1 (4.5-7.0); hindlimb length, males 199.1
(185.0-—222.2), females 205.8 (168.0-—225.2); tibia length, males 62.2 (54.1-
76.4), females 62.3 (51.0—68.1). No marked sexual dimorphism is evident
from these measurements, although the relative tympanum heights are
slightly distinctive.

Karyotype.—Bogart (1970) described the karyotype of a specimen re-
ferred to this species from Monteverde, Costa Rica. We have karyologically
examined one example (CRE 4130), a female, from near Cerro Punta, Pan-
ama, using the technique of Patton (1967) as modified by Lowe et al. (1966).
7 spreads were counted, 3 were photographed and the individual chromo-
somes were measured (Table 1).

The diploid number (2N) is 22 and the nombre fundamental (N.F.) is 36.
The chromosomes form a gradually descending series with no obvious size
groupings. Chromosome pairs (centromere placement according to the sys-
tem of Levan et al., 1964) 1, 2, 5, 6 and 7 are metacentric; 4 is submeta-
centric; 3 is subtelocentric; and 8—11 are acrocentric (Fig. 3).

VOLUME 93, NUMBER 4 937

Ld
(—=)

a=
. — ]

Fa
=
2)
y 4
[aad
oo)
=
Zz
hal
5)
Be
hal
0.

lined 234 98 Oe a B90 TT
CHROMOSOME NUMBER

Fig. 3. A representative idiogram for Eleutherodactylus melanostictus.

Bogart’s (1970) description and figure differ slightly from our data, with
2N = 22 and N.F. = 38. Chromosome pairs are 1, 5, 6 and 8 metacentric;
3 and 4 submetacentric; 2 and 11 subtelocentric; and 7, 9 and 10 acrocentric,
according to Bogart. Essentially his interpretation is that there is one less
acrocentric and one more subtelocentric pair, one less metacentric and one
more submetacentric and that 11 is subtelocentric rather than acrocentric.
This latter interpretation or observation explains the difference in N.F. be-
tween Bogart’s example and our data. These apparent differences may be
due to 1) differing techniques of preparation and measurement of the
spreads; 2) variation, either individual or populational; or 3) misidentifica-
tion of the Monteverde specimen by Bogart. Although we have been unable
to locate Bogart’s voucher specimen to confirm the identification, E. me-
lanostictus is so distinctive in color pattern that the last alternative seems
unlikely. Until additional karyological preparations are available, no deci-
sion between the other two alternative explanations is possible.

Jaw musculature.—The depressor mandibulae has a single slip originating
on the dorsal fascia with a few fibers coming from the squamosal (dfsq).
The adductor mandibulae externus superficialis is present (e). The jaw mus-
cle formula dfsq + e, follows the terminology of Starrett (1968).

Ecological notes.—Taylor (1952:761) reported the collection of a number

938 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of examples of this form from bromeliads, where they may hide during the
day. Most specimens in our samples (La Palma, La Hondura and Monte-
verde) are males taken at night while calling from herbaceous vegetation 1.0—
2.5 m above the ground in dense rainforest habitats. USC examples from
the Talamanca range are mostly males found under rocks or logs during the
day.

Distribution.—Evergreen forests of the upper portion of the premontane
and lower montane zones on both Atlantic and Pacific slopes of the Cor-
dilleras de Tilaran, Central, and Talamanca of Costa Rica and the extension
of the latter as the Chiriqui massif of western Panama, 1,128—2,483 m (Fig. 1).

Locality records. COSTA RICA: no other data; ALAJUELA: Isla Bon-
ita; Hda. Cayuga; Cinchona; above Cinchona (1,590 m); E slope Volcan
Poas (1,590 m) 1 km W Poasito, Rio Poasito; CARTAGO: 1.6 km NE Casa
Mata; 2.5 km W Sanatorio Duran, Volcan Irazu; El Empalme and 16 km S;
2 km NW Pastora, Volcan Turrialba; Volcan Irazu (2,000 m); Rio Quiri,
Tapanti; nr. El Sesteo; 1-2 km SE Santa Teresa; Trinidad; CARTAGO-
SAN JOSE: 1 km S El Empalme; Cerro de la Muerte; HEREDIA: Volcan
Barba (1,828 m); Cerro Chompipe; LIMON: Cerro Utyum (2,135 m); PUN-
TARENAS: 1.75, 2.6 km ESE, 1.25, 2.25 km SE and Monteverde; SAN
JOSE: Boquete Camp; La Hondura; 0.5 km W Bajo La Hondura; 2 km E
Las Nubes; 0.5 km E, N, S and La Palma; 1.4 km S Alto La Palma; 3 km
SE and Rancho Redondo. PANAMA: BOCAS DEL TORO: N slope Cerro
Pando (1,200 m); CHIRIQUI: 2 km W and 0.5 km N Cerro Punta.

Remarks.—We have examined holotypes of both Lithodytes melanostic-
tus and Hylodes platyrhynchus. The former (USNM 30608) is now in very
poor condition and some of the features noted when it was examined 14
years ago by Savage are no longer distinguishable. The distinctively barred
thighs are still apparent and all other characteristics and the original de-
scription (Cope, 1875) confirm its identity with the montane populations of
the Cordillera Talamanca—Chiriqui axis of Costa Rica and Panama.

Gunther (1900:226, 236) ascribed a smooth ungranulated venter to this
specimen, doubtless based on Cope’s (1875:109) statement ‘‘skin every-
where smooth.’’ Cope probably was referring to the upper surfaces of the
body. However, the rugosely granulate venter of this and many other
Eleutherodactylus is distorted in poorly fixed specimens or those long in
preservative.

The type of AHylodes platyrhynchus (BM 1905-7-18,1/1947.2.15.81) re-
mains in excellent condition. It is an adult female 57 mm in standard length
and morphologically agrees in all ways with the sample (3) from the pass
between Volcanes Barba and Irazu. The dorsal color pattern is of type i)
described above. The head markings and thigh barring as illustrated by
Gunther (1900, pl. 67A) confirm the identification.

VOLUME 93, NUMBER 4 939

The most consistent and distinctive characteristic of E. melanostictus is
the continuation of the dark thigh bars onto the posterior surface of the
thighs. Since the light interspaces between the dark bars are usually brightly
colored in life, the contrasting dark versus brilliant yellow, yellow-green,
orange, magenta, or red makes this species readily recognizable in the field.
Two examples here referred to this species exhibit a considerable reduction
and suffusion of the light interspaces with dark pigment to obscure the
usually contrasting pattern. An adult male (CRE 3296), 37 mm in standard
length, from near Bajo La Hondura, Costa Rica, has the light interspaces
reduced to several small light spots while the margins of the dark bars are
obscured by the suffusion of dark pigment. On close inspection the typical
dark thigh bars may be observed. This example also has the throat, venter
and undersides of limbs heavily covered with dark pigment, a condition
unusual in other examples of E. melanostictus from this locality. As a matter
of fact it has the greatest degree of ventral dark pigmentation within all
samples of this species. In all other features this frog agrees with our concept
of the species and we conclude that it is a variant with much more extensive
development of melanin than is usual.

A second extremely dark example of this form is an adult female (CRE
3985), 56 mm in standard length, from Monteverde, Costa Rica. In all fea-
tures but coloration this example is a typical E. melanostictus. Essentially,
the Monteverde frog has a very dark dorsum and extensive dark pigmen-
tation over all the under surfaces. The light interspaces on the thigh surfaces
are very heavily suffused with dark so that the dark thigh bars appear out-
lined by a very narrow light line. The extensive dark suffusion makes the
dark bars obscure on the posterior thigh surface, since even the bordering
remnant of the interspaces is lost as the bar proceeds downwards. The bars
are barely distinct from the heavily suffused interspaces in this region, but
can be distinguished by their heavier concentration of black pigment.

Relationships

Evaluation of evolutionary relationships among the multitudinous species
of Eleutherodactylus has long defied the capabilities of herpetologists. Most
attempts to group clusters of the 400 or so nominal species now placed in
the genus have been based primarily on external ‘‘key’’ characteristics of
dubious aid in discerning all but the most closely related species similarities.
Recently Savage (1976) and Lynch (1976) have applied two somewhat dif-
ferent approaches to establishing divisions within Eleutherodactylus. The
former recognized species groups comprised of species that are extremely
similar morphologically and are probably evolutionarily closely allied.
Lynch follows a second alternative of forming groups that correspond to
sub-genera, without giving them formal nomenclatural status. Both methods

940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

have thus far proven inadequate to cope with the complex mosaic of su-
perficial morphological characteristics that have been combined over and
Over again in distantly related species of Eleutherodactylus for the reasons
already pointed out by us (Savage and DeWeese, 1979).

The, for the most part, unpublished studies on jaw musculature (Starrett,
1968), serum proteins (Harris, 1973) and karyology (DeWeese, 1976) of
Eleutherodactylus suggest that at least six major evolutionary lineages are
recognizable within the limits of the genus (sensu lato). In most cases these
lineages do not conform to the groupings recognized by Lynch (1976) based
on external and skeletal morphology. That the problem is real is suggested
by the fact that Savage (1976) included E. melanostictus in the melanostic-
tus group, its synonym platyrhynchus in the cruentus group; Lynch (1976)
regarded the latter taxon as belonging to his wnistrigatus group in which he
included cruentus and its allies.

Eleutherodactylus melanostictus morphologically, but apparently only
superficially, resembles the members of the cruentus group (Savage, 1976)
in having a strongly granulate venter, large digital disks, and no toe webbing
or tarsal fold. Other Central American species included in this group by
Savage are: altae, pardalis, cruentus, cerasinus, ridens, frater, caryophal-
laeceus, moro and monnichorum. Studies by DeWeese (1976) on the kary-
ology of most of these forms and the less extensive serum protein data of
Harris (1973) indicate that the cruentus group is a composite of species from
rather diverse origins that happen to share a series of trivial morphologic
features in common. Lynch (1976) included all of these disparate forms
among the 92 species placed in his wnistrigatus group.

In terms of jaw muscle features, E. melanostictus shows the greatest
similarity to the fitzingeri and rugulosus groups of Savage (1975, 1976),
which are included together in the fitzingeri group of Lynch (1976). As
pointed out by Starrett (1968) the most consistent and evolutionarily signif-
icant of these muscles is the condition of the adductor mandibulae. Three
conditions may be recognized: 1) both an adductor mandibulae posterior
subexternus and an externus superficialis present (s + e); 2) only the poste-
rior subexternus present (s); or 3) only the externus superficialis present (e).
Eleutherodactylus melanostictus, like the members of the fitzingeri-rugu-
losus groups now examined from Central America, has the e condition. All
known members of the uwnistrigatus group of Lynch (1976) have the s con-
dition.

In addition, the features of the depressor mandibulae of E. melanostictus
support a similar relationship. Starrett (1968) recognized 10 patterns of de-
pressor mandibulae muscle origins. Six of these conditions occur within the
family Leptodactylidae and four are known to occur in Eleutherodactylus.
These conditions are: 1) a single slip principally from the dorsal fascia, but
with a few fibers from the squamosal (dfsq); 2) a single slip principally from
the dorsal fascia, but with a few fibers from the squamosal and annulus

VOLUME 93, NUMBER 4 941

tympanicus (dfsqat); 3) three slips, one each from fascia, squamosal and
annulus tympanicus (DFSQAT); and 4) three slips, with superficial slip from
fascia and annulus tympanicus and a deep slip (internal to the others) from
the squamosal (DFSQ,AT). We regard conditions 1 and 2 as representing
only slight variation in a single character.

The depressor mandibulae in E. melanostictus are of condition 1 (dfsq).
This condition is typical of the fitzingeri-rugulosus groups of Savage (1976).
Some species in this complex (brocchi and rugulosus) have condition 2
(dfsqat). All members of the wnistrigatus group of Lynch (1976) for which
this character is known have condition 4 (DFSQ,AT).

Karyologically, E. melanostictus agrees in diploid number (2N = 22) and
N.F. (36) with E. talamancae of Costa Rica and Panama, E. berkenbuschii
(recently revived by Savage and DeWeese, 1979) and E. vocalis of Mexico,
and E. rayo of Costa Rica, all members of the fitzingeri-rugulosus complex.
Other leptodactylids with the same numbers are E. decoratus (alfredi group)
and Hylactophryne augusti (of Mexico); and Leptodactylus podicipinus,
and L. wagneri of South America. Of these species the chromosome fea-
tures of E. melanostictus most closely approach those of E. berkenbuschii.
Members of the cruentus group of Savage (1976) and the wnistrigatus group
of Lynch (1976) for which the karyology is known have 2N = 26, 32, 34 and
N.F. = 32, 36, 46 (Bogart, 1970; DeWeese, 1976).

What a dilemma! A frog that in external and skeletal morphology clearly
belongs with one major section of the genus (the wnistrigatus-cruentus
stock), but in extremely important features of jaw musculature and karyol-
ogy belongs elsewhere (fitzingeri-rugulosus complex)! Obviously our knowl-
edge of this marvelously complex and perverse genus is far from complete.
It may be that E. melanostictus represents a primitive stock ancestral to the
two very distinctive major groups (unistrigatus-cruentus and fitzingeri-ru-
gulosus). Full investigation of this possibility awaits more extensive data on
jaw musculature and karyology. Until these studies are completed it seems
wise to retain melanostictus in a distinct monotypic group.

Acknowledgments

Materials used in this study has been made available through the kindness
of the authorities at the American Museum of Natural History; Academy
of Natural Sciences, Philadelphia; British Museum (Natural History) (BM):
Field Museum of Natural History; Museum of Comparative Zoology, Har-
vard College; University of Kansas (KU); University of Michigan and the
United States National Museum of Natural History (USNM). Also studied
were specimens in the University of Southern California collections (CRE).

Field assistance was provided by several workers from the University of
Southern California, especially Norman J. Scott and James L. Vial. James
Hanken and David B. Wake of the University of California provided a

942 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

critical live specimen for karyological analysis. P. H. Starrett confirmed the
condition of the jaw musculature for E. melanostictus. Our work in Costa
Rica was expedited by the air and encouragement of the Facultad de Bio-
logia, Universidad de Costa Rica, and the Organization for Tropical Studies.
To all of the individuals and institutions we express our deep appreciation.

Literature Cited

Bogart, J. P. 1970. Los cromosomas de anfibios anuros del genero Eleutherodactylus.—Act
IV Congr. Latin Zool. 1:65—78.

Boulenger, G. A. 1882. Catalogue of the Batrachia Salientia S. Ecaudata in the collection of
the British Museum. 2nd. ed.—Taylor and Francis, London, Engl., xvi + 503 pp.

Brocchi, P. 1881. Etude des Batraciens de 1’ Amerique Centrale.—Miss. Sci. Mex. 1 Amer.
Centr. 3(2):1-56.

Cope, E. D. 1875. On the Batrachia and Reptilia of Costa Rica.—J. Acad. Nat. Sci. Phila.
ser. 2, 8:93-157.

DeWeese, J. E. 1976. The karyotypes of Middle American frogs of the genus Eleutherodac-
tylus (Anura: Leptodactylidae): A case study of the significance of the karyologic meth-
od.—Ph.D. diss., University of Southern California, 210 pp.

Dunn, E. R. 1937. The amphibian and reptilian fauna of bromeliads in Costa Rica and Pan-
ama.—Copeia 1937(3): 163-167.

Gunther, A. C. L. G. 1900. Reptilia and Batrachia.—Biol. Centr.-Amer. London. Sign.
30:229-236.

Harris, R. T. 1973. Comparative serum-protein electrophoresis and protein characterization
in the systematics of Costa Rican frogs genus Eleutherodactylus (Anura: Leptodactyl-
idae). Ph.D. diss., University of Southern California, 247 pp.

Levan, A., D. Fedga, and A. A. Sandberg. 1964. Nomenclature for centromere positioning
on chromosomes.—Hereditas 52:201-220.

Lowe, C. H., J. W. Wright, and C. J. Cole. 1966. Chromosomes and karyotypes of sceloporine
iguanid lizards in the North American southwest.—Mamm. Chrom. Newsletter, Hous-
ton 2(2):201-20S.

Lynch, J. D. 1976. The species groups of the South American frogs of the genus Eleuthero-
dactylus (Leptodactylidae).—Occ. Pap. Mus. Nat. Hist., Univ. Kansas 61:1-24.
Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus (Rodentia,

Heteromyidae).—J. Mamm. 48(1):27-37.

Savage, J. M. 1975. Systematics and distribution of the Mexican and Central American stream

frogs related to Eleutherodactylus rugulosus.—Copeia 1975(2):254—306.

. 1976. A preliminary handlist of the herpetofauna of Costa Rica. II edition.—Editonial

Univ. de Costa Rica:1-19.

and J. E. DeWeese. 1979. A new species of leptodactylid frog, genus Eleutherodac-

tylus, from the Cordillera de Talamanca, Costa Rica.—Bull. South. Calif. Acad. Sci.

78(2): 107-115.

Starrett, P. H. 1968. The phylogenetic significance of the jaw musculature in anuran amphib-
ians. Ph.D. diss., Univ. of Michigan, 179 pp.

Taylor, E. H. 1952. A review of the frogs and toads of Costa Rica.—Univ. Kansas Sci. Bull.
35(1-S):577-942.

Allan Hancock Foundation and Department of Biological Sciences, Uni-
versity of Southern California, Los Angeles, California 90007.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 943-946

PHILANDER AND FOUR-EYED OPOSSUMS
ONCE AGAIN

Philip Hershkovitz

Abstract.—Despite attempts by Pine (1973) and Husson (1978) to suppress
Philander Tiedemann, 1808, as generic name for the four-eyed pouch opos-
sum, Didelphis opossum Linnaeus (1758), it is shown that the type-species,
P. virginianus Tiedemann, 1808, is a renaming or objective synonym of
Didelphis opossum Linnaeus, with identical lectotype, the female four-eyed
pouch opossum described and figured by Seba (1734).

Philander Tiedemann, 1808, with type Philander virginianus Tiedemann,
1808 = Didelphis opossum Linnaeus, 1758, was established as the name for
the four-eyed pouch opossum in 1949 when I reported its availability and
designated the type-species. In 1973, Pine raised the question that the type-
species, originally described as ‘“‘rotlich braun ...’’ could well be the
pouchless four-eyed opossum known as Metachirus nudicaudatus E. Geof-
froy 1803. In Pine’s opinion, the pouched form was grayish, never brownish.
Thereupon, he suggested suppression of the name Philander Tiedemann.
However, grayish and brownish individuals occur among both pouched and
non-pouched four-eyed opossums (Hershkovitz, 1976), and only pouched
opossums were known to Tiedemann and all authors cited by him in his
description of Philander virginianus.

Husson (1978:27) agreed with Pine regarding suppression of Philander
Tiedemann but “‘for different reasons.’’ None are given unless Husson re-
fers to the anticipated effect of his use of a nomenclatural strategem.

In my discussions of the status of Philander Tiedemann, 1808, I (1949: 11)
noted that P. virginianus ‘‘is merely a new name for the four-eyed pouch
opossum, Didelphis opossum Linnaeus,”’ and (1949:12, 1976:302) designat-
ed the type-species of Philander Tiedemann as the equation ‘‘Philander
virginianus Tiedemann = Didelphis opossum Linnaeus.’’ Notwithstanding,
Husson (1978:27) claims I ‘‘made the mistake to consider the name Philan-
der virginianus Tiedemann, 1808, a replacement for Didelphis opossum Lin-
naeus 1758. It is true that Linnaeus’s name is given as a synonym in Tie-
demann’s account, but there is no indication that it should be treated
differently from the other references given under Philander virginianus. In
fact Philander virginianus Tiedemann is a composite species based in part
on Didelphis opossum Linnaeus ... partly on Metachirus nudicaudatus
(E. Geoffroy, 1803) . . . and partly on Didelphis marsupialis Linnaeus, 1758

944 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. . and the mention of Virginia as one of the type localities.’’ Husson then
goes on to say that “so far as I know no lectotype has ever been designated
for Philander virginianus Tiedemann, 1808, and therefore I now select as
such the female specimen from Virginia discussed by Tyson (1698). In this
way Philander virginianus Tiedemann, 1808, becomes a junior synonym of
Didelphis virginiana Kerr (1792:103), and the generic name Philander Tie-
demann, 1808, disappears in the synonymy of Didelphis Linnaeus, 1758.”

Husson errs in his judgment of a work he did not see (Husson, 1978:546).
There is more than one indication that Tiedemann proposed the name Phi-
lander virginianus as a replacement for Didelphis opossum Linnaeus which,
as senior objective synonym, becomes the correct name for the taxon. Tie-
demann, it is noted (see Hershkovitz, 1949:11, for extract) introduced the
two names in direct apposition thus: **P{hilander]. virginianus (Did. opos-
sum L.).’’ The construction, as used here, is more than a mere indication.
It is an equation consisting of two objective synonyms. This interpretation
is supported by the fact that Tiedemann described no specimens before him
or new species and he mentions no other scientific name. He simply re-
placed the old name Didelphis opossum Linnaeus for four-eyed pouch opos-
sums by the new name combination Philander virginianus. The reference
to Virginia was made in ignorance. Neither Tiedemann nor his contempo-
raries knew the distributional limits of the species they described.

The construction of paired binomials in direct apposition is employed by
Tiedemann (1808:427—428) for the two remaining species of his genus Phi-
lander, but with only the new generic name used as replacement in the
equated binomials, thus:

‘2. Das mausartige Opossum P. murinus (Did. murina L.)....
3. Das kurzgeschwantze Opossum. P. brachyurus (Did. brachyuros
Penns)h. eck

It is evident here and throughout his text that Tiedemann replaced older
name combinations with new ones, whether his own or of other authors.
Where alternate interpretations are possible, the decision of the first reviser
holds.

A second indication is Tiedemann’s diagnosis of Philander virginianus,
‘‘Korper rotlich braun. Ueber jedem Auge ein gelblich/weiser Flecken.
Schwanz so lang als der Leibe./ 1 Fuss und 3 Zoll lang ohne den Schwanz.’’
References to Buffon and Schreber’s figures leave no doubt that Tiedemann
defined only the four-eyed pouch opossum. The description is followed by
bibliographic references to works on opossum anatomy by Tyson (1698) and
Cowper (1704).

Third, Tiedemann (1808:426, footnote b, [a reference not included in my
1949 extract]) based on the generic name Philander on the ‘‘Philandri’’ of
the non-binomialist Brisson (1762:207). The diagnosis for the species Bris-

VOLUME 93, NUMBER 4 945

son names ‘‘Philander’’ derives from the same animals described by Seba
(1734) under the name ‘‘Philander’’ and which served Linnaeus (1758:55)
for the following description of Didelphis opossum:

3. Dfidelphis}]. cauda semipilosa, superciliorum regione pallidiore, mam-
mis binis.
Philander, Opossum s. Carigueja. Seb. mus. 1. p. 56. t. 36. f. 1. 2.
Habitat in America
Abdomen circa mammas contrahitur in marsupium, polices postici mutici.

The repeated tautonomy in generic and specific names and the other in-
dications pointed out above definitely equate Philander virginianus Tiede-
mann with Didelphis opossum Linnaeus. On the other hand, the genus
Philander Tiedemann, 1808, is construed as a new taxon containing two of
the five nominal species of the genus Didelphis Linnaeus, 1758, and ‘‘P.
brachyurus (Did. brachyuros Penn.) currently Monodelphis brevicaudata
Erxleben, a species not included in the original Linnaean Didelphis.

Husson’s motive for designating Tyson’s animal as lectotype of Philander
virginianus Tiedemann for the deliberate purpose of sinking the name Phi-
lander Tiedemann in the synonymy of Didelphis Linnaeus, is obscure. He
could as readily have designated any of the four-eyed pouch opossums cited
by Tiedemann. As it stands, Husson’s action violates the letter and spirit
of Recommendation 74A of the International Code of Zoological Nomen-
clature that “‘in designating a lectotype, a zoologist should in general act
consistently with, and in any event should give great weight to, previous
valid restrictions of the taxonomic species, in order to preserve stability of
nomenclature.’ More to the point, Article 74(C) of the Code requires that
‘‘each designation of the lectotype must be made specifically for an individ-
ual nominal species [in this case Philander virginianus| and must have as
its object the definition of that species.’’ Tiedemann had already clearly
defined his species as a four-eyed pouch opossum. Tyson’s opossum is not
that species and cannot serve for its definition. Such specimens not seen
but cited by the author in or appended to the description of a species, are
properly treated as misidentified referrals and not in the sense of syntypes.
In any event, my (1976:297) prior designation of the female four-eyed pouch
opossum described and figured by Seba (1734:56, pl. 36, fig. 2) as the lec-
totype of Didelphis opossum Linnaeus, 1758, makes it ipso facto the lec-
totype of the junior objective synonym Philander virginianus Tiedemann,
1808. Metachirops Matschie, 1916, also with type Didelphis opossum Lin-
naeus, is an objective junior synonym of Philander Tiedemann, 1808.

In conclusion, I find no zoological support for Pine’s (1973) opinion that
Philander virginianus Tiedemann is conspecific with Metachirus nudicau-
datus E. Geoffroy, and no zoological or nomenclatural justification for Hus-
son’s (1978) attempt to convert the name Philander virginianus Tiedemann,

946 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

already restricted to four-eyed pouch opossums, into a synonym of Didel-
phis virginianus Kerr, for the Virginia opossum.

Literature Cited
(For a complete bibliography see Husson (1978) cited below.)

Brisson, A. D. 1762. Regnum animale in classes IX. Distributum sive synopsis methodica.—
Theodore Haak, Leiden, pp. 1-6, 1-296.

Hershkovitz, P. 1949. Generic names of the four-eyed pouch opossum and the woolly opossum

(Didelphidae).—Proc. Biol. Soc. Washington 62:11-12.

. 1976. Comments on generic names of four-eyed opossums (family Didelphidae).—

Proc. Biol. Soc. Washington 89(23):295—304.

Husson, A. M. 1978. The mammals of Suriname.—Bnill, Leiden, pp. i-xxxiv, 1-569.

Linnaeus, C. 1758. Systema naturae per regna tria naturae secundum classes, ordines, genera,
species cum characteribus, differentiis, synonymis, locis. (Ed. 10.)—L. Salvia, Stock-
holm, 1:1-824.

Pine, R. H. 1973. Anatomical and nomenclatural notes on opossums.—Proc. Biol. Soc. Wash-
ington 86(33):391—402.

Seba, A. 1734. Locupletissimi rerum naturalium thesauri accurata descriptio et iconibus ar-

tificiosissimis expressio, per universam physices historiam opus . . . .—Amsterdam 1,
38 unnumbered, pp. 1-178, pls. 1-111, 2 unnumbered (frontispiece), 6 ornamental en-
gravings.

Tiedemann, D. F. 1808. Zoologie. Zu seinen Vorlesungen entworfen. Vol. 1, Allgemeine
Zoologie, Mensch und Saugthiere.—Weberschen Buchhandlung, Landshut, pp. i-xvi,
1-610.

Field Museum of Natural History, Chicago, Illinois 6060S.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 947-962

POLYDORA AND BOCCARDIA SPECIES (POLYCHAETA:
SPIONIDAE) FROM WESTERN MEXICO,
CHIEFLY FROM CALCAREOUS HABITATS

James A. Blake

Abstract.—Nine species of spionid polychaetes are reported from the Gulf
of California. Most species are borers in various gastropod shells. The de-
scription of Polydora nuchalis is clarified and P. heterochaeta is rede-
scribed. Polydora barbilla is new to science. Polydora anophthalma is
transferred to Boccardia and redescribed.

A list of polychaetes from the Gulf of California was provided by Reish
(1968), who recorded 20 species of Spionidae. Additional species were
added by Light (1970), Fauchald (1972), Kudenov (1973, 1975) and Blake
(1979a) bringing the total number to about 30. In general, the shell boring
spionids from the Gulf of California have not been studied. Recently, while
conducting revisionary work on the Spionidae, it was possible to examine
several gastropod shells from various localities in the Gulf of California.
Spionids were recovered following dissolution of the shells in an acidified
alcohol solution (70% isopropanol + HCl). These forms are reported herein
along with some additional records from benthic habitats.

The collections are deposited in the Allan Hancock Foundation, Univer-
sity of Southern California (AHF), National Museum of Natural History,
Smithsonian Institution (USNM) and the California Academy of Sciences
(CAS).

The following spionid species from Mexico are included in this report:
Polydora barbilla, new species; Polydora convexa Blake and Woodwick,
1972; Polydora socialis (Schmarda, 1861); Polydora giardi Mesnil, 1896;
Polydora nuchalis Woodwick, 1953; Polydora websteri Hartman, 1943; Po-
lydora heterochaeta Rioja, 1939; Boccardia anophthalma (Rioja, 1962), new
combination; Boccardia tricuspa (Hartman, 1939).

Polydora barbilla, new species
Fig. 1

Material examined.—MEXICO, Gulf of California, 3—S km west of Puerto
Penasco, 15 m, from shell of Muricanthus nigritus, 20 March, 1971, coll. J.
D. Kudenov, holotype (AHF Poly 1296), 10 paratypes (AHF Poly 1297) and
5 paratypes (USNM 58978).

Description.—A moderate sized species, up to 12 mm long and 0.5 mm

948 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

G I

--20um— +

J

Le 20am

Fig. 1. Polydora barbilla (paratype, AHF Poly 1297): A, Anterior end in dorsal view; B,
Posterior end in dorsal view; C, Notopodial acicular spine from posterior setiger showing
basally directed barbs; D, Posterior parapodium in dorsal view showing location of acicular
spines; E—G, Hooded hooks; H, Dorsal geniculate seta from setiger 5; I, Major spine from
setiger 5; J, Companion seta from setiger 5; K, Fascicle of major spines and companion setae
from setiger 5.

VOLUME 93, NUMBER 4 | 949

wide for about 130 segments. Color: light tan in alcohol, some specimens
with dorsal intersegmental pigment bands. Intersegmental glands present in
middle and posterior setigers (Fig. 1D).

Prostomium deeply incised on anterior margin (Fig. 1A), with caruncle
extending to posterior margin of setiger 3; no occipital tentacle; eyes 0-4.
Peristomium enlarged, truncate on anterior margin; palps long, thin, reach-
ing posteriorly for about 20 segments.

Setiger 1 with long, fingerlike notopodial lobes; notosetae short, with two
tiers of thin capillaries; setigers 2-4, 6 and subsequent setigers with noto-
setal fascicles of bilimbate capillaries arranged in two tiers, setae of anterior
tier being shorter than posterior tier; posterior setigers with 3—4 emergent
or imbedded acicular spines and 3-5 long capillaries (Fig. 1B—D), each spine
with minute barbs basally directed on shaft (Fig. 1C). Neurosetae of setigers
1-4 and 6 with capillaries arranged in two tiers as described for notosetae;
bidentate hooded hooks from setiger 7, numbering 4—6 throughout most of
body, then diminishing to 2 in far posterior segments; hooks accompanied
by capillaries for only 5 or 6 segments; hooks lacking constriction or manu-
brium on shaft, with angle between teeth being less than 90° but about 150°
between main fang and shaft (Fig. 1E-F); apical tooth difficult to see in
some angles (Fig. 1G), but always present; hood opening with fine bristles
(Fig. 1E-G).

Setiger 5 modified, larger than either 4 or 6 (Fig. 1A); setae including
superior dorsal fascicle of 5 or 6 finely bristled geniculate setae (Fig. 1H),
curved row of major spines (Fig. 11, K) with bilimbate companion setae
(Fig. 1J—K) and ventral fascicle of bilimbate capillaries; major spines falcate
with finely bristled collar on convex side.

Branchiae from setiger 9-10, small at first, reaching full-size by setigers
12-14, absent from posterior one-half of body.

Pygidium with 4 lobes, dorsal pair being distinctly smaller than ventral
pair (Fig. 1B). :

Etymology.—barbilla: Spanish for barb, as on a fish.

Remarks.—Polydora barbilla is closely related to P. langerhansi Mesnil,
1896, from Madeira, P. convexa Blake and Woodwick, 1972, from California
and P. pilocollaris Blake and Kudenov, 1978, from Victoria, Australia in
having a flange or collar on the convex side of the major spines of setiger
5. Only P. barbilla and P. convexa, however, have spines in posterior
notopodia. In P. barbilla, these spines are of a unique acicular type bearing
minute, basally directed barbs. P. convexa, on the other hand, has small
posterior needles arranged in flattened packets. P. barbilla has branchiae
from setiger 9-10 and bidentate hooded hooks occurring throughout, while
P. convexa has branchiae from setiger 8 and unidentate hooded hooks in
posterior neuropodia.

Rioja (1939) described a larval stage (Polydora sp.) from Acapulco having

950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

similarly appearing major spines on setiger 5 as occur in P. barbilla and P.
convexa. Since both species are now known to occur in the Gulf of Cali-
fornia, it is not possible at this time to assign Rioja’s record to either species
until their larval morphologies are described.

Distribution.—Mexico: Gulf of California, boring into gastropod shells.

Polydora convexa Blake and Woodwick, 1972
Polydora convexa Blake and Woodwick, 1972:73-74, fig. 1.

Material examined. —MEXICO, Gulf of California, 3-5 km west of Puerto
Penasco, 15 m, from shell of Muricanthus nigritus, 20 March 1971, coll. J.
D. Kudenov, | specimen (AHF).

Remarks.—TYhe Mexican specimen agrees with the description of speci-
mens from California by Blake and Woodwick (1972), except that the nature
of the posterior spines was originally misconstrued. They were said to be
tight fibril bundles, but are instead flattened packets of needles, similar to
those reported for Polydora flava and P. latispinosa by Blake and Kudenov
(1978). These structures glisten and shine in the posterior segments of the
animal and make it possible to separate the species from others in a sample.

Distribution.—Mexico and California. A borer of mollusk shells; also re-
ported from algal holdfasts, sponges, Dodecaceria colonies, and bryozoa
by Blake and Woodwick (1972).

Polydora socialis (Schmarda, 1861)

Polydora socialis.—Blake, 1971:20-23, fig. 13 [Synonymy]; 1979b:607—
609.—Light, 1978:179-181, fig. 180a—1 [Synonymy].—Rioja, 1947:207-—
208.

Polydora socialis plena.—Reish, 1968:82.

Material examined.—MEXICO, Bahia de Los Angeles, April 1962 and
October 1963, coll. Beaudette Foundation, Sta. 3, 2 specimens (USNM
58983); Sta. 5, 1 specimen (USNM 58984); Sta. 9, 1 specimen (USNM
58985); Sta. 28, 1 specimen (USNM 58986); Sta. 36, 2 specimens (USNM
58987); Sta. 146, 1 specimen (USNM 58988). San Felipe, near mouth of
Arroyo Estralla, boring in shell of Muricanthus nigritus occupied by hermit
crab, 12 April 1952, coll. L. O. Miles, 10+ specimens (AHF).

Remarks.—Polydora socialis is widespread and may prove to be cos-
mopolitan once sufficient material is examined. The species appears to be
highly opportunistic and occupies benthic habitats as well as the shells of
mollusks. The present specimens agree well with published descriptions.

Distribution.—East, west, and gulf coasts of North America; Chile; Aus-
tralia.

VOLUME 93, NUMBER 4 951

Polydora giardi Mesnil, 1896
Fig. 2A—B

Polydora giardi Mesnil, 1896:195-202, pl. 13, figs. 1-12.—Fauvel, 1927:50-
52, fig. 17h-m.—Hartman, 1941:309, pl. 48, fig. 43; 1961:29; 1969:135-
136, figs. 1-6.—Rioja, 1941:727; 1943:230.— Rainer, 1973:560, fig. 9.—
Read, 1975:413.—Blake and Kudenov, 1978:252, fig. 38i—k.

Material examined.—MEXICO, Sonora, off Cholla, 16.7 m, boring into
shell of gastropod, 26 Dec. 1966, coll. T. A. Burch, 3 specimens (CAS
010131).

Remarks.—Polydora giardi is a small species, seldom exceeding 6-7 mm
in length, and characterized by having an incised prostomium, a tooth on
the major spines of setiger 5, branchiae usually from setiger 9, and by lacking
a constriction on the hooded hooks and posterior notopodial spines. Its
closest relative appears to be P. tridenticulata Woodwick from the central
Pacific, which differs only in having 2 distinct teeth on the major spines of
setiger 5. Two undescribed species from the eastern Pacific have been dis-
covered which appear to be closely related to P. giardi and P. tridenticu-
lata. Both forms, however, have posterior notopodial spines.

The setal morphology of P. giardi and P. tridenticulata poses some in-
teresting questions with regard to the consistency of having 1 or 2 accessory
teeth on the major spines of setiger 5S. Some specimens of P. giardi from
the eastern Pacific appear to have a second spur on the opposite side from
the large accessory tooth. The large accessory tooth is clearly seen with the
scanning electron microscope (SEM) (Fig. 2A). The same figure also reveals
a thin, closely adhering protuberance on the opposite side. This structure
is difficult to detect, but appears to flare at its apex into several bristles. It
is not known how common this structure is among P. giardi populations,
or whether it is homologous to the large second accessory tooth of the
related species, P. tridenticulata. Rice and Simon (1980) have observed that
a small percentage of some Florida populations of P. ligni have an extra
accessory tooth on the major spines of setiger 5. This suggests to me that
such structures among species of Polydora may represent small scale ge-
netic variation rather than wide scale species level morphological criteria.
Whether or not such a system is operative in P. giardi and its relatives
remains to be determined.

Figure 2A also reveals that the companion setae of setiger 5 are distinctly
bristled. The hooded hooks are also bristled on their apical ends, with the
teeth only barely protruding through the hood opening (Fig. 2B).

Polydora nuchalis Woodwick, 1953
Figs. 2C-D, 3

Polydora nuchalis Woodwick, 1953:381-383, fig. la—f; 1960:122—128, pls.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Polydora giardi: A, Fascicle of major spines from setiger 5, arrows denotes locations
of small adhering secondary spurs; B, Hooded hook.—Polydora nuchalis: C, Two hooded
hooks in apical view; D, Hooded hook in dorsofrontal view. SEM micrographs.

VOLUME 93, NUMBER 4 953

L_500um_, "—40um |

Fig. 3. Polydora nuchalis: A, Anterior end of Mexican specimen in dorsal view; B, Hooded
hook; C, Fimbriated capillary notoseta from setiger 5; D, Group of companion setae and major
spines from setiger S.

1-3.—Hartman, 1961:29; 1969:143-144, figs. 1-5.—Blake, 1975:215.—
Light, 1978:177.—Kudenov, 1973:116—-117; 1975:206.

Material examined.—MEXICO, Puerto Penasco, Bahia Cholla, high in-
tertidal sand flats, occurring in dense mats, 19 Jan. 1972, coll. J. D. Kuden-
ov, 2 specimens (AHF). CALIFORNIA, Duxbury Reef, near Bolinas, from
high intertidal rock platform, in sand on bottom of pool, 2 Feb. 1978, coll.
J. A. Blake (CAS 010130).

Description.—Specimens from Cholla Bay up to 19 mm long and | mm
wide for approximately 110 segments. Color: light tan in alcohol, no pig-
ment.

Prostomium incised on anterior margin, continuing posteriorly as caruncle
to setiger 3 (Fig. 3A); two pairs of eyes: anterior pair cup-shaped, posterior
pair oval-shaped and less widely spaced; occipital tentacle present posterior
to eyes.

Setiger 1 with dorsally elevated, short notopodial lobes, notosetae absent;
capillary notosetae of setigers 2-4, 6 and subsequent setigers unilimbate,

954 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

arranged in 2 tiers, with anterior tier bearing shorter setae; posterior setigers
with 4—5 long capillaries, posterior spines lacking. Capillary neurosetae of
setigers 1-4 and 6 similar to corresponding notosetae, those of setiger 1
thinner; bidentate hooded hooks replace capillaries from setiger 7; hooks
with constriction and manubrium on shaft, hood with fine striations (Fig.
3B) evident in light microscopy, but seen to be minute bristles with SEM
(Fig. 2C-D).

Setiger 5 modified, with setae including superior dorsal fascicle of fim-
briated capillaries bearing punctations on shaft (Fig. 3C), curved row of
simple major spines alternating with bristled companion setae bearing nu-
merous fine bristles (Fig. 3D), and ventral fascicle of unilimbate capillaries.

Branchiae from setiger 7, continuing to near posterior end; gills long,
meeting at midline. Pygidium disclike, with wide dorsal gap.

Remarks.—There is confusion in the literature concerning the true nature
of the companion setae of setiger 5 for P. nuchalis. Woodwick (1953, fig.
le) termed these setae plumose and figured them as having the end of the
shaft covered with numerous bristles. Light (1978:177), on the other hand,
indicated that these setae are geniculate with a distinct limbus. He also
mentioned that the hooded hooks lacked a manubrium, although Wood-
wick’s fig. If suggests that a constriction is present on the shaft. In the
present study, P. nuchalis has been observed to have companion setae on
setiger 5 which are very similar to those depicted by Woodwick. The ends
of the shaft are covered with numerous fine bristles, observed both with the
light microscope and SEM. At low magnifications (400) with light micros-
copy, the setae appear to be bilimbate, with the bristles visible under oil
immersion (1,000). The hooded hooks have a manubrium and constriction
on their shafts. The bristles observed on the hoods have not been described
previously, but have been observed on most other spionids when examined
under SEM.

Distribution.—Mexico: Gulf of California; California.

Polydora websteri Hartman, 1943

Polydora websteri.—Blake, 1971:6-8, fig. 3 [Synonymy].—Foster, 1971:26—
27, figs. 30-36 [Synonymy].
?Polydora ciliata.—Rioja, 1943:229 [not Johnston, 1838].

Material examined.—_MEXICO, Gulf of California, 3—5 km west of Puerto
Penasco, 15 m, from shell of Muricanthus nigritus, 20 March 1971, coll. J.
D. Kudenov, 15 specimens (AHF); Sonora, off Cholla, 16.7 m, with bryozoa
and shell fragments, 26 Dec. 1966, coll. T. A. Burch, | specimen (CAS
000114).

Remarks.—These specimens agree well with the published descriptions.
It is probable that the record of P. ciliata by Rioja (1943) from Guymas may

VOLUME 93, NUMBER 4 955

Fig. 4. Polydora heterochaeta: A, Anterior end in dorsal view; B, Dorsal capillary notoseta
from setiger 5; C, Ventral capillary neuroseta from setiger 5; D, Giant falcate spine from setiger
5; E, Inflated spine from setiger 5; F-G, Normal major spines and hastate companion setae
from setiger 5; H, Hooded hook; I, Capillary notoseta from anterior tier of anterior setiger; J,
Capillary notoseta from posterior tier of same; K, Posterior end with pygidium in lateral view.

be P. websteri, but since Rioja’s specimens are no longer available, new
materials will need to be collected.
Distribution.—East, west, and gulf coasts of North America; Australia.

Polydora heterochaeta Rioja, 1939
Fig. 4

Polydora heterochaeta Rioja, 1939:308-309, figs. 6-10; 1962:185.—Reish,
1968:82.

956 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Material examined.—MEXICO, Bahia de Los Angeles, in silty sands,
Oct. 1963, coll. Beaudette Foundation, Sta. 2, 1 specimen (USNM 58981);
Sta. 102, 1 specimen (USNM 58982).

Description of postlarval juvenile.—Both specimens small, 2.3 and 2.5
mm long for 28 and 26 setigers, respectively, both 0.5 mm wide. Color in
alcohol: brown with paired pigment bars on dorsum of setigers 3 and 4,
medial chromatophores on dorsum of setigers 5-11, with chromatophores
of setigers 7-11 arranged as 2 tandem spots (Fig. 4A). Parapodia, branchiae
and much of body with conspicuous glandular appearance.

Prostomium broad, entire on anterior margin; caruncle extending poste-
riorly to end of setiger 2; eyes numerous with pair of large oval ones located
medially and about 4 pairs of small ones more laterally and anteriorly located
(Fig. 4A). Peristomium reduced, fused to setiger 1; palps short, thin, pos-
teriorly directed to setiger 5.

Setiger 1 reduced, with small noto- and neuropodial lobes; with 4—5 long
notosetae on one specimen, lacking on second specimen; neurosetae deli-
cate capillaries. Setigers 2—4, 6 and subsequent setigers with 2-tiered fas-
cicles of unilimbate, fimbriated capillary notosetae, those of posterior tier
being longer (Fig. 41, J); capillaries of posterior setigers also fimbriated, but
longer and more robust than those of anterior setigers imparting spinous
appearance to posterior end (Fig. 4K). Neurosetae of setigers 2—4 and 6-8
include fimbriated capillaries, arranged in 2 tiers; bidentate hooded hooks
replace anterior tier from setiger 9, posterior tier of capillaries remaining
intact throughout body; hooks numbering 2—4 throughout, with striations
on hood and weak constriction on shaft (Fig. 4H).

Setiger 5 enlarged and greatly modified; setae include 2 simple dorsal
capillaries (Fig. 4B), 3 types of major spines with slender hastate companion
setae (Fig. 4D-G) and a ventral fascicle of 2—3 inflated capillary neurosetae
(Fig. 4C); major spines include a single giant falcate spine with long lateral
groove (Fig. 4D) [tip end of this spine usually broken off, imparting bilobed
appearance to end of shaft], a single pointed spine with subterminal inflated
portion (Fig. 4E) and 2-3 falcate spines bearing a large accessory tooth (Fig.
4F, G), these accompanied by hastate companion setae.

Branchiae from setiger 7; gills slender, glandular, continuing for 10 seg-
ments. Pygidium cufflike, with distinct striations (Fig. 4K).

Remarks.—The fully developed adult form of Polydora heterochaeta has
never been discovered and the present postlarval specimens from Bahia de
Los Angeles differ somewhat from the original description of planktonic
larvae by Rioja (1939) from Acapulco. In Rioja’s form, the hooded hooks
were Said to begin on setiger 7 instead of 9 and the dorsal chromatophores
were depicted as paired throughout the body, while the present specimens
have paired chromatophores only on setigers 3 and 4, with medial chro-
matophores on subsequent segments. The major spines of setiger 5 are gen-

VOLUME 93, NUMBER 4 957

erally of the same form as Rioja described, although he did not observe the
inflated spine.

Blake (1969) commented on the validity of P. heterochaeta and other
species which have been based upon larval or postlarval forms. Polydora
heterochaeta exhibits several features which are typical of such postlarval
forms. These include the poorly developed peristomium, undifferentiated
and broadened prostomium, posteriorly directed palps, retention of larval
pigment and the occurrence of multiple major spine types on setiger 5. Extra
spine types, similar to those described here for P. heterochaeta, have been
described for the larvae or postlarvae of P. ciliata (Johnston) and P. hoplura
Claparéde by Wilson (1928), P. hermaphroditica Hannerz (1956), P. bioc-
cipitalis Blake and Woodwick (1972) and P. websteri Hartman by Blake
(1969). These setae are represented in P. heterochaeta by the giant falcate
spine (Fig. 4D) and the inflated spine (Fig. 4E). In other species, these setae
are the first modified spines to appear on setiger 5 during development and
the first to drop off, being entirely replaced by the typical form found on
adults. In P. heterochaeta the adult form of major spine is that represented
in Fig. 4F, G. These spines have an enlarged sheathlike accessory tooth,
which should make it possible to identify the adults of this species when
other characters, such as the hooded hooks beginning on setiger 9, are
considered.

Distribution.—Mexico: Acapulco and the Gulf of California.

Boccardia anophthalma (Rioja, 1962), new combination
Fig. 5

Polydora anophthalma Rioja, 1962:185—188, figs. 89-93.

Material examined.—MEXICO, Gulf of California, 3-5 km west of Puerto
Penasco, 15 m, from shell of Muricanthus nigritus, 20 March 1971, coll. J.
D. Kudenov, 4 specimens (AHF). '

Description.—A moderate-sized species, up to 14 mm long and 1.0 mm
wide for 85 segments. Color in alcohol: light tan, pigment lacking.

Prostomium and first 5 setigers dorsoventrally compressed; subsequent
body segments normal-appearing, oval to rounded in cross-section. Prosto-
mium truncate on anterior margin (Fig. 5A); caruncle narrow, twisted, con-
tinuing posteriorly to end of setiger 4; no eyes; no occipital tentacle. Palps
thin, short, extending posteriorly for 11 or 12 segments.

Setiger 1 reduced, with notopodial lobe, lacking notosetae; with short,
thin capillary neurosetae. Setigers 2—4, 6 and subsequent segments with
unilimbate capillary notosetae arranged in 2 tiers, first tier with short thick-
ened setae, those of second tier longer and thinner; notosetae of posterior
segments reduced to 5 or 6 simple capillaries; last 10 to 12 segments with
elevated parapodia bearing 2-4 long capillaries and 1-2 stout recurved

958 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

————
==

f LiOum
: AF -300um-
iy day,
Y; aH Hi me | oe
Os, Wh
: eG i Hy
p “3 AY K\
3 eL\ ‘A i
7 BN
aie

Fig. 5. Boccardia anophthalma: A, Anterior end in dorsal view; B, Hooded hook; C,
Notopodial boathook from posterior setiger; D, Unilimbate capillary neuroseta from setiger 5;
E, Group of major spines from setiger 5; F, Posterior end in dorsal view.

VOLUME 93, NUMBER 4 959

boathooks (Fig. 5C, F) projecting into shallow medial channel. Neuropodia
of setigers 24, 6 with fascicles of unilimbate capillaries; bidentate hooded
hooks from setiger 7 numbering 6—8 throughout most of body, then reduced
to 2-3 in far posterior segments; accompanied by 1-3 inferior capillaries;
hooks exhibiting wide angle between teeth (Fig. 5B).

Setiger 5 with ventral fascicle of 5—6 unilimbate capillaries (Fig. SD);
major spines of 2 types: (1) falcate with flange along curved edge (Fig. SE);
(2) bristle-topped with small tooth among bristles (Fig. 5E).

Branchiae on setigers 2-4, 6 and subsequent segments (Fig. 5A), con-
tinuing for most of body length, absent from those posterior segments having
boathooks; gills short on setigers 2, 3, 4, and 6, reaching full-size on setiger
8, overlapping at midline, then becoming short again in posterior one-half
of body.

Pygidium reduced to 4 minute lobes surrounding anal opening (Fig. 5F).

Remarks.—Polydora anophthalma Rioja (1962) was described and figured
as having branchiae from setiger 8, but the posterior end was not described.
In most respects, however, the present specimens agree well with Rioja’s
description. The major spines of setiger 5 are unique. There is no other
example of a Boccardia species having an accessory structure on the falcate
unbristled spines. The flange occurring on these spines in B. anophthalma
was illustrated by Rioja for his specimens. Since the anterior branchiae are
considerably smaller than those following setiger, it seems probable that
Rioja overlooked them, as did Hartman (1939:16) for B. tricuspa (see Wood-
wick, 1963:209). I strongly feel that the evidence indicates that the present
specimens represent the form which Rioja described as Polydora ano-
phthalma.

With the addition of B. anophthalma, Boccardia, as emended by Blake
and Kudenov (1978) now has 17 species. Boccardia anophthalma is most
closely related to B. berkeleyorum Blake and Woodwick (1971) in having
a small tooth on the bristle-topped spines of setiger 5, posterior notopodial
spines and 4 small pygidial lobes. In B. anophthalma, however, the falcate
spines of setiger 5 have an accessory flange and the posterior spines are
recurved. B. berkeleyorum, on the other hand, has simple falcate spines on
setiger 5 and posterior acicular spines. The unusual flattened appearance of
the anterior end of B. anophthalma is caused by dorsal-ventral compression
of the prostomium and first 5 setigers. The functional significance of this
flattening is not understood, but it may be related to its shell-boring habit.

Distribution.—Mexico, Gulf of California; Asuncion Island, a borer into
mollusk shells.

Boccardia tricuspa (Hartman, 1939)

Polydora tricuspa Hartman, 1939:16-17, fig. 3c—k; 1961:29.—Rioja,
1939:304—308, figs. 22-31; 1941:727; 1943:238; 1962:185.

960 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Boccardia tricuspa.—Woodwick, 1963:209—212, fig. 1.—Hartman, 1969:99—
100, figs. 1-5.

Boccardia proboscidea.—Carrasco, 1974:185—187, figs. 1-4 [not Hartman,
1940], fide Blake and Kudenov, 1978:238.

Material examined.—MEXICO, Gulf of California, 3—5 km west of Puerto
Penasco, 15 m, from shell of Muricanthus nigritus, 20 March 1971, coll. J.
D. Kudenov, | specimen (AHF). Guadelupe Island, Melbomene Cove, in-
tertidal in shell of Haliotis sp., Velero Sta. 1912-49, 17 Dec. 1949, 5 spec-
imens (AHF). Guadelupe Island, Camp Arroyo, Lava Point, intertidal, 10
July 1975, coll. D. Sutton, 1 specimen (CAS 009331), 3 specimens (CAS
009371).

Remarks.—The major spines of setiger 5 of B. tricuspa have a more
complex structure than previously believed. The spines are of 2 types: (1)
a simple falcate spine, (2) a cusped spine generally noted by previous in-
vestigators to have 3 teeth. The end of the cusped spine is produced laterally
to form a tooth at either end. Between these teeth is a narrow ridge which
bears a thin curtain. This structure is what is generally considered to be a
medial or third cusp. Being very thin and fragile, it is easily worn into
different forms and configurations, but is usually gone from the oldest and
most worn spines in a setal fascicle series. On the inner edge of the ridge
between the lateral teeth are numerous fine bristles. These, too, are best
seen on the youngest spines, being mostly worn away on the older ones.
The presence of bristles on these teeth allies B. tricuspa more closely to
other species of the genus which have the ends of their spines entirely
covered with bristles. Detailed comparative studies of these spines of the
various species is in progress, using the SEM, and it is expected that these
studies will help clarify such relationships.

Distribution.—California, Mexico, Galapagos Islands, Chile.

Acknowledgments

I am grateful to the following persons and their institutions for the loan
of materials on which this report is based: Mr. Dusty Chivers, California
Academy of Sciences, Dr. Jerry D. Kudenov, while he was at the University
of Arizona, Dr. Kristian Fauchald, while he was at the Allan Hancock
Foundation and Dr. Keith H. Woodwick, California State University, Fres-
no. An early draft of the manuscript benefited from comments by Dr. Wood-
wick.

Literature Cited

Blake, J. A. 1969. Reproduction and larval development of Polydora from northern New
England (Polychaeta: Spionidae).—Ophelia 7: 1-63.

VOLUME 93, NUMBER 4 961

. 1971. Revision of the genus Polydora from the east coast of North America (Poly-

chaeta: Spionidae).—Smithsonian Contrib. Zool. 75:1-32.

1975. Phylum Annelida: Class Polychaeta. Pp. 151-243 in R. I. Smith and J. A.
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. 1979a. Four new species of Carazziella (Polychaeta: Spionidae) from North and South

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. 1979b. Revision of some polydorids (Polychaeta: Spionidae) described and recorded

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, and J. D. Kudenov. 1978. The Spionidae (Polychaeta) from southeastern Australia

and adjacent areas with a revision of the genera.—Mem. Nat. Mus. Victoria 39:171-

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, and K. H. Woodwick. 1971. A review of the genus Boccardia Carazzi (Polychaeta:

Spionidae) with descriptions of two new species.—Bull. Southern California Acad. Sci.

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. 1972. New species of Polydora (Polychaeta: Spionidae) from the coast of California.—

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Carrasco, F. D. 1974. Spionidae (Polychaeta) provenientes de la Bahia de Concepcion y
lugares adyacentes.—Bol. Soc. Biol. Concepcion 48:185-—201.

Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and
adjacent areas in the eastern Pacific Ocean.—Allan Hancock Monogr. Mar. Biol. 7:1-
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Fauvel, P. 1927. Polychetes Sédentaires. Addenda aux Errantes, Archiannélides, Myzosto-
maires.—Faune de France 16:494 pp.

Foster, N. 1971. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea.—Stud.
Fauna Curagao Caribb. Islands 36:183 pp.

Hannerz, L. 1956. Larval development of the polychaete families Spionidae Sars, Disomidae
Mesnil and Poecilochaetidae n. fam. in the Gullmar Fjgrd (Sweden).—Zool. Bidrag
Uppsala 31:1-204.

Hartman, O. 1939. The polychaetous annelids collected by the Presidential cruise of 1938.—

Smithsonian Misc. Coll. 98:1—22.

. 1940. Boccardia proboscidea, a new species of spionid worm from California.—Jour.

Wash. Acad. Sci. 30:382—387.

. 1941. Some contributions to the biology and life history of Spionidae from Califor-

nia.—Allan Hancock Pacific Exped. 7:289-324, pls. 45-48.

. 1943. Polydora websteri Hartman. In V. L. Loosanoff and J. Engle, Polydora in

oysters suspended in the water.—Biol. Bull. 85:69-78.

. 1961. Polychaetous annelids from California.—Allan Hancock Pacific Exped. 25:1-

226, 34 pls.

. 1969. Atlas of the sedentariate polychaetous annelids from California.—Allan Han-

cock Foundation, Univ. of Southern California, Los Angeles, 812 pp.

Johnston, G. 1838. Miscellanea zoologica. Arcadae.—Mag. Zool. Bot. Edinburgh 2:62-73.

Kudenov, J. D. 1973. Annelida (Polychaetes). Pp. 76-131 in R. C. Brusca, ed., A handbook

to the common intertidal invertebrates of the Gulf of California.—Univ. Arizona Press,

Tucson, Arizona.

. 1975. Sedentary polychaetes from the Gulf of California, Mexico.—Jour. Nat. Hist.

9:205—231.

Light, W. J. 1970. A new spionid (Annelida, Polychaeta) from the Gulf of California.—Bull.
Southern Calif. Acad. Sci. 69:74-79.

962 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

——. 1978. Invertebrates of the San Francisco Bay Estuary System. Family Spionidae
(Annelida, Polychaeta).—The Boxwood Press, Pacific Grove, California, xii + 211 pp.

Mesnil, F. 1896. Etudes de morphologie externe chez les annélides. I. Les spionidiens des
cotes de la Marche.—Bull. Sci. France Belg. 29:110-—287, pls. 7-15.

Rainer, S. 1973. Polydora and related genera (Polychaeta: Spionidae) from Otago waters.—
Jour. Roy. Soc. New Zealand 3:545-564.

Read, G. B. 1975. Systematics and biology of polydorid species (Polychaeta: Spionidae) from
Wellington Harbour.—Jour. Roy. Soc. New Zealand 5:395-419.

Reish, D. J. 1968. A biological survey of Bahia de Los Angeles, Gulf of California, Mexico.
II. Benthic polychaetous annelids.—Trans. San Diego Soc. Nat. Hist. 15:67—106.

Rice, S. A., and J. L. Simon. 1980. Intraspecific variation in the pollution indicator polychaete
Polydora ligni (Spionidae).—Ophelia 19 1:79-115.

Rioja, E. 1939. Estudios anelidologicos I. Observaciones acerca de varias formas larvarias y

postlarvarias pelagicas de Spionidae, precedentes de Acapulco, con descripcion de una

especie neuva del género Polydora.—Ann. Inst. Biol. México 10:297-311.

. 1941. Estudios anelidologicos III. Datos para el conocimiento de la fauna de Poli-

quetos de las costas del Pacifico de México.—Ann. Inst. Biol. México 12:669-746.

. 1943. Estudios anelidologicos VIII. Datos acerca de las especies del genero Polydora

Bosc, de las costas Mexicanas del Pacifico.—Ann. Inst. Biol. México 14:229-241.

1947. Estudios anelidologicos XVII. Contribucién al conocimiento de los anélidos

poliquetos de Baja California y Mar de Cortés.—Ann. Inst. Biol. México 18:197—224.

. 1962. Estudios anelidologicos XXVI. Algunes anelidos poliquetos de las costas del

Pacifico.—Ann. Inst. Biol. México 32:131-229.

Wilson, D. P. 1928. The larvae of Polydora ciliata Johnston and Polydora hoplura Cla-
parede.—Jour. Mar. Biol. Assoc. U.K. 15:567-603.

Woodwick, K. H. 1953. Polydora nuchalis, a new species of polychaetous annelid from
California.—Jour. Wash. Acad. Sci. 43:381-383.

—. 1960. Early larval development of Polydora nuchalis Woodwick, a spionid poly-

chaete.—Pacific Sci. 14:122-128.

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Battelle's New England Marine Facility, The William F. Clapp Labora-
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!

|

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 963-970

A NEW FISH SPECIES OF THE GENUS
VANDERHORSTIA (TELEOSTEI: GOBIIDAE)
FROM THE AMIRANTE ISLANDS,
INDIAN OCEAN

Ernest A. Lachner and James F. McKinney

Abstract.—Vanderhorstia praealta, a new species of the fish family Go-
biidae, is described from the Amirante Islands, Indian Ocean. Vanderhorstia
praealta resembles the Cryptocentrus-like gobies, a large group of more
than 60 species comprising at least seven genera. We provisionally place V.
praealta in the genus Vanderhorstia because it lacks vertical rows of cu-
taneous papillae on the cheek, has a moderate number of scales in the lateral
series (54), a moderate gill opening, and an elongate caudal fin. Vanderhor-
stia praealta has an extremely elongate first dorsal fin; spines I-III are
longest, about equal in length and 62.5% of the standard length. The species
is known by only one specimen.

The new species was captured by the field crew of the Seychelles Islands
Program, 1964, of the U.S. National Science Foundation International In-
dian Ocean Expedition organized by James E. Bohlke, Academy of Natural
Sciences, Philadelphia. We provisionally place this species in the genus
Vanderhorstia Smith (1949:103). Superficially our new species resembles
the Cryptocentrus-like gobies, a large group of more than 60 species of
which the following genera are probably interrelated: Cryptocentrus Va-
lenciennes, Amblyeleotris Bleeker, Mars Jordan and Seale, Vanderhorstia
Smith, Flabelligobius Smith, Ctenogobiops Smith, and Eilatia Klausewitz.

RA single character is often used to distinguish among these genera. The most
_ frequently used generic (or group) characters are: number of vertical and
| horizontal rows of cutaneous papillae on the cheek; length of caudal fin;

extent of gill opening; size and number of rows of teeth in the jaws; kind
and number of scales on the trunk; presence of tubular nares; shape of head
(depressed, compressed); length of snout; and width and contour of gape.
One can demonstrate with ease that a particular gobiid species is new, but

| to allocate it properly to a genus is difficult. This problem has resulted in

the description of many new genera without any meaningful, comparative
analysis of generic characters and their generic group affinities. Our new
species lacks vertical rows of papillae on the cheek and therefore does not
agree with the generic concept that Hoese and Steene (1978:382) presented for
Cryptocentrus and Amblyeleotris, each having variously developed vertical

964 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

or transverse rows of papillae on the cheek. On the basis of the development
of papillae on the cheek, the genus Mars (including Obtortiophagus Whitley
as a synonym) should be linked with the Cryptocentrus-Amblyeleotris
group. The four other nominal genera—Vanderhorstia, Flabelligobius,
Ctenogobiops and Eilatia—have no vertical rows of papillae on the cheek.
Flabelligobius has a long jaw, a pointed snout, minute scales numbering
over 100 in the lateral series, and a narrow gill opening restricted to the
pectoral fin base (Smith, 1959:205, fig. 22). Ctenogobiops has ctenoid scales,
and the gill opening is extensive (Smith, 1959:191). Eilatia has cycloid and
ctenoid scales, and a restricted gill opening; the caudal fin is longer than the
head length, and the head is depressed (Klausewitz, 1974:206). Vanderhor-
stia has about 50-65 scales in the lateral series, and cycloid and ctenoid
scales; the gill opening is wide (Smith, 1959:192). Our new species has no
vertical rows of papillae on the cheek, has 54 scales in the lateral series, all
cycloid scales, a moderate gill opening, head moderately round, caudal fin
longer than length of head, teeth of jaws caninoid, in 2 to 4 rows, and the
anterior nares opening on a short tube. The first dorsal fin spines I-IV are
very long and spines I-III are nearly equal in length. The combination of
these characters does not agree with the characters listed for any of the
above seven genera. Some characters, such as the width of the gill opening,
may be less variable among the genera and species than the number of
scales on the trunk or the length of the caudal fin. However, all of the
characters mentioned above must receive a comparative evaluation among
all of the related nominal genera and included species in order for us to
ascertain practical or natural generic groupings. We place our new species
in Vanderhorstia because of the moderate number of scales in the lateral
series, the moderate gill opening and the elongate caudal fin. Also, the genus
Vanderhorstia has priority among the nominal Cryptocentrus-like gobies
that lack vertical rows of cheek papillae.

The methods of taking and recording counts and data are given in Lachner
and McKinney (1974, 1978).

Important comparative material studied for generic determination in-
cludes: Amblyeleotris guttata, USNM 220084, Philippines; Amblyeleotris
randalli, USNM 220085, Philippines; Cryptocentrus cryptocentrus, USNM
220078, St. Brandon Shoals, Indian Ocean; Ctenogobiops crocineus,
USNM 220080, Philippines; Mars caeruleomaculatus, Field Mus., sta. no.
LW-5, Gulf of Mannar, S. India; Mars strigilliceps, USNM 220083, Phil-
ippines, and USNM 220077, off Somali coast, Indian Ocean; Vanderhorstia
delagoae, Acad. Nat. Sci. Philadelphia, sta. no. F-119, Seychelles; Van-
derhorstia ornatissima, Acad. Nat. Sci., Philadelphia, sta. no. F-44, Sey-
chelles.

VOLUME 93, NUMBER 4 965

Vanderhorstia praealta, new species
Figs. 1-2

Holotype.—USNM 215290, female, 35.5 mm SL, from Amirante Islands,
D’Arros Island, off E side, at a depth of 18.3—30.5 m, 9 Mar. 1964, collected
by D. Dockins, R. Rosenblatt, W. Starck, and J. Tyler, sta. F-109, Inter-
national Indian Ocean Expedition, Seychelles Islands Program 1964, Acad-
emy of Natural Sciences of Philadelphia.

Diagnosis.—Related to Cryptocentrus-like gobies but variously different
from them in the following characters: first dorsal fin very elongate and far
forward on body, its length 62.5 percent of SL, spines I-III longest and
about equal in length; scales of moderate size, about 54 in lateral series,
cycloid; three horizontal rows of papillae on cheek; gill opening moderate;
caudal fin longer than head length; dorsal fin VI-I,10; anal fin I,10; pectoral
rays 18; head, nape, base of pectoral fin, and breast with numerous, small
light spots; trunk plain, light brown; the fins mostly dark.

Description.—Dorsal fin rays VI-I,10; anal fin rays I,10; pectoral fin rays
18-18; pelvic fin rays I,5; segmented caudal fin rays 17; branched caudal fin
rays 13; lateral scale rows 54; transverse scale rows 24.

Scales cycloid, smaller anteriorly on trunk; scales absent on cheek, oper-
cle, base of pectoral fin, breast, and predorsal area; scales eccentric, focal
area narrow, about 9-11 primary radii and 2-3 secondary radii in large
anterior field, radii absent in posterior field.

Vertebrae 10 + 16; pterygiophore formula 3(22110).

The following measurements are expressed as thousandths of the SL:
head length, 250; snout length, 54; bony interorbital width, 6; greatest di-
ameter of orbit, 73; upper jaw length, 115; predorsal length, 290; greatest
depth of body, 189; pectoral fin length, 299; pelvic fin length, 245; caudal
fin length, 324; pelvic fin insertion to anal fin origin, 270; first to third spines
of spinous dorsal fin longest and about equal, the second spine slightly
longer, its length 625. :

Head cylindrical; trunk somewhat compressed; interorbital narrow, much
less than diameter of eye; lower jaw equal to upper jaw, the gape wide;
length of jaw extending posteriorly nearly to vertical through posterior mar-
gin of eye; anterior and posterior nares close together, the anterior nares on
a short tube; tongue truncate, the tip free; gill opening moderate, extending
just below and anterior to lower insertion of pectoral fin, not reaching pos-
terior margin of preopercle; first dorsal fin extremely elongate, about five
times higher than second dorsal fin; first dorsal fin far forward, the insertion
above pectoral fin base; first dorsal fin membrane free from second dorsal
fin; origin of second dorsal fin anterior to a vertical through anal opening;

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

966

010N) ‘UBIO ULIPU] UWJd}soM ‘spuUe]s] BURIIWY

X

Ay | OY A FY a TS Te

067S1Z WNSN ‘PyvavId vUSsoYysap

UDA

‘| ‘31

VOLUME 93, NUMBER 4 967

eof
ee

eogveol

e©ecece

/ my

Fig. 2. Diagrammatic lateral view of head and anterior body of Vanderhorstia praealta,
showing cephalic sensory pore and canal system (dashed lines and black spots) and sensory
papillae (small open circles). Interorbital pores not shown. See Lachner and McKinney (1978)
for pore and papillae terminology.

adpressed pectoral fin extends posteriorly to a vertical through anal opening;
pectoral fin longer than deep, the posterior margin round; pelvic fin, when
adpressed, reaches anal opening; pelvic frenum well developed, the inner
pelvic fin rays connected to tip, the rays multibranched; anal fin as high as
second dorsal fin; caudal fin longer than deep, longer than head, the pos-
terior margin pointed; genital papilla bulbous, wider than long.

Teeth in upper jaw caninoid in four rows medially, reduced to two rows
laterally; teeth in outer row about three times longer than those of inner
row; teeth of lower jaw caninoid, in a patch of 3-4 rows medially and two
rows laterally; those on outer row and laterally on inner row longest, about
two times as long as the other teeth of lower jaw; no vomerine or palatine
teeth.

The cutaneous papillae system consists of small structures in a few rows
on the head and trunk (Fig. 2). The important diagnostic papillae consist of
three short horizontal rows on the cheek and a suborbital row beneath the
posterior half of the eye, terminating at the AOT sensory pore. There are
no vertical papillae rows on the cheek. Other obvious rows of papillae are:
a row above margin of upper jaw; short rows on snout anterior to NA pores
and between them; two preoperculo-mandibular rows; a short lateral ce-
phalic row extending posteriorly from the IT pore and a longer, secondary
lateral cephalic row extending posteriorly from about the POT pore to above

968 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the mid-opercular area; a short row extending posteriorly from SOT pore;
a short row on each side of midline of nape; short, scattered vertical rows
above insertion of upper pectoral fin; short, staggered, vertical rows on
trunk, along midline and belly, each row encompassing about 2-3 scales; a
row bordering the lower margin of the sixth and ninth segmented caudal fin
rays and one on the upper margin of the twelfth segmented ray. The three
rows on the opercle common to most gobies are present.

The cephalic sensory pore system consists of the following pores: paired
NA, SOT, AOT, POT, IT and three POP pores, and unpaired AITO and
PITO pores on the midline of the head. There is a deep, transverse groove
immediately behind the eyes.

Color in preservation.—All of head except dorsal portion of occipital area
with numerous, small, pale to white, circular spots over a light brown back-
ground. The pale spots are best developed on snout, lips, cheek and opercle.
Pale spots are also on the breast and base of the pectoral fin. The trunk is
uniformly light brown with fine, dark brown, pepper-like specks. The spi-
nous dorsal fin is mottled in light and dark brown irregular markings; the
lower half is light brown and the outer half darker brown. There are alter-
nating, circular, pale spots and brownish areas on the lower two-thirds of
the first five interradial membranes of the soft dorsal fin; the posterior, lower
two-thirds portion has pale elongate spots in one or two rows near the
midportion of the fin; the remaining membrane and rays are brownish; the
membrane and rays of the outer third of the soft dorsal fin are transparent;
the interradial membrane between the posterior-most two rays is brownish
to black, much darker than the rest of the fin. The interradial membranes
of the anal fin are brownish to black, the rays are light brown. The outer
margin of the anal fin has a narrow light border. The caudal fin has a pale,
elongate mark distally on the upper five branched rays, and two small pale
spots on the upper margin of the procurrent rays; the remainder of the fin
membrane is brownish-black. The membrane between the longest rays in
the middle portion of the caudal fin is darker brown; the rays of the fin are
light brown. The pectoral fin has an alternating series of elongate brown
marks and light areas basally on the rays, and these spots extend about two-
thirds outward on the rays of the central portion of the fin; the membrane
of the fin is pale; the outer half of the upper six rays is pale; the outer third
of the rest of the lower portion of the fin is pale. The pelvic fin rays are pale
and the interradial membranes are brownish black. The first dorsal, second
dorsal, pelvic, anal, and caudal fins are darker than portrayed in Fig. 1.

Etymology.—The specific epithet is taken from the Latin, praealtus, in
reference to the unusually long first four elements of the spinous dorsal fin.

Discussion.—Vanderhorstia praealta superficially resembles two other
species with high spinous dorsal fins, Cryptocentrus pretoriusi Smith
(1959:193, fig. 5) and Vanderhorstia mertensi Klausewitz (1974:210, figs. S—

VOLUME 93, NUMBER 4 969

7). Cryptocentrus pretoriusi has small scales, 100 in the lateral series; the
third dorsal spine is longer than the first and second spines; the dorsal fin
elements number VI-I,11 and the anal fin elements I,11; and the body col-
oration consists mainly of some large light spots on the head and midlaterally
on the trunk. Vanderhorstia mertensi has intermediate sized scales, about
60 in the lateral series; the fourth spine of the spinous dorsal fin is longer
than spines I-III and a large outer portion is free from the interspinous
membrane; the dorsal and anal fins are long, dorsal fin elements number VI-
I,16 and the anal fin elements I,17—18; and the body coloration consists of
moderately sized light spots on the head, nape and on the dorsal fins, and
several weak, dusky bars on the trunk. Vanderhorstia praealta has about
54 scales in the lateral series; spines I-IV are elongate, the tips connected
by the interspinous membrane, and spines I-III are longest and about equal
in length; the soft dorsal and anal fin elements number I,10. The dominant
body coloration consists of small light spots on the head, nape, breast, and
base of pectoral fin; the trunk is uniformly light brown and the fins are
mostly dark colored, with the outer portion of the second dorsal fin and
upper portion of the caudal fin light. Three species referred to the genus
Vanderhorstia by several authors are treated in a key by Klausewitz
(1974:211), namely V. delagoae (Barnard), V. mertensi Klausewitz and V.
ornatissima Smith. Vanderhorstia praealta has a strikingly different color
pattern from these species as well as having certain differences in meristic
characters.

Polunin and Lubbock (1977:92), while working in the Seychelles, ob-
served but did not collect a dark colored “‘Cryptocentrus’’ that had a high
first dorsal fin; they stated that it may have been C. pretoriusi. This species
may have been V. praealta.

Acknowledgments

Figure 1 was rendered by Ms. Keiko H. Moore, NMFS, U.S. National
Museum of Natural History, Washington, D.C. We thank Dr. James E.
Bohlke, Academy of Natural Sciences, Philadelphia, for the exchange of
the holotype resulting from this cooperative, international field collecting
effort. Comparative material of various nominal genera of gobies was also
received from the Academy of Natural Sciences, Philadelphia, and the Field
Museum of Natural History, Chicago. Ms. Susan J. Karnella, U.S. National
Museum of Natural History, provided valuable comments on the manu-
script.

Literature Cited

Hoese, D. F., and R. Steene. 1978. Amblyeleotris randalli, a new species of gobiid fish living
in association with alphaeid shrimps.—Rec. West. Australian Mus. 6(4):379-389.

970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Klausewitz, W. 1974. Fische aus dem Roten Meer. XIV. Eilatia latruncularia n.gen. n.sp.
und Vanderhorstia mertensi n.sp. vom Golf von Aqaba.—Senck. Biol. 55(4/6):205—212.
Lachner, E. A., and J. F. McKinney. 1974. Barbuligobius boehlkei, a new Indo-Pacific genus
and species of Gobiidae (Pisces), with notes on the genera Callogobius and Pipidonia.—

Copeia 1974(4):869-879.

. 1978. A revision of the Indo-Pacific fish genus Gobiopsis with descriptions of four

new species (Pisces: Gobiidae).—Smithsonian Cont. to Zool. 262:1-S2.

Polunin, Nicholas V. C., and Roger Lubbock. 1977. Prawn-associated gobies (Teleostei: Go-
biidae) from the Seychelles, Western Indian Ocean: Systematics and ecology.—J. Zool.,
London (1977)183:63-101.

Smith, J. L. B. 1949. Forty-two fishes new to South Africa, with notes on others.—Ann. Mag.

Nat. Hist. 12(2):97-111.

1959. Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae,

Taenioididae, and Kraemeriidae of the Western Indian Ocean.—Ichthyol. Bull. Rhodes

Univ., 13:185—225.

(EAL) Department of Vertebrate Zoology (Fishes), National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560;
(JFMcK) 102 Gilstrap Drive, Greenville, South Carolina 29609.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 971-981

SYMETHINAE, NEW SUBFAMILY, AND SYMETHIS
GARTHI, NEW SPECIES, AND THE TRANSFER
OF RANINOIDES ECUADORENSIS TO
NOTOSCELES (RANINIDAE:
BRACHYURA: GYMNOPLEURA)

Gary D. Goeke

Abstract.—A new subfamily, Symethinae, of the frog crab family Ran-
inidae proposed for the reception of the genus Symethis, is detailed and
compared with Ranininae in which Symethis formerly had been included.
A Pacific species Symethis garthi is described and compared with the Gulf
of Mexico species Symethis variolosa. The spermathecal pits of the genus
are compared with other genera of Raninidae and illustrated. The taxonomic
status of Raninoides ecuadorensis Rathbun is reviewed and it is transferred
to Notosceles Bourne.

The family Raninidae is a unique group of decapod crustaceans, charac-
terized by exposed thoracic epimera, abdomen incompletely flexed under
the cephalothorax and elongate thoracic nerve ganglionic mass. Other mor-
phological features found within this group have recently been reviewed by
Stevcic (1974). This family of crabs possesses many specialized adaptations
which, combined with a superficial resemblance to albuneid “‘crabs,’’ has
led to a long history of confusion surrounding the systematic position of the
group. They have been alternately classified among most of the major taxa
of brachyuran crabs. This paper deals with the taxonomic status of 2 frog
crabs which occur in the eastern Pacific. :

I wish to express my appreciation to Drs. R. B. Manning (National Mu-
seum of Natural History, Smithsonian Institution), Gilbert Voss (Rosenstiel
School of Marine and Atmospheric Science, University of Miami) and Janet
Haig (Allan Hancock Foundation) for the loan of specimens. I am also
indebted to Dr. Daniele Guinot (Muséum National d’ Histoire Naturelle) for
examining specimens of Cyrtorhina granulosa. Darryl Felder (University
of Southwestern Louisiana), Richard W. Heard (Gulf Coast Research Lab-
oratory), and John S. Garth (Allan Hancock Foundation) kindly reviewed
the manuscript and Linda Lutz aided in preparation of the illustrations.

In a recent attempt to arrive at a satisfactory phylogenetic classification
of the family, Serene and Umali (1972) recognized 2 subfamilies: Notopinae
and Ranininae. Morphological features which define the subfamilies are rel-
ative sizes of the first and second male pleopods and the orientation of the

972 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

orbital sinuses. The crabs of the genus Symethis do not fit into either of the
available subfamilies for the reasons enumerated below. A new subfamily,
Symethinae, subfam. nov., is proposed for the reception of the Atlantic and
Pacific species of Symethis.

Symethis is restricted to American waters and until now has contained a
single species, S. variolosa (Fabricius, 1793), which ranges from North
Carolina south to Bahia, Brazil, and through the Gulf of Mexico. This
species was listed by Rathbun (1937:18) with a geographic range into the
Pacific Ocean along the Panamanian coast. However, examination of spec-
imens collected from that area has shown they represent a previously un-
described species formerly confused with S. variolosa.

Symethinae, subfam. nov.

Diagnosis.—Eyes greatly reduced; spermathecal pits widely separated
with overarching hoods; 7 pairs of gills; palms of cheliped greatly inflated.

Symethis Weber, 1795

Symethis Weber, 1795:92.—Rathbun, 1937:24.
Zanclifer Henderson, 1888:34.—Bourne, 1922:66.

Symethis garthi, sp. nov.
Figs. 1; 2a, b; 3a—c
Symethis variolosa Rathbun, 1937:26, fig. 10, pl. 5, figs. 7, 8 (in part).—
Correa, 1970:10, pl. 5, figs. 38-47, pl. 6, figs. 48-55, pl. 7, figs. 60-61 (in
part).

Diagnosis.—Carapace heavily eroded, hepatic spines blunt, well devel-
oped; abdominal segments coarsely granulate anteriorly; dactyls of ambu-
latory legs 1 and 2 with flattened spine at base of lunate curvature; dorsal
teeth of carapace blunt.

Description.—Carapace eroded, generally oval in shape, convex front to
back and side to side; length of carapace 1.64 times width. Rostrum pro-
duced beyond general outline of carapace into trilobed process with dimin-
utive eyes at base; eyestalks retract slightly into cup-shaped socket formed
by small lateral and ventral outfoldings of carapace and upper margin of
basal antennal article; eyes calcified anteriorly, completely protected and
concealed when retracted. Rostrum deeply eroded medially at base, floor
of depression with numerous raised tubercles. Eroded depressions of car-
apace spread posteriorly and laterally, forming symmetric depressions over
anterior two-thirds of carapace; floor of depressions occasionally with well-
developed tubercles. Fronto-orbital region demarcated by elevated trans-
verse ridge (often interrupted by depressions) which connect hepatic spines;

VOLUME 93, NUMBER 4 973

Fig. 1. Symethis garthi, holotype d, dorsal view.

rostrum lying on lower level than remainder of carapace. Anterolateral mar-
gin of carapace with well developed blunt hepatic spines behind which setae
become evident laterally. Posterolateral margins of carapace bordered by
slight ridge, obvious in dorsal view, terminating posteriorly at base of ab-
domen. Carapace widest about two-thirds of way from front. Abdomen with
7 segments, only slightly deflexed so that 4 or 5 segments are evident in
dorsal view; all segments with very dense setae laterally. First abdominal
segment four-fifths as wide as posterior margin of carapace, ornamented
with coarse granules anteriorly. Remaining somites of abdomen decreasing
in width, all with prominent tubercles; setae forming row dorsally on pos-
terior 5 segments. Penultimate abdominal somite with short, posterolateral
extensions, last somite rounded.

Merus of first leg massive with concave inner border, covered with setae
proximally. Carpus heavy and minutely eroded with small tubercles; distally
with rounded mound of small tubercles forming complex network on bul-
bous palm of chela; fixed finger slightly deflexed with granules forming par-
allel longitudinal rows. Broad dorsal carina extending distally on dorsal
surface of manus, becoming much broader than base of movable finger.
Propodus and fixed finger with numerous large teeth which mesh with tip
of propodus interior to fixed finger. Propodus moderately developed, also
with parallel rows of granules.

Second leg fringed extensively with setae; carpus tuberculate dorsally

974 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with 2 distal outfoldings to form spatulate processes. Propodus short, with
4 spatulate processes on distal margin; dactyl of second leg flat, lunate, with
spine at base. Third leg similar except carpus with 2 small processes and
numerous larger rounded spines; propodus with 1 spatulate process and
series of spines extending out onto posterior extension of segment; dactyl
flat and lunate with spine produced at base. Merus and carpus of fourth leg
dorsally tuberculate; propodus with large curved spatulate process on in-
terior margin, dactyl flat, lunate, fringed with setae. Fifth leg greatly re-
duced, dorsal in position; all segments heavily tuberculate dorsally with
serrate margins; posterior margin of propodus with curved spatulate pro-
cess, dactyl flat, lunate, fringed with setae.

Ischium of third maxilliped greatly elongated, 2.5 times length of merus,
paved with low granules; merus short, slightly longer than wide, covered
with low tubercles except in central region where floor of depression is
clear; palp not evident in ventral view. Exognath of third maxilliped slightly
shorter than ischium. Sub-hepatic region of carapace with eroded depres-
sions, floor of depressions with numerous tubercles, produced anteriorly
into 2 large mounds, | forming lateral margin of pocket surrounding eye.
Antennal peduncle massive, produced into 2 lobes, | projecting ventrally
and | extending anteriorly almost to tip of rostrum. Antennule very small
and not obvious in ventral aspect.

Material examined.—Holotype: USNM 173452, 31 March 1939, M. S.
Stranger, collector, Gulf of California, Espiritu Santo 2, Mexico, 24°26'50”
N, 110°18’W, 11-36 m, male 33.3 mm. Paratypes: USNM 173453, collected
with holotype, 10 males, 10 females, 1 carapace, 13.8-35.0 mm. USNM
155089, 14 April 1939, M. S. Stranger, collector, Gulf of California, Puerto
Refugio, 29°32'17”N, 113°33'50”W, 18-27 m, 3 males, | female, 22.8—25.5
mm. AHF 3513, 6 February 1935, Velero III station No. 458-35; W. L.
Schmitt, collector, Secas Isle, Panama, 7°57'55”N, 82°02’ W; 9-27 m, | male,
13.6 mm.

Etymology.—This species is named in honor of Dr. John S. Garth (Allan
Hancock Foundation), whose work has done much to further crustacean
knowledge.

Remarks.—Symethis garthi and S. variolosa are closely related species
restricted to American waters. In the Gulf of Mexico, S. variolosa occurs
only on substrates of shell and shell hash (Goeke, unpublished data). S.
garthi may also occur most frequently on shell bottoms although specimens
from Panama were collected from substrates of mud and sand. Sediment
information obtained from original collection tag of the type specimen in-
dicates a sand and shell bottom close inshore. Offshore, coral and rocky
bottom is reported; however, it is not known from which bottom type the
material was collected. The very few specimens of S. garthi available sug-
gest highly restricted populations in the Pacific or the species is not easily
collected by usual methods.

VOLUME 93, NUMBER 4 975

The genus Symethis has been placed within the subfamily Ranininae by
Serene and Umali (1972:25) along with Ranina Lamarck, 1801, Lyreidus de
Haan, 1841, Notopoides Henderson, 1888, Raninoides H. Milne-Edwards,
1834, Notosceles Bourne, 1922 and Cyrtorhina Monod, 1956. The charac-
ters which define the subfamily are: eye peduncle folded obliquely or lon-
gitudinally forward; and male pleopod 2 with elongated, tapering shaft short-
er than pleopod 1 (Serene and Umali, 1972:34); apex of male pleopod 2
usually ornamented distally.

Recent work by Hartnoll (1979) has centered on the structure of the sper-
mathecal pits of the raninids and casts some doubt on the validity of the 2
subfamilies. The work has shown no correlation between the subfamilies
Ranininae and Notopinae and spermathecal structure of the females. The
spermathecal pits of this family have been commonly characterized as a
single, unpaired median pit. Gordon (1966) noted the presence in Ranina
and Notopoides of what she thought to be an unpaired spermathecal pit.
Hartnoll (1979) has shown that the spermathecal pits of this group are indeed
paired but often recessed to the bottom of a pit-like depression present in
the anterior part of sternite 7. The depression had mistakenly been described
as a single spermathecal pit. The structure of spermathecal pits in this family
has been thought to form a continuum, beginning with that characterized by
Lyreidus through the form exemplified by Ranilia to Raninoides and No-
tosceles (Hartnoll, 1979:80). However, the spermathecal pits of the genus
Symethis must be inserted within this grouping. It can be modified so that
the progression now proceeds from Symethis, through Lyreidus and Ranilia
to the recessed form of Raninoides or Notosceles.

Of the 10 genera currently recognized within the family Raninidae, the
spermathecal pits have been described in 5 by Hartnoll (1979). Gordon
(1966) illustrated the spermathecae of Notopoides and those of the 4 genera
which remain are described here.

The spermathecae of Ranilia agree closely with those described for Ran-
ina. The sternal shield between pereiopods 3 and 4 is constricted with paired
spermathecae obvious upon close examination. The anterior part of sternite
7 contains a median pit with a trough-like depression proceeding posteriorly.
The anterior walls of the spermathecal depression are steep-sided with the
pits covered by a flexible membrane as in Ranina.

Notosceles conforms with that form of spermatheca described for Ran-
inoides. The spermathecae are not obvious, but dissection shows them to
be located at the bottom of the deep pit-like depression. The pit is circular
in ventral view with steep-sided walls on all sides. No trough is evident
posteriorly and the median depression occupies most of the area of the
sternal shield between pereiopods 3 and 4.

Specimens of the genus Cyrtorhina were not available but at the request
of the author were examined by Dr. Daniele Guinot (Muséum National
d’Histoire Naturelle). She has described a single median pit located on the

976 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

anteriormost section of sternite 7. Similar to that found in Ranina, the pit
is steep-sided anteriorly with indications of a trough-like depression poste-
riorly.

The spermathecae of S. garthi (Fig. 2a) differ greatly from those described
for the other genera of frog crabs. The spermathecal openings are situated
partly on the suture of sternite 7. There is no indication of a median trough
as in Lyreidus or Ranilia and the spermathecal openings are large and
widely separate. Obvious features are the calcified hoods which arch over
the spermathecal openings in such a manner as to make them approachable
only from the posterior. A decalcified flap is present below the “‘hood”’ in
Symethis as in most of the remaining genera. Symethis variolosa from the
Gulf of Mexico agrees very closely with this description in the structure of
the spermathecal pits. Slight differences in the calcified hood over the open-
ing may be of specific value.

In the Gulf of Mexico, females of S. variolosa are often collected with
the spermathecal openings completely occluded by an amorphous plug of
hardened material. In alcohol, the hardened mass completely occupies the
area of the sternal shield between pereiopods 3 and 4 obscuring the sper-
mathecae. This “‘plug’’ effectively seals off the pits and may be the result
of the packing of the spermathecae with sperm packets by the male after
mating. Excess sperm packets transferred during copulation would remain
outside the spermathecal openings and may harden on exposure to sea
water, effectively preventing the female from mating with a second male.
Over 60% of the adult females collected in the Gulf show this condition
(Goeke, unpublished data). Eight of the 11 females of the Pacific species
examined are also plugged in this manner.

Crabs of the genera Cyrtorhina Monod and Symethis Weber resemble
each other and are considered most closely related by Serene and Umali
(1972:49). No evidence was cited by Serene and Umali to support the state-
ment, although the dactyls of the pereiopods and the frontal regions of the
carapaces are very similar. Monod (1956:49) states that Cyrtorhina is closer
to Ranina and details how the genera differ. It is my opinion that the re-
semblance between Cyrtorhina and Symethis is superficial. Considerable
differences are immediately obvious in a comparison of the pleopods and
female spermathecae of Symethis and Cyrtorhina. The first male pleopods
of Cyrtorhina (Monod, 1956:52) are short and stout, much more closely
resembling those of Ranina as figured by Barnard (1950:398) than the very
broad type of pleopod of Symethis (Fig. 3).

The spermathecae of Cyrtorhina and Ranina, as described previously,
are also in close agreement. In both genera, the structure of the spermathe-
cal pits consists of a deep, steep-sided pit in the anterior part of sternite 7.
A depression extends posteriorly which becomes increasingly shallower to-
ward the eighth sternal plate. This form of spermatheca lies about midway

VOLUME 93, NUMBER 4 977

2mm

Fig. 2. Symethis garthi: A, Spermathecal openings, paratype ° (g, genital opening; p3, p4,
pereiopods 3 and 4; so, spermathecal openings; 6, 7, 8, sternites 6, 7, and 8); B, Dactyls of
pereiopods 2-5 (left to right).

978 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in the series of structures found within this family. The spermathecae of
Symethis are described above and can be considered a basically different
form of spermatheca.

As noted by Stevcic (1974) the first male gonopods of the subfamily Ran-
ininae are characteristically ornamented distally. This is well documented
as the male pleopods of all the genera of the subfamily have been illustrated:
Raninoides and Notosceles by Serene and Umali (1972), Lyreidus by Griffin
(1970). Ranina by Barnard (1950), Notopoides by Gordon (1966), Cyrto-
rhina by Monod (1956) and Symethis by Correa (1972) and present paper
(Fig. 3a. b). In all genera of the subfamily except Symethis, the distal part
of the first male pleopod is ornamented with either broad spine-like projec-
tions or complex folds of tissue. In Symethis, however, the first male pleo-
pod has no spines or folds distally. It is highly compressed laterally with
the distal portion accuminate. The abundant tufts of setae on the anterior
and posterior margins almost obscure the sharply tapering tip. This char-
acter supports the separation of the genus Symethis from the remaining
genera of the subfamily.

Perhaps the most astonishing difference between Symethis and the other
genera of the family is a difference in the number of gills. In Symethis the
number of pairs of gills is only 7, all of which lie in the normal vertical
position, with the anteriormost pair greatly reduced. In the remaining genera
of frog crabs the gills number 8 pairs. Raninoides louisianensis Rathbun
possesses 8 pairs of gills as does Lyreidus, Ranilia and Notosceles. In these
forms, the anteriormost 2 pairs of gills are reduced in size but are large
enough to be easily observed. The anterior 2 pairs are most likely still
effective in oxygen exchange.

The reduced number of gills present in the genus Symethis represents a
major departure from the remaining genera of the family Raninidae. This
feature is of considerable taxonomic value and is a conservative character
often used in the diagnosis of families. Much importance is attached to the
respiratory mechanisms in this family and was stressed by Bourne (1922) in
his account of the physiology of this group. Very little information is avail-
able on the life histories and ecology of the raninids: however, it is probable
that the reduced number of gills is an adaptation to the niche occupied by
this genus. Stevcic (1974) stated the reduced number of gills in this and
other groups of crabs (i.e., Calappidae, Leucosiidae) is a modification for
burrowing.

The differences in the number of gills and the spermathecae are sufficient
to warrant the establishment of the proposed subfamily Symethinae. Future
research may show that the subfamily deserves elevation to the rank of
family. Subsequent work on this group should concentrate on the larval
development of Symethis. The structure of the nervous system and fossil

VOLUME 93, NUMBER 4 979

—_—
=_
—

A |
AGHA AY
OAR
VR |
YARN y I)
Wee WH
\ i 7]

Fig. 3. Symethis garthi: A, First pleopod; B, Second pleopod: C, Lateral view of anterior
part of carapace of holotype ¢.

history should be examined for support of the separation of this genus from
the others within the family.

Notosceles ecuadorensis Rathbun, nov. comb.

The genus Notosceles was erected by Bourne (1922) for N. chimmonis
in his account of the physiology of the family Raninidae. Other species
subsequently assigned to the genus are N. serratifrons (Henderson, 1893),
and N. viadari Ward, 1942. In establishing the genus, Bourne (1922) listed
many characters which he used to differentiate Notosceles from the closely
related Raninoides. Some features he listed were later questioned by Cho-
pra (1933) who doubted the validity of the genus and suggested that it should
be considered a synonym of Raninoides. Serene and Umali’s (1972) work
on the Philippine raninids dealt with this matter and attempted to define
characters separating the 2 genera. For this study, Notosceles ecuadorensis
was compared with 2 species of Raninoides: the type-species of the genus,

980 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

R. loevis; and R. louisianensis. Some of the following characters are those
recognized by Bourne (1922) and others were utilized by Serene and Umali
(1972). All represent characters that can be of value to separate Notosceles
from Raninoides: 1) anterior end of carapace roughly granulate; 2) eyes of
Notosceles shorter and stouter; 3) fronto-orbital width half of the extraor-
bital width of the carapace; 4) eyes slightly longer than rostrum; 5) no spine
on ischium of cheliped; 6) truncate third maxilliped in Notosceles with
shorter ischium to merus length than Raninoides (1.58 in N. ecuadorensis
and 1.33 in R. loevis); 7) propodus of cheliped with double crested carina
dorsally; 8) sternal shield not produced into acute spine between bases of
pereiopods 1 and 2; 9) rostrum pointed with 2 flat lateral teeth at base; 10)
first abdominal tergum approximating width of posterior margin of carapace,
with second shield about as broad; 11) fifth pereiopod less massive than that
of Raninoides; and 12) basal segment of the antennular peduncles broader
than in Raninoides.

Raninoides ecuadorensis was described from material gathered off La
Plata Island, Ecuador in 82-101 m. Originally included within the genus
Raninoides Milne-Edwards, a recent examination of type-material has
shown the species would be more properly included in the genus Notosceles
Bourne, 1922. The material examined consisted of the type male (USNM
#69319) and two lots of specimens collected from the type-locality (USNM
#69320 and 81933). Supplemental material from the University of Miami
(32:3437) was also examined. Raninoides ecuadorensis more closely agrees
with the generic description of Notosceles given by Bourne (1922) and
amended by Serene and Umali (1972). It is here designated Notosceles
ecuadorensis (Rathbun, 1935), nov. comb.

With the transfer of R. ecuadorensis to Notosceles, 5 species of the genus
Raninoides are known to occur in American waters. Four of the species are
present in the western Atlantic with a single species, R. benedicti Rathbun,
1933, in the eastern Pacific. Two additional species, R. fossor A. Milne-
Edwards and R. nitidus A. Milne-Edwards, 1880, have been recently syn-
onymized and removed from species known to occur in American waters.
Manning (1975) has shown R. fossor to be a junior synonym of Notosceles
chimmonis Bourne, 1922, and Goeke (1980) removed R. nitidus to the
genus Lyreidus as a synonym of L. bairdii Smith, 1881.

Literature Cited

Barnard, K. H. 1950. Descriptive catalogue of South African decapod crustacea.—Ann. South
African Mus. 38: 1-837.

Bourne, G. C. 1922. The Raninidae: A study in carcinology.—J. Linn. Soc. Zool. 35:25-—75.

Chopra, B. 1933. On two new species of oxystomatous crabs from the Bay of Bengal. Further
notes on the Crustacea Decapoda in the Indian Museum, IV.—Rec. Ind. Mus. 35:77—
87.

VOLUME 93, NUMBER 4 981

Correa, M. M. G. 1970. Crustaceos Braquiuros brasiluros da familia Raninidae.—Boletin do
Museu Nacional no. 276:1-21.

Goeke, G. E. 1980. Range extensions of six western Atlantic frog crabs (Gymnopleura;
Raninidae) with notes on the taxonomic status of Lyreidus bairdii.—Proc. Biol. Soc.
Wash. 93(1):145-152.

Gordon, I. 1966. On the spermatheca in the Raninidae (Crustacea: Decapoda).—Jn H. Barnes,
ed., Some contemporary studies in marine sciences.

Griffin, D. J.G. 1970. A revision of the Recent Indo-West Pacific species of the genus Lyreidus
De Haan.—Trans. Roy. Soc. New Zealand, Biol. Sci. 12(10):89-112.

Hartnoll, R. G. 1979. The phyletic implications of spermathecal structure in the Raninidae
(Decapoda: Brachyura).—J. Zool., Lond. 187:75-83.

Henderson, J. R. 1888. Report on the Anomura collected by H. M. S. Challenger during the
years 1863—1876.—Rept. Scientific Results Explor. Voyage H. M. S. Challenger 27(1):1-
VIA

Manning, R. B. 1975. The identity of Raninoides fossor A. Milne-Edwards and Bouvier, 1923
(Decapoda).—Crustaceana 29:297-298.

Monod, Th. 1956. Hippidae et Brachyura ouest-africains.—Mem. I.F.A.N. 45:1-674, 1-884.

Rathbun, M. J. 1937. The oxystomatous and allied crabs of America.—Bull. U.S. Nat. Mus.
No. 166, 272 pp.

Serene, R., and A. Umali. 1972. The family Raninidae and other new and rare species of
brachyuran decapods from the Philippines and adjacent regions.—Philippine J. Sci.
99:21-105.

Stevcic, Z. 1974. The systematic position of the family Raninidae.—Syst. Zool. 22:625-632.

Barry A. Vittor & Associates, Inc., 8100 Cottage Hill Road, Mobile,
Alabama 36609.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 982-988

PARAMUNNA QUADRATIFRONS , NEW SPECIES,
THE FIRST RECORD OF THE GENUS IN THE
NORTH PACIFIC OCEAN (CRUSTACEA:
ISOPODA: PLEUROGONIIDAE)

Ernest W. Iverson and George D. Wilson

Abstract.—Paramunna quadratifrons n. sp., described from the southern
California borderland, represents the first record of the genus in the north
Pacific Ocean.

The asellote isopod genus Paramunna is predominantly a southern hemi-
sphere Antarctic and sub-Antarctic genus with a few north Atlantic Ocean
representatives. Most records of Paramunna are from intermediate depths
of the continental shelf and slope regions while a few species occur in the
deep sea. Pacific Ocean records were previously limited to those reported
by Menzies (1962) from off southern Chile. The species described here is
the first Paramunna reported from the north Pacific Ocean. This single
record is not surprising in view of the relative rarity of pleurogoniids in both
population sizes and number of species in the northern hemisphere (Wilson,
1980).

This paper is based on material collected during the second year benthic
survey of the southern California borderland (Fauchald and Jones, 1978) in
connection with the Southern California Baseline Studies and Analysis pro-
gram (FY 1977). All specimens were collected from a single spade corer
sample (0.16 m? surface area) taken at a depth of 197 meters on North-West
Tanner Bank, approximately 130 kilometers southwest of Los Angeles, Cal-
ifornia. Drawings were made with the aid of camera lucida and checked for
detail at magnifications up to 430 power.

Pleurogoniidae Nordenstam, 1933, sensu Wilson (1980)
Paramunna G. O. Sars, 1866

Type-species.—Paramunna bilobata Sars, 1866.

Diagnosis.—Body broad, flat, tergal plates extending laterally well be-
yond coxae. Cephalon often with anterior protuberances: knobs, spines, or
flattened curved plates. Lateral margin of pleotelson always with spines.
Antenna article 3 with spines or knobs. Mandibular molar distally truncate.

Remarks.—This diagnosis should be regarded as preliminary until the
genus is properly revised. However, the above concept is more restricted
than the one currently recognized in the literature (e.g., Menzies, 1962).

VOLUME 93, NUMBER 4 983

Fig. 1. Paramunna quadratifrons: A, Male holotype; B, Female allotype.

Consequently, some species should probably be returned to Austrimunna
(Wilson, 1980).

Paramunna quadratifrons, new species
Figs. 1-4

Material examined.—Station 81832, 197 m, 33°53.24’N, 119°23.35’W, 4
males and 3 females from a light brown, coarse sand and shell sediment,
collected 24 August 1978 by the R/V Thompson. Holotype male 1.23 mm
body length (USNM 173915), allotype female 1.29 mm body length with 4
eggs (USNM 173916) deposited in the National Museum of Natural History,
Smithsonian Institution, Washington, D.C. Paratypes have been deposited
as follows: 1 male (CASIZ 015902) and 1 female (CASIZ 015903), California
Academy of Sciences, San Francisco, California, and 2 males and | female
(AHF 781), Allan Hancock Foundation, University of Southern California,
Los Angeles, California.

984 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Paramunna quadratifrons, male: A, Lateral habitus; B, Lateral margin of cephalon
with antennula, antenna and ocular peduncle; C, Ocular peduncle; D, Right uropod; E, Ce-
phalon, anterior oblique view, cuticular structures shown only on rostral lobe.

Diagnosis.—Cephalon anterior margin expanded into large plate-like
shelf, somewhat concave dorsally, anterolateral angles rounded, more ro-
bust in male. Eyes absent, ocular peduncles very reduced. Lateral margins
of pereonites heavily denticulate, a small hooked seta dorsally near apex of
each pereonite. Antenna third article with 4—5 (typically 4) teeth dorsally.

Supplemental description.—Body broadest anteriorly at pereonite 2, ta-
pering posteriorly. Pereonite 1 more robust in male compared to female,

VOLUME 93, NUMBER 4 985

Fig. 3. Paramunna quadratifrons, female: A, Pereopod 1; B, Inner margins of pereopod 1
dactylus and propodus. Male: C, Pereopod 1; D, Pereopod 1 dactylus; E, Pereopod 7.

lateral margin truncate; pereonites 2—4 also sexually dimorphic, noticeably
lengthened in ovigerous female, laterally truncate, in male acutely pointed;
pereonites 5-7 laterally acute in both sexes. Pleotelson pyriform, somewhat
concave at point of uropod attachment; apex rounded.

986 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Paramunna quadratifrons: A, Maxilliped; B, Second maxilla; C, First maxilla; D,
Left mandible; E, Incisor process and lacinia mobilis of same; F, Male pleopod 1; G, Male
pleopod 2.

Antennula with 6 articles; third, fifth and sixth articles subequal in length,
fourth shortest; terminal article tipped with several setae and single long
aesthetasc. Antenna flagellum with 6-8 articles.

Maxilliped: Endite with 2 coupling hooks, palp articles 2 and 3 much
broader than articles 4 and S, distal article about three-fourths length of
fourth article. First maxilla outer lobe apically with 8 stout setae, inner lobe
apically with 2 long and 2 short setae. Second maxilla bilobed, inner lobe
with 4 apical and 2 subapical setae; inner lappet of outer lobe with 4 setae,
outer lappet with 2.

Left mandible: Incisor process and lacinia mobilis each with 4 teeth, setal
row with 4 simple setae; molar process cylindrical, distally truncate; distal
article of palp short, about half length of middle article, with 2 short setae
apically.

VOLUME 93, NUMBER 4 987

Pereopod 1 strongly subchelate, more robust in male; claw of dactylus
chisel-like, propodus inner margin armed with about 4 small unequally bifid
setae, carpus armed with 2 large unequally bifid setae. Pereopods 1-3 with
denticles on bases. Pereopods 2-7 similar, not notably lengthened poste-
riorly.

Male first pleopod sagittate, medially fused for about three-fourths length,
distal tip broadened laterally into bifurcate lateral corners each with tuft of
several short setae, apex posteriorly pointed. Uropods biramous, basal ar-
ticle absent, inserted on lateral margin.

Remarks.—The extreme reduction of the eyes and ocular peduncles in-
dicates that this species has its evolutionary roots in the deep sea (Hessler
et al., 1979). It is likely that this species or its ancestors migrated through
the deep sea from the Antarctic center of pleurogoniid diversity (Wilson,
1980) to its present location. By this reasoning Paramunna quadratifrons
is an emergent species.

The fact that seven individuals were collected in only one of the 8 replicate
cores taken at the North-West Tanner Bank station suggests a very patchy
distribution. Isopods collected in the same spade core sample with Para-
munna quadratifrons include: Ilyarachna acarina Menzies and Barnard, an
upper continental slope representative of a predominately deep-sea genus,
and Silophasma geminatum (Menzies and Barnard), a continental shelf
species which is widespread throughout the southern California bight. The
pleurogoniid, Pleurogonium californiense Menzies was collected in another
replicate.

Etymology.—The specific name quadratifrons is a combination of the
Latin words quadratus meaning squared and frons meaning forehead, re-
ferring to the anterior margin of the cephalon.

Acknowledgments

This study was, in part, supported by the Southern California Baseline
Studies and Analysis program (FY 1977) funded by the Bureau of Land
Management (U.S. Department of the Interior), contract number AA550-
CT6-40 with Science Applications, Inc., La Jolla, California. This is con-
tribution No. 378 of the Allan Hancock Foundation.

Literature Cited

Fauchald, K., and G. F. Jones. 1978. Variation in community structure of shelf, slope and
basin macrofaunal communities of the southern California bight.—Jn Final report on
the baseline study, 1977-78, prepared by Science Applications, Inc. for the Bureau of
Land Management. 2(19):1—-167.

Hessler, R. R., G. D. Wilson, and D. Thistle. 1979. The deep-sea isopods: a biogeographic
and phylogenetic overview.—Sarsia 64(1—2):67—75.

Menzies, R. J. 1962. The zoogeography, ecology, and systematics of the Chilean marine

988 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

isopods. Reports of the Lund University Chile Expedition 1948—49:42.—Lunds Univ.
Arsskr., N.F. Avd. 2, 57(11):1-162.
Sars, G. O. 1866. Beretning omeni Sommeren 1865 foretagen zoologisk Reise ved kysterne
af christianias og christiansands stifter—Nyt. Mag. Naturvid. Christiania 15:84—-128.
Wilson, G. D. 1980. New insights into the colonization of the deep sea: Systematics and
zoogeography of the Munnidae and the Pleurogoniidae nov. comb. (Isopoda: Janiro-
idea).—Journal of Natural History 14:215-236.

(EWI) Allan Hancock Foundation, University of Southern California,
University Park, Los Angeles, California 90007; (GDW) A-002 Scripps In-
stitution of Oceanography, La Jolla, California 92093.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 989-996

PARAPERCIS DIPLOSPILUS (PISCES:
MUGILOIDIDAE), A NEW SPECIES
FROM THE PHILIPPINE ISLANDS

Janet R. Gomon

Abstract.—Parapercis diplospilus n. sp., from the Visayan Sea between
northern Negros and Masbate islands, is most similar to Parapercis om-
matura Jordan and Snyder, 1902, from which it differs most notably in
having two prominent spots on the caudal-fin base instead of one, 22 to 23
instead of 23 to 26 scale rows around the caudal peduncle, and 9 to 11
instead of 12 to 13 scale rows below the lateral line. It also differs in certain
body proportions and in having the caudal fin truncate with its dorsalmost
rays slightly prolonged rather than rounded as in P. ommatura.

In 1978, the Smithsonian Institution’s expedition to the Philippine Islands
collected 21 specimens of an undescribed species of Parapercis. This
species is herein described and compared with P. ommatura, to which it is
most similar. P. ommatura is known from Japan, Korea, and China.

Cantwell (1964) last revised the Indo-Pacific Parapercis, in which he treat-
ed 26 species. Schultz (1968) added six more: four newly described, one he
had described in 1966, and one described by Kamohara (1960). He failed to
include P. elongata Fourmanoir (1965) and P. guezei Fourmanoir (1966).!
Since that time, four additional Indo-Pacific species have been described:
P. dockinsi McCosker (1971), P. gushikeni Yoshino (1975), P. biordinis
Allen (1976), and P. cephalus Kotthaus (1977). Current work by G. Stroud
and J. E. Randall is expected to change the status of some of these species
and add new ones.

Methods

Methods and terminology are those of Hubbs and Lagler (1958) except
for the following as modified by Cantwell (1964): in the pectoral fin the
unbranched, dorsalmost ray is indicated by a lower case Roman numeral
and the branched rays by Arabic numerals; each dorsal- and anal-fin ray
with a separate external base was counted as one; lateral-line scale counts

' Parapercis elongata was first described in 1965 under the name Parapercis sp., although
referred to as P. elongata in the text of the description and in the index. Fourmanoir rede-
scribed the species in 1967.

990 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

include all pored scales in the series; counts of scale rows above the lateral
line were begun at the origin of the first segmented dorsal-fin ray; scale
counts below the lateral line were made along an oblique row from the
anterior margin of the anus dorsoposteriorly to the lateral line. Vertebral
counts, not including the urostylar vertebra, and median-fin ray counts were
taken from radiographs. Fleshy orbit diameter is a horizontal measurement.
Measurements (to the nearest 0.1 mm) were made with needlepoint dial
calipers or with an ocular micrometer.

Type material has been deposited in the following collections: British
Museum (Natural History), London (BMNH); Australian Museum, Sydney
(AMS); Bernice P. Bishop Museum, Honolulu (BPBM); California Academy
of Sciences, San Francisco (CAS); National Museum of Natural History,
Smithsonian Institution, Washington, D.C. (USNM).

Parapercis diplospilus, new species
Figs. 1, 2

Holotype.—USNM 220470 (75.0 mm SL), Philippine Islands, Visayan Sea
between northern Negros and Masbate islands, southeast of Sicogon Island,
11°22'00’N, 123°19'48”E, 38.4 meters, 9 June 1978, Smithsonian Philippines
Expedition 1978, sta. T-29, L. Knapp and party.

Paratypes.—All collected by the Smithsonian Philippines Expedition
1978, L. Knapp and party, in the Visayan Sea between northern Negros
and Masbate islands. BPBM 22770 (3, 56.6-—78.0), AMS I.21362-001 (3, 64.0-
76.9), BMNH 1979.11.2.1-3 (3, 56.9-73.4), CAS 44715 (3, 61.7-70.1), all
same data as holotype; USNM 220406 (1, 56.5), southeast of South Gigante
Island, 11°30’23"N, 123°23’45”E, 38.4 meters, 8 June 1978, sta. T-27; USNM
220407 (1, 48.7), east of South Gigante Island, 11°31'38”N, 123°31'00’E, 38.4
meters, 8 June 1978, sta. T-26; USNM 220409 (6, 51.6-66.6), east of Sicogon
Island, 11°27’4S’N, 123°23'45”E, 47.6 meters, 4 June 1978, sta. T-3.

Diagnosis.—A species of Parapercis with: palatine teeth absent; usually
7 or 8 teeth in outer row of lower jaw; dorsal-fin rays V,22 (rarely VI,22);
fourth dorsal-fin spine longest (fifth in a specimen with six spines), 6.4—7.8%
SL; last dorsal-fin spine connected by membrane to base of first dorsal-fin
ray (Fig. 2); anal-fin rays I,18; pectoral-fin rays i,13-i,15; scales below lateral
line 9-11; scale rows around caudal peduncle 22-23; two prominent dark
spots on caudal base, the upper one appearing ocellated in most specimens,
the lower of equal size or slightly longer, but not ocellated.

Description.—Meristic values for the holotype are indicated by asterisks
with the number of specimens having a given value placed in parentheses;
morphometric values are expressed as percentages of SL, with values for
the holotype given in parentheses after the ranges. The range of SL is 48.7
to 78.0 mm.

VOLUME 93, NUMBER 4 991

Fig. 1. Parapercis diplospilus n. sp., holotype, USNM 220470, 75.0 mm SL.

Dorsal fin V,22*(20) or VI,22(1); anal fin I,18; principal caudal rays 8 +
7; pectoral fin i1,13*(7), 1,14(13) or 1,15(1); pelvic fin I,5; vertebrae 10 +
19*(19) or 10 + 18(1) (one additional specimen abnormal with some verte-
brae fused); gill rakers 8(3), 9*(5), 10(10), 11(2) or 12(1); lateral-line scales
58(3), 59*(8) or 60(6); scales above lateral line 3(1) or 4*(16); scales below
lateral line 9-11*; scale rows around caudal peduncle 22—23*.

Body elongate, greatest depth, at about level of anal opening, 14.5—-17.6
(17.1); least depth of caudal peduncle 7.7-8.5 (7.9); tip of lower jaw to anal
origin 39.6—43.5 (41.7). Head length 26.4—28.7 (26.4); snout length to fleshy
orbit 6.5—7.1 (6.5); fleshy orbit diameter 6.0-7.9 (6.0); least fleshy interor-
bital width 2.2—2.5 (2.5); mouth oblique, lower jaw projecting beyond upper;
maxilla reaching slightly posterior to a vertical through anterior edge of
orbit; snout tip to rear edge of maxilla 7.2—9.1 (8.4); preopercle with 1-12
small spines on margin of angular portion; posterior margin of operculum
with 1 strong spine at upper angle, not covered by skin posteriorly, and 11-
21 small spines along ventral margin. Dorsal-fin base 62.0—68.0 (65.2); dor-
sal-fin spines increasing in length to fourth spine, fourth spine 6.4—7.8 (6.4),
length of fifth spine about equal to third; dorsal-fin rays longer than spines;
anal-fin base 45.3—49.9 (49.2); anal-fin origin approximately below fifth seg-
mented dorsal-fin ray; posterior anal-fin base below posterior base of dorsal
fin, appressed tips of posterior rays of both fins reaching almost to caudal-
fin base; caudal fin truncate, uppermost rays slightly prolonged; pectoral fin
acutely rounded, length 17.3-—20.0 (18.8), reaching to or slightly posterior to
vertical through anal-fin origin; pelvic fin pointed, fourth ray longest, reach-
ing vertical through anus or between anus and anal-fin origin, fin length
19.9-23.6 (19.9).

Based on a Student’s t test for allometry at the 95% level of significance,
fleshy orbit diameter and length of fourth dorsal spine are negatively allo-
metric. Greatest body depth is positively allometric. The remaining mor-
phometric characters are isometric.

Teeth in 2 series on both jaws. Outer series of upper jaw a single row of

992 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Semidiagrammatic drawing of dor-
sal-fin spine arrangement of Parapercis diplo-
spilus.

large, curved, conical teeth decreasing in size posteriorly; inner series a
band of dense villiform teeth with depth of band decreasing posteriorly.
Usually 8 (6-11) large, curved, conical teeth in outer row of lower jaw; inner
series a band of villiform teeth mesially and a single row of small conical
teeth laterally. Palatine teeth absent. Vomer with a chevron-shaped patch
of small conical teeth in 2 to 3 rows.

Scales ctenoid; small scales extending onto basal half of caudal fin and
basal sixth of pectoral fin in some specimens; snout, interorbital space, and
occiput naked.

Color in preservation.—Background color of body and head yellowish to
tan; free edges of body scales and scale pockets above lateral line with dark
brown melanophores giving a tesselated, overall dark appearance to upper
half of body; six or seven indistinct V-shaped markings of a darker brown
also on upper half of body, relatively evenly spaced between head and
caudal base, the more anterior markings extending slightly below lateral
line; ventral half of body paler, with a row of eight dark-brown spots alter-
nating small and large in size, in some specimens these spots extending
ventrad to anal fin as faint vertical or oblique bars; a row of smaller, paler,
and less distinct spots slightly above larger spots and just below lateral line;
faint vertical bar extending from axilla to ventral body midline in some
specimens; anal opening with U-shaped ring of dense, dark brown mela-
nophores. Scaled portion of dorsal head surface with same tesselated ap-
pearance as upper body; cheek and operculum with 2 to 3 irregular dark-
brown blotches; upper naked areas of head with small brown spots; lips and
chin with scattered dark-brown melanophores, upper lip darkest mesially;
lower surface of head, branchiostegal membranes, and isthmus without me-
lanophores. Spinous dorsal fin with large, diffuse, dark spots on membranes

VOLUME 93, NUMBER 4 993

forming a band between first and fifth spines, membranes between first and
third spines almost entirely covered with melanophores; soft dorsal fin with
2 to 3 rows of elongate, dark spots on membranes; anal fin with dark me-
lanophores on anterior halves of membranes between consecutive rays, pos-
terior portions of membranes without melanophores; caudal-fin base with
two prominent, dark spots, each approximately pupil size, the upper spot
appearing ocellated in some specimens, the lower spot often slightly more
elongate and non-ocellate; very faint, irregular, narrow, dark bars posterior
to prominent, dark spots; tips of mesial caudal-fin rays in some specimens,
and all caudal-fin ray tips in others, dark brown; pectoral fin with scattered
large, dark, melanophores at distal margin, fin base with an oblique dark
bar; pelvic fin dark except for spine and distal margin.

Reproduction.—Hermaphroditism in mugiloidids was first suggested by
Marshall (1950). He believed P. hexophthalma and P. polyophthalma to be
males and females, respectively, of the same species, with males invariably
being larger than females. One specimen intermediate in color pattern be-
tween the hexophthalma and polyophthalma forms appeared to have both
Ovarian and testicular tissue. G. Stroud (in litt.) has confirmed this sugges-
tion of protogynous hermaphroditism, and along with J. E. Randall (in litt.)
indicated that other species of Parapercis also reverse sex.

Gross examination of the gonads of the 21 type specimens (48.7-78.0 mm
SL) of P. diplospilus showed all to have well developed ova. Examination
of serial histological sections from the gonad of a 78 mm specimen (the only
specimen sectioned) revealed the presence of both ovarian and testicular
tissue with spermatids. Although examination of a series of specimens is
needed for final determination of sexual mode, it is possible that P. diplo-
spilus represents another example of protogynous hermaphroditism in Par-
apercis.

Distribution.—Known only from the type specimens collected by trawl
in the Visayan Sea, Philippine Islands, at 38.4 to 47.6 meters depth. Scuba
and rotenone collecting efforts to the south (Mindanao Sea and Tonon Strait)
and to the east (Cuyo Islands) of the Visayan Sea failed to take this species.

Etymology.—The specific name is a noun in apposition taken from the
Greek diplo, double, and spilos, spot, in reference to the two prominent
caudal-fin spots.

Comparisons.—Parapercis diplospilus keys to P. ommatura Jordan and
Snyder (1902), in Kamohara’s (1960) review of Japanese parapercids, in
Cantwell’s (1964) revision of the genus, and in Schultz’s (1968) review con-
taining an expanded, modified key. Diagnostic characters shared by these
two species are: dentition, as described above; spinous dorsal fin with mid-
dle rays longest, last spine connected by membrane to base of first seg-
mented dorsal-fin ray (Fig. 2); caudal vertebrae 19; dorsal fin V,22; anal fin
1,18 (rarely 1,19 in P. ommatura); pectoral fin i,13 or i,14 (rarely i,15); dark,

994 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ss
2
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z
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S =
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SL SL
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50 60 70 80 90 100
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Fig. 3. Relationship between standard length and (A) 4th dorsal-fin spine length, (B) length
from tip of lower jaw to anus, (C) snout length and fleshy interorbital width, (D) greatest body
depth, in Parapercis diplospilus (closed circles) and P. ommatura Jordan and Snyder (open
circles).

VOLUME 93, NUMBER 4 995

ocellate spot at upper caudal-fin base, and 2 to 3 rows of elongate spots on
membrane between segmented dorsal rays. Parapercis diplospilus and P.
ommatura also have similar lateral-line scale counts (58-60 in P. diplospilus,
58-61 in P. ommatura), number of scales above lateral line (usually 4), and
gill-raker counts (8-12 in P. diplospilus, 9-13 in P. ommatura).

The following characters serve to distinguish P. diplospilus from P. om-
matura: prominent dark spot on lower caudal-fin base (absent in P. om-
matura); blotches on cheek (versus 2 stripes); 11-21 small spines along
posteroventral edge of operculum (versus 1—18, usually 1-11, though spines
may be worn down); scale rows around caudal peduncle 22-23 (versus 23-
26); scales below lateral line 9-11 (versus 12-13); caudal fin truncate, upper
rays slightly longer (versus broadly rounded). In body proportions, P. dip-
lospilus has a lower spinous dorsal fin, fourth dorsal-fin spine length 6.4—
7.8% SL (versus 8.1-9.6%; Fig. 3A), shorter length from tip of lower jaw
to anus (39.6—43.5% versus 43.5—47.5%; Fig. 3B), and shorter snout length
(6.5—7.1% versus 6.4-8.2%; Fig. 3C, upper). Greatest body depth and least
fleshy interorbital width are smaller in P. diplospilus than in P. ommatura
at small sizes, but P. ommatura is more slender with a narrow interorbital
space at larger sizes (Fig. 3D and C, lower). Analysis of covariance of the
linear regressions using the BMD10V program indicates significant or highly
significant differences between the two species in all except greatest body
depth, in which the regressions are curvilinear.

Comparative Material

Parapercis ommatura.—JAPAN: Nagasaki, USNM 179803 (5, 56.6—75.3
mm, paratypes), USNM 50260 (3, 61.1—79.0, paratypes); Tsuruga, USNM
50258 (3, 73.2-92.7); Toba Mkt., USNM 151813 (2, 72.9-76.6); Tokyo,
USNM 50261 (2, 75.7-89.0). KOREA: USNM 37776 (2, 75.6-79.0).
CHINA: USNM 6867 (3, specimens damaged).

Acknowledgments

I thank G. Stroud (James Cook University of North Queensland), J. E.
Randall (BPBM), and R. H. Gibbs and V. G. Springer (USNM) for critically
reviewing an earlier draft of the paper. Susan Karnella photographed the
holotype. Serial histological sections were prepared by Margaret Melville.

Literature Cited

Allen, G. R. 1976. Descriptions of three new fishes from Western Australia.—J. Roy. Soc.
Western Australia 59(1):24—30.

Cantwell, G. E. 1964. A revision of the genus Parapercis, Family Mugiloididae.—Pac. Sci.
18(3):239-280.

996 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fourmanoir, P. 1965. Liste complementaire des poissons de Nha-Trang.—Cahiers

O.R.S.T.O.M., Oceanogr., Contrib. 84:1-114.

. 1966. Trois especes nouvelles de poissons perciformes du Vietnam et de la Reunion.—

Bull. Mus. Nat. d’Hist. Nat. 38(3):217-222.

. 1967. Sur cing nouvelles especes de poissons du Vietnam.—Bull. Mus. Nat. d’Hist.

Nat., 2nd Ser. 39(2):267-274.

Hubbs, C. L., and K. F. Lagler. 1958. Fishes of the Great Lakes Region.—Univ. Michigan
Press, Ann Arbor, pp. i-xv, 1-135.

Jordan, D. S., and J. O. Snyder. 1902. A review of the trachinoid fishes and their supposed
allies found in the waters of Japan.—Proc. U.S. Nat. Mus. 24:461-497.

Kamohara, T. 1960. A review of the fishes of the family Parapercidae found in the waters of
Japan.—Rep. Usa Mar. Biol. Sta. 7(2):1-14, 2 pls.

Kotthaus, A. 1977. Fische des Indischen Ozeans. A. Systematischer Teil, X VIII. Percomorphi
(8).—*‘Meteor’’ Forsch.-Ergebnisse, Reihe D, no. 24:37-S53.

Marshall, N. B. 1950. Fishes from the Cocos-Keeling Islands.—Bull. Raffles Mus. 22:166—-
205.

McCosker, J. E. 1971. A new species of Parapercis (Pisces: Mugiloididae) from the Juan
Fernandez Islands.—Copeia 1971(4):682—686, 3 figs., 2 tables.

Schultz, L. P. 1966. Parapercis kamoharai (family Mugiloididae), a new fish from Japan with

notes on other species of the genus.—Smithsonian Misc. Coll. 151(4):1-4, 1 pl.

. 1968. Four new fishes of the genus Parapercis with notes on other species from the
Indo-Pacific area (family Mugiloididae).—Proc. U.S. Nat. Mus. 124:1-16.

Yoshino, T. 1975. Parapercis gushikeni, a new mugiloid fish from the Ryukyu Islands.—Publ.
Seto Mar. Biol. Lab. 22(5):343-346.

Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 997-1015

THE SOUTH AMERICAN FISH GENUS RACHOVISCUS,
WITH A DESCRIPTION OF A NEW SPECIES
(TELEOSTEI: CHARACIDAE)

Stanley H. Weitzman and Carlos Alberto Goncalves da Cruz

Abstract.—Rachoviscus graciliceps, a new species of characid fish, is
described from small coastal blackwater streams near Prado, Bahia, Brazil.
The relationships of this species appear close to Rachoviscus crassiceps
Myers (1926) which is redescribed on the basis of the types and new spec-
imens from small blackwater streams near the Atlantic coast 50 to 60 km
south of Paranagua, Parana, Brazil. The new species, R. graciliceps appears
to be less derived than R. crassiceps. The type locality of R. crassiceps,
the neighborhood of Rio de Janeiro, is discussed and found to be question-
able. Rachoviscus is redefined; its relationships to other characids remain
obscure.

Rachoviscus Myers (1926) was established for two aquarium specimens
of a new species, Rachoviscus crassiceps Myers (1926), sent to Myers by
Arthur Rachow of Germany. The specimens were said to have been im-
ported from the neighborhood of Rio de Janeiro but no specimens with
verified locality information have ever been recorded. All the later aquarium
reports cited below seem to be based on the account of Rachow in Holly
Meinken, and Rachow (1939). The species remained known only from the
specimens examined by Myers and from Rachow’s account until September
of 1975 when Persio de Santos Filho, a student at the Universidade de Sao
Paulo, collected specimens about 50 km south of Paranagua, Parana, Brazil
in small blackwater streams and ponds emptying into the Atlantic Ocean.
In December of that same year, two small specimens of Rachoviscus cras-
siceps were obtained by N. Menezes and W. L. Fink from a small black-
water stream near the Atlantic Ocean not far from the locality visited by de
Santos Filho. In spite of attempts to collect additional specimens, none was
found. At that time many of the coastal blackwater streams and ponds were
dry, including those visited by de Santos Filho. The two specimens, a pair,
were brought alive to Washington, D.C. where they grew to adults but never
spawned. Both were preserved after surviving three and one half years in
aquaria and are now USNM 220732.

The new species, Rachoviscus graciliceps, was collected by Carlos Cruz
in October 1977 from a small blackwater stream near the Atlantic Ocean
about one km north of the town of Prado in Bahia, Brazil. Live specimens
of this species were brought to Rio de Janeiro where they spawned in aquaria.

998 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The coastal blackwater streams of eastern Brazil have been little collected
and the distributions here reported for these two species may be greatly
extended in the future.

The methods used here for counting and measuring specimens were those
described for characoids by Fink and Weitzman (1974). All morphometric
values in the descriptions are expressed as a percentage of standard length
(SL) except where otherwise designated. The value for the holotype or
lectotype is given first followed by values for the remaining males in paren-
theses ( ) and for the females in brackets [ ]. In some cases of meristic
values, only a series of figures, in parentheses, is given after the holotype;
these include counts for both males and females. Specimens have been
deposited in the following museums: California Academy of Sciences (CAS)
(note, SU following CAS means the number is a former Stanford University
Natural History Museum number with the specimen now deposited at CAS),
Museu de Zoologia da Universidade de Sao Paulo (MZUSP), Museu Na-
cional do Rio de Janeiro (MN), and the National Museum of Natural His-
tory, Smithsonian Institution (USNM).

Rachoviscus Myers

Rachoviscus Myers, 1926:1, original description, type by monotypy Ra-
choviscus crassiceps Myers 1926.

Diagnosis.—The following series of characters will distinguish Rachov-
iscus from all other genera of the Characidae: pelvic fins 1,5 with only second
and third branched rays of males bearing bony hooks or spines on ventral
surface of fin; one bony spine to each ray segment that bears a spine.

Rachoviscus Myers

Rachoviscus Myers, 1926:1, original description, type by monotypy Ra-
choviscus crassiceps Myers 1926.

Diagnosis.—The following series of characters will distinguish Rachov-
iscus from all other genera of the Characidae: pelvic fins 1,5 with only second
and third branched rays of males bearing bony hooks or spines on ventral
surface of fin; one bony spine to each ray segment that bears a spine. Other
important characters for Rachoviscus are as follows. Premaxillary teeth in
two rows, teeth of outer row one or two, unicuspid or tricuspid. Those of
inner row five to six and mostly tricuspid, some bicuspid or unicuspid. Teeth
of these two rows sometimes closely approximate, and giving appearance
of a single undulating row of teeth. Lateral line incomplete, of about four
to five perforated scales placed anteriorly. Maxillary bone set with teeth for
about one-third to two-thirds of its length. These teeth unicuspid or anterior
first to third teeth may be tricuspid, others unicuspid. Sub- and postorbital

VOLUME 93, NUMBER 4 999

area nearly completely covered by four infraorbital bones. Third infraorbital
(=great or second suborbital of Eigenmann) with its ventral border in con-
tact with preopercle. In life both known species have bright red adipose fins
in both sexes and the distal one-quarter to one-third of anal fin pigmented
with a stripe of red or yellow, especially in males.

Discussion.—The relationships of this genus to other characid genera are
obscure. Myers (1926) originally placed it in the nominal subfamily Chei-
rodontinae because he believed the type species had a single row of pre-
maxillary teeth. Myers further suggested the relationships might be with
Prionobrama Fowler.

Géry (1977:347) placed Prionobrama, Rachoviscus, Paragoniates Stein-
dachner, Leptagoniates Boulenger, Xenagoniates Myers, and Phenagon-
iates Eigenmann and Wilson in a group because they are reported to have
a ‘“‘very compressed, usually elongate body with a long anal fin.’’ He further
stated that ‘“‘they usually have a single series of teeth on the upper jaw.”’
Géry placed Paragoniates, Rachoviscus, and Prionobrama in a single group
in his key because they share the following characters: the presence of an
adipose fin, a relatively short body (compared to the other genera in his
Paragoniatinae), an incomplete lateral line and less than 50 anal-fin rays. He
associated the genera Prionobrama and Rachoviscus in his key by their
common posession of 29 to 37 anal-fin rays, 10 to 16 maxillary teeth, and
35 to 41 scales. Géry was uncertain about this placement and remarked that
both of these genera could be placed in the Aphyocharacinae.

The problem of the putative relationships among these six genera needs
much study and will be considered here only as it may concern the rela-
tionships of Rachoviscus. The important questions that must be considered
here are as follows. What are the unique characters shared by the species
of Rachoviscus that can used to define the genus and phylogenetically relate
the species? The same question must be asked for Prionobrama and Par-
agoniates. Finally are there any shared derived characters that might indi-
cate a relationship between Rachoviscus and Prionobrama, or between
Rachoviscus and Paragoniates?

The characters utilized by Géry (1977:347) are difficult to evaluate from
a phylogenetic point of view. One character utilized by Géry, the posession
of an adipose fin, can be eliminated for phylogenetic analysis because it is
almost undoubtedly primitive for the family. Outgroup comparison of the
Characidae with other characoids and the Otophysi as a whole shows that
this character is not unique to the Characidae. The character probably oc-
curred in the family’s ancestor and therefore its presence cannot be used to
relate genera or species in monophyletic lines within the family. The loss
of the adipose fin appears to have occurred independently several times
within the family making its absence difficult to use in phylogenetic analysis.

The other characters as listed by Géry (1977) occur commonly in diverse

1000 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

groups of characids. Some of these characters, such as the reduced lateral-
line count and a single row of premaxillary teeth, are “‘simple’’ loss char-
acters which may have evolved independently and repetitively in characids.
Attention to morphological detail of jaw modification and tooth arrangement
may, in some cases at least, make a reduction in tooth rows useful in phy-
logenetic analysis. Other characters used by Géry are apparently genetically
labile characters such as fin-ray counts and tooth counts, or body shape.
The primitive versus derived nature of these kinds of characters is very
difficult and perhaps often impossible to evaluate since the advanced versus
plesiomorphic polarities of similar appearing characters may or may not be
the same in these instances of labile characters. Therefore, these characters
are often useless for phylogenetic analysis except in certain specific cases
wherein a large series of such characters may correlate in their distributions
among a series of taxa when processed by outgroup comparison and par-
simony.

The species of Rachoviscus seem related by the unique pelvic-fin struc-
ture and color pattern described above in the generic description. The two
species of Prionobrama, P. filigera Cope and P. paraguayensis Eigenmann,
seem related to each other by a large series of characters none of which is
unique to these two species but which occur nowhere else in this particular
combination, producing a series of correlated synapomorphies that undoubt-
edly has real phylogenetic significance. They have large, white-tipped an-
terior anal-fin lobes and elongate strong pectoral fins with large pectoral-fin
muscles and pectoral girdle. The dorsal-fin origin and anal-fin origin are
nearly approximate or the dorsal-fin origin is a little anterior to the anal-fin
origin. They have an elongate, compressed body with an elongate anal fin
of 29-37 branched anal-fin rays, a nearly fully toothed maxillary bone with
mostly unicuspid teeth, a single series of tricuspid teeth on the premaxillary
bone, an incomplete lateral line and an oblique mouth upturned distally. A
few of these derived characters were used by Myers (1926) to relate Prion-
obrama and Rachoviscus. They were stated to share a distally upturned
mouth, a well toothed maxillary, an incomplete lateral line, and a single row
of premaxillary teeth. New information shows that Rachoviscus has two
rows of premaxillary teeth and that a well toothed maxillary occurs only in
one species, R. crassiceps, the most derived. We believe than many more
than the two remaining shared characters, a distally upturned mouth and an
incomplete lateral line, are needed if Prionobrama and Rachoviscus are to
be considered close relatives. This is especially convincing since both of
these characters occur commonly in characoids of no apparently close phy-
logenetic relationship to either of these genera. We fail to find unique syn-
apomorphic characters uniting Rachoviscus and Prionobrama more closely
to each other than to any other characids.

The problem of a possible relationship of Rachoviscus with Paragoniates

VOLUME 93, NUMBER 4 1001

4 Salvador

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NY

Vitonia

“Rio de Janeiro

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Si Asie :

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é

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‘

ONCE

Joinville

y
ae He ae reported for Rachoviscus graciliceps and Rachoviscus crassiceps. Black
disk within two circles, type locality of R. graciliceps, Prado, Bahia. Black star, putative type
locality for R. crassiceps, neighborhood of Rio de Janeiro; see text. Black star within two

circles, new records for R. crassiceps, near Guarituba and Brejotuba, Parana. All localities are
in Brazil.

1002 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

is more complex but involves much the same kinds of considerations. Par-
agoniates iS a very compressed deep-bodied, rather elongate fish with a long
anal fin of about 34-48 rays, according to Géry (1977). It has a combination
of a distally upturned mouth and long slender jaw bones of a shape found
nowhere else in characoids. This is associated with the two tooth rows of
the premaxillary in at least some populations of Paragoniates being com-
pressed together, producing an appearance somewhat similar to that of the
premaxillary tooth rows in Rachoviscus crassiceps. The premaxillary tooth
rows of Paragoniates were reported as one row by Eigenmann (1915), but
two specimens of Paragoniates sp. from the Rio Tiznados, Guarico, Ven-
ezuela (MBUCV-V-7108), have one or two teeth in an outer row which are
more or less pressed against the inner main row, giving the impression of
a single undulating row. Six specimens of Paragoniates alburnus Stein-
dachner from the Rio Pachitea at Porto Inca, Peru (FMNH 83874) appear
to have a short inner row of two teeth instead of an outer row of two teeth.
The main row of teeth is present as a continuous outer row. Paragoniates
has a short lateral line of 13-16 perforated scales in the specimens recorded
above. The dorsal fin originates well posterior to the origin of the anal fin
and the pectoral fins are elongate, with enlarged pectoral-fin muscles and
girdles. Paragoniates has a deep, short caudal peduncle relative to most
characid species in genera that are presumably relatively primitive such as
Brycon Miller and Troschel, Astyanax Baird and Girard, and Moenkhausia
Eigenmann. The Venezuelan specimens of Paragoniates reported above
have the pelvic-fin rays 1,6 whereas the Peruvian population sample had 1,7,
the usual count for characids, including the primitive genera in the family.

Of the apomorphic characters recorded above for Paragoniates, Rachov-
iscus has a relatively deep caudal peduncle (especially in the more derived
Species, indicating an origin independent from that of Paragoniates), a re-
duced lateral line, and a distally upturned mouth. All these characters
occur commonly elsewhere in the Characidae and probably represent in-
dependent derivations for these two genera. The trend toward a reduction
in pelvic-fin rays in Paragoniates is undoubtedly independent of the reduc-
tion in Rachoviscus, and although both genera appear to have a trend to-
ward compression of premaxillary tooth rows, in detail the premaxillary
bones are very different. Furthermore, it is only the more derived species
of Rachoviscus that has this character.

Gery (1977:347) suggests that Rachoviscus might be allied to Aphyochar-
ax Gunther, but discussed no evidence for this opinion. Although all species
of Aphyocharax appear to have their premaxillary teeth in a single row,
there is much morphological divergence in the jaws of these fishes (Eigen-
mann, 1915). We could find no synapomorphic character between species
of Aphyocharax and Rachoviscus. Eigenmann’s definition of Aphyocharax
is extensive, but most of the characters listed are either primitive for the

VOLUME 93, NUMBER 4 1003

Characidae or, when derived, are found in a number of apparently more or
less remotely related, diverse characid genera. An examination was made
of the pelvic fins of eight morphologically diverse species of Aphyocharax
from localities in Paraguay, Argentina, Brazil, Bolivia and Venezuela. These
specimens (USNM numbers 220869, 220870, 220871, 220872, 220873,
220874, 220875 and 220878) are unidentified because no recent and adequate
study of the species and their relationships is available. In all those species
that had no pelvic-fin spines, the males had an 1,7 pelvic-fin ray count. Of
those that had i,7 or 1,6 pelvic-fin rays, some males had spines on all the fin
rays except the anterior undivided ray (one species), spines on all soft rays
(one species), on all soft rays except the last, or on the first, second, third
and fourth soft rays but not on the three terminal soft rays. All specimens
that had spines in the pelvic fin had the spines extending ventrally. The
pelvic fins of the species of Aphyocharax examined are not remarkably
different from those found in many other characids and do not show any
synapomorphies with Rachoviscus. A more complete evaluation of the pos-
sible relationship of Aphyocharax and Rachoviscus must await a detailed
phylogenetic study of the species of Aphyocharax.

This discussion indicates that the relationships of Rachoviscus within the
Characidae remain unknown. This genus was placed with the *‘cheirodon-
tin’ characids by Myers (1926) following the traditional hypothesis estab-
lished by Eigenmann (1915) that all small characoids with a single row of
premaxillary teeth are phylogenetically related in a single characid subfam-
ily, the Cheirodontinae. The phylogenetic unity of the Cheirodontinae was
challenged by Fink and Weitzman (1974). In the present study, Rachoviscus
was found to have two rows of premaxillary teeth, a fact which would place
it in the characid subfamily Tetragonopterinae, following a traditional con-
cept best expressed by Eigenmann (1917). The phylogenetic relationships
among the tetragonopterin characids are essentially unknown and unstudied
and the group is probably a paraphyletic assemblage or a series of paraphy-
letic assemblages with undetermined relationships.

Rachoviscus graciliceps, new species
Figs. 1-2

Holotype.—MZUSP 14387, SL 44.4 mm, Brazil, State of Bahia; taken
from one of three small creeks about 1 km north of Prado and about 500
meters from Atlantic Ocean, 39°14’W, 17°19’S, 18 October 1977 by Carlos
Alberto Gongalves da Cruz.

Paratypes.—3, MZUSP 14388, 14389, and 14390, SL 39.8-42.9 mm; 2,
USNM 220355, 40.3-47.6 mm; both lots with same data as holotype. 1
additional specimen, USNM 220355, SL 35.1 mm, is young of specimens
collected with holotype. 2, MN 10585 and 10586, SL 33.2-37.2 mm, col-

1004 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lected at same locality as holotype, 1-4 December 1978 by E. Izecksohn,
O. L. Peixoto and C. A. G. da Cruz.

Diagnosis.—This species differs markedly from R. crassiceps in shape of
head and caudal peduncle, configuration of premaxillary and maxillary teeth
as well as in number of scale rows around caudal peduncle and predorsal
scale count. Rachoviscus graciliceps has a more slender head, interorbital
width quite variable but 29.1-35.0% of head length (n = 5); R. crassiceps
has an interorbital width 37.5-43.8% of head length (n = 12). The caudal
peduncle of R. graciliceps is much more slender than that of R. crassiceps.
Caudal peduncle length of R. graciliceps nearly same as caudal depth
length, 93.0-100% of caudal peduncle depth; In R. crassiceps, caudal pe-
duncle length is 62.7—74.1% of caudal peduncle depth. Rachoviscus gracil-
iceps has 14 scale rows around caudal peduncle; R. crassiceps has 18.
Predorsal scale count in R. graciliceps (n = 5) 14 to 16, * = 15, SD =
0.707, and in R. crassiceps (n = 11) 17 to 19, * = 18.2, SD = 0.751. Lateral
series scale counts not significantly different; when tested with a two-tailed
Student’s ¢ test using square root transformations to compute value of f,
t = 1.88 and P = 0.04. In R. graciliceps (n = 5), always 35 scales in a
lateral series and R. crassiceps (n = 11) 34-39 scales, « = 36.5, SD =
1.695.

Premaxillary tooth rows of R. crassiceps “*‘compressed’’ together (see
description below) but remain separate in R. graciliceps as in most other
characids with 2 premaxillary tooth rows. Rachoviscus graciliceps with
fewer teeth (3-8) on maxillary, occurring on about anterior one-fourth
to one-third of its length, whereas more (8-14) teeth occur along about one-
half to two-thirds of that length in R. crassiceps.

Description.—Morphometric values based on 2 males and 3 females un-
less otherwise designated. Specimens spawned and raised in aquaria were
not utilized in taking morphometric or meristic values. Eight specimens with
locality data were in good enough condition for some of the meristic values.
Body moderately compressed, relatively elongate, greatest depth most often
at dorsal-fin origin, occasionally anterior to that origin. Greatest depth 35.5
(32.4—35.5, ¢ = 34.0) [32.0-35.2, « = 33.7]. Depth at dorsal-fin origin same
as greatest depth in males, in females [31.7—-34.0, « = 32.6]. Predorsal body
profile slightly convex, slightly concave over nape, continuing anteriorly to
dorsal to eye. Snout slightly convex. Body profile along base of dorsal fin
slightly convex, nearly straight between posterior dorsal-fin termination and
adipose fin. Posterior to adipose fin, body profile slightly convex up to origin
of anterior procurrent rays of dorsal lobe of caudal fin. Dorsal-fin origin
about equidistant between snout tip and caudal-fin base or slightly nearer
to the latter. Distance between snout tip and dorsal-fin origin 54.1 (53.2—
54.1, * = 53.7) [S2.6—-53.8, ¢ = 53.3]. Distance between dorsal-fin origin and
caudal-fin base 50.9 (50.9-51.3, * = 51.1) [49.7-5S0.4, * = 50.2]. Distance

VOLUME 93, NUMBER 4 1005

Fig. 2. Above: Rachoviscus graciliceps, new species, MZUSP 14387, SL 44.4 mm, holo-
type, male. Below: Rachoviscus graciliceps, MZUSP 14389, SL 42.9 mm, paratype, female.
Both specimens from 1 of 3 small blackwater streams about one km north of Prado, Bahia,
Brazil, 18 October 1977.

between posterior border of eye and dorsal-fin origin as,a percentage of
distance between dorsal-fin origin and caudal-fin base 89.7 (82.0-89.7, « =
85.9) [83.4-85.4, * = 84.2]. Ventral body profile moderately convex from
symphysis of lower jaw to posterior termination of anal-fin base. Distance
between snout tip and pectoral-fin origin 27.7 (26.9-27.7, * = 27.3) [26.3-
26.6, < = 26.5]. Distance between snout tip and pelvic-fin origin 49.5 (48.5—
49.5, x = 49.0) [48.7-50.5, x = 49.4]. Distance between snout tip and anal-
fin origin 59.9 (57.1-59.9, x = 58.5) [56.3-63.8, « = 60.2]. Caudal peduncle
depth 14.2 (12.6-14.2, « = 13.4) [12.3-13.3, * = 12.9]. Caudal peduncle
fengthels=o1(1276—13-5, « — 13,3 =13-25 x — 1226)

Head deep, relatively short, bony head length 25.0 (23.9-25.0, « = 24.5)
[23.3-25.9, x = 24.6]. Snout moderately acute, not blunt, lower jaw pro-
truding slightly beyond upper jaw. Mouth gape angled ventrally. Posterior
ventral border of maxillary bone reaching to or somewhat beyond a vertical

1006 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

line drawn ventrally from anterior border of pupil of eye. Horizontal eye
diameter 31.5% (31.5-32.5, x = 32.0) [28.2-34.0, x = 30.8] of bony head
length. Snouth length 26.1% (26.1-27.2, x = 26.7) [21.4-27.0, « = 24.6] of
bony head length. Least bony interorbital width 33.3% (30.7-33.3, « = 32.0)
[29.1-35.0, « = 31.8] of bony head length.

Dorsal-fin rays 11,9 in all specimens (last ray not split to its base). Dorsal-
fin length (=dorsal-fin origin to distal tip of longest ray when fin adpressed
to back) 25.0 (25.0-25.2, « = 25.1) [23.1-25.6, = 24.0]. Distal margin of
dorsal fin strongly convex in both sexes (Fig. 2).

Adipose fin present. Anal-fin rays iv,26 (iv,24—-iv,25) [iv,23 in one and
iv,24 in two]. Last anal-fin ray split to its base. Margin of anal fin slightly
concave, nearly straight (Fig. 2). Pectoral-fin rays 1,13 in all specimens.
Posterior tip of longest ray not reaching pelvic-fin origin in either males or
females.

Anal-fin spines of a male specimen, SL 47.6 mm, USNM 220355 as fol-
lows: second branched ray through twenty-fifth branched ray with spines,
these about five in number for about anterior 12 fin rays, gradually dimin-
ishing in number to one spine at twenty-fifth anal-fin ray. One spine for each
side of each ray segment in all cases. Spines in a continuous series with
distal 5 or 6 ray segments free of spines. All spines extend laterally and
curve somewhat dorsally. Females without anal-fin spines.

Caudal fin 10/9 in all specimens, its dorsal and ventral lobes equal in
length. Caudal fin and dorsal fin without bony hooks.

Pelvic-fin rays 1,5 in all specimens. Pelvic fin length 12.2 (12.2-12.6, x =
12.4) [12.1-12.8, ¥ = 12.4]. Second branched pelvic-fin ray of a male spec-
imen, SL 47.6 mm, USNM 220355, with 11 thick conical spines with their
apices pointing medially. Third branched pelvic-fin ray of this specimen
bears 8 similar spines. All these spines confined to ventral surface of fin.
Females without pelvic-fin spines.

Scales cycloid. Lateral line incomplete, perforated lateral-line scales 5
(3 or 5 in males, 5 in all females). Scales in a lateral series 35 in all specimens.
Scale rows between dorsal-fin origin and pelvic-fin origin 15 in all speci-
mens. Predorsal scales 16 (14-16, « = 15 in all specimens).

Premaxillary teeth in 2 very distinct rows, outer row teeth tricuspid and
1 to 2 in number, not placed far back and pressed against or between inner
row teeth. Outer row teeth lie anterior to space between first and second,
and second and third inner row teeth. Inner row teeth 5 in all specimens,
all tricuspid. Medial tooth with a very small medial cusp. Maxillary bone
with 3-8 teeth along approximately anterior one-fourth to one-third of its
total length. Anterior 1 to 3 teeth tricuspid, others unicuspid. Dentary with
a single row of teeth. Anterior 5 teeth large and tricuspid; posterior 5 to 9
teeth unicuspid and small. Small specimens with fewer dentary teeth than
large specimens.

VOLUME 93, NUMBER 4 1007

Total vertebrae including Weberian apparatus and terminal complex cen-
trum (35-36, « = 35.4, n = 8). Gill rakers 7/10 in holotype, (6—7 on upper
limb, 10-11 on lower limb, never more than a total of 17 or less than 16
rakers, t = 16.6, n = 8).

Color in alcohol.—Body and head a pale brown, immaculate, shading to
dark borwn along dorsum and to white on belly. Cheeks and opercle pale
brown. Top of head and snout dark brown. Color much like that of R.
crassiceps except none of specimens at hand are nearly black or very dark
brown. Fins hyaline except for dorsal and anal fins. Dorsal fin with a row
of dark pigment spots between first or second through fourth or fifth fin
rays. This pigment located in about mid-length of these rays and producing
a horizontal dark line across fin (Fig. 2, above, MZUSP 14387). A similar
line of dark pigment occurs along length of anal fin at about distal two-thirds
of fin-ray length from base of fin.

Color in life. —This color description is taken from color slides of speci-
mens kept in aquaria. Pigment distribution and colors very similar to those
in R. crassiceps. Back olive brown with a considerable amount of yellow.
Sides of body silver but with lemon yellow in abdominal area. Caudal pe-
duncle and area dorsal to anal fin reflect a light pink silvery color. Caudal
fin greenish lemon yellow, tips of lobes, especially ventral lobe, pale red,
sometimes tipped white. Adipose fin deep red, especially in male. Dorsal
fin white to greenish or reddish white, distal to dark streak or line across its
middle length, hyaline ventral to that streak. Anal fin with distal one-quarter
to one-third of its length a dusky red or sometimes yellow, especially in
males. Basal portion of anal fin hyaline. Pelvic fins of males with a distal
reddish or yellowish spot, otherwise hyaline. Pectoral fins hyaline.

Sexual dimorphism.—Sexual differences not as apparent in R. graciliceps
as in R. crassiceps. Females a little paler in life colors than males, and red
of adipose fins likely to be less extensive. Difference in caudal-peduncle
depth noted below for males and females in R. crassiceps appears absent
or at least not as obvious in R. graciliceps. Males have this depth 12.6 to
14.2, x = 13.4% of standard length, females 12.3 to 13.3, « = 12.9%. Only
males have hooks on pelvic and anal fins.

Etymology.—From Latin gracilia, slender or thin, and ceps, head. The
name is used in reference to the fact that this species has a more slender
head than R. crassiceps.

Rachoviscus crassiceps Myers
Figs. 1, 3, 4

Rachoviscus crassiceps Myers, 1926:389, original description, figure, Brazil,
neighborhood of Rio de Janeiro.—Rachow, 1928:18, aquarium descrip-
tion, figure.—Arnold and Ahl, 1936:115, aquarium description, figure.—

1008 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Rachoviscus crassiceps, USNM 220732, SL 38.5 mm, male, small blackwater stream
just south of Guarituba, Parana, Brazil, 28 December 1975. Bent fin rays of caudal, dorsal,
anal, and pectoral fins are due to regrowth from damage in an aquarium.

Rachow in Holly et al., 1939:284, restatement of original description,
aquarium description, figure, breeding habits in aquaria.—Sterba,
1959:120, aquarium description, figure.—Géry, 1977:350, rediscovery
near Rio de Janeiro; reference actually to specimens (USNM 220732)
recorded below from near Guarituba, Parana, Brazil.

Comments.—This incomplete synonymy lists the major aquarium publi-
cations in which this fish has appeared. This fish has never been accorded
systematic treatment subsequent to its original description. The species is
little known to aquarists and to systematic ichthyologists. All of the above
aquarium reports appear to have been taken from Myers (1926), Rachow
(1928) and Rachow in Holly et al. (1939). The last reference contains a brief
comment on its original importation as an aquarium fish into Germany. The
stated type locality can be considered questionable. The fish was first im-
ported by M. Gregor of Hamburg in 1926 from the *‘Umgelbung von Rio de
Janeiro.’ At that time the German aquarium import trade was receiving
fishes from coastal or near coastal cities in southern Brazil such as Rio de
Janeiro, Santos, Paranagua and Porto Alegre. There is no certain record of
this species from anywhere except the two localities cited below, both with-
in 10 kilometers of Guarituba, a small town about 50 km south of Paranagua.
Although this species may occur (or may have occurred) in small blackwater
streams as far as or further north than Rio de Janeiro, there is no firm

VOLUME 93, NUMBER 4 1009

evidence that the type locality is correct. The fish could have been imported
from Paranagua and transshipped to Rio de Janeiro before shipment to Ger-
many.

Material examined.—Lectotype, USNM 92971, 32.7 mm SL, male, pelvic
rays with hooks; lectotype here selected from two syntypes originally in
collection of G. S. Myers, number 86. Second original syntype, 1, CAS(SU)
18146, 28.1 mm SL, same locality data as lectotype. USNM 220732, 2, SL
34.1 mm and 38.5 mm, Brazil, State of Parana, small blackwater stream just
south of Guarituba, about 25 to 50 meters from Atlantic Ocean, 24°37’W,
25°55'S, collected by N. Menezes and W. L. Fink, 28 December 1975.
MZUSP 14635, 4, SL 21.8—30.5 mm, largest a male, others females, Brazil,
State of Parana, stream pond at beach of Brejotuba, 10 km south of Guar-
ituba (24°37’W, 25°55’S), collected by Persio de Santos Filho, September
1975. USNM 220756, 2, SL 21.5—24.4 mm, females, same locality data as
MZUSP 14635. USNM 220757, 2, SL 23.7 and 29.5 mm, both cleared and
stained with alizarin, both with same locality data as MZUSP 14635.

Diagnosis.—See diagnosis above under R. graciliceps.

Description.—Morphometric values based on 4 males and 7 females un-
less otherwise designated. Body moderately compressed, relatively deep,
greatest depth usually anterior to dorsal-fin origin 38.5 (36.7—40.0, « = 38.6)
[37.5—39.3, « = 38.5]. Depth at dorsal-fin origin 37.0 (36.7-39.0, « = 37.8)
[35.8-38.5, *« = 37.1]. Predorsal body profile slightly convex, somewhat
concave at nape and again convex over eye and snout. Base of dorsal fin
and body profile from base of dorsal fin to origin of adipose fin origin nearly
straight. Body profile posterior to adipose fin straight to origin of caudal fin
at beginning of anterior procurrent caudal-fin rays. Dorsal-fin origin nearer
to caudal-fin base than to snout tip. Distance between snout tip and dorsal-
fin origin 55.7 (55.1—58.7, « = 56.3) [S6.2-59.5, x = 58.1]. Distance between
dorsal-fin origin and caudal-fin base 49.2 (48.6—49.8, « = 49.1) [45.4-50.8,
X = 47.6]. Distance between posterior border of eye and dorsal-fin origin as
a percentage of distance between dorsal-fin origin and caudal-fin base 94.4
(93.3-97.3, « = 95.1) [90.7-102.0, « = 97.1]. Ventral body profile moder-
ately convex from symphysis of lower jaw to anterior procurrent fin ray of
caudal fin. Distance between snout tip and pectoral-fin origin 32.4 (26.5-
32.4, x = 28.8) [26.6—28.4, x = 27.8]. Distance between snout tip and pel-
vic-fin origin 51.7 (49.1-51.7, « = 50.0) [49.2-53.5, « = 50.9]. Distance be-
tween snout tip and anal-fin origin 62.1 (60.3-62.1, = 61.1) [S9.8-65.1, =
62.4]. Caudal peduncle depth 16.5 (15.3-16.5, « = 15.9) [13.5-15.0, % =
14.2]. Caudal peduncle length 11.3 (9.9-11.3, « = 10.7) [9.4-10.6, « = 10.0].

Head deep, short; bony head length 24.2 (24.1—25.1, * = 24.5) [23.5-28.4,
X = 25.5]. Snout blunt, lower jaw protruding beyond upper jaw. Mouth gape
angled posteroventrally. Posterior ventral border of maxillary bone reaching
to or somewhat beyond a vertical line drawn ventrally from anterior border

1010 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Rachoviscus crassiceps, USNM 220732, SL uncertain at time of photograph, live
adult male, same specimen as Fig. 3. Note black color of adipose fin, due to its deep red color
in life.

of pupil of eye. Horizontal eye diameter 30.5% (30.5-33.3, x = 32.0) [32.8-
37.9, x = 35.3] of bony head length. Snout length 17.1% (16.9-20.4, x =
17.9) [11.9-16.3, « = 14.1] of bony head length. Least bony interorbital
width 39.0% (38.7-42.3, « = 40.5) [37.5-43.8, x = 40.4] of bony head
length.

Dorsal-fin rays 11,9 in all specimens (last ray not split to its base). Dorsal-
fin length (=dorsal-fin origin to distal tip of longest ray when fin adpressed
to back) damaged (31.2—33.1, * = 32.1, n = 3) [29.3-32.1, « = 30.4]. Distal
margin of dorsal fin convex in both sexes (Figs. 3, 4).

Adipose fin present. Anal-fin rays iv,25 (iv,25 in two, iv,27 in one, and
iv,28 in one, males) [iv,25 in one, iv,26 in one, iv,27 in three, and iv,28 in
two, females]. Last anal-fin ray split to its base. Anterior distal margin of
anal fin convex, posterior distal margin concave (Fig. 4).

Anal-fin spines as follows on a male alizarin preparation, SL 29.5 mm,
USNM 220757. Second branched anal-fin ray first to bear spines. This ray
and next 8 rays bear 5 spines on each ray, | for each ray segment that bears
spines. Spines and their respective segments occur consecutively 6 or 7
segments from distal end of fin rays. Spines occur only on posterior seg-
ments of branched portions of rays when they occur on branched portions
of rays. Eleventh branched ray bears 4 spines, twelfth and thirteenth
branched rays with 3 spines and fourteenth with 1 spine. Each spine rela-
tively short, conical and extends dorsally and laterally. In a large, old male

VOLUME 93, NUMBER 4 1011

specimen, SL 38.5 mm, USNM 220732, about 17 rays bear spines. First
anal-fin ray to bear spines is posteriormost unbranched ray. Number of
spines per ray about same as in smaller specimen but spines extend in a
more lateral direction than in smaller specimen. Spines occur on both sides
of anal fin. Females without anal-fin spines.

Pectoral-fin rays 1,14 (i,12 in one, 1,13 in three) [i,12 in one, 1,13 in three,
i,14 in two, and i,15 in one]. Posterior tip of longest pectoral-fin ray reaching
beyond pelvic-fin origin. Pectoral-fin length 24.2 (21.8-24.6, « = 23.3) [21.1-
24.2, x = 22.2]. Distal pelvic-fin tip reaching to or somewhat beyond anal-
fin origin in both sexes. Pelvic-fin rays 1,5 in all specimens.

Pelvic-fin length 14.7 (11.7-14.7, « = 13.6) [12.4-14.2, « = 13.3]. Second
branched ray of pelvic fin bears 8 spines and third ray bears 6 spines in a
male specimen, SL 29.5 mm, USNM 220757. These spines relatively elon-
gate, curved, and conical, with their sharp apices directed medially. These
spines only occur on ventral surface of fin. A male specimen, SL 38.5 mm,
USNM 220732, bears 11 spines on its second branched pelvic-fin ray and
9 spines on its third branched ray. Females without pelvic-fin spines.

Caudal fin 10/9 in all specimens, its dorsal and ventral lobes equal in
length. Caudal fin, dorsal fin, and pectoral fin without bony hooks.

Scales cycloid. Lateral line incomplete, perforated lateral-line scales 5,
(4-S, x = 4.5, n = 12, in both sexes). Scales in a lateral series 36 (33-39,
x = 36.5, n = 12, in both sexes). Scale rows between dorsal-fin origin and
pelvic-fin origin 15 (14-15, « = 14.4, n = 12, in both sexes). Predorsal scales
18 (17-19, « = 18.2, n = 12, in both sexes).

Premaxillary teeth in 2 rows. Outer row teeth unicuspid and 1 to 2 in
number. Usually these teeth placed far back, against and nearly between
inner row teeth, causing Myers (1926) to describe premaxillary teeth as
occurring in a single row. Occasionally an outer row tooth well forward of
inner row teeth. Outer row teeth lie between first and second inner row
teeth and/or second and third inner row teeth. Inner row teeth 5 to 6, me-
dialmost bicuspid or tricuspid but with a very tiny medial cusp. Proceeding
laterally and posteriorly next 1 to 3 teeth tricuspid or sometimes | or 2 of
these bicuspid or unicuspid. Posteriormost 1 to 2 teeth usually unicuspid,
occasionally bi- or tricuspid. Maxillary bone with 8 to 14 mostly unicuspid
teeth, occasionally anteriormost tooth tricuspid. Teeth occur along about
anterior one-half to nearly two-thirds of maxillary length. Dentary with a
single row of teeth. Anterior 5 (sometimes only 4) teeth large and tricuspid;
posterior 6 to 8 teeth small and unicuspid. Ectopterygoid and palatine with-
out teeth.

Total vertebrae including Weberian apparatus and terminal complex cen-
trum 34-36, x = 35.4, n = 10. Gill rakers 7/10 in lectotype, 3—7 on upper
limb, 10-12 on lower limb, never more than a total of 18 or less than 13
fakerss x — 15.) me — 10:

1012 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Color in alcohol.—Body dark, sometimes almost black in freshly pre-
served specimens caught in black acid waters with a black muddy substrate.
Dark chromatophores most dense on back, becoming less dense ventrally
(Fig. 3). Scattered, relatively dark chromatophores on all fins except adipose
fin which is pale with very small scattered dark chromatophores. Area of
medial rays of pectoral fins with dark chromatophores on membranes be-
tween fin rays. Sides of head, eye, and operculum with scattered dark chro-
matophores. Pigment on sides of body and head silvery beneath dark chro-
matophores.

Color in life. —This color description is from an aquarium specimen kept
in dark acid water with a dark sandy substrate. Brown to black pigment
about same as described above for color in alcohol. Back olive brown,
especially in area anterior to dorsal fin, top of head, along dorsal part of
back, ventral and posterior to dorsal fin. Dorsal part of caudal peduncle
olive brown; see dark area along back in Fig. 4. Adipose fin bright deep red
in both sexes. Eye silvery yellow with dorsal portion rusty red. Snout and
distal portion of lower jaw olive brown. Pale areas of cheek and operculum
in Fig. 4 yellowish silver in color. Dark areas of operculum brown. Abdom-
inal area silvery yellow on sides, ventrally silvery white. Area dorsal to anal
fin silvery blue with much pale pink or pale purple pink color. Pectoral fin
hyaline with pale yellow color in male. Pelvic fin yellow brown distally.
Anal fin hyaline except about one-third of its distal length forms a yellowish
brown border with darker pigment along dorsal portion of this bordering
band of color (Fig. 4). Distal tips of anal-fin rays and dorsal-fin rays white.
Dorsal fin hyaline except for some brown and yellow pigment in branched
portions of first two branched rays and a little of this same pigment distally
in succeeding two rays. Caudal peduncle without distinct dark spot but
bases of dorsal and ventral caudal-fin lobes with darkened brown pigment.
Caudal fin otherwise hyaline except for some yellow color in dorsal portions
of dorsal lobe and ventral portions of ventral lobe. A small wild specimen
caught in black acid water with a black mud substrate had a very dark
brown olive back and sides with scales reflecting golden green along sides
of body. Adipose fin intensely blood red. Other pigment of fins and body
more intense and darker than aquarium specimen described above.

Sexual dimorphism.—In life, males more deeply colored than females and
adipose fin a deeper red. Sometimes red of adipose fin slightly less extensive
in female than in male. Males may grow larger; in single pair kept in aquaria
for three and a half years, male grew to a standard length of 38.5 mm and
the female to 34.1 mm. Depth of caudal peduncle different in sexes, 15.3-
16.5, « = 15.9% of standard length in males and 13.5-15.3, « = 14.2% in
females. Only males have bony hooks in anal and pelvic fins.

Etymology.—From Latin crassus, thick or stout, and ceps, head, in ref-

VOLUME 93, NUMBER 4 1013

erence to the stout head of this fish relative to the head of most other
characids (Myers, 1926).

Remarks.—Jaws, head shape, and caudal peduncle depth all appear to be
more derived in R. crassiceps than in R. graciliceps.

Acknowledgments

The following persons have loaned specimens for examination: Heraldo
A. Britski and Naérico A. Menezes (MZUSP); William Eschmeyer, Cali-
fornia Academy of Sciences (CAS); and Francisco Mago-Leccia, Museo de
Biologia, Instituto de Zoologia Tropical, Universidad Central de Venezuela
(MBUCV-V). Heraldo Britski, Naércio Menezes, William L. Fink, Sara V.
Fink, Marilyn Weitzman, Oswaldo L. Peixoto and Eugenio Izecksohn aided
in the field collecting. Paulo Vanzolini provided aid and transportation for
part of the field collecting as did the Smithsonian Institution Amazonian
Ecosystems Research Program under the direction of Clifford Evans. We
are especially grateful to Persio de Santos Filho for relinquishing his spec-
imens of Rachoviscus crassiceps for study. Rosario LaCorte brought the
existence of the new species of Rachoviscus to the senior author’s attention.
Eugenio Izecksohn, Oswaldo L. Peixoto and Richard Vari read the manu-
script and offered helpful comments.

Summary

Rachoviscus graciliceps, a new species of characid fish, is described from
small coastal blackwater streams near Prado, Bahia, Brazil. The relation-
ships of this species appear close to Rachoviscus crassiceps Myers (1926),
which is redescribed on the basis of a new examination of the types and
recently discovered specimens from small blackwater streams along the
Atlantic coast about 50 to 60 km south of Paranagua, Parana, Brazil. The
head is more slender in R. graciliceps and the least bony interorbital width
is 29.1—-35.0% of the head length while R. crassiceps has a*bony interorbital
width 37.5—43.8% of the head length. The caudal peduncle of R. graciliceps
is much more slender, being 93.0—100% of its depth, while in R. crassiceps
the caudal peduncle length is 62.7—74.1% of its depth. Rachoviscus gracil-
iceps has 14 transverse scale rows around the caudal peduncle whereas R.
crassiceps has 18. Other differences are noted in the text.

Myers (1926) tentatively considered Rachoviscus to be related to Prion-
obrama, and Géry (1977) suggested that it might be related to Prionobrama
or Paragoniates. Rachoviscus and Paragoniates might be related but we
think it doubtful. Too little is known about these fishes for useful hypotheses
about their relationships. The genus Rachoviscus is redefined and the type
locality of R. crassiceps, stated as being the neighborhood of Rio de Janeiro,

1014 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

is discussed. Since the original specimens described by Myers (1926) were
imported into Germany for the aquarium trade and since the fish has never
been found again in the vicinity of Rio de Janeiro, it may be that the fish
was originally imported from Paranagua and transshipped to Germany
through Rio de Janeiro.

Resumo

Rachoviscus graciliceps, uma nova espécie de caracideo, é descrita de
um pequeno riacho litoraneo, de agua preta, proximo a cidade de Prado,
Estado da Bahia, Brasil. Esta espécie esta intimamente relacionada com
Rachoviscus crassiceps Myers (1926) a qual é aqui redescrita com base em
novo exame dos tipos e exemplares recentemente colecionados em pequen-
os corregos, de agua preta, ao longo da costa atlantica, distando cerca de
50 a 60 quilometros ao sul da cidade de Paranagua, Estado do Parana, Brasil.
As duas espécies podem ser distinguidas pelos seguintes caracteres: a forma
da cabeca é mais delgada em R. graciliceps e a memor largura Ossea inter-
orbital e de 29,1 a 35,0% do comprimento da cabeca enquanto que R. cras-
siceps tem uma largura Ossea interorbital de 37,5 a 43,8% do comprimento
da cabeca. O pedunculo caudal de R. graciliceps € muito mais delgado,
sendo 0 seu comprimento cerca de 93,0 a 100% da sua altura enquanto que
em R. crassiceps 0 comprimento do pedunculo caudal é de aproximada-
mente, 62,7 a 74,1% de sua altura. R. graciliceps possui 14 fileiras trans-
versais de escamas em torno do pedunculo caudal enquanto R. crassiceps
possui 18 fileiras. Outras diferencas sao assinaladas no texto.

Myers (1916) tentativamente considerou Rachoviscus relacionado com
Prionobrama, e Géry (1977) sugeriu um relacionamento entre Rachoviscus
e Prionobrama ou Paragoniates. Rachoviscus e Paragoniates podem ser
relacionados mas achamos isso duvidoso. Contudo, pouco se sabe suas
esses peixes para que se possa formular hipotesis consistentes sobre suas
afinidades. O género Rachoviscus é redefinido e a localidade tipica de R.
crassiceps, tida como sendo nos arredores do Rio de Janeiro, é discutida.
Tendo em vista que os exemplares originais descritos por Myers (1926)
foram importados para a Alemanha pelo comércio de aquario e visto que o
peixe nunca foi reencontrado nas vizinhangas do Rio de Janeiro, podemos
considerar provavel que aquele material fosse originalmente importado de
Paranagua e, atraves do Rio de Janeiro, transportado para a Alemanha. A
localidade tipica deve ser os arredores de Paranagua, onde os peixes foram
agora colecionados, ao invés do Rio de Janeiro.

Literature Cited

Arnold, J. P., and E. Ahl. 1936. Fremdlandische Stisswasserfische.—Braunschweig, 592 pp.
Eigenmann, C. H. 1915. The Cheirodontinae, a subfamily of minute characid fishes of South
America.—Mem. Carnegie Mus. 7:1—99, pls. 17-48.

VOLUME 93, NUMBER 4 1015

. 1917. The American Characidae, part 1.—Mem. Mus. Comp. Zool. Harvard College,

43(1):1-102, pls. 1-8, 12, 14-16, 95, 100, 101.

Fink, W. L., and S. H. Weitzman. 1974. The so-called cheirodontin fishes of Central America
with descriptions of two new species (Pisces Characidae).—Smithsonian Contrib. Zool.
(172): 1-46.

Géry, J. 1977. Characoids of the world.—Neptune City, New Jersey, 672 pp.

Holly, M., H. Meinken, and A. Rachow. 1939. Die Aquarienfische in Wort und Bild.—Stutt-
gart-Winnenden, Lieferung 27/28:284—285.

Myers, G. S. 1926. Eine neue Characinidengattung der Unterfamilie Cheirodontinae aus Rio
de Janeiro, Brasilien.—Blat. fiir Aquarien- und Terrarienkunde, 37 Jahrgang 24: 1-2.

Rachow, A. 1928. Handbuch der Zierfischkunde.—Stuttgart, pp. I-VIII, 1-247, 168 pls.

Sterba, G. 1959. Siisswasserfische aus aller Welt.—Berchtesgaden, 638 pp.

(SHW) Department of Vertebrate Zoology, National Museum of Natural
History, Washington, D.C. 20560; (CAGC) Departamento de Biologia An-
imal, Universidade Federal Rural do Rio de Janeiro, km 47, da Antiga Ro-
dovia Rio, Sao Paulo, 23.460, Seropadica, RJ, Brasil.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1016-1034

STUDIES ON DECAPOD CRUSTACEA FROM THE
INDIAN RIVER REGION OF FLORIDA.

XIX. LARVAL DEVELOPMENT IN THE LABORATORY
OF LEPIDOPA RICHMONDI BENEDICT, 1903,
WITH NOTES ON LARVAE OF AMERICAN
SPECIES IN THE GENUS
(ANOMURA: ALBUNEIDAE)

Robert H. Gore and Cindy Lee Van Dover

Abstract.—The complete larval development of a sand crab species at-
tributed to Lepidopa richmondi (based on morphological characters seen in
the megalopal stage) is described and illustrated, using a first stage zoea
obtained from the plankton and cultured in the laboratory. Three zoeal and
one megalopal stage were obtained. Zoeal features are compared with those
seen in two other western Atlantic and two eastern Pacific species. Data
suggest that the zoeal stages may be segregated along the same lines of
species groupings established for adults, with Atlantic zoeae differing in
telsonal characters from those known in eastern Pacific larvae. This report
is the first to describe the complete development for any Atlantic species
in the laboratory.

The genus Lepidopa, commonly known as sand crabs, consists of small,
filter-feeding intertidal and subtidal marine anomurans which have the car-
apace and pereopods modified for burying in the substratum. Five species
occur in the western North Atlantic, 3 primarily in the Caribbean Sea, and
2 reaching continental waters of the United States. The complete larval
development of these crustaceans remains unknown. Johnson and Lewis
(1942) and Knight (1970) described larvae of an eastern Pacific species at-
tributed to Lepidopa myops. Stimpson, 1860 [since considered by Efford
(1971) to be larvae of L. californica]. In the same paper, Knight described
and illustrated the planktonic larvae of a second species she called Lepidopa
species *‘B.’’ In the Atlantic, the only description of Lepidopa larvae is
that of Sandifer and Van Engel (1972) who described and illustrated 3 zoeal
stages of a species obtained from Chesapeake Bay plankton, which they
attributed to L. websteri Benedict 1903, because it was the only species
known to reach Virginian coastal waters. The second continental species,
L. benedicti Schmitt, 1935, occurs in the Gulf of Mexico and along the
southeastern and central eastern Florida coastline (Holthuis, 1960; Efford,
1971; Gore, unpublished).

VOLUME 93, NUMBER 4 1017

On 19 September 1972, a single albuneid zoeal stage I was collected from
the plankton 2 miles east of Jupiter Inlet, Martin County. on the central
eastern Florida coast. This specimen, originally maintained in the laboratory
as a matter of curiosity, completed its zoeal development and attained me-
galopal stage, but unfortunately died without molting to crab stage I. Mor-
phological features of the frontal margin of the carapace, the walking legs,
third maxillipeds and second abdominal somite, were sufficiently developed
in the megalopa to suggest that the species observed was Lepidopa rich-
mondi Benedict, 1903 (see Discussion). In view of the limited amount of
knowledge on larval development in the Albuneidae we provide herein a
description of the zoeal and megalopal stages of what we believe to be L.
richmondi. If our identification of the megalopa is correct, it indicates that
larvae of L. richmondi occur in neritic waters along the eastern Floridan
continental shelf, although adults have yet to be discovered there.

Materials and Methods

The single first zoeal stage was maintained in the laboratory in a glass
finger bowl with 250 cc of seawater (35%o salinity). Fresh Artemia nauplii
and Chlorella sp. algae were provided and water was changed daily. Room
temperature of about 24°C (+0.5°C) was obtained using closed circuit air-
conditioning. Upon attaining megalopal stage the specimen was transferred
to a 500 cc glass bowl containing sieved, autoclaved quartzite sand in hopes
that crab stages could be obtained in order to positively identify the species.
As noted, the specimen died before this occurred. The description that
follows is based on the molted carapaces and dissected appendages of the
zoeal stages, and the partial dissection of the megalopal stage. Methodology
was identical to that used by Gore (1973).

Results and Discussion of the Rearing Experiment

When collected on 19 September 1972 the specimen was obviously in the
first zoeal stage as determined by the fixed eyes, and the presence of 4
natatory setae on the maxillipeds. The second and third zoeal stages were
attained on 22 and 28 September, followed by the megalopal stage on 6
October. Although the duration of the first zoeal stage is unknown (certainly
at least 4 days, and probably 7-8 days if second stage duration can be used
as an equivalent; see Knight, 1970), the overall developmental time in the
plankton was at least 18 days based on molting occurrence in the laboratory.
Examination of the megalopa which died in day 4 of that stage gave no
indication of imminent molt, so that we may extrapolate a planktonic and
postlarval duration of at least 3 weeks or so for the species at 24°C, before
the first crab stage is attained.

This is substantially less time than the 45—5S3 days required for L. cali-

1018 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fornica to reach first crab (Knight, 1970). However, that species, and the
larvae attributed to L. websteri by Sandifer and Van Engel (1972) possessed
4 zoeal stages, so that extended duration would be suspected. Unfortu-
nately, because they were working with larvae obtained from the plankton,
the latter authors provided no duration-in-stage data. It thus appears that
L. richmondi differs from other known members of the genus in its devel-
opmental sequence, possessing 3 instead of 4 zoeal stages. This assumes,
of course, that laboratory development in this species did not accelerate the
zoeal sequences and cause a late stage to be skipped. In this respect, our
third stage zoea resembled that figured by Sandifer and Van Engel in pos-
sessing well-formed uropods and six abdominal somites, but differed in hav-
ing well-developed pleopod buds which were lacking in Sandifer and Van
Engel’s third stage zoea. Presence of pleopod buds usually indicates the
molt to megalopa is imminent, although some species of galatheid zoeae
may pass through one additional stage before reaching postlarvae (Gore,
1979).

If L. richmondi does pass through only 3 zoeal stages it might explain in
part the relative distribution of the species which seems to be confined to
the Caribbean Sea and southward to Brazil. Shortened zoeal development,
perhaps larvae entrained in local current gyres, would tend to hold the
Species in a more limited geographic area. The species’ farthest occurrence
northward is the islands of Puerto Rico and Jamaica (Holthuis, 1960; Efford,
1971), and the single larva used in this study could have come from either
locality.

Description of the Larvae

First zoea.—(Carapace length 1.6, rostral spine 4.6, posterior carapace
Spine 2.6 mm.)

Carapace (Fig. 1A, B): Smooth, somewhat inflated, ovoid; with elongate
rostral and posterior carapace spines, former 2.9, latter 1.6 carapace
length, all smooth, unarmed. No setae observed on carapace. Eyes un-
stalked.

Antennule (Fig. 1C): Flabellate rod, 3 aesthetascs, | seta.

Antenna (Fig. 1D): Protopodite a slender, dagger-like process armed on
distal 34 with rows of serrated teeth. Exopodite an elongate tapering process,
1.6x longer than protopodal spine, a.single plumose seta medially. Basal
segment with a single short spine distally at junction with exopodite. En-

=

Fig. 1. Lepidopa richmondi, first zoea: A, Lateral view; B, Dorsal view; C, Antennule; D,
Antenna; E, Mandible; F, Maxillule; G, Maxilla; H, Maxilliped 1; I, Maxilliped 2; J, Abdomen
and telson. Scale lines in mm.

VOLUME 93, NUMBER 4 1019

1020 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dopodite a short bud adjacent to protopodal spinous process. General form
of antenna remains similar through subsequent stages, but relative propor-
tions of processes change.

Mandible (Fig. 1E): Asymmetrical dentate processes, no palp.

Maxillule (Fig. 1F): Endopodite unsegmented, 2 terminal setae, a subter-
minal hair; basal endite with 2 strong spines plus a small hair laterally; coxal
endite with 5 strong setae plus a lateral hair.

Maxilla (Fig. 1G): Endopodite unsegmented, 4 terminal, 1 subterminal
seta (2 quite long); basal endite proximal and distal lobes each with 4 setae;
coxal endite with 3 and 2 long setae on distal and proximal lobes, respec-
tively; scaphognathite an elongate sublanceolate lobe with 17 setae.

Maxilliped 1 (Fig. 1H): Coxopodite with a single spinule; basipodite setal
formula progressing distally 3, 3, 3, 3; distal triads each with heavy serrate
seta as shown; endopodite 5-segmented, setal formula 3, 2, 1, 2 + I, 5;
exopodite a single segment, 4 natatory setae. (Note: Roman numeral de-
notes dorsal seta.)

Maxilliped 2 (Fig. 11): Coxopodite with a single spinule; basipodite setal
formula 1, 2; endopodite 4-segmented, setal formula 3, 2,2 + I, 5; exopodite
single segment with 4 natatory setae.

Maxilliped 3 and pereopod buds: Present but too diaphanous to illustrate
from molt.

Abdomen and telson (Fig. 1J): Five somites, third to fifth with postero-
lateral spines increasing in size toward telson; latter a roundly spatulate
process, with two pairs lateral spines, distal 3x length of proximal, former
with a small spine plus thin anomuran hair in axil; telson width 1.8 x length,
posterior margin with 29 spines of varying length, ninth or ninth and tenth
from exterior noticeably longer than others; no dorsal or ventral setae noted
on telsonal surface. In the terminology used by Gore (1979) for this type of
telson, the setal formula reads: I + 11 + 3 + IV + 5-18.

Color: Not noted; no chromatophores visible in molts.

Second zoea.—(Carapace length 1.9; rostral spine 5.7; posterior carapace
spine 2.6 mm.)

Carapace (Fig. 2A, B): Similar to stage I, but larger; rostral spine 3x,
posterior carapace spine 1.4, carapace length, all smooth, unarmed. Car-
apace now with small anterolateral spinule subocularly; eyes mobile.

Antennule (Fig. 2C): Two-segmented, distal with 1 apical, 2 subapical, 2
lateral aesthetascs; | thin apical, 2 basal setae, latter at junction of proximal
segment as shown.

—

Fig. 2. Lepidopa richmondi, second zoea: A, Dorsal view; B, Dorsolateral view; C, An-
tennule; D, Antenna; E, Mandible; F, Maxillule; G, Maxilla; H, Maxilliped 1; I, Maxilliped 2;
J, Abdomen and telson. Scale lines in mm.

VOLUME 93, NUMBER 4

MY

mi

L (25
= VV ACAT AYR

!

1021

1022 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Antenna (Fig. 2D): Exopodite 1.6 length of protopodal spine, 3.1
length of endopodite; latter about half length of protopodal spine; basal
Spine at junction of exopodite slightly larger.

Mandible (Fig. 2E): Relatively unchanged from stage I, teeth sharper,
more numerous. |

Maxillule (Fig. 2F): Endopodite and coxal endite unchanged from stage
I, except latter with additional fine seta laterally; basal endite with additional
strong spine.

Maxilla (Fig. 2G): Endopodite with 5 terminal, 1 subterminal setae; basal
endite unchanged from first stage; coxal endite distal lobe with 3 terminal,
1 subterminal setae; proximal lobe unchanged; scaphognathite with 19 mar-
ginal setae.

Maxilliped 1 (Fig. 2H): Coxopodite without spinule; basipodite and en-
dopodite unchanged from first stage; exopodite with 10 natatory setae.

Maxilliped 2 (Fig. 21): As in first stage, now lacking coxopodal spinule;
exopodite with 10 natatory setae.

Abdomen and telson (Fig. 2J): Five somites, configuration and armature
similar to stage I, but spine on somite 3 reduced; telson process formula
remains I + 11 + 3 + IV + 5-18; movable spines 9-10 larger than preceding
or succeeding spines; a reduced median spinule present; telson otherwise
without surface setae, its width 1.4x< length.

Third zoea.—(Carapace length 2.9; rostral spine 7.7; posterior spine 4.2
mm.)

Carapace (Fig. 3A, B): Similar to previous stage but larger, more inflated;
rostral spine 2.7, posterior carapace spine 1.4x, carapace length, un-
armed. Anterolateral spinule larger.

Antennule (Fig. 3C): Distal segment more elongate relative to proximal,
bearing 7 aesthetascs, 3 setae progressing distally as 2, 2, 2, 1 + 3 setae; 2
small, 2 larger thin setae at junction of proximal and distal segments; former
with single seta medially.

Antenna (Fig. 3D): Exopodite 1.8x protopodal spine, 2.0x endopodite
length, latter just slightly shorter than protopodal spine; basal spine at junc-
tion of exopodite unchanged.

Mandibles (Fig. 3E): Becoming distinctly scoop-shaped, with larger teeth;
no palp.

Maxillule (Fig. 3F): Endopodite with 3 terminal setae plus usual subter-
minal hair; basal endite with 4 strong spines plus lateral seta; coxal endite
with 5 terminal, 1 subterminal, and 2 lateral setae as illustrated.

=

Fig. 3. Lepidopa richmondi, third zoea: A, Lateral view; B, Dorsal view; C, Antennule; D,
Antenna; E, Mandibles; F, Maxillule; G, Maxilla; H, Maxilliped 1; 1, Maxilliped 2; J, Maxilliped
3 and pereopods; K, Abdomen and telson; L, Detail, telsonal lateral spines. Scale lines in mm.

|

|

VOLUME 93, NUMBER 4 1023

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f f v4 / Yt ; LP 5

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— oa E.FGL

1024 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Maxilla (Fig. 3G): Endopodite setae unchanged from previous stage, now
noticeably separated into lobes, progressing apically as 1 + 3 + 2; basal
endite proximal lobe with additional strong seta, all other lobes unchanged;
scaphognathite with 41 marginal setae.

Maxilliped 1 and 2 (Fig. 3H, I): Unchanged except for exopodites, bearing
14 and 13 natatory setae, respectively; serrate basal spines quite noticeable.

Maxilliped 3 and pereopods (Fig. 3J): Undivided and amorphous buds,
latter without evidence of chelation or segmentation.

Abdomen and telson (Fig. 3K, L): Now with 6 somites, spine on somite
3 absent; pleopod buds on somites 2-5; somite 6 with uniramous, two-seg-
mented setose uropods; telson 1.5x wider than long, with 5 pairs of thin
setae in longitudinal row dorsally; posterior marginal formula I + ii + 3 +
IV + 5-19, ninth process noticeably larger than adjacent movable spines;
under high magnification (20) a series of very small spinules interspersed
among all processes (Fig. 3L).

Megalopa.—(Carapace length < width 3.0 x 2.5 mm.)

Carapace (Fig. 4A, B): Dorsoventrally depressed, subrectangular, more
or less smooth, areas poorly delineated, marginally setose from anterolateral
to branchial regions; anterolateral and outer orbital spines distinct; rostral
spine thin, elongate, reaching distal margin of second antennal segment;
orbits deeply excavate, concave, with marginal setae; remnants of posterior
carapacial spines on posterolateral margin of carapace; a transverse row of
fine hairs on frontal region, a similar but shorter row on cardiac area.

Abdomen (Fig. 4C): Six somites, 2-5 with expanded, setose pleura, de-
creasing in size distally; paired biramous pleopods on somites 2 (Fig. 5K)
through 5 (Fig. 5L), appendices internae on endopodites of 3, 4, 5, and
varying numbers of setae present; exopodites becoming more setose pro-
gressing toward telson. Somite 6 with biramous uropods, setose as illus-
trated, but protopodite naked (Fig. 5M).

Telson (Fig. 4C): Width 1.4x length; subcircular, bearing remnants of
larger lateral spines of zoeal telson, plus marginal setae as shown; dorsal
longitudinal row of 5 pairs of thin setae still present, additional transverse
and grouped setae along anterior and anterolateral margin as illustrated.

Antennule (Fig. SA, B): Peduncle 3-segmented, a minute palp-like ventral
flagellum, an elongate 24-segmented setose dorsal flagellum, latter with
aesthetascs on terminal 3 segments, placed as shown in detail (Fig. 5B);
other setae as illustrated.

Antenna (Fig. 5C): Three-segmented, enlarged peduncle, basal segment
with lamellar setose lobe; third with short proximally setose scaphocerite,
a longer 9-segmented dorsal flagellum; fourth segment from tip with 2 small
palp-like processes.

Mandible (Fig. 5D): Marginally dentate scoop-shaped process; an en-

1025

VOLUME 93, NUMBER 4

><

e"

Cr

oY

N

LEE

ZZ
=

1.0
0.5

Fig. 4. Lepidopa richmondi, megalopa: A, Lateral view; B, Dorsal view; C, Abdominal

somites and tail fan. Scale lines in mm.

1026 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

larged 3-segmented palp, armed with strong setae and sharp spines as
shown.

Maxillule (Fig. SE): Endopodite elongate, basally inflated, folded back on
itself distally, a single seta from protuberance on proximal lobe; basal endite
with 15 strong spines and setae plus a thin seta laterally; coxal endite 12
apical, 2 lateral strong spines and setae; protopodal lobe basally with short
sharp spine, an elongate plumose seta, plus a short stout seta.

Maxilla (Fig. SF, G): Endopodite distally naked, with 3 minute setae ba-
sally; basal and coxal endites with varying number of setae, not exactly
determined because of fragility, appearing as 11 + 1, 5 + 1 on distal and
proximal lobes of former, 3 + 1, 3 + 2? + 2? on respective lobes of latter;
what appears to be either a foreshortened epipod, or modified arthrobranch
bud present on basal lobe; a second, rectangular setose lobe apparent proxi-
mally; scaphognathite with 101 marginal setae in addition to scattered short
hairs on lateral surface.

Maxilliped 1 (Fig. 5H): Endopodite 5 segmented, setae progressing dis-
tally 3, 1, 1, 1, 0; exopodite 2-segmented, distalmost flattened, ovoid, with
plumose marginal setae; protopodite with expanded, setose and spinose
distal basal lobe, plus somewhat amorphous coxal lobe bearing 2 setae; a
recurved epipod present.

Maxilliped 2 (Fig. 5I): Endopodite 5-segmented, setose and spinose as
illustrated; exopodite 2-segmented, proximal segment about 4x length of
distal, with 4 long setae from distomedial lobe of former, 7 plumose apical
setae on latter; protopodite sparsely setose, with rudimentary epipod.

Maxilliped 3 (Fig. 5J): Endopodite 5-segmented, clothed with long bristle-
like setae; antepenultimate segment developed distally into rounded lobe
reaching about half length of penultimate segment; exopodite weakly cal-
cified, naked; no epipod observed.

Pereopods (Fig. 6A—G): Pereopod 1 chelate, heavily setose; chela height
0.8x length, carpus 1.6, merus 2.1 longer than wide; pereopods 2-5 as
illustrated, merus of third with plumose seta plus scattered small spinules,
carpus with longer spine-like setae; dactyls of all walking legs variably fal-
cate; fifth pereopod indistinctly chelate, armed apically with several short
sharp teeth, plus long setae, entire appendage considerably reduced in size
relative to other pereopods.

—> ] |

Fig. 5. Lepidopa richmondi, megalopa: A, Antennule; B, Detail, terminal segments of an-
tennule; C, Antenna; D, Mandible; E, Maxillule; F, Maxilla; G, Maxilla, detail of scaphog-
nathite seta; H, Maxilliped 1; I, Maxilliped 2; J, Maxilliped 3; K, Pleopod, somite 2; L, Pleopod,
somite 5; M, Uropod, somite 6. Scale lines in mm.

1027

VOLUME $3, NUMBER 4

1028 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Discussion

The zoeal larvae of the Albuneidae are quite as distinctive as those of the
Porcellanidae, another anomuran family, to which they show obvious mor-
phological relationships. Larvae in both families possess elongate rostral
and paired posterior carapacial spines. However, albuneid larvae and par-
ticularly those assigned to the genus Lepidopa are quickly distinguished
from porcellanid zoeae by the possession of an extremely wide roundly or
triangularly spatuliform telson, often armed with 2 pairs (instead of one pair)
of fixed lateral spines. In addition, the posterior margin of the telson is
heavily spinulose, a feature reminiscent of galatheid larvae (also Anomura)
in the genus Munidopsis (see Samuelsen, 1972; Gore, 1979). Lepidopa lar-
vae are further distinguished by having paired, elongate, often recurved
spines on the posterolateral margin of the fifth abdominal somite (Knight,
1970), these being absent or reduced in porcellanids.

As presently delineated by Knight (1970) the larvae of Lepidopa may be
separated from those of Albunea most easily by the telson (2 pairs lateral
spines, i.e. processes I and IV fixed and enlarged in Lepidopa; | lateral
pair, i.e. only process I fixed in Albunea); rostral and posterior carapacial
spines greatly lengthened in Lepidopa, relatively short in Albunea; and the
antennal scaphocerite more spine-like in Lepidopa, but more blade-like in
Albunea. Based on these characters, the larvae illustrated by Gurney (1942)
as Species A, that identified by Menon (1937) as Albunea symmista, and
the telson figured by Gurney (1924) can all be considered as Albunea. In
fact, if Species A is an Albunea, and if Gurney’s statement is correct that
only Albunea oxyophthalma [=A. paretii fide Monod, 1956] occurs in Ber-
muda, then Gurney’s (1942) larva is a first zoea of A. paretii Guerin, 1853.

In the genus Lepidopa, larvae from the eastern Pacific (L. californica and
L. species B) possess triangularly spatuliform telsons (see Johnson and
Lewis, 1942, pl. 5, fig. 5; Knight, 1970, figs. 40-43, 61-64), whereas those
from the Atlantic (see Gurney, 1942, fig. 110A—D; Sandifer and Van Engel,
1972, figs. 1B-3B; and this study) have a more roundly trigonal telson.
Efford (1971) in reviewing the American species of Lepidopa diagnosed 3
groupings, viz. a myops group (including L. californica), a benedicti group
(which contains L. richmondi), and a venusta group to which L. websteri_ |
belongs. Presently available data are admittedly scanty, but offer the pos- |
sibility that larvae in the myops grouping may be distinguishable from those |
of the remaining two groups on telsonal characters alone. If the larvae of L.
websteri and L. richmondi are correctly assigned, then both the benedicti
and venusta group larvae may exhibit similar telsonal characters, i.e. round-
ly and broadly trigonal, as opposed to more narrowly and triangularly spa-
tuliform telsons seen in known larvae of the myops group.

The first zoeal larvae of Lepidopa species B [as Albunea sp. B] illustrated |
by Gurney (1942) provides some support for this hypothesis. Although sim- |

VOLUME 93, NUMBER 4 1029

Es
Ne O 7

0.5 AE

—~>—! |S

0.1

0.1 G

Fig. 6. Lepidopa richmondi, megalopa: A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D,
Pereopod 4; E, Pereopod 5; F, Pereopod 5, detail of distal segments; G, Pereopod 5, Detail of
cheliform segment. Scale lines in mm.

1030 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ilar in most features to first zoeae of L. websteri and L. richmondi, Gurney’s
larva is immediately distinguishable by having 3 setae on the antennal ex-
opodite (scaphocerite), whereas the larvae of the 2 other species have but
1 seta there. Inasmuch as only 5 species of Lepidopa are presently known
from the western Atlantic, with larvae of 2 species more or less completely
described, Gurney’s larva would presently be assignable to either Lepidopa
benedicti, L. venusta, or L. distincta. The first species is known in the Gulf
of Mexico from Vera Cruz to western Florida, and in the Atlantic along the
Floridan east coast (Efford, 1971; Gore, unpublished) and thus seems the
most likely candidate for Gurney’s larva. Although the remaining 2 species
are not yet known from continental waters of the United States, both are
widely distributed throughout the eastern Caribbean and along the eastern
coast of South America. While neither L. venusta nor L. distincta can be
completely excluded from consideration, especially because their larvae
might possibly become entrained in Antillean/Gulf Stream currents (as ap-
parently do larvae of L. richmondi), and thus be carried to Florida or even
Bermuda, we can provisionally eliminate L. distincta because it belongs to
the myops group and its larvae may exhibit the triangularly spatulate telson
of larvae known from that group. But until more data become available on
the species the assignment to groupings remains conjectural for the most
part.

In order to provide a comparison among the known larvae of Lepidopa,
the salient morphological features of the 4 well-described species are pre-
sented in Table 1. All that can be said for Gurney’s (1942) Lepidopa species
B is that it shows some similarities to L. websteri in maxillary and maxil-
lipedal setation, but differs most noticeably in the scaphocerite of the an-
tenna having 3, instead of 1, marginal setae. In this respect it differs from
all known first zoeae in the genus. As can be seen in the Table, maxillipedal
setal formulae and antennal endopodite:protopodal spine lengths provide
the easiest distinction among the larvae. Other features of some value, but
requiring dissection, include maxillulary and maxillary setation, mandibular
palp budding, and presence or absence of a coxopodal spine on the maxil-
lipeds. Pleopodal buds appear in stage III of L. richmondi (which has no
stage IV), but not until the subsequent stage in the other species. Except
for the telsonal configuration previously noted, the species are more or less
similar in many respects.

Megalopal comparisons are of limited value at present. The only mega-

lopal stages available are those for L. californica and the single specimen

in our study. A comparison of Knight’s (1970) illustration with ours shows
the differences between the 2 species most clearly, but morphological fea-

tures such as setal formulae may eventually prove to be of value when more

species become known.
We made specific identification for our study using the following char-

|
|

VOLUME 93, NUMBER 4

1031

Table 1.—Comparison of selected morphological characters in Lepidopa zoeal stages.

L. californica* L. sp. B L. websteri L. richmondi
ZOEA I
Carapace length 1.2 mm 1.3 mm ca. 1.0 mm 1.6 mm
Rostral spine 2.4x cl 2.6x cl 2-3 cl 2.9x cl
Dorsal margin 2 medial knobs’ No data Smooth Smooth
Antennule 3 aesthetascs 3 aesthetascs 3 aesthetascs 3 aesthetascs
3 setae 3 setae 1-3 setae 1 seta
Antenna
Endopodite Absent Absent Absent Short bud
Maxilliped 1
Coxopodite Unarmed Unarmed Unarmed 1 spine
Endopodite Soleus 25) ZW bite 2 30 Seeleel sale) Borsa hese rata (bas
Maxilliped 2
Coxopodite Unarmed Unarmed Unarmed 1 spine
Endopodite 322s 5 SoA 4) Sea 3, 2, 2+I1, 5
Pereopods Small buds Small buds Minute buds Not seen
Abdomen Dorsal setae + Dorsal setae + No setae No setae

Lateral spine

Telson formula

ZOEA II
Carapace length
Rostral spine
Antennule

Aesthetasc-
setal formula

Basal segment

Antenna
Endopodite

Maxillule
Coxal endite
Maxilla
Scaphognathite
Maxilliped 1

Coxopodite
Endopodite
Exopodite

3rd pair curved
at tips

I+ u+3+
IV + 5-10
(to 13)

1.4 mm
3x cl

Di Do Mets)
setae
2 setae

‘*Distinct bud’’

5—6 setae

13-18 setae

Unarmed
SED ollie Dees
8 natatory

3rd pair curved
at tips

Il+nu+3+
IV + 5-12
(to 15)

1.5 mm
3.3x cl

2,2,1+3
setae
2 setae

Similar

Similar

13-20 setae

Unarmed
Similar
9-10 natatory

3rd pair curved
at tips

I+nu+3+
IV + 5-14
(to 16)

ca. 1.9 mm
No data

252s bates

setae c

0-1 setae

‘*Small bud’’

6 setae

20-23 setae

Unarmed
35 Ay dhs Sells S
10 natatory

3rd pair straight
at tips

JPsr it se 3) ap IN
+ 5-18

1.9 mm
3x cl

2, 2, 1, +1 seta

2 setae

Ys Exopodite
length

7 setae

19 setae

Unarmed
3.2. 1, 2+1,.5
10 natatory

1032

Maxilliped 2

Coxopodite
Endopodite
Exopodite

Abdomen

Uropods

Telson formula

ZOEA III

Carapace length
Rostral spine

Antennule
Prox./dist.
jet.
Antenna
Endopodite

Mandible

Maxillule

Endopodite
Basal endite

Maxilla

Basal endite
Coxal endite

Scaphognathite

Maxilliped 1

Endopodite
Exopodite

Maxilliped 2

Endopodite
Exopodite

Pereopods
Abdomen

Pleopods

Table 1.—Continued.

L. californica*

Unarmed
Bean, PAaes)
8 natatory

No change

Not apparent

I+n+3+
IV + S-15
(to 18)

1.8 mm
3x cl

2 seta

ca. 0.5x
protopodal
spine

Small palp bud

2 + 1 setae
3-4 spines, 1
seta

4, 4 setae
2, 2 setae
13-18 setae

32571 25 5
10-11 natatory

3,.2,2+1, 5
10-11 natatory
P, cheliform
Somite 3
spine —

Absent

L. sp. B

Unarmed
Similar
Similar

No change

Similar

l+it+3+
IV + S-17
(to 19)

1.8 mm
3.2x cl

Similar

Similar

Similar

Similar
Similar

4-5, 4-5 setae
2-4, 2 setae
26—33 setae

35 ay Why arlle dS
11-13 natatory

Similar
12-13 natatory

No data

Similar?

Similar?

L. websteri

Unarmed
3, 2, 2+I, 5
10 natatory

No change

Anlage visible

l+n+3 +
IV + 5-14
(to 16)

ca. 2.0 mm
No data

None?

“Nearly as
long’’ as
protopodite

No palp

2 + 1 setae
3 spines, 1 seta

4, 4 setae
3, 2 setae
33-36 setae

35 45 Ibs Barly, Dd
13-14 natatory

Sy Ap @arlly
13-14 natatory

P, cheliform

Somite 3
spine +

Absent

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

L. richmondi

Unarmed
3, 2, 2+I, 5
10 natatory

Somite 3 spine
reduced

Not apparent

sei ap 3 ae JY
+ 5-18

2.9 mm
2.9x cl

4 setae

0.9x protopodal
spine

No palp

3 + 1 setae
4 spines, | seta

4, 5 setae
4, 2 setae
41 setae

32 op ae etalon
14 natatory

3, 2, 2+
13 natatory

P, undivided

Somite 3
spine —

Buds, somites
MS

VOLUME 93, NUMBER 4 1033

Table 1.—Continued.

L. californica* L. sp. B L. websteri L. richmondi
Uropod Endopod Similar? Endopod No bud
bud + bud + observed
Telson formula Unchanged from previous stage in all species

* As L. myops in Knight, 1970.

acters in the megalopa: fifth abdominal somite with pleural expansion sep-
arates Lepidopa from Albunea, Zygopa, and Stemonopa (the latter is not
yet known from the western Atlantic); carpus of maxilliped 3 not longer
than propodus eliminates the myops group of species; absence of subrostral
spine eliminates venusta grouping; a distinct rostrum, deeply concave or-
bital margins, abdominal somite 2 truncately rectangular and not directed
posteriorly, and the shape of dactylii on pereopods 2, 3, 4, all suggest L.
richmondi (Efford, 1971, especially figs. 6, 7), and not L. benedicti. Regret-
tably, the groove around the posterior margin of the carapace, an important
diagnostic character in adults (and presumably megalopae) of the species,
was undeveloped in our specimen. We also feel certain that we were not
dealing with L. websteri because features in larvae of that species differ
from those seen in ours (see Sandifer and Van Engel, 1972; Table 1, this
study). Even though the distributional range of L. benedicti encompasses
the central eastern Florida coast where our first zoeal stage was originally
collected (suggesting it was that species) the Caribbean range of L. rich-
mondi could easily allow larvae of this species to be entrained in the Florida
Current and swept along the Atlantic coast of Florida.

Literature Cited

Efford, I. E. 1971. The species of sand crabs in the genus Lepidopa (Decapoda: Albunei-
dae).—Zool. Anz. 186:59-102, figs. 1-12.

Gore, R. H. 1973. Pachycheles monilifer (Dana 1852): The development in the laboratory of

larvae from an Atlantic specimen with a discussion of some larval characters in the

genus (Crustacea: Decapoda; Anomura).—Biol. Bull. 144:132-150, figs. 1- 6.

. 1979. Larval development of Galathea rostrata under laboratory conditions, with a

discussion of larval development in the Galatheidae (Crustacea Anomura).—Fishery

Bull. U.S. Fish Wildl. Serv. 76:781-806, figs. 1-9.

Gurney, R. 1924. Crustacea. Part IX.—Decapod larvae.—British Antarctic (“‘Terra Nova’’)
Exped., Zoology 8:37-—202, figs. 1-78.

Gurney, R. 1942. Larvae of decapod crustacea.—Ray Society No. 129:i-viii, 1-306, figs. 1-122.

Holthuis, L. B. 1960. Notes on American Albuneidae (Crustacea Decapoda, Anomura) with
the description of a new genus and species.—Koninkl. Nederl. Akad. Wetenschappen,
Amsterdam 64:21-36, figs. 1-S.

Johnson, M. W., and W. M. Lewis. 1942. Pelagic larval stages of the sand crabs Emerita

1034 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

analoga (Stimpson), Blepharipoda occidentalis Randall, and Lepidopa myops Stimp-
son.—Biol. Bull. 83:67-87, text-fig. 1, pls. 1—S.

Knight, M. W. 1970. The larvae development of Lepidopa myops Stimpson, (Decapoda,
Albuneidae) reared in the laboratory, and the zoeal stages of another species of the
genus from California and the Pacific coast of Baja California, Mexico.—Crustaceana
19: 125-156, figs. 1-65.

Menon, M. K. 1937. Decapod larvae from the Madras plankton. I. Bull. Madras Govt. Mus.
new ser. (Nat. Hist.) 3:1—55, 9 pls.

Monod, Th. 1956. Hippidea et Brachyura ouest-africains.—Mém. Inst. Francais Afrique Noir
45:1-674, figs. 1-884.

Samuelsen, T. J. 1972. Larvae of Munidopsis tridentata (Esmark) (Decapoda, Anomura)
reared in the laboratory.—Sarsia 48:91—98, figs. 1-2.

Sandifer, P. A., and W. A. Van Engel. 1972. Lepidopa larvae (Crustacea, Decapoda, Albu-
neidae) from Virginia plankton.—J. Elisha Mitchell Sci. Soc. 88:220-225, figs. 1-4.

Smithsonian Institution, Fort Pierce Bureau, Ft. Pierce, Florida 33450.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1035-1079

REVIEW OF HYSTEROTHYLACIUM AND
IHERINGASCARIS (BOTH PREVIOUSLY
=THYNNASCARIS) (NEMATODA: ANISAKIDAE)
FROM THE NORTHERN GULF OF MEXICO

Thomas L. Deardorff and Robin M. Overstreet

Abstract.—The genus Hysterothylacium Ward and Magath (type-species
H. brachyurum Ward and Magath) is resurrected to include those species
previously considered as members of the junior synonym Thynnascaris
Dollfus and others described in the genus Contracaecum Railliet and Henry
that mature in fishes. [heringascaris Pereira is resurrected for I. inquies
(Linton), the senior synonym of I. iheringascaris Pereira. Consequently,
we establish 47 new combinations plus present lists of species that may later
be shown to be members of Hysterothylacium. At least six species of Hys-
terothylacium |H. fortalezae (Klein), H. incurvum (Rudolphi), H. chaunaxi
(Olsen), H. reliquens (Norris and Overstreet), H. ogcocephali (Olsen), and
H. corrugatum sp. n.] occur in the Gulf of Mexico, and morphological data
and a key are presented for these. Hysterothylacium corrugatum infecting
the swordfish from the Atlantic and Pacific oceans can be characterized
primarily by having modified preanal annules ventrally in males and a tail
without ornamentation in conjunction with lacking both deep interlabial
grooves and flaring cervical alae.

Several adult nematodes infecting the digestive tracts of finfishes have
been described as species of Thynnascaris Dollfus, 1933 or Contracaecum
Railliet and Henry, 1912. Species of Contracaecum sensu stricto, however,
mature in birds and mammals. Reexamination of Hysterothylacium brachy-
urum Ward and Magath, 1917, as discussed below, reveals it to have the
same generic features as species of Thynnascaris. Before the genus Thyn-
nascaris gains more acceptance, especially in light of potential public health
significance of some species (Petter, 1969a, b; Norris and Overstreet, 1976;
Norris and Ebert, MS in preparation; Overstreet and Meyer, MS in prep-
aration), we wish to establish the priority of Hysterothylacium Ward and
Magath, 1917 and Iheringascaris Pereira, 1935, transfer known species, and
provide data on species of those genera from the Gulf of Mexico.

Most specimens that we collected were removed from hosts, fixed in
glacial acetic acid, stored in a solution of 5 parts glycerin and 95 parts 70%
ethyl alcohol, and examined in lactic acid or in glycerin after the evaporation
of the alcohol. A few others from hosts in museums had been fixed in

1036 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

formalin and transferred to ethyl alcohol. Many of the common names for
fishes follow the American Fisheries Society’s list (Bailey, 1970). Sections
of worms were stained with Harris’ hematoxylin and eosin. Measurements
locating the position of the nerve ring are taken from the anterior extremity
of the worm to the center of the nerve ring. In spicule ratios, the length of
the left spicule equals one. All measurements are in micrometers unless
stated otherwise, and figures were drawn with the aid of a drawing tube.

Abbreviations for repositories of examined nematodes or host fish are
BMNH—British Museum (Natural History), London, England; GCRL—
Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi;
MPM—Meguro Parasitological Museum, Tokyo, Japan; OCI—Oswaldo
Cruz Institute, Helminthology Collection, Rio de Janeiro, Brazil; SIO—
Scripps Institution of Oceanography, La Jolla, California; TABlL—South-
east Fisheries Center, NMFS, Miami, Florida; UMML—Rosenstiel School
of Marine and Atmospheric Science, University of Miami, Florida;
USNM—National Museum of Natural History, Smithsonian Institution (for
fish); USNM Helm. Coll.—United States National Museum, Helmintho-
logical Collection, Beltsville, Maryland; VIMS—Virginia Institute of Marine
Science, Gloucester Point, Virginia; ZM—Zoologisches Museum, Hum-
boldt-Universitat, Berlin, Germany.

Hysterothylacium Ward and Magath

Hysterothylacium Ward and Magath, 1917 (type-species H. brachyurum
Ward and Magath, 1917).

Thynnascaris Dollfus, 1933 (type-species T. legendrei Dollfus, 1933, junior
synonym of Ascaris cornuta).

Thynnascaris.—Dollfus, 1935 (relegated to subgenus of Contracaecum).

Simplexonema Kreis, 1952 (as subgenus of Contracaecum with C. (S.)
cyclopteri Kreis, 1952 as type-species).

Erschovicaecum Mozgovoi in Skrjabin et al., 1951 (as subgenus of Contra-
caecum with C. [E.] aduncum [Rudolphi, 1802] as type-species).

Acollaris Araujo, 1970 (as subgenus of Contracaecum with C. [A.] marinum
[Linnaeus, 1767] as type-species).

Diagnosis.—Body elongate, reaching greatest width near midbody. Cu-
ticle with annulations moderately or weakly defined or lacking (presumably
unreported). Cuticular alae distinct or indistinct. Lips approximately equal
in size, usually broader than long, bearing transparent cuticular flanges on
lateral margins; flanges with or without indentations; internal pulp usually
pedunculated; dorsal lip with 2 lateral doubled papillae; subventral lips with
amphid, adjacent mediolateral doubled papilla, and single lateral papilla.
Dentigerous ridges absent (not present as indicated by Jackson et al., 1978).
Interlabia present; interlabial grooves present or absent. Ventriculus nearly

VOLUME 93, NUMBER 4 1037

spherical; ventricular appendage saclike or cylindrical, with septum dividing
structure into 2 equal longitudinal pouches; intestinal cecum usually shorter
than ventricular appendage. Excretory system with excretory pore located
at or near level of nerve ring, with duct extending to at least left lateral cord
in which primary excretory canal extending posteriorly (at least in species
examined). Rectal glands present or absent (presumably unreported). Spic-
ules similar, alate, equal or slightly unequal in length. Gubernaculum absent.
Vulva anterior to midbody. Uterus didelphic, opisthodelphic. Tail conical;
tip with or without ornamentation. Medioventral preanal organ usually dis-
tinct on preanal fold. Phasmids usually distinct. Parasites of marine, estu-
arine, and freshwater fishes. Type-species: Hysterothylacium brachyurum
Ward and Magath, 1917.

Comparisons.—Species of Contracaecum, especially larval forms, are
often confused with those of Hysterothylacium primarily because some au-
thors still accept Contracaecum in its broad sense. In the strict sense as
followed here, even larvae can be distinguished because the excretory pore
empties near the ventral interlabium in species of Contracaecum, adults of
which mature in birds and mammals. Most species related to those in Hys-
terothylacium mature in fishes. Those of Goezia Zeder, 1800 have plicated
cuticular annulations possessing spines along their rear border, at least as
adults, and that of [heringascaris to be reviewed later in this paper has
similar annulations but without spines. Other nematodes with a ventricular
appendage, intestinal cecum, and excretory pore emptying near level of the
nerve ring include members of several genera that need further investiga-
tion. Heterotyphlum Spaul, 1927 and Paraheterotyphlum Johnston and
Mawson, 1948 (see Schmidt and Kuntz, 1973; Sprent, 1978) have members
with no interlabia; Lappetascaris Rasheed, 1965 has one species, and it has
no interlabia and considerably modified lips; and Pulchrascaris Vicente and
dos Santos, 1972 also has one species, and it has no interlabia and its lips
have dentigerous ridges. The positicn of the excretory pore and whether a
ventricular appendage exists both need to be determined. Species of Ra-
phidascaris Railliet and Henry, 1915, presumably without interlabia, and
Raphidascaroides Yamaguti, 1941, with them, also are similar to those of
Hysterothylacium except they lack the intestinal cecum.

Remarks.—Dollfus (1933), assuming the type-species lacked a ventricular
appendage, erected Thynnascaris as a genus, but later (1935a) reduced it to
a subgenus of Contracaecum Railliet and Henry, 1912 when he realized the
appendage was present. Hartwich (1957) resurrected Thynnascaris, distin-
guishing it from Contracaecum on the basis of having species that lack a
ribbonlike widening of excretory system, have an excretory pore near the
level of the nerve ring rather than the ventral interlabium, and mature in
fishes rather than in birds and mammals. Yamaguti (1962), basing his clas-
sification scheme mainly on the presence of both intestinal and ventricular

1038 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

diverticula, recognized Contracaecum as the only valid genus. In other
recent classifications, Chabaud (1965) accepted both Thynnascaris and Con-
tracaecum and placed them in different tribes of the subfamily Anisakinae,
whereas Hartwich (1974) placed Thynnascaris in Raphidascariinae (as
Raphidascaridinae), and Contracaecum in Anisakinae.

None of the above authors accepted Hysterothylacium as valid. Yamaguti
(1962) followed Van Cleave and Mueller (1934) and considered it a junior
synonym of the older Contracaecum, whereas Chabaud (1965) and Hart-
wich (1974), following Yorke and Maplestone (1926) and Baylis and Daub-
ney (1926), considered it a junior synonym of Raphidascaris Railliet and
Henry, 1915. Gibson (1975), in a review of Hartwich’s key, questioned the
synonymy, but did not resurrect Hysterothylacium nor did he intend to do
so (Gibson, personal communication). Margolis and Arthur (1979) briefly
discussed the problem. Hysterothylacium was erected for H. brachyurum,
which was considered to lack an intestinal cecum, from the stomach of
black bass (=largemouth bass) in Lake St. Clair, Michigan (Ward and Magath,
1917). When Van Cleave and Mueller (1934) examined additional specimens
from bass in Oneida Lake, they noted an intestinal cecum, the primary char-
acter separating Contracaecum sensu lato from Raphidascaris. Consequent-
ly, they considered Hysterothylacium a synonym of Contracaecum rather
than Raphidascaris. Hartwich (1957) assumed Van Cleave and Mueller (1934)
had a species different from Ward and Magath’s. We, however, examined the
type (USNM Helm. Coll. No. 52120) of H. brachyurum plus additional
other specimens (USNM Helm. Coll. No. 55995) and confirm the presence
of an intestinal cecum, ventricular appendage, and excretory pore located
near the nerve ring. We therefore consider Thynnascaris a junior synonym
of Hysterothylacium in accordance with replaced sections a and b of Article
23 (Anonymous, 1974) of the International Code of Zoological Nomencla-
ture. The name Thynnascaris cannot be considered ‘‘long-established,’’ and
application of the Law of Priority will not disturb stability or cause confu-
sion, points confirmed by W. I. Follett (California Academy of Sciences,
personal communication). The junior synonyms of Hysterothylacium other
than Acollaris listed in the synonymy have been discussed by Hartwich
(1957). Araujo (1970) established that subgenus to include those species
(without naming them) that we consider Hysterothylacium.

By accepting the name Hysterothylacium, a number of new combinations
must be established (Table 1). For a number of other species, some based
on adults and some on larvae, we have not seen type-material and are
uncertain of their generic relationships or their validity (Tables 2, 3). Even
less information is known for several additional species mentioned by Baylis
(1923), Dollfus (1935a), and Punt (1941), but not listed in this paper. Tché-
prakoff (1966) described C. prevosti Tchéprakoff, 1966 from the emperor
penguin and reported the excretory pore as opening posterior to the nerve

VOLUME 93, NUMBER 4 1039

ring. Since we know of no species of Hysterothylacium infecting a bird,
that observation should be confirmed and evaluated.

Hensley and Nahhas (1975) reported C. brachyurum from the intestine
and ceca of four fishes in the Sacramento—San Joaquin Delta, California,
but their specimens were not readily available for us to examine (Nahhas,
personal communication). Numerous other references to records from the
northern U.S. and Canada are listed by Hoffman (1967) and Margolis and
Arthur (1979).

The only other species described as a member of Hysterothylacium is H.
cayugensis Wigdor, 1918 (see Wigdor, 1918), based on immature females
infecting Esox lucius Linnaeus and Ictalurus nebulosus (Lesueur) (as Ameiu-
rus n.) in Cayuga Lake, New York. Van Cleave and Mueller (1934) assumed
it was conspecific with H. brachyurum. No designated type was located,
but specimens (USNM Helm. Coll. Nos. 52121 and 52122) from Esox lucius
in Lake Cayuga and Douglas Lake, Michigan, deposited in 1917 by Wigdor
and by A. R. Cooper, respectively, were identified by Wigdor and Magath
as Hysterothylacium sp., followed by **(=Raphidascaris sp.)’’ on the label.
We keyed those to Raphidascaroides Yamaguti, 1941 in Hartwich’s (1974)
generic key, and consider H. cayugensis a nomen dubium.

Hysterothylacium fortalezae (Klein), new combination
Figs. 1-13

Contracaecum fortalezae Klein, 1973:200—210, figs. 1-7 (original descrip-
tion; type-host Scomberomorus brasiliensis; type-locality Ceara, Brazil).

Diagnosis.—Cuticle with prominent cervical alae; lips with ratio of length
to width 1:0.9-1.0; caudal papillar pairs: preanal 13-25; postanal 8; spicules
3-5% of body length; spicule-ratio 1:1.0; tuft of 12-14 relatively large pro-
jections at posterior extremity.

General (based on specimens from Mississippi). —Body reaching greatest
width near midbody. Cuticle with inconspicuous annulations. Alae extend-
ing entire length of body; cervical alae prominent, wing-like, extending from
base of lips to level of posterior 4% of esophagus, up to maximum of 30-68
thick (on 5 specimens); caudal alae most prominent at posterior extremity.
Lips approximately equal in size, as wide as long; flanges widest near pos-
terior; pulp slightly pedunculated. Interlabia with height equal to or slightly
greater than width at base; interlabial grooves lacking. Esophagus 6.4—
10.0% of body length. Ventriculus narrower than widest level of esophagus,
usually longer than wide; ventricular appendage descending without angu-
lation from posterior portion of ventriculus. Nerve ring located adjacent to
anterior 22-34% of esophagus. Excretory pore immediately posterior to
level of nerve ring. Tail with tuft of projections at posterior extremity.

Male (based on 3 specimens): Body 12.0-23.4 mm long by 141-432 at

1040

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—List of new combinations in the genus Hysterothylacium parasitizing fishes of the

world.

New combination

H. aduncum

AH. amoyensis

H, arii

H.. assi

H. auctum

H. baylisi

H. bidentatum

H.. carangis

H. chaunaxi

H. clavatum

HZ. coiliae

H. cornutum

H. cyclopteri

H. dollfusi

H. epinepheli

H. fabri

Original name

Ascaris adunca Rudolphi, 1802

Contracaecum amoyensis Hsu,
1933

Contracaecum arii Yamaguti,
1954

Contracaecum assi Parukhin,
1973

Ascaris aucta Rudolphi, 1802

Contracaecum baylisi
Yamaguti, 1941

Ascaris bidentata Linstow,
1899

Thynnascaris carangis
Kalyankar, 1971

Contracaecum chaunaxi Olsen,
1952

Ascaris clavata Rudolphi, 1809

Contracaecum coiliae
Yamaguti, 1941

Ascaris cornuta Stossich, 1904

Contracaecum (Simplexonema)
cyclopteri Kreis, 1952

Thynnascaris dollfusi Schmidt,
Leiby, and Kritsky, 1974

Contracaecum epinepheli
Yamaguti, 1941

Ascaris fabri Rudolphi, 1819

Pertinent reference(s)

Hartwich, 1975 (compare
Punt, 1941; Petter, 1969b;
and other cited references)

Hsu, 1933a, b

Yamaguti, 1954

Parukhin, 1973

Dollfus, 1953; Hartwich, 1975
Yamaguti, 1941

Mozgovoi, 1953 (A. stroumovi
Golowin, 1900, as a junior
synonym)

Kalyankar, 1971

Olsen, 1952; this paper

Kahl, 1936; Osmanov, 1940;
Kreis, 1952

Yamaguti, 1941

Baylis, 1923; Dollfus, 1935a;
Berland, 1961; Petter,
1969b; Hartwich, 1975 (T.
legendrei Dollfus, 1933, as
a junior synonym)

Kreis, 1952 (considered junior
synonym of C. aduncum by
Berland, 1961)

Schmidt et al., 1974

Yamaguti, 1941

Compare Baylis, 1923;
Zhukov, 1960 (A.
biuncinata Molin, 1858, as
a junior synonym)

VOLUME 93, NUMBER 4

Table 1.—Continued.

1041

New combination

Original name

Pertinent reference(s)

H. fortalezae

Sa

Se

ee

Sa

Se

Se

mS

Sa

Sa

my

Se

x

Se

. gadi

. gracile

. habena

. hapalogenyos

. haze

histiophori

. ilishae

increscens

incurvum

. longispiculum

. magnum

. marinum

. melanogrammi

. melichthysi

. murrayense

. ogcocephali

Contracaecum fortalezae Klein,
1973

Ascaris gadi Miller, 1776

Contracaecum gracile
Yamaguti, 1935

Ascaris habena Linton, 1900

Contracaecum hapalogenyos
Yamaguti, 1961

Thynnascaris haze
Machida, Takahashi, and

Masuuchi, 1978

Contracaecum histiophori
Yamaguti, 1935

Contracaecum ilishae
Yamaguti, 1941

Ascaris increscens Molin, 1858

Ascaris incurva Rudolphi, 1819

Contracaecum longispiculum
Fujita, 1940

Contracaecum magnum
Smedley, 1934

Ascaris marina Linnaeus, 1767

Contracaecum melanogrammi
Smedley, 1934

Contracaecum melichthysi
Olsen, 1952

Contracaecum murrayense
Johnston and Mawson, 1940

Contracaecum ogcocephali
Olsen, 1952

Klein, 1973; this paper

Johnston and Mawson, 1945;
Hartwich, 1975

Yamaguti, 1935, 1941

Chandler, 1943; Norris and
Overstreet, 1975; this paper

Yamaguti, 1961

Machida et al., 1978

Yamaguti, 1935; Olsen, 1952;
this paper

Yamaguti, 1941

Molin, 1858

Baylis and Daubney, 1922;
Dollfus, 1935b; Hartwich,
1975; this paper

Fujita, 1940

Smedley, 1934 (considered
probable synonym of H.
aduncum by Margolis and
Arthur, 1979)

Yamaguti, 1935

Smedley, 1934 (possible
synonym of AH. aduncum,
see Margolis and Arthur,
1979)

Olsen, 1952

Johnston and Mawson, 1940

Olsen, 1952; this paper

1042 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

New combination Original name Pertinent reference(s)

H. okadai Contracaecum okadai Fujita, Fujita, 1940
1940
H. pagrosomi Contracaecum pagrosomi Yamaguti, 1935

Yamaguti, 1935

H. paralichthydis Contracaecum paralichthydis Yamaguti, 1941
Yamaguti, 1941

Hi. rectum Contracaecum rectum Yamaguti, 1961
Yamaguti, 1961

H. reliquens Thynnascaris reliquens Norris Norris and Overstreet, 1975;
and Overstreet, 1975 this paper

H. rhacodes Thynnascaris rhacodes Deardorff and Overstreet,
Deardorff and Overstreet, 1978
1979

H. rigidum Ascaris rigida Rudolphi, 1809 Punt, 1941; Berland, 1961;
Petter, 1969b; Hartwich,
1975 (considered a junior
synonym of C. marinum by
Yamaguti, 1935)

H. saba Contracaecum saba Yamaguti, Yamaguti, 1941
1941

H. salvelini Contracaecum salvelini Fujita, Fujita, 1940
1940

H. scomberomori Contracaecum scomberomori Yamaguti, 1941

Yamaguti, 1941

H. seriolae Contracaecum seriolae Yamaguti, 1941; Brunsdon,
Yamaguti, 1941 1956; Machida et al., 1978
AH. trichiuri Contracaecum trichiuri Thwaite, 1927

Thwaite, 1927

H. zenis Contracaecum zenis Baylis, Baylis, 1929
1929

H. zenopsis Contracaecum zenopsis Yamaguti, 1941; Brunsdon,
Yamaguti, 1941 1956

greatest width; ratio of greatest width to length 1:54—85. Lips 43-81 long by
43-80 wide. Esophagus 0.9-1.5 mm long or 6.4—7.8% of body length. Ven-
triculus 65-127 long by 50-100 wide; ventricular appendage 506-835 long by
24-38 wide; ratio for lengths of ventricular appendage to esophagus 1:1.6—
1.8. Intestinal cecum 144-296 long by 43-84 wide; ratio of cecal to ventric-
ular appendage lengths 1:2.1—3.9; ratio of cecal to esophagus lengths 1:5.0-
6.5. Nerve ring located within anterior 25.5-29.4% of esophageal, 24—38 in
breadth. Spicules 2.9-4.7% of body length, 560-680 long by 14-17 wide;

VOLUME 93, NUMBER 4 1043

Table 2.—Species of uncertain status*, but possibly referable to Hysterothylacium because
they contain an intestinal cecum and ventricular appendage and mature in the alimentary canal
of fishes.

Contracaecum filiforme (Stossich, 1904)

Contracaecum macquariae Johnston and Mawson, 1940
Contracaecum macrozarcium Nigrelli, 1946

Contracaecum mulli (Wedl, 1855)

Contracaecum naitoi Fujita, 1932

Contracaecum ochotense Fujita, 1932

Contracaecum serrani Kalyankar, 1972

Contracaecum synpapillus Bilqees, Khanum, and Jehan, 1971

* See text for references to other nominal species with status more uncertain.

spicule-ratio 1:1.0. Caudal papillae 21-33 pairs, becoming closer together
and more medial as approaching anus; preanal pairs 13-25; postanal pairs
8; para-anal pairs lacking. Medioventral preanal organ indistinct. Phasmids
anterior and lateral to last papillar pair. Tail 129-173 long including 12-14
rounded and pointed projections measuring 9-19 long plus several shorter
ones near posterior nonconical extremity.

Female (based on 5 specimens): Body 10.0-15.4 mm long by 144-189
wide at greatest width; ratio of greatest width to length 1:69-92. Lips 48-63
long by 48-63 wide. Esophagus 1.0—1.1 mm long or 7-10% of body length.
Ventriculus 72—107 long by 48-86 wide; ventricular appendage 556-661 long
by 24-49 wide; ratio for lengths of ventricular appendage to esophagus
1:1.5—2.1. Intestinal cecum 120-166 long by 50-60 wide; ratio of cecal to
ventricular appendage lengths 1:3.4—5.4; ratio of cecal to esophageal lengths
1:6.4-8.3. Nerve ring located within anterior 22-34% of esophagus, 31-41
in breadth. Vulva without salient lips, opening 5.0-6.0 mm or 30-38% of
body length from anterior extremity. Ovaries not extending beyond anterior
level of vulva. Phasmids inconspicuous. Tail 247-284 long including 12-14
rounded and pointed projections measuring 7-18 long plus several shorter
ones near posterior nonconical extremity.

Hosts. —Scomberomorus brasiliensis Collette, Russo, and Zavala-Camin,
serra Spanish mackerel, type-host; S$. maculatus Mitchill, Spanish mack-
erel; S. cavalla (Cuvier), king mackerel (Scombridae); Oligoplites saurus
(Bloch and Schneider), leatherjacket (Carangidae); Mycteroperca bonaci
(Poey), black grouper (Serranidae).

Sites of infection.—Stomach and intestine.

Localities.—Horn Island, Mississippi; Biscayne Bay, Florida; Ceara, Bra-
zil (type-locality).

Specimens deposited.—USNM Helm. Coll. No. 75838 (pair); BMNH
Reg. No. 1980.359-360 (pair); OCI 31.812a—b (pair).

Remarks.—Klein (1973) collected Hysterothylacium (=Contracaecum)
fortalezae at Ceara, Brazil, from the stomach and intestine of Scombero-

1044 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Larval ascaridoids of uncertain status, some of which probably will be shown as
referable to Hysterothylacium and some probably will be junior synonyms.

Parasite

Contracaecum aori
Khan and Yaseen, 1969

Contracaecum benimasu
Fujita, 1932

Contracaecum brevicaecum
Khan and Yaseen, 1969

Contracaecum collieri
Chandler, 1935

Contracaecum hippoglossi
Fujita, 1932

Contracaecum hypomesi
Fujita, 1932

Contracaecum mesopi
Fujita, 1940

Contracaecum (Thynnascaris)
nototheniae Johnston and
Mawson, 1945

Contracaecum oshoroensis
Fujita, 1940

Contracaecum tati
Fujita, 1932

Contracaecum (Thynnascaris)
tasmaniense Johnston and
Mawson, 1945

Contracaecum vittatii
Khan and Begum, 1971

Remarks

In body cavity of Mystus aor; excretory \pore not
mentioned; intestinal cecum shorter than ventricular
appendage; probably fourth stage larva.

In body cavity of Oncorhynchus spp.; location of
excretory pore not mentioned; vagina opening to
rectum (?); probably fourth stage larva.

In body cavity of ““sawfish’’; excretory pore slightly
posterior to level of nerve ring; third stage larva.

In body cavity of Cyprinodon variegatus; no excretory
pore mentioned; intestinal cecum longer than
ventricular appendage; third stage larva.

In body cavity of several fishes; location of excretory
pore not mentioned; intestinal cecum longer than
ventricular appendage; vulva opens to rectum (7);
probably developing fourth stage larva within third
stage molt.

In body cavity of Hypomesus spp., Clupea pallasi,
and Oncorhynchus nerka (as well as intestine);
location of excretory pore not mentioned; intestinal
cecum longer than ventricular appendage; probably
fourth stage larva.

In body cavity of Mesopus olidus; excretory pore near
nerve ring; males unknown; probably fourth stage
larva.

Immature specimens from Notothenia rossi and N.
macrocephala; broad cervical alae of unequal length;
location of excretory pore not mentioned; probably
fourth stage larva.

In body cavity of Oncorhynchus keta and Mesopus
olidus; excretory pore ‘‘indistinct’’; fourth stage
larva.

In body cavity of Taius tumifrons; excretory pore not
mentioned; probably developing fourth stage larva.

Site of infection not mentioned; from Notopogon liliei
and Coclorhynchus australis; location of excretory
pore not mentioned; intestinal cecum longer than
ventricular appendage; fourth stage larva.

In body cavity of Upeneus vittatus; excretory pore
near nerve ring; third stage larva.

VOLUME 93, NUMBER 4 1045

f
en Senay

eS ieeieeeeet emer finnoial

500
100

Sa oo oa ee

~acalad

FeeGGncene

30
3
La

(CCC ecco SER CULCLLLU LLL tt

150

ae

Figs. 1-7. Hysterothylacium fortalezae: 1, Anterior end showing lateral alae and spacial
relationship between excretory pore and nerve ring; 2, Dorsal view of lips; 3, Ventral view of
lips showing interlabium; 4, En face showing lateral alae; 5, Body at level of intestinal-ven-

tricular junction, lateral view; 6, Posterior extremity of female tail showing tuft of projections,
lateral view; 7, Female tail, lateral view.

1046 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

WW

Gl

I

200

50

12

iW

Figs. 8-13. Hysterothylacium fortalezae: 8, Ventral view of lips (paratype No. OCI
30.810c); 9, Posterior end of male paratype showing caudal papillae, lateral view; 10, Posterior
end of male showing post- and para-anal papillae, ventral view; 11, Lateral ala at esophageal-
ventricular junction; 12, Lateral ala immediately anterior to anus (scale same for Figs. 11, 12);
13, Posterior end of male from Florida showing caudal papillae, lateral view.

VOLUME 93, NUMBER 4 1047

morus cavalla and the type-host, S. grasiliensis (as S. maculatus) (Vicente,
personal communication). Although most characteristics reported by Klein
are comparable to those of our specimens collected from S$. maculatus and
Oligoplites saurus in the northern Gulf of Mexico and Biscayne Bay, Florida,
discrepancies exist among her description, figures, and a paratype (OCI
30.810c) loaned by Vicente. She reported 21 pairs of caudal papillae made up
of 5 preanal, 11 postanal, and 5 para-anal papillae. However, the paratype we
examined, which was stained and permanently mounted on a slide, pos-
sessed 28 pairs of caudal papillae: 18 preanal, 8 postanal, and no para-anal
(Fig. 9). Additionally, the cloacal opening was incorrectly located in the
illustration, leading to difficulty assessing the papillae. Klein noted the
prominent cervical alae, but did not include them in the en face illustration.

Hysterothylacium fortalezae and H. chaunaxi (Olsen, 1952) are distin-
guished from all other members of the genus in the Gulf of Mexico by having
a tuft of projections at their posterior extremity rather than dense, evenly
distributed spines on a conical tip; most of these projections were rounder
on H. chaunaxi. What appears to be similar projections are illustrated for
other species including specimens identified by Zhukov (1960) as C. fabri
from Scomber japonicus (as Pneumatophorus japonicus), also a mackerel,
from Putyatin Island, USSR. Those latter specimens and the status of C.
fabri need further investigation. The conspicuous cervical alae on H. for-
talezae separates it from H. chaunaxi. Those alae also occur on H. adun-
cum as shown by Punt (1941), H. trichiuri, and specimens identified as H.
clavatum by Kreis (1952) and as H. histiophori by Olsen (1952). They may
occur also on other species.

Hysterothylacium ogcocephali (Olsen), new combination
Figs. 14-21

Contracaecum ogcocephali Olsen, 1952:177-178, pl. I, figs. 12-17 (original
description; type-host Ogcocephalus radiatus; type-locality Tortugas,
Florida).

Thynnascaris ogcocephali.—Norris and Overstreet, 1975:333 (new combi-
nation).

Diagnosis.—Cuticle without conspicuous annulations; lips with ratio of
length to width 1:0.5—0.7, with flanges indented at anterior % of lip; caudal
papillar pairs: preanal 28-31, postanal 3—4 with 3rd or 4th papilla from pos-
terior extremity doubled; spicules 1-2% of body length; spicule ratio 1:0.8-
1.3; tail with short digitiform process lacking ornamentation on posterior
extremity.

General.—Body reaching greatest width near midbody. Cuticle with in-
conspicuous annulations; lateral alae distinct on each side, becoming more
apparent at levels posterior to rectum; supports nearly flat anteriorly and

1048 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

near midbody, with slight V-shape (in cross-section) in tail but not reaching
external surface of cuticle. Lips approximately equal in size, 1.4—1.8 times
longer than wide; flanges constricted at anterior 4 of lip; pulp narrowest at
base, gradually becoming wider anteriorly. Interlabia with height equal to
or slightly greater than width at base; interlabial grooves lacking. Esophagus
10-12% of body length. Ventriculus narrower than widest level of esopha-
gus, nearly spherical; ventricular appendage descending without angulation
from posterior portion of ventriculus. Nerve ring located at anterior 10-14%
of esophagus. Excretory pore immediately posterior to level of nerve ring.
Tail with short digitiform process; process lacking spines.

Male (based on 10 mature specimens for most characters and 5 tail mounts
for postanal papillae): Body 25.5-33.5 mm long by 0.4-0.6 mm at greatest
width; ratio of greatest width to length 1:49-60. Lips 180-236 long by 103-
139 wide. Nerve ring 482-593 from anterior extremity, 24—41 in breadth.
Esophagus 2.6—4.0 mm long by 166-216 wide. Ventriculus 108—166 long by
129-185 wide; ventricular appendage 0.9-1.5 mm long by 49-111 wide; ratio
for lengths of ventricular appendage to esophagus 1:2.4—3.2. Intestinal ce-
cum 185-309 long by 74-148 wide; ratio of cecal to ventricular appendage
lengths 1:3.3—7.3; ratio of cecal to esophageal lengths 1:9.1-18.5. Spicules
1-2% of body length, unequal; left spicule 330-500 long by 19-28 wide; right
spicule 350-660 long by 19-28 wide, longer than left spicule in 5 specimens;
spicule ratio 1:0.8-1.3. Caudal papillae 31-34 pairs; preanal papillae pairs
28-31, with posteriormost one on right side doubled on one specimen; post-
anal pairs 3, except 4 papillae on right side and 3 on left of one specimen;
with 3rd papilla from posterior extremity doubled except for one specimen
on which left 2nd papilla doubled; para-anal papillae lacking. Medioventral
preanal organ papillated, simple, at middle of preanal fold, single, directed
posteriorly. Tail flexed ventrad, 139-185.

Female (based on 10 mature specimens): Body 25.6—48.1 mm long by 0.6—
1.0 mm wide; ratio of greatest width to length 1:41-61. Lips 168-309 long
by 130-166 wide. Nerve ring 552-704 from anterior extremity, 30-55 in
breadth. Esophagus 3.1-5.3 mm long by 234-339 wide. Ventriculus 154-247
long by 185-241 wide; ventricular appendage 0.8—1.4 mm long by 74-123
wide; ratio for lengths of ventricular appendage to esophagus 1:2.8—4.7.
Intestinal cecum 185—444 long by 123-247 wide; ratio of cecal to ventricular

=

Figs. 14-21. Hysterothylacium ogcocephali: 14, Dorsal view of lips; 15, Ventral view of
lips showing interlabium; 16, En face; 17, Body at level of intestinal-ventricular junction,
lateral view; 18, Posterior extremity of female tail, lateral view; 19, Posterior end of female
tail, lateral view; 20, Posterior end of male showing caudal papillae, lateral view; 21, Posterior
end of male showing postanal papillae with atypical double second papilla on left side, ventral
view.

VOLUME 93, NUMBER 4

1050 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

appendage lengths 1:2.7—-5.0; ratio of cecal to esophageal lengths 1:11-17.
Vulva without salient lips, opening 9-13 mm or 24-40% of body length from
anterior extremity. Ovaries rarely extending beyond anterior level of vulva.
Eggs 30-55 in diameter. Tail 472-587 long including minute process.

Hosts.—Ogcocephalus radiatus (Mitchill), polka-dot batfish GCRL 7002,
O. cubifrons (Richardson) (Ogcocephalidae).

Site of infection.—Stomach and upper intestine.

Localities.—Biscayne Bay, Tortugas, and Lee County (26°52'00"N,
82°18'35"W), Florida.

Specimens deposited. —USNM Helm. Coll. No. 75839 (pair); BMNH
Reg. No. 1980.361—362 (pair); OCI 31.604.

Remarks.—Our report is the second for H. ogcocephali. Olsen (1952)
originally described it from Ogcocephalus radiatus using data and figures
from an unpublished thesis by B. Rees with most measurements given for
one male and one female. Consequently, his description did not provide
morphological variation among individuals. Additionally, Olsen did not treat
some specific characters, and some differed from what we found. He re-
ported 18-26 pairs of preanal papillae and 1-2 pairs of para-anal ones,
whereas we counted a total of 31-34 pairs including the 3 postanal ones. We
examined the holotype (USNM No. 37241) and confirmed that the lateral
cephalic flanges had indentations near the anterior 4 similar to those for
Theringascaris inquies. The holotype’s poor condition, however, resulted
in many features being obscured. The terminal end of the tail lacked a
multispinous structure (Fig. 21); a structure present on all species with sim-
ilar lip morphology. This multispinous structure may prove to be a valuable
diagnostic character in ascaridoid nematodes and should be noted when
present.

Hysterothylacium reliquens (Norris and Overstreet), new combination
Figs. 22-39

T. reliquens Norris and Overstreet, 1975:330-332, figs. 1-11 (original de-
scription; type-host Archosargus probatocephalus; type-locality Missis-
sippi Sound, Mississippi).

Diagnosis.—Cuticle without conspicuous annulations; lips with ratio of
length to width 1:0.6—1.0, with flanges indented equatorially; caudal papillar
pairs: preanal 23-29, postanal 4-5 with 3rd pair from posterior extremity
doubled; spicules 3-8% of body length; spicule-ratio 1:0.6—1.3; multispinous
conical process on posterior extremity.

General (based on specimens from Ogcocephalus cubifrons).—Body
reaching greatest width near midbody. Cuticle with inconspicuous annula-
tions and minute lateral alae; alae becoming more apparent at levels pos-
terior to rectum. Lips approximately equal in size, all longer than wide;

VOLUME 93, NUMBER 4 1051

400

600

Figs. 22-28. Hysterothylacium reliquens: 22, Anterior end showing spacial relationship
between excretory pore and nerve ring; 23, Dorsal view of lips; 24, Ventral view of lips showing
interlabium; 25, Multispinous process on female tail, lateral view; 26, En face; 27, Body at
level of intestinal-ventricular junction, lateral view; 28, Posterior end of male showing caudal
papillae, lateral view.

1052 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

flanges constricted near middle of lip; pulp pedunculated. Interlabia with
height equal to or slightly greater than width at base; interlabial grooves
lacking. Esophagus 11-13% of body length. Ventriculus narrower than wid-
est level of esophagus, usually longer than broad; ventricular appendage
departing without angulation from posterior portion of ventriculus. Nerve
ring located at anterior 16-28% of esophagus. Excretory pore at or imme-
diately posterior to level of nerve ring; excretory system with duct extending
posteromesial from pore toward esophagus before entering left lateral cord
as lateral canal; lateral (=longitudinal) canal gradually increasing in diameter
until termination within tail. Tail with spined conical mucronate extremity.

Male (based on 11 mature specimens, but 9 tail mounts for postanal pa-
pillae): Body 25-40 mm long by 0.5-0.9 mm at greatest width; ratio of
greatest width to length 1:30—57. Lips 234-291 long by 234-285 wide. Nerve
ring 646-977 from anterior extremity, 49-69 in breadth. Esophagus 3.3—4.5
mm long by 150-230 wide. Ventriculus 141-300 long by 110-236 wide; ven-
tricular appendage 1.2-1.7 mm long by 80-130 wide; ratio for lengths of
ventricular appendage to esophagus 1:2.5-2.8. Intestinal cecum 315-598
long by 260-310 wide; ratio of cecal to ventricular appendage lengths 1:2.2-
3.5; ratio of cecal to esophageal lengths 1:7-10. Spicules 3-8% of body
length, 1.7-2.4 mm long by 9-20 wide; right spicule longer than left one in
5, equal in 5, and shorter in 1 specimen; spicule-ratio 1:0.6—1.3. Caudal
papillae 27-33 pairs, becoming closer together and more medial as ap-
proaching anus; preanal pairs 23-29; postanal pairs 4-5, with 3rd pair from
posterior extremity doubled; para-anal papillae lacking. Medioventral pre-
anal organ distinct, papillated. Tail flexed ventrad, 123-185 long including
a multispinous process 12—46 long.

Female (based on 18 mature specimens): Body 21-44 mm long by 0.5-1.1
mm wide; ratio of greatest width to length 1:33-58. Lips 222-340 long by
204-309 wide. Nerve ring 646-977 from anterior extremity, 21-57 in
breadth. Esophagus 2.9-5.0 mm long by 180-280 wide. Ventriculus 142-286
long by 126-331 wide; ventricular appendage 1.0-—2.9 mm long by 95-140
wide; ratio for lengths of ventricular appendage to esophagus 1:1.7-3.0.

=>

Figs. 29-39. Hysterothylacium reliquens from Batrachoides surinamensis (29, 30, 31),
Ogcocephalus cubifrons (33), and Archosargus probatocephalus (32, 34-39). Lateral ala at
anterior (29), middle (30), and posterior (31) parts of male. 32, Lateral ala at level of base of
esophagus equivalent to region in Fig. 29 (scale applies to all figures to its left); 33, Posterior
end of male showing postanal papillae, ventral view; 34, Female tail, lateral view; 35, Posterior
end of male showing postanal papillae, ventral view; 36-39, Cross-sections of excretory system
showing excretory canal anterior to nerve ring (36), excretory pore (37), excretory duct (37-
38), and excretory canal at esophageal-ventricular junction (39). Abbreviations: ec = excretory
canal, ed = excretory duct, ep = excretory pore, es = esophagus, Ic = lateral cord.

VOLUME 93, NUMBER 4 1053

<ijil, j ‘es

1054 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Intestinal cecum 394-788 long by 265-322 wide; ratio of cecal to appendage
lengths 1:1.6—5.2; ratio of cecal to esophageal lengths 1:5—8. Vulva without
salient lips, opening 6.5-17.0 mm or 16-45% of body length from anterior
extremity. Ovaries rarely extending beyond anterior level of vulva. Tail
370-574 long including multispinous process 12-60 long.

Hosts.—Archosargus probatocephalus (Walbaum), sheepshead (Spari-
dae) type-host; Opsanus beta (Goode and Bean), Gulf toadfish, Batra-
choides surinamensis (Bloch and Schneider), B. pacifici (Gtinther), B. bou-
lengeri Gilbert and Starks, and Batrachoides sp. (new species from El
Salvador and the Pacific coast of Mexico being described by Collette and
Russo, in press) (Batrachoididae); Ogcocephalus cubifrons (Richardson),
Ogcocephalus pantostictus Bradbury, (GCRL 564) (Ogcocephalidae);
Arius felis (Linnaeus), sea catfish (Ariidae); Chilomycterus schoepfi (Wal-
baum), striped burrfish (Diodontidae); Micropogonias undulatus (Lin-
naeus), Atlantic croaker, Sciaenops ocellata (Linnaeus), red drum (Sciae-
nidae); Gymnothorax nigromarginatus (Girard), blackedge moray
(Muraenidae).

Sites of infection.—Stomach and intestine.

Localities.—Mississippi Sound, Mississippi (Archosargus probatoce-
phalus, Chilomycterus schoepfi, and Micropogonias undulatus); offshore in
northern Gulf of Mexico (Sciaenops ocellata, 30°10'’N, 87°45’ W, 11 meters;
M. undulatus and Gymnothorax nigromarginatus, 29°33'N, 88°25'W, 46
meters; C. schoepfi and A. probatocephalus, 29°58'—59'N, 88°44'W, 15
meters; Ogcocephalus sp., offshore from Horn Island); Galveston, Texas
(Arius felis); Biscayne Bay, Florida (Opsanus beta); Miami, Florida (Ogco-
cephalus cubifrons); Honduras, Guyana, and northern Brazil at 4°48’N,
51°19'W (Batrachoides surinamensis, TABL 104319, 104839, USNM
219457); Panama (Miraflores Locks, San Francisco Beach, and 8°60’N,
79°31'W) and Colombia (B. pacifici, USNM 111862, 111865, UMML 26705,
USNM 219385, B. boulengeri, GCRL 8402); and El Salvador (Bahia de
Jiquilisco) and Mexico (Gulf of Tehuantepec) (Batrachoides sp. being de-
scribed by Collette and Russo, in press, USNM 219788, SIO 63-517).

Specimens deposited.—from Ogcocephalus cubifrons (2 males and fe-
males) USNM Helm. Coll. No. 75840; from Batrachoides surinamensis (6
specimens) USNM Helm. Coll. No. 75498; from Ogcocephalus pantostictus
(6 specimens) USNM Helm. Coll. No. 75841.

Remarks.—In addition to adding new host and locality records to those
provided by Norris and Overstreet (1975), we present and discuss minor
morphological variations among specimens from several hosts and provide
descriptive data from specimens from a batfish for comparisons with de-
scriptions of other species.

Worms from all hosts are similar, but we note minor differences in fea-
tures such as postanal papillae. Norris and Overstreet (1975) reported 4—6

VOLUME 93, NUMBER 4 1055

pairs of postanal papillae for H. reliquens from Archosargus probatoce-
phalus. We examined 14 ventral tail mounts of the same and additional
specimens from A. probatocephalus from Mississippi Sound and also count-
ed 4—6 pairs; however, all but three males had uneven postanal papillar
patterns. Six worms had 5 papillae on the right side and 4 on the left, and
four worms had 6 on the right and 5 on the left. Additionally, the presence
of doubled papillae was not originally noted. The fourth postanal papilla on
the right side and third on the left from the posterior extremity of the worm
exhibited this doubled condition. Of those worms with paired postanal pa-
pillae, one worm had 5 pairs with the fourth papilla doubled and two worms
had 4 pairs with the third papilla doubled.

Papillae on three males from the sea catfish were like those from the
batfish with 4 or 5 even pairs. Either the third or fourth pair from the worm’s
posterior end was doubled. However, the sclerotized supports of the lateral
alae protruded beyond the cuticle, especially at the esophageal level.

Numerous worms were examined from several species of Batrachoides
collected from Atlantic coasts of South America, the Panama Canal, and
Colombia to Mexico in the Pacific Ocean. The papillae and alae extension
(Figs. 29-31) agreed with those of catfish specimens indicated above. Spec-
imens from Opsanus beta, also a batrachoidid, from Biscayne Bay, Florida,
were the same, suggesting that the feature may be either host- or temper-
ature-induced, since the worms in most other fishes as represented by the
sheepshead, or type-host, in the northern Gulf of Mexico appeared to have
indistinct alae by comparison (Fig. 32).

The presence of H. reliquens in the Pacific Ocean is unusual because few,
if any, other ascaridoids in nonpelagic fishes occur in both the Atlantic and
Pacific oceans. Hysterothylacium aduncum, considered a junior synonym
of H. gadi by Hartwich (1975) and others, has been reported from the
Pacific, but specimens reported from both oceans need critical examination.

Hysterothylacium reliquens heavily infects different hosts in different re-
gions. As examples, heavy infections occurred in Ogcocephalus cubifrons
from Miami, Florida, Archosargus probatocephalus from Mississippi, Arius
felis from Galveston Bay, Texas, and Batrachoides surinamensis from
northern Brazil. Collette and Russo (in press) revised the genus Batra-
choides and examined a few intestines of each of the nine species, finding
H. reliquens in four species. Batrachoides manglae Cervigon from hyper-
saline regions of Venezuela and B. goldmani Evermann and Goldsborough
from freshwater streams in Mexico and Guatemala were two species that
did not harbor H. reliquens. We speculate that fish acquire H. reliquens in
relatively high-salinity regions or from intermediate hosts that recently in-
habited such regions based on 1) infections in batrachoidids restricted to
locations with salinities above 20 ppt but not hypersaline; 2) heavy infections
in some fishes caught offshore or in high-salinity habitats; 3) large numbers

1056 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of noninfected susceptible fishes in relatively low-salinity water of Missis- |

sippi Sound and elsewhere; and 4) findings in some infected fish from rel-
atively low salinities of Mississippi Sound of additional concurrent hel-
minths that depend on high-salinity habitats to complete their cycles.
Infections also relate to food habits and perhaps to physiological races of

worms. In any event, we neither encountered infected A. felis in Mississippi |
Sound nor encountered or heard about conspicuously infected sheepshead |
in Texas (including Galveston Bay) from numerous Texas sources. Speci- |

mens of Chilomycterus schoepfi and Micropogonias undulatus occasionally

had infections in Mississippi Sound or near the barrier islands, but never |
heavy ones like in individual sheepshead which often harbored more than |

350 worms, half of which were mature and many of which exited the host

following removal of the fish from water. Other than the single Atlantic |
croaker (M. undulatus) listed from 46 meters deep off the Chandeleur I[s- |
lands in 1973, we did not collect any other specimens of H. reliquens from |

offshore croaker. Between 1974 and 1977, we examined 1,700 croaker col-

lected by the R/V OREGON II and GEORGE M. BOWERS from 42 stations |

with depths ranging from 11-90 meters and did not find H. reliquens.

Four species of Hysterothylacium, H. reliquens and H. ogcocephali from {|
the Gulf of Mexico, H. melichthysi from the Hawaiian Islands, and 4H. |
rhacodes from the eastern Mediterranean Sea, have the diagnostically elon- |
gated lips. The lips contain elongated pulp bordered by distinctly indented
lateral flanges. Other species also have indented flanges, but not in con- |
junction with elongated lips. Flanges on H. reliquens are indented equato-
rially, whereas the position on those of H. ogcocephali and H. melichthysi \
occurs about 4 of the length from the anterior end. Hysterothylacium rha- |
codes has similar flanges to H. reliquens, but does not have any doubled |

postanal papillae.

Hysterothylacium chaunaxi (Olsen), new combination
Figs. 40-45

Contracaecum chaunaxi Olsen, 1952:178, pl. II, figs. 18-22 (original de- |

scription; type-host Chaunax sp.; type-locality Tortugas, Florida).

Diagnosis.—Cuticle without conspicuous annulations; lips with ratio of |
length to width 1:0.8, with pulp not pedunculate; caudal papillar pairs: |:
preanal 23, postanal 7 with doubled papilla 6th from posterior extremity on

right side; tuft of projections at posterior extremity.
General.—Body reaching greatest width near midbody. Cuticle with in-
conspicuous annulations. Lips approximately equal in size; pulp not pedun-

culate, widest at base. Interlabia with width at base equal to or slightly |

greater than height; interlabial grooves lacking. Ventriculus narrower than

widest level of esophagus, usually longer than wide; ventricular appendage |

VOLUME 93, NUMBER 4 1057

Figs. 40-45. Hysterothylacium chaunaxi: 40, Dorsal view of lips; 41, Ventral view of lips
showing interlabium; 42, Posterior extremity of female tail showing terminal tuft with projec-
tions, lateral view; 43, En face; 44, Body at level of intestinal-ventricular junction, lateral
view; 45, Posterior end of male showing postanal papillae, ventral view; note broken spicules.

1058 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

descending without angulation from posterior portion of ventriculus. Ex-
cretory pore just posterior to level of nerve ring. Tail with tuft of rounded
and pointed projections at posterior extremity.

Male (based on | specimen plus holotype USNM Helm. Coll. No. 37243
with its measurements in parentheses): Body 34.7 (39.5) mm long by 947
(709) at greatest width; ratio of greatest width to length 1:36 (1:56). Lips 253
(120) long. Esophagus 2.8 (3.3) mm long by 333 wide or 8 (8)% of body
length. Ventriculus 284 (259) long by 174 (279) wide; ventricular appendage
1.2 mm long (not visible) by 173 wide; ratio for lengths of ventricular ap-
pendage to esophagus 1:2.3. Intestinal cecum 819 (1.0 mm) long by 488 (247)
wide; ratio of cecal to ventricular appendage lengths 1:1.4; ratio of cecal to
esophagus lengths 1:3.4 (1:3.3). Nerve ring located within anterior 26% of |
esophagus, 50 in breadth. Excretory pore immediately below posterior level |
of nerve ring. Spicules broken at cloaca, 1.5 mm long by 31 wide. Caudal |
papillae 30 pairs (not able to count), becoming closer together and more |
medial as approaching anus; preanal pairs 23; postanal pairs 7, doubled |
papilla 5th from posterior extremity on right side only; para-anal papillae |
lacking. Medioventral preanal organ distinct, papillated. Tail 185 (203) long |
including tuft with projections up to 12 long on posterior extremity.

Host.—Chaunax pictus Lowe (Chaunacidae) GCRL 679.

Site of infection.—Intestine. |

Localities.—Tortugas, Florida (type-locality) and 29°13'N, 87°55.5’W (460 |
meters) R/V OREGON Station No. 3680. |

Specimen deposited.—USNM Helm. Coll. No. 75842, body-fragments in |}
vial plus en face and tail mounted in glycerin jelly on separate slides. '

Remarks.—Olsen (1952) noted the tuft on the tail of H. chaunaxi, al- §
though he did not illustrate it. On the basis of this distinct structure at the |
posterior extremity (Figs. 42, 45), H. chaunaxi and H. fortalezae differ |
from all other examined members of the genus. Hysterothylacium chaunaxi |
can be distinguished from H. fortalezae by body length, ratio of greatest |
width to length, ratio of cecal to ventricular appendage lengths, and number |
and arrangement of pre- and postanal papillae. Additionally, H. chaunaxi |
lacks conspicuous anterior cervical alae. ;

One of five, fixed, museum specimens of Chaunax pictus collected from |
the same locality in the Gulf of Mexico harbored the single male H. chaun- |
axi. Olsen described H. chaunaxi from Chaunax sp. at Tortugas, Florida, |
also from a single male. The holotype is rather opaque; consequently, we
do not include data on all its features above. The type-host is presumably |
C. pictus. No other records of an infection exist. |

Hysterothylacium incurvum (Rudolphi), new combination
Figs. 46-56

Ascaris incurva Rudolphi, 1819:51, 292-293 (original description; type-host |)
Xiphias gladius; type-locality Baltic Sea). |

VOLUME 93, NUMBER 4 1059

Contracaecum incurvum.—Baylis and Daubney, 1922:282—284, figs. 17-18
(new combination).

Contracaecum (Thynnascaris) incurvum.—Dollfus, 1935:89-90 (new com-
bination).

Thynnascaris incurva.—Hartwich, 1957:240 (new combination).

Diagnosis.—Cuticle with enlarged and modified annules on ventral sur-
face of male near cloaca; alae bifurcated at tips; lips with ratio of length to
width 1:1.0-1.6, with pulp deeply constricted at base, with interlabial
grooves deep; caudal papillar pairs: preanal 27-28, postanal 4, para-anal 1,
doubled; spicules 12-25% of body length; spicule-ratio 1:0.9-1.0; tail grad-
ually tapering, lacking ornamentation on posterior extremity (note exception
below).

General (based on specimens from Florida).—Body reaching greatest
width near midbody. Cuticle with inconspicuous annulations except those
enlarged and modified on ventral surface of male near cloaca. Lateral alae
bifurcate at tips throughout worm; supports in cross-section nearly meeting
before diverging, typically external to cuticle; cervical alae not flaring. Lips
approximately equal in size, all broader than long; flanges widest near base,
indented at anterior 4 of lip. Pulp conspicuously narrow near base. Inter-
labia with height 2 times greater than width at base, with rounded tips;
interlabial grooves deep; adjacent grooves nearly merging at base of each
lip. Esophagus clavate, 7—-16% of body length. Ventriculus narrower than
widest level of esophagus, generally longer than broad. Nerve ring located
between anterior 7-19% of esophagus. Excretory pore immediately poste-
rior to level of nerve ring. Tail gradually tapering.

Male (based on 11 mature specimens): Body 17-34 mm long by 606-772
at greatest width; ratio of greatest width to length 1:29—-49. Lips 86-154 long
by 108-212 wide. Nerve ring 376-593 from anterior extremity, 33-55 in
breadth. Esophagus 2.3—5.2 mm long by 92-166 wide. Ventriculus 117-166
long by 129-216 wide; ventricular appendage 1.7—2.6 mm long by 61-195
wide; ratio for lengths of ventricular appendage to esophagus 1:1-3. Intes-
tinal cecum 1.9-4.5 mm long by 166—233 wide; ratio of lengths of cecum to
ventricular appendage 1:0.4-1.0; ratio for lengths of cecal to esophageal
length 1:1.0-1.5. Spicules 12-25% of body length, 2.6-8.7 mm long by 9-18
wide, equal in 8 specimens; left spicule longer in 2 specimens; spicule-ratio
1:0.9-1.0. Caudal papillae 31-32 pairs, becoming closer and more medial as
approaching anus; preanal pairs 27-28; postanal pairs 4; para-anal papilla
1, doubled. Medioventral preanal organ distinct, papillated. Tail flexed ven-
trad, 79-197 long, without ornamentation. Modified annules on ventral sur-
face beginning near anus, extending anteriorly 2.0-6.1 mm or 10-18% of
body length; each annule with 45—57 minute longitudinal supports; supports
similar, lamellar, semicircular, 8—17 apart.

Female (based on 21 mature specimens): Body 25-69 mm long by 0.5—1.7

1060 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mm wide; ratio of greatest width to length 1:28-61. Lips 110-214 long by
132-222 wide. Nerve ring 414-646 from anterior extremity, 26-48 in
breadth. Esophagus clavate, 2.6—7.2 mm long by 100-278 wide. Ventriculus
173-362 by 210-551; ventricular appendage 0.6—4.0 mm long by 0.2-1.1 mm
wide; ratio for lengths of ventricular appendage to esophagus 1:1.5-4.5.
Intestinal cecum 1.5—5.1 mm long by 80-488 wide; ratio of lengths of cecum
to ventricular appendage 1:0.3-18.0; ratio of cecal to esophageal lengths
1:1.0-2.0. Vulva opening 7.3—-35.1 mm or 25-40% of body length from an-
terior extremity, not protruding. Ovaries rarely extending anteriad beyond
level of vulva. Eggs 18-63 in diameter. Tail 677—945 long, with tip usually
without spines but single spine on one worm.

Hosts.—Xiphias gladius (Linnaeus), swordfish (Xiphiidae) type-host; Tet-
rapturus albidus (Poey), white marlin; T. brevirostris (Playfair), striped mar-
lin (from Tanzania) (Istiophoridae).

Site of infection.—Stomach.

Localities.—Offshore from Destin (T. albidus) and Miami (X. gladius),
Florida; Baltic Sea; off India; near Zanzibar, Tanzania.

Specimens deposited.—From T. albidus (2 pairs) USNM Helm. Coll. No.
75843; (pair) BMNH Reg. No. 1980.363-364; (pair) ZM No. 7053.

Remarks.—We examined six male and six female syntypes of H. incur- |
vum (ZM No. 468) deposited by Rudolphi in October 1818, and, even though
their condition was poor, we determined the presence of diagnostic deep |
interlabial grooves and modified ventral annules on the males. They lacked |

prominent cervical alae. Because of their condition and because we did not |

see all the syntypes, we refrain from establishing a lectotype, but neverthe- |
less consider the specimens conspecific with our material, some of which |
came from the type-host. With the permission of G. Hartwich, who counted |
the same number and kind of postanal papillae on two syntypes that we did |

(personal communication), we deposited a female and the tail and two an- }-

terior fragments of male syntypes in the U.S. National Museum (USNM j'
Helm. Coll. No. 75151). 1
In our material the vulva did not protrude as reported by Berland (1961), |
suggesting that such a feature results from a shrinkage or fixation artifact |
as Berland proposed.

We also examined a male specimen (BMNH No. 1923.1.22.3) from a |
swordfish caught off India and identified by Baylis and Daubney (1922) as |
Contracaecum incurvum and 21 females and 5 males (BMNH No. 1962.40- |
47) from Tetrapturus brevirostris (as Makaira audax) caught near Zanzibar.

—

Figs. 46-52. Hysterothylacium incurvum: 46, Anterior end showing spacial relationship |
between excretory pore and nerve ring, lateral view; 47, Dorsal view of lips; 48, Ventral view _ |
of lips showing interlabium; 49, En face; 50, Posterior end of male showing both caudal papillae |
and annules on ventral surface, lateral view; 51, Posterior end of male showing postanal pa- |
pillae, lateral view; 52, Posterior end of male showing postanal papillae, ventral view.

1061

VOLUME 93, NUMBER 4

WP
nd

1062 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

All appeared to be H. incurvum. In fact, the species probably occurs
throughout the range of swordfishes. Unconfirmed reports of C. incurvum
indicate the worm’s abundance in white marlin off Ocean City, Maryland
(Wallace and Wallace, 1942) and the southern Gulf of Mexico (Nikolaeva
and Parukhin, 1968) and in 7. audax (as Makaira mitsukurii Jordan and
Snyder) from Brett Cape to the Cavalli Islands, New Zealand (Morrow,
1952). Hosts other than swordfish such as the sailfish (as Histiophorus gla-
dius), the Spanish mackerel, and the sand-bar shark, Carcharhinus plumbe-_ |
us (Nardo) (as Carcharias cerulcus) have been reported as hosts of mature }
specimens (e.g., Linton, 1901; Walton, 1927; Nikolaeva and Parukhin,
1968), but such identifications also require confirmation. |

Nikolaeva and Parukhin (1968) reported a maximum of 1,787 specimens |
from a single white marlin. We never encountered such heavy infections, |
but numerous individuals often exited from the mouth and opercula of dead }
fish to the disgust of fishermen. |

Only two species of Hysterothylacium have been reported from sword- |
fishes and billfishes: H. incurvum and H. histiophori. Rudolphi (1819) orig- |
inally described Ascaris incurva from the swordfish, Xiphias gladius Lin- |
naeus, in the Baltic Sea, a rather rare locality for the host (A. Wheeler, |
personal communication). Yamaguti (1935) described C. histiophori from |
the sailfish Istiophorus platypterus (Shaw and Nodder) (as Histiophorus |
orientalis) in Toyama Bay, Japan. Neither Rudolphi nor Yamaguti provided |
many diagnostic features in their descriptions. |

The status of H. histiophori has not been adequately ascertained. Ya- |
maguti (1935), who based the description on one mature female and an |
immature pair, differentiated it from H. incurvum, which he reported from |
Xiphias gladius in the same paper, on the basis of body length and positions |
of both the nerve ring and vulva. We have been unable to locate type spec- |
imens of H. histiophori or any other specimens from the same host from |
the Meguro Parasitological Museum (Atsuo Ichihara, personal communi- |
cation) or elsewhere. Olsen (1952) reported C. histiophori from the sailfish |
I. platypterus (as I. americanus) caught off the ‘‘Florida coast.’’ We do not |
know whether these specimens (6 mature and 3 immature males, plus 3 |
mature and 14 immature females), not deposited in the U.S. National Mu- |
seum, represent H. histiophori or a new species. In any event, even though |
it has deep interlabial grooves and modified ventral annules in the male, we |
consider its prominent cervical alae sufficient to differentiate the species |
from H. incurvum.

Hysterothylacium corrugatum, new species
Figs. 57-63

Diagnosis.—Cuticle with enlarged and modified annules on ventral sur- !
face of male near cloaca; lips with ratio of length to width 1:1.0-1.3, with |

VOLUME 93, NUMBER 4 1063

A We WEP
AW 28) =

200

f
ee Tp OPS
nl Kea aud ti ip, Ijin lee
ae \9 eee,
ae t

Figs. 53-56. Hysterothylacium incurvum: 53, Cross-section of worm through level of ven-
tricular appendage; 54, Cross-section of worm showing intestinal cecum; 55, Cross-section
through modified annules on ventral surface of males; 56, Posterior end of female tail, lateral
view. Abbreviations: a = lateral alae, c = intestinal cecum, e = esophagus, i = intestine, ic =
inner cuticular layer, Ic = lateral cord, Is = longitudinal supports, m = muscle, oc = outer
cuticular layer, and va = ventricular appendage.

1064 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

interlabial grooves lacking; caudal papillar pairs: preanal 24-26, postanal 4,
para-anal 1, doubled; spicules 4—-7% of body length; spicule-ratio 1:1.0; tail
gradually tapering, terminating without ornamentation.

General (based on specimens from Florida).—Body reaching greatest
width near midbody. Cuticle with inconspicuous annulations except ven-
trally on male’s posterior. Cervical alae not flaring. Lips approximately
equal in size, occasionally broader than long; flanges widest near base,
indented at anterior 2/5. Pulp for each lip not narrowed at base, each with
2—4 lobular projections often a few times longer than wide. Interlabia with
height equal to or slightly less than width at base, with rounded tips; inter-
labial grooves lacking. Esophagus clavate, 5-13% of body length. Ventric-
ulus narrower than widest level of esophagus, generally longer than broad;
ventricular appendage descending without angulation from posterior portion
of ventriculus. Nerve ring located between anterior 6-12% of esophagus.
Excretory pore immediately posterior to level of nerve ring. Tail gradually
tapering, terminating without ornamentation.

Male (based on 5 mature specimens): Body 26-50 mm long by 0.4—-1.0
mm at greatest width; ratio of greatest width to length 1:49-69. Lips 96-145
long by 89-168 wide. Nerve ring 284-593 from anterior extremity, 30—48 in
breadth. Esophagus 3.0-5.6 mm long by 60-191 wide. Ventriculus 36-154
long by 57-185 wide; ventricular appendage 2.4—3.8 mm long by 43-60 wide;
ratio for lengths of ventricular appendage to esophagus 1:1.0—2.3. Intestinal
cecum 2.3—4.6 mm long by 48-204 wide; ratio of cecal to ventricular ap-
pendage lengths 1:0.6-0.9; ratio of cecal to esophageal lengths 1:1.0-1.3.
Spicules 4—7% of body length, similar; left spicule 1.2—2.5 mm long by 12-
18 wide; right spicule 1.2-2.6 mm long by 12-18 wide; spicule-ratio 1:1.0.
Caudal papillae 29-31 pairs, becoming closer together and more medial as
approaching anus; preanal pairs 24-26; postanal pairs 4; para-anal pair 1,
doubled. Medioventral preanal organ distinct, papillated. Tail flexed ven-
trad, 191-245 long. Modified annules on ventral surface, beginning near
anus, extending anteriorly 1.0-4.1 mm or 3-15% of body length.

Female (based on 6 mature specimens): Body 100-142 mm long by 1.7-
2.6 mm wide; ratio of greatest width to length 1:40-75. Lips 203-259 long
by 210-247 wide. Nerve ring 709-756 from anterior extremity, 43-61 in
breadth. Esophagus clavate, 5.5-8.5 mm long by 283-315 wide. Ventriculus
154-472 by 220-283; ventricular appendage 3.5—-4.3 mm long by 80-98 wide;
ratio of ventricular appendage to esophagus lengths 1:1.5-2.0. Intestinal
cecum 5.3—7.2 mm long by 346-583 wide; ratio of lengths of cecum to ven-
tricular appendage 1:0.6-0.7; ratio of cecal to esophagus lengths 1:1.0—1.2.
Vulva opening 35-40 mm or 33-39% of body length from anterior extremity.
Ovaries rarely extending anteriad beyond level of vulva. Eggs 40-80 in
diameter. Tail 0.8—1.1 long.

Host.—Xiphias gladius (Linnaeus), swordfish (Xiphiidae).

VOLUME 93, NUMBER 4 1065

Site of infection.—Stomach.

Localities. —Offshore from Miami (type-locality); Panama City, Florida;
and Manta, Ecuador.

Specimens deposited.—Holotype, male, USNM Helm. Coll. No. 75844;
Allotype, female, No. 75845; Paratypes, No. 75846 (pair), British Museum
(Natural History) Reg. No. 1980.365-366 (pair); Additional specimens,
ZM No. 7054 (female and anterior of male).

Etymology.—The Latin corrugatum means ‘‘ridged,’’ referring to the
modified annules on the ventral surface of the males.

Remarks.—We examined three mature specimens, a male and two fe-
males (USNM Helm. Coll. No. 60051), from the swordfish caught at Manta,
Ecuador, which were identified by M. Chitwood as Contracaecum incur-
vum. They lacked interlabial grooves, the male had 30 preanal papillae, and
we consider them H. corrugatum.

Of the more than 300 nematodes from 22 specimens of white marlin caught
offshore from Destin, Florida, to Orange Beach, Alabama, and the 85 from
4 specimens of the swordfish caught offshore from Miami and Panama City,
Florida, only H. incurvum infected the white marlin, but both H. incurvum
and H. corrugatum occurred in the swordfish.

By possessing modified annules on the ventral surface of the male, H.
corrugatum most closely resembles H. incurvum and H. histiophori of Ol-
sen (1952), but differs conspicuously by lacking deep interlabial grooves.

Hysterothylacium habena (Linton), new combination
Figs. 64-67

Ascaris habena Linton, 1900:302-303, figs. 109-115 (original description;
type-host Opsanus tau; type-locality Woods Hole, Massachusetts).

Contracaecum habena.—Linton, 1934:125 (distribution and prevalence rec-
ords; new combination).

Thynnascaris habena.—Hartwich, 1957 (new combination).-

Diagnosis.—Cuticle with inconspicuous annulations; lips approximately
as wide as long, narrowest at base, with ratio of greatest length to width
1:0.9-1.1, with labial flanges appearing as broad triangular projections slight-
ly indented at anterior % of lip; pulp pedunculate; caudal papillar pairs:
preanal 21-31; postanal 3-4, with 3rd pair from posterior extremity doubled;
process on posterior extremity conical, multispinous.

Remarks.—Since Linton’s (1900) original description of H. habena from
the oyster toadfish, Opsanus tau (Linnaeus), from Massachusetts, it has
been redescribed by Chandler (1943) and by Norris and Overstreet (1975).
We reexamined 2 males and 1 female from O. tau collected at Gloucester
Point, Virginia, reported by Norris and Overstreet and supplement that
description below. A pair of prominent phasmids occurred laterally on the

1066 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

spined distal process and three pairs of postanal papillae occurred on one
specimen and four on the other. In both cases, the third pair from the
posterior extremity was doubled. Designating papillae as para-anal is sub-
jective for many species, including H. habena. Even though two pairs of
papillae occur lateral to the lip anterior to the anus of H. habena, these
appear as a continuum of preanal papillae as evident in ventral view (Fig.
64) rather than being clearly set-off and could therefore be considered pre-
anal papillae. Ventral tail mounts are especially helpful in determining pa-
pillar patterns.

Hysterothylacium habena probably does not occur in the northern Gulf
of Mexico as assumed by Hutton et al. (1962) and others. Their assumption
was based on the presence of a larval species of Hysterothylacium (=Con-
tracaecum) in penaeid shrimps from North Carolina to Central America and
the Caribbean (actually at least three species infect these shrimps, Norris
and Overstreet, 1976, personal observations) which have a ratio for the
lengths of intestinal cecum to ventricular appendage similar to that for H.
habena. Hysterothylacium habena extends from at least Massachusetts to
Georgia in a batrachoidid from high-salinity habitats. We have not seen it
among abundant ascaridoid material from batrachoidids collected in Florida,
the northern Gulf of Mexico, and Central and South America.

Theringascaris Pereira

Theringascaris Pereira, 1935 (type-species I. iheringascaris Pereira, 1935,
junior synonym of I. inquies).

Diagnosis.—Body relatively narrow. Cuticle with distinctly plicated an-
nulations lacking spines. Cuticular alae present. Lips approximately equal
in size, bearing transparent cuticular flanges on lateral margins; dorsal lip
with two lateral doubled papillae; subventral lips with amphid, adjacent
mediolateral doubled papilla, and single lateral papilla. Dentigerous ridges
absent. Interlabia present, without deep interlabial grooves. Ventriculus
nearly spherical; ventricular appendage cylindrical with septum dividing
Structure into 2 equal longitudinal pouches; intestinal cecum shorter than
ventricular appendage. Excretory system with duct off central commissure
opening as pore near level of nerve ring, with excretory canals extending

—=>

Figs. 57-63. Hysterothylacium corrugatum: 57, Dorsal view of lips; 58, Ventral view of
lips showing interlabium; 59, En face; 60, Posterior end of male showing modified annules and
postanal and para-anal papillae, ventral view; 61, Posterior end of male showing postanal and
para-anal papillae, lateral view; 62, Posterior end of female tail, lateral view; 63, Posterior end
of male showing entire view of spicules, lateral view.

VOLUME 93, NUMBER 4

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1068 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

posteriorly in both lateral cords. Spicules similar and approximately equal.
Gubernaculum absent. Lateral row of caudal papillae in addition to regular
pre- and postanal papillae. Medioventral preanal organ present. Vulva an-
terior to midbody. Uterus didelphic, opisthodelphic. Tail conical. Phasmids
present. Parasites of marine fish. Type-species: [heringascaris iheringas-
caris Pereira, 1935 Gunior synonym of I. inquies, see below).

Comparisons: [heringascaris has long been considered a junior synonym
of Thynnascaris or its synonym Contracaecum (Johnston and Mawson,
1945). Its only species can be easily differentiated from those of Hysterothy-
lacium by possessing distinctly plicated cuticular annulations and an ad-
ditional lateral pair of caudal papillar rows. However, H. cornutum does
have irregular lateral papillae extending anteriorly and posteriorly from the
anal level. Once all species of Hysterothylacium are more critically evalu-
ated, they may additionally differ if none have a similar excretory system.

Biological differences may also occur between I. inquies and species of
Hysterothylacium. Adult I. inquies live abundantly free in the lumen of the
pyloric ceca. Those in the stomach occur free, entwined in digesting prey,
and embedded in the host’s mucosal tissue. Fourth stage larvae embed in
nodules generally causing much less host response than do related species
of Goezia (see Deardorff and Overstreet, 1980). Iversen and Kelley
(1974) suggested gastric ulcers invaded by “‘Contracaecum sp.?’’ in marlin
resulted from mechanical injury by sharply pointed food items, possibly
aggravated by the nematodes. We observed no H. incurvum or any other
species of Hysterothylacium to embed in the alimentary tract of their hosts.
Kalyankar (1972) reported larval J. inquies from gills of sea-crabs in Rat-
nagira, India, but that identification requires confirmation. Adults of Goezia
are much wider than J. inquies, have spines on the plicated annulations,
and have angulated overhanging lips.

Theringascaris inquies (Linton), new combination
Figs. 68-84

Ascaris inquies Linton, 1901:452, pl. 6, figs. 46—SO (original description;
type-host Rachycentron canadum; type-locality Buzzard’s Bay, Massa-
chusetts).

Theringascaris iheringascaris Pereira, 1935:56—58, figs. 10-16 (original de-
scription; type-locality Natal, Estado do Rio Grande do Norte, Brazil).

Contracaecum (Thynnascaris) inquies.—Johnston and Mawson, 1945:132-
133 (new combination).

Thynnascaris iheringascaris.—Hartwich, 1957:240—241 (new combination).

Contracaecum iheringascaris.—Yamaguti, 1961:28, pl. 5, fig. 36 (new com-
bination).

Thynnascaris inquies.—Rasheed, 1965:323-329, figs. 18-28 (new combina-

VOLUME 93, NUMBER 4 1069

Figs. 64-74. Hysterothylacium habena (64-67) and Iheringascaris inquies (68-74): 64, Male
tail of H. habena, ventral view; 65-67, Lateral ala at anterior (65), middle (66), and posterior
(67) parts of female; 68-70, Lateral ala of J. inquies at anterior (68) and midbody (69) of fourth
stage larva and midbody (70) of mature male; 71-74, Cross-sections of excretory system show-
ing excretory pore (71), excretory commissure (72), and excretory canals immediately anterior
to intestinal cecum (73), and immediately posterior to esophageal-ventricular junction (74).
Abbreviations: c = commissure, ec = excretory canal, ed = excretory duct, ep = excretory
pore, es = esophagus, i = intestine, Ic = lateral cord, va = ventricular appendage.

tion; redescription based on Linton’s specimens and others from Karachi,
West Pakistan).

?Neogoezia elacateiae Khan and Begum, 1971:12—14, fig. 8 (original de-
scription; type-host Rachycentron canadum [as Elacate nigra]; type-lo-
cality Karachi Coast).

1070 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Diagnosis.—Cuticle with deep transverse annulations; lips with ratio of
length to width 1:1.1—1.6; caudal papillar pairs: preanal 25-29, medial-post-
anal 6, lateral 6; spicules 23-25% of body length; spicule-ratio 1:0.8-1.2;
digitiform process lacking ornamentation on posterior extremity.

General (based on specimens from Mississippi and Alabama).—Body
reaching greatest width near midbody. Cuticle with deep transverse annu-
lations along entire length. Lateral alae extremely reduced or absent in
adults (relatively prominent in fourth stage larva). Lips approximately equal
in size, wider than long; flanges widest near posterior, indented at anterior
14; pulp pedunculated. Interlabia with height equal to or slightly greater than
width at base; interlabial grooves lacking. Cervical papillae small, near
nerve ring. Esophagus 12-15% of body length. Ventriculus narrower than
widest level of esophagus, usually longer than broad; ventricular appendage
descending without angulation from posterior portion of ventriculus. Nerve
ring at anterior 10-23% of esophagus. Excretory pore immediately posterior
to level of nerve ring; excretory canal in right lateral cord terminating slight-
ly anterior to level of tip of intestinal cecum; canal in left lateral cord ex-
tending considerably farther posteriorly. Tail conical.

Male (based on 20 mature specimens): Body 20—27 mm long by 216-709
at greatest width; ratio of greatest width to length 1:54-94. Lips 72-132 long
by 96-144 at greatest width. Nerve ring 204-442 from anterior extremity,
31-55 in breadth. Esophagus 2.0—4.0 mm long by 116-144 wide. Ventriculus
74-166 long by 92-135 wide; ventricular appendage 1.3—1.7 mm long by 50-
70 wide; ratio for lengths of ventricular appendage to esophagus 1:1.9-2.5.
Intestinal cecum 330-649 long by 75-105 wide; ratio of cecal to ventricular
appendage lengths 1:1.7-4.2; ratio of cecal to esophageal lengths 1:5-8.
Spicules bifurcate at tip, covered by thin sheath, 23-25% of body length,
unequal; left spicule 5.3—9.2 mm long by 12-15 wide; right spicule 5.8-10.7
mm long by 12-15 wide, usually longer than left spicule; spicule ratio 1:0.8—
1.2. Caudal papillae 37-41 pairs, becoming closer together and more medial
as approaching anus; preanal pairs 25-29; medial-postanal pairs 6 with 3rd
from posterior end doubled; lateral pairs 6. Medioventral preanal organ
simple, distinct, papillated. Tail flexed ventrad, 79-132 long.

Female (based on 20 mature specimens): Body 14-35 mm long by 253-
709 wide; ratio of greatest width to length 1:54—70. Lips 86-127 long by 122-
151 wide. Nerve ring 450-550 from anterior extremity, 16-34 in breadth.
Esophagus 2.0-3.5 mm long by 80-140 wide. Ventriculus 110-170 long by
62-130 wide; ventricular appendage 1.2—2.0 mm long by 77-105 wide; ratio
for lengths of ventricular appendage to esophagus 1:1.6—2.5. Intestinal ce-
cum 260-505 long by 50-100; ratio of cecal to ventricular appendage lengths
1:1.7-4.3; ratio of cecal to esophageal lengths 1:6.0-7.1. Vulva without sa-
lient lips, opening 10.0-11.7 mm or 31-38% of body length from anterior
extremity. Ovaries rarely extending anteriorly to level of vulva. Eggs 40-60

VOLUME 93, NUMBER 4 1071

in diameter. Tail 250-611 long. Phasmids 1 pair, located laterally, equato-
rially between anus and posterior extremity of tail.

Host.—Rachycentron canadum (Linnaeus), cobia (Rachycentridae).

Sites of infection.—Stomach and pyloric ceca.

Localities.—Mississippi Sound, Mississippi; Alabama Point, Alabama;
Buzzard’s Bay, Massachusetts; Lower Chesapeake Bay, Virginia; Natal,
Brazil; and Karachi, West Pakistan.

Specimens deposited.—{2 pairs) USNM Helm. Coll. No. 75847; (1 pair)
OCI 31.605; (1 pair) BMNH Reg. No. 1980.367—368.

Remarks.—Linton (1901) originally described Ascaris inquies from ma-
terial removed from the stomach of Rachycentron canadum near Woods
Hole, Massachusetts. He reported 24 pairs of preanal papillae and 4 pairs
of postanal ones. Because Linton’s description and illustrations were in-
adequate, Rasheed (1965) redescribed it using one of Linton’s deposited
specimens and those removed from the same host species in West Pakistan.
She transferred it to Thynnascaris, agreed with the presence of 24 pairs of
preanal and 4 pairs of medial-postanal papillae, and noted an additional 4
lateral pairs of papillae which Linton had missed. Since the holotype was
not available, we examined two male and five female specimens (USNM
Helm. Coll. No. 6622) deposited by Linton in 1899 from the type-host and
type-locality. On the best male specimen, we counted 6 lateral, 6 medial-
postanal, and 24 pairs of preanal papillae. We also examined specimens
(VIMS Host No. S-498) from a cobia caught in Lower Chesapeake Bay, off
Seaford, Virginia, with similar papillar counts; the doubled papillae were
obvious. Our specimens from the northern Gulf of Mexico showed no
marked differences with those or with Rasheed’s specimens (BMNH
1964.1523—1532) which we also examined and are consequently considered
conspecific with them. With the permission of Rodney Bray, Parasitic Worm
Section, British Museum (Natural History), we deposited one pair of Rash-
eed’s specimens in the U.S. National Museum (USNM Helm. Coll. No.
75545).

We also examined four specimens, 3 males and | female, of [heringascaris
theringascaris loaned to us by J. J. Vicente (OCI No. 17.157) from R.
canadum in Brazil. Pereira (1935) listed the type-host as “‘bijupira’’ which,
according to Vicente, is a common name for Rachycentron canadum. The
elongated specimens we examined were fragments or had stripped cuticles.
Nevertheless, we counted at least 18 pairs of preanal, 6 pairs of medial-
postanal, and 6 pairs of lateral papillae and confirmed the characteristic
annulations. These features in addition to the measurements in the original
description reported by Pereira (1935) prompt us to consider [heringascaris
iheringascaris a junior synonym of J. inquies.

Species of Hysterothylacium purported to possess conspicuous trans-
verse annulations are H. cornutum from Thunnus thynnus (Linnaeus) (as

1072 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

T. vulgaris) at Trieste, Italy, and H. carangis from the intestine of Caranx
malabaricus Day at Panjim, India. Hysterothylacium cornutum was origi-
nally described by Stossich (1904), and Baylis (1923) later added to that
description using Stossich’s original material. Baylis noted that the cuticular
striations had ‘‘rather prominent edges, like minute sawteeth.’’ We exam-
ined these specimens of H. cornutum, 2 males and 2 females (BMNH No.
1931.10.20.80-89), but observed fine cuticular annules such as typically seen
on adult specimens of some species of Hysterothylacium. Kalyankar (1971)
stated that the striations of H. carangis were “‘not so distinct at the anterior
and posterior extremities of the male, but very distinct in the middle of the
body.’’ In female specimens these striations were apparently very distinct
with the exception of the ‘‘anterior-most region.’’ Cuticular annulations of J.
inquies are consistently conspicuous at all regions of both male and female
worms. Hysterothylacium carangis is reported to have 28 caudal papillae in
contrast to the 36-42 on I. inquies. Kalyankar purported two spicules of equal
length; yet, his illustration showed the left spicule slightly longer than the right.
In I. inquies the right is usually slightly longer than the left. Kalyankar stated
that H. carangis differed from I. inquies by possessing a straight blunt tail
and a postequatorial vulva. Hysterothylacium carangis needs critical re-
examination. In any event, J. inquies differs from H. cornutum and appar-
ently from H. carangis.

Khan and Begum (1971) described Neogoezia elacateiae from Elacate
nigra (Day) (=Rachycentron canadum) off the coast of Karachi, West Pa-
kistan. That host and locality are the same as Rasheed reported for H.
inquies. Also, those authors’ description and illustrations show the con-
spicuous transverse cuticular annulations and other diagnostic features of
I. inquies. They, however, did not observe interlabia or a ventricular ap-
pendage, and they reported 11 preanal papillae without mentioning lateral,
para- or postanal papillae. To date we have been unsuccessful in corre-
sponding with Khan. Consequently, until Khan and Begum’s specimens can
be reexamined, we suggest that Neogoezia elacateiae may be a junior
synonym of I. inquies.

In addition to the previous diagnoses, the following key should permit

—>

Figs. 75-84. Iheringascaris inquies: 75, Anterior end showing conspicuous annulations and
spacial relationship between excretory pore and nerve ring, lateral view; 76, Dorsal view of
lips; 77, Ventral view of lips showing interlabium; 78, Posterior extremity of female tail, lateral
view; 79, En face; 80, Male tail showing lateral and medial postcloacal papillae, ventral view;
81, Tip of spicule, lateral view; 82, Body at level of intestinal-ventricular junction, lateral view;
83, Female tail, lateral view; 84, Posterior end of male showing caudal papillae, lateral view.
(Portions of the annulations in Figs. 80, 83, and 84 were omitted for clarity of other structures.)

1073

VOLUME 93, NUMBER 4

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1074 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rapid identification of adults of species of Hysterothylacium reported in this
paper.

Key to the Species of I[heringascaris and Hysterothylacium
Parasitizing Fishes in the Northern Gulf of Mexico

1. Cuticle with deep transverse annulations (Fig. 75); 25—29 pairs of
preanal, 6 pairs of medial-postanal, and 6 pairs of lateral-postanal
papillae. Host: Rachycentron canadum. Western North Atlantic and
Gulf of Mexico (also Brazil and Pakistan) ...................000-

— Cuticle lacking deep transverse annulations ..................... 22
2. Cervical alae prominent (Figs. 1, 4); 16-25 pairs of preanal, 8 pairs
of postanal, and no para-anal papillae; tuft of spinous-like projec-
tions at posterior extremity. Hosts: Scomberomorus brasiliensis, S.
maculatus, S. cavalla, and Oligoplites saurus. Gulf of Mexico (also

Atlantic coasts of Florida and Brazil) .... H. fortalezae (Klein, 1973)

Cervical alae not prominent’ (2.2... 2. ee ee 3
3. Cuticle with ventral crests (enlarged, modified annules [Figs. 50—52])

anterior’tovanus on’ male. 2.52 ee. ee 4
= Cuticle lacking ventral crests on males ...................-. 9006 5

4. Interlabial grooves deep (Figs. 47, 48); spicules 12-25% of body
length. Hosts: Xiphias gladius and Tetrapturus albidus plus other
marlin. Gulf of Mexico (also in Baltic Sea and off Floridian Atlantic
coast, India, Tanzania, and probably Japan, New Zealand, and else-
WET Khe mere enor ent cc Me en nnn ant teen H. incurvum (Rudolphi, 1819)

— Interlabial grooves lacking; spicules 4—7% of body length. Host:
Xiphias gladius. Gulf of Mexico (also off Floridian Atlantic coast
ANGdPE CUAGOR)E. che ok es oe eee ok Meee ees H. corrugatum, sp. n.

5. Tail tip with relatively large projections on tuft (Figs. 42, 45); 23
pairs of preanal and 7 pairs of postanal papillae; pulp of lips not
pedunculate. Host: Chaunax pictus. Gulf of Mexico .............

Bo js SS, High tase oe din 5, aa ko Aaa tee si H. chaunaxi (Olsen, 1952)

— Tail tip with dense covering of minute spinous structures or lacking
Omamentation 4.50% b. vesk sw oasye vse owls aes cs oe Ie ee eee 6

6. Lateral flanges of lips indented equatorially (Figs. 22—24); spicules
3-8% of body length; multispinous conical process at posterior ex-
tremity. Hosts: numerous fishes (see text). Gulf of Mexico (also
Pacific and Caribbean coasts of Central America and Atlantic coast
of South America to Brazil) .. H. reliquens (Norris and Overstreet, 1975)

— Lateral flanges of lips indented at anterior %4 of lip (Figs. 14, 15);
length of spicules 1-2% that of body; digitiform process on posterior
extremity of tail lacking ornamentation. Hosts: Ogcocephalus ra-

VOLUME 93, NUMBER 4 1075

diatus and O. cubifrons. Gulf of Mexico (and Floridian Atlantic coast)
Sys i Oe 2s ee nga eee H.. ogcocephali (Olsen, 1952)

Acknowledgments

We gratefully acknowledge the assistance of the following persons: D. I.
Gibson and R. A. Bray for loaning specimens deposited in the British Mu-
seum (Natural History); G. Hartwich at the Zoologisches Museum, for loan-
ing syntypes of H. incurvum; J. R. Lichtenfels for loaning specimens de-
yosited in the U.S. National Museum Helminthological Collection; C. D.
Dawson (GCRL) for permission to remove worms from stomachs of several
museum hosts; B. B. Collette at the National Marine Fisheries Service
Systematics Laboratory (USNM) for most specimens of H. reliquens from
Batrachoides spp.; C. T. Fontaine previously at NMFS Biological Labo-
ratory of Galveston, Texas, for specimens of H. reliquens from Arius felis;
R. G. Yaeger at Tulane University School of Medicine for specimens of H.
reliquens collected off the R/V OREGON II (we also thank the NMFS
Laboratory at Pascagoula, Mississippi, for these plus samples of uninfected
Atlantic croaker from offshore in the Gulf of Mexico); J. J. Vicente at the
Oswaldo Cruz Institute for loaning specimens of IJ. iheringascaris and H.
fortalezae; E. M. Burreson at Virginia Institute of Marine Science for pro-
viding specimens of J. inquies; D. E. Norris at University of Southern
Mississippi for providing specimens of H. habena; P. J. Pristas at NOAA,
NMES (Panama City Laboratory) for aid in collecting white marlin; C. A.
Hamilton and Mobile Big Game Fishing Club, Orange Beach, Alabama, and
Pflueger Marine Taxidermist, Hallandale, Florida, for providing cooperation,
space, and permission to examine billfish collected by sportsfishermen; E. W.
Cake and T. E. Mattis of GCRL, S. H. Deardorff and L. A. Covington of
Ocean Springs, Mississippi, and F. Crooke of Pensacola, Florida, for aiding
in the collection of hosts and parasites; and R. Buxton, J. D. Durfee, R. G.
Palmer, R. A. Krol, and K. E. Richards for technical assistance.

This study was conducted in cooperation with the U.S. Department of
Commerce, NOAA, National Marine Fisheries Service, under PL 88-309
Project No. 2-325-R and HEW/Public Health Service, Food and Drug
Administration Contract No. 223-76-2141.

Literature Cited

Anonymous. 1974. The Bulletin of Zoological Nomenclature.—The International Trust for
Zoological Nomenclature 31:76-89.

Araujo, P. 1970. Consideragoes sobre 0 género Contracaecum Railliet & Henry, 1912 (Nem-
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Bailey, R. M., chairman. 1970. A list of common and scientific names of fishes from the
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1076 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Baylis, H. A. 1920. On the classification of the Ascaridae. I. The systematic value of certain
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—. 1923. Report on a collection of parasitic nematodes, mainly from Egypt. Part I.

Ascaridae and Heterakidae.—Parasitology 15:1-13.

. 1929. Parasitic Nematoda and Acanthocephala collected in 1925-1927.—Discovery

Rep. 1:541-560.

, and R. Daubney. 1922. Report on the parasitic nematodes in the collection of the

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. 1926. A Synopsis of the Families and Genera of Nematoda.—Trustees of the British

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Berland, B. 1961. Nematodes from some Norwegian marine fishes.—Sarsia 2:1—50.

Brunsdon, R. V. 1956. Studies on nematode parasites of New Zealand fishes (2 volumes).—
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Chabaud, A. G. 1965. Systématique des Ascaridada.—Traité Zoologie 4:988—997.

Chandler, A. C. 1943. A redescription of Contracaecum habena (Linton, 1900) Linton,
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Collette, B. B., and J. L. Russo. In press. A revision of the scaly toadfishes, genus Batra-
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Deardorff, T. L., and R. M. Overstreet. 1978. Thynnascaris rhacodes sp. n. (Nematoda:

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Dollfus, R. Ph. 1933. Thynnascaris legendrei n. gen., n. sp. de lestomac du germon, Germo
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. 1975. Parasitic nematodes of vertebrates. I. Rhabditida und Ascaridida.—Die Tierwelt

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VOLUME 93, NUMBER 4 1077

Hutton, R. F., T. Ball, and B. Eldred. 1962. Immature nematodes of the genus Contracaecum
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1078 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 93, NUMBER 4 1079

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Gulf Coast Research Laboratory, Ocean Springs, Mississippi 39564.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1080-1172

SYLLIDAE (POLYCHAETA), PRINCIPALLY FROM
FLORIDA, WITH DESCRIPTIONS OF A NEW
GENUS AND TWENTY-ONE NEW SPECIES

Thomas H. Perkins

Abstract.—The following new species are described: Brania gallagheri,
Dioplosyllis octodentata, Exogone arenosa, E. atlantica, Odontosyllis lon-
gigulata, Pionosyllis gesae, Plakosyllis quadrioculata, Sphaerosyllis acic-
ulata, S. bilobata, S. brevidentata, S. glandulata, S. magnidentata, S. pir-
iferopsis, S. riseri, S. taylori, Streptosyllis pettiboneae, Syllides bansei, S.
floridanus, Trypanosyllis inglei, T. parvidentata, and T. savagei. Dentati-
syllis, n. gen., is proposed for Syllis carolinae Day. Brania swedmarki Gid-
holm is newly reported for the western Atlantic. Brania wellfleetensis Pet-
tibone is newly reported from Tampa Bay, Florida. Exogone longicirris
(Webster and Benedict) is removed from synonymy with E. dispar (Web-
ster), and both and E. lourei Berkeley and Berkeley are additionally de-
scribed. Parapionosyllis longicirrata (Webster and Benedict) is newly re-
ported from both coasts of Florida, and Pionosyllis manca Treadwell from
Virginia is reported as a synonym. Presence of Pionosyllis uraga Imajima,
originally described from Japan, is confirmed for North Carolina and newly
reported from Florida. Florida specimens of Trypanosyllis coeliaca Cla-
paréde are described. Sphaerosyllis longicauda Webster and Benedict is
removed from synonymy with S. erinaceus Claparede and redescribed.
North American specimens previously referred to S. hystrix Claparede are
referred to S. taylori, n. sp. Specimens from North Carolina previously
referred to S. pirifera are referred to S. glandulata, n. sp. Characters of
systematic importance to Sphaerosyllis Claparede are discussed, and a key
is given for 13 species of the genus from the northwestern Atlantic Ocean.

This report is one of several (Perkins, 1979) based primarily on specimens
collected between September 1971 and July 1973 in an environmental base-
line study of marine biota near the Florida Power and Light Company nu-
clear power plant at Hutchinson Island, St. Lucie County, Florida. Speci-
mens representing significant range extensions and numerous new species
from these collections are reported herein. Some Syllidae collected at
Hutchinson Island are not included in this report, but will be included in a
complete report on the Polychaeta which is in preparation. Additional spec-
imens collected later at Hutchinson Island and other specimens from Maine
to the Florida Keys, Bimini, Bahamas, the eastern Gulf of Mexico and the
Northeast Pacific are included.

VOLUME 93, NUMBER 4 1081

The study area was characterized and methods of collection were de-
scribed by Gallagher and Hollinger (1977). Sediments were described by
Gallagher (1977); other aspects of the physical and chemical environment
were reported by Worth and Hollinger (1977). Brief descriptions of sampling
stations and methods were also given by Perkins (1979).

Types and other specimens available for study are deposited in the Allan
Hancock Foundation, University of Southern California (AHF), the Inver-
tebrate Reference Collection of the Florida Department of Natural Re-
sources Marine Research Laboratory (FSBC I), the U.S. National Museum
of Natural History, Smithsonian Institution (USNM), the Virginia Institute
of Marine Science Invertebrate Collection (VIMS), and the Zoologisches
Museum, Hamburg (ZMH).

Drs. Marian H. Pettibone, Kristian Fauchald and Meredith Jones
(USNM) loaned specimens. Dr. Nathan W. Riser, Marine Science Institute,
Northeastern University; Dr. Norman J. Blake, Department of Marine Sci-
ence, University of South Florida; Mr. Robert G. Ernest, Applied Biology,
Inc., Jensen Beach, Florida; Mr. Gary R. Gaston (VIMS); Mr. Richard J.
Helbling and Dr. Douglas Farrell, Florida Department of Environmental
Regulation; and Dr. Joseph L. Simon, University of South Florida, donated
specimens. Drs. Kristian Fauchald, Joseph L. Simon, Gesa Hartmann-
Schroder (ZMH), Nathan W. Riser and Ms. Joan M. Uebelacker, Dauphin
Island Sea Lab commented on earlier drafts and provided information, and
Dr. Fauchald critically reviewed the manuscript. Many individuals from the
Marine Research Laboratory and Applied Biology, Inc., participated in the
Hutchinson Island Study for which Florida Power and Light Co., Inc., pro-
vided partial funding. All are gratefully thanked.

Brania Quatrefages, 1865
Brania swedmarki Gidholm, 1962

Brania swedmarki Gidholm, 1962:256—258; fig. 3.—Hartmann-Schréder,
1974a:195, 196.

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6’N,
80°13.2’W, about 11 m, coarse calcareous sand; 11 specimens (USNM
60475; FSBC I 20576). Sta. IV, 27°20.7’N, 80°12.8’W, about 11 m, coarse
calcareous sand; 2 specimens (FSBC I 20577, 20578). Sta. V, 27°22.9’N,
80°13.9’W, about 11 m, coarse calcareous sand; 3 specimens (FSBC I 20579-
20581).

Remarks.—Brania swedmarki was described from Roscoff, France, and
has since been reported from the northwestern North Sea (Hartmann-Schro-
der, 1974a). Hutchinson Island specimens are in excellent agreement with
Gidholm’s (1962) description except for variations attributable to differences
in preservation or state of contraction.

1082 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Specimens from the Florida east coast represent the first Western Atlantic
records for the species.

Brania wellfleetensis Pettibone, 1956
Fig. 1

Brania wellfleetensis Pettibone, 1956:282, fig. 2a—c; 1963:134, 135, fig. 35h.
Brania sp. Taylor, 1971:223-225, fig. Sa, b.—Hall and Saloman, 1975:12
[list].

Material examined.—_MASSACHUSETTS: Wellfleet Harbor, Cape Cod
Bay side, sandy bottom, among tubes of Diopatra cuprea (Bosc); 25 Aug.
1953; holotype (USNM 27783). Chappaquoit, Buzzards Bay, muddy sand;
paratype (USNM 27784). FLORIDA, GULF OF MEXICO: Hillsborough
Bay, Tampa Bay; S. Santos col. and det., 1975-1979; 32 specimens (USNM
60477; FSBC I 22498; AHF 367; J. L. Simon). Same, 27°49'03'N,
82°26'07"W, sand, 4 m; J. Taylor and C. Saloman cols., 4 Sep. 1963; 3
specimens (FSBC I 15458). Upper Tampa Bay, 27°49’29"N, 82°33’50"W,
shelly sand, 1 m; J. Taylor and C. Saloman cols., 27 Aug. 1963; 24 specimens
(FSBC I 15456). Same, 27°49'28"N, 82°33'24’W, sand, 5 m; J. Taylor and
C. Saloman cols., 27 Aug. 1963; 2 specimens (FSBC I 15457). Lower Tampa
Bay, 27°36'56’N, 82°41'05’W, sand, 9 m; J. Taylor and C. Saloman cols.,
25 Oct. 1963; 2 specimens (FSBC I 17924). Same, 27°34’54"N, 82°43'01"W,
sand, 7m; J. Taylor and C. Saloman cols., 30 Oct. 1963; 1 specimen (FSBC
I 15459). Same, 27°34’24"N, 82°42'53”W, sand with algae cover, 6 m; J.
Taylor and C. Saloman cols., 4 Nov. 1963; 1 specimen (FSBC I 15460).
Same, 27°35’26"N, 82°45'27”W, sand with algae and Thalassia cover, 3 m;
J. Taylor and C. Saloman cols., 9 Oct. 1963; 2 specimens (FSBC I 13187).

Additional description.—Holotype: mature male with sperm in setigers
12-34 of 41 setigers, without natatory setae, with notoacicula between dor-
sal cirri and parapodial lobes first visible at about setiger 20 and continuing
to setiger 33. Prostomium with anteromedian part forming rounded, obtuse
angle. Antennae, tentacular cirri and dorsal cirri slightly pseudoarticulated
[preserved]. Parapodia mostly with 4 compound setae but 1-2 additional
setae on anterior parapodia (Fig. 1c); upper one with blade edge oriented
ventrally; lower 3 with blade edges oriented dorsally; blades with about 5
long, slender serrations; shafts with few indistinct serrations below tip. Su-
perior simple setae beginning on setiger 34 (Fig. 1a, b), with pointed, slightly
hooked tips, mostly with 2 large serrations below tip and indistinct serra-
tions below. Inferior simple setae on last 3 or 4 setigers (Fig. 1d), slightly
curved, with pointed tips, without serrations or hairs below tips. Acicula
solitary, with slightly nobbed tips. Pygidium with 3 anal cirri (Pettibone,
1963). Pharynx coiled, with small, indistinct middorsal tooth slightly back

VOLUME 93, NUMBER 4 1083

wit GC]

Fig. 1. Brania wellfleetensis: a, Superior simple seta, setiger 37; b, Same, setiger 38; c,
Compound seta, setiger 37; d, Inferior simple seta, setiger 38; e, f, Superior simple setae,
posterior setigers; g, Upper compound seta, middle setiger; h, Same, posterior setiger; i, Middle
compound seta, posterior setiger; j, Lower compound seta, middle setiger; k, Same, posterior
setiger; 1, Inferior simple seta, posterior setiger; m, Anterior end with pharynx everted, lateral
view, pharyngeal tooth darkened (a—d, holotype; e-m, Tampa Bay specimens, J. L. Simon).

from rim. Proventriculus cylindrical, 5 times longer than diameter, with 27
rows of muscle cells with anterior 6 rows small.

Tampa Bay, Florida, specimens: mature specimens larger than holotype,
with up to about 55 (39-55) setigers. Pharynx brown on some specimens.
Eyes of some mature specimens enlarged, with lenses; small, without lenses
on others; easily damaged, often not visible (Fig. 1m). Superior simple setae
of mature specimens similar to those of holotype (Fig. le, f), beginning on

1084 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

more anterior setigers (13-30) on mature specimens. Compound setae (Fig.
lg—k) with blades slightly stouter than those of holotype, with similar num-
ber of stouter serrations; shafts similar, often stouter below. Inferior simple
setae on several posterior segments (Fig. 1 1) with tips pointed and few small
serrations below tips. Pharynx longer than proventriculus, extending from
posterior half of setiger 1 to posterior part of setiger 6 when inverted; an-
terior rim surrounded by 10, soft, ciliated papillae; middorsal tooth small;
tip back from rim by about !/. of total length (Fig. 1m). Proventriculus long,
cylindrical, about 3 segments long in setigers 6-9, with about 30 (28-32)
rows of muscle cells in mature specimens.

Sexually mature specimens with sperm and eggs or ventrally attached
embryos beginning on setigers 12-14 and extending to setigers 27-43. No-
toacicula in sexual segments; one female with long natatory setae; males
with natatory setae not seen. Six-setiger stage embryos without dorsal cirri
on setiger 2 attached by pygidial end below ventral cirri of one female
(USNM 60477).

Remarks.—Specimens from the Tampa Bay area differ from the holotype
in having a larger number of segments on most mature specimens; however,
some mature specimens are about the same size. They also differ from the
holotype in having superior simple setae of most specimens beginning on
more anterior segments and in always having inferior simple setae with a
few small serrations below the tips.

Brania wellfleetensis has previously been reported only from Massachu-
setts. Gulf of Mexico specimens reported herein may constitute a disjunct
population, with only slight differences from the more northern form.

Brania gallagheri, new species
Figs. 2, 3

Material examined.—FLORIDA: Hutchinson Island Sta. IV, 27°20.7’N,
80°12.8’W, about 11 m, coarse calcareous sand, holotype (R. Gallagher col.,
Jan. 1972; USNM 60209), 13 paratypes (USNM 54503, 54504; AHF Poly
1305; ZMH P-16386-16388, FSBC I 20597, 20598, 20603-20605). Sta. II,
27°21.6’N, 80°13.2'W, about 11 m, coarse calcareous sand; 3 paratypes
(USNM 54502; AHF Poly 1306; FSBC I 20583). Sta. III, 27°22.0'N,
80°12.4’W, about 7 m, medium calcareous sand; 19 paratypes (AHF Poly
1307; FSBC I 20584-20593, 20595, 20596).

Description.—Well-preserved, sexually mature specimens of 50-60 seti-
gers 5—7 mm long, 0.2-0.5 mm wide with parapodia. Body without color
markings; eyes orange to dark red; pharynx often brown with ring of yel-
lowish brown cells anterior to rim. Prostomium (Fig. 2a—c) about twice as
wide as median length, anteromedian part forming rounded, obtuse angle;
lateral margins rounded; posterior margin straight. Two pairs of eyes near

VOLUME 93, NUMBER 4 1085

lateral margins of posterior half; outer, anterior pair more lateral; small,
composed of aggregated pigment spots on most specimens; large, with
lenses, with outer pair larger on some sexually mature males and females.
Median antenna originating near posterior margin, extending about to tips
of palps or beyond, similar to upper tentacular cirri. Lateral antennae orig-
inating on anterior margin anterior to eyes, extending about to tips of palps,
similar to lower tentacular cirri. Posterior third of palps fused, each about
as long as prostomial width on noncompressed specimen (Fig. 2a). Ten-
tacular segment similar but slightly shorter than following segment, upper
tentacular cirri subequal to median antenna, extending about to tips of palps.
Lower tentacular cirri shorter, similar to lateral antennae. Dorsal cirri of
setiger 1 slender, longer than median antenna and upper tentacular cirri.
absent on setiger 2; slenderer than those on setiger 1, extending to or beyond
tips of setae on other setigers; all originating on short cirrophores (Fig. 2a,
d). Parapodial lobes distally truncate, bilabiate. Ventral cirri originating
from lower margins of parapodial lobes, slightly slenderer than dorsal cirri,
extending about to tips of parapodial lobes. Solitary, superior simple setae
(Fig. 3a—c, 1, j) above acicula beginning on setiger 14-21 of mature speci-
mens, 8-20 on other specimens, with tips bluntly-pointed, 1-2 blunt to point-
ed teeth near tips above indistinct, irregular serrations on large specimens;
tips pointed on juveniles. Usually 4 compound setae (Fig. 3d-f, k-m), 1-2
additional ones on anterior setigers; upper ones with blade edges oriented
ventrally; lower 3 with blade edges oriented dorsally; blades short, falci-
gerous, unidentate, with up to 7 relatively stout serrations on larger speci-
mens, fewer serrations on juveniles (Fig. 3k—m); shafts with few indistinct
serrations on protuberance below tips; lower shafts greatly inflated on some
middle and posterior parapodia of mature specimens (Fig. 3f), slender on
juveniles (Fig. 3m). Solitary, bidentate inferior simple setae on few posterior
segments (Fig. 3g, n). Acicula solitary, with nobbed tips (Fig. 3h). Pygidium
with 3 anal cirri; long, subcylindrical lateral pair about as long as last 3 or
4 segments; shorter, slenderer median one originating véntrally (Fig. 2e).
Males with sperm beginning in setiger 15S—18, extending to setiger 49 in
specimen of 59 setigers; with short, natatory setae and notoacicula (Fig. 2d)
between bases of dorsal cirri and parapodial lobes, or with only notoacicula,
beginning on setiger 20 or 21 and extending to about end of sexual region.
Females with eggs beginning in setigers 16 or 17 and long, natatory setae
and notoacicula, or with only notoacicula, beginning on setiger 18 and ex-
tending to about 10 segments from posterior end.

Pharynx relatively thick-walled, extending to about setiger 6, with long,
broad, anterior middorsal tooth (Fig. 2b, c), anterior end surrounded by 10
soft, ciliated lobes. Proventriculus of adults cylindrical, slightly longer than
pharynx, 5—6 times longer than diameter, in 7-9 segments, with about 45
rows of muscle cells, with anterior 6 rows small. Ventricle convoluted in

1086 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Brania gallagheri: a, Anterior end, dorsal view; b, Same, slightly flattened showing
internal structure, pharyngeal tooth darkened; c, Same, lateral view; d, Parapodium of setiger
37 of male, slightly turned anterior view; e, Pygidium, ventral view (a, d, USNM 54502; b,
FSBC I 20601; c, e, USNM 54503).

figured specimen (Fig. 2b), 1-2 segments long depending on state of con-
traction.

Remarks.—Brania gallagheri is very similar to B. wellfleetensis Pettibone
in general shape. However, B. gallagheri has longer and slenderer antennae
and cirri and a larger, broader middorsal pharyngeal tooth. The proventric-
ulus of B. gallagheri is about 6 segments long with about 45 rows of muscle
cells, while that of B. wellfleetensis is about 3 segments long with about 30
rows of muscle cells; inferior simple setae of B. gallagheri have bidentate
tips, while those of B. wellfleetensis are unidentate; and sperm and eggs
begin in setigers 15—18 and notoacicula and natatory setae begin in setigers
18-21 of B. gallagheri, while sexual products, natatory setae and notoacic-
ula begin in setigers 12-14 of B. wellfleetensis.

VOLUME 93, NUMBER 4 1087

n \ \m f

ae
15 ym

Fig. 3. Brania gallagheri: a—c, Superior simple setae, middle segments; d, Upper compound
seta of setiger 37; e, Middle compound seta of same; f, Lower compound seta of same, blade
somewhat turned; g, Inferior simple seta of posterior setiger; h, Aciculum of setiger 37; i, j,
Superior simple setae, posterior setigers of juvenile; k, Upper compound seta of same; 1,
Middle compound seta of same; m, Lower compound seta of same; n, Inferior simple seta of
same (a-—h, USNM 54502; i-n, FSBC I 20584).

Etymology.—The species is named in honor of Mr. Robert M. Gallagher,
who was instrumental in providing the excellent specimens from the Hutch-
inson Island study.

Dioplosyllis Gidholm, 1962
Dioplosyllis octodentata, new species
Fig. 4

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6’N,
80°13.2'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,

1088 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wit OO!

d

ae
15 um

Fig. 4. Dioplosyllis octodentata: a, Posterior parapodium, posterior view; b—e, Compound
setae of same: b, Upper; c, Upper middle; d, Lower middle; e, Lower; f, Compound seta,
oblique view; g, Acicula, anterior parapodium; h, Tip of pharynx, ventral view, teeth darkened,
midventral tooth obscuring tip of middorsal tooth which is outlined by dashed line and stippled
(FSBC I 23501).

col., Nov. 1972; USNM 54509), 2 paratypes (FSBC I 23500, 23501). Sta.
IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 1 paratype
(USNM 54508). Sta. V, 27°22.9’N, 80°13.9'W, about 11 m, coarse calcar-
eous sand; | paratype (FSBC I 23502).

Description.—Holotype, largest adult specimen, incomplete posteriorly,
in 2 pieces, 8 mm long, 0.5 mm wide without parapodia, with about 50
setigerous segments; complete juvenile relatively long, slender, with about
35 setigers. Prostomium pentagonal, about twice as wide as long, with point-
ed anteromedial lobe comprising anterior half; sides of lateral half rounded;
posterior edge convex, with very slight medioposterior notch. Antennae on
short ceratophores, lengths 3-5 times prostomial width, median slightly
longer; laterals originating from sides of anteromedial lobe; median origi-
nating between posterior eyes. Palps longer than prostomial lobe, stouter

VOLUME 93, NUMBER 4 1089

distally, probably free to base. Eyes 3 pairs; 2 pairs on posterior half in
trapezoidal arrangement open to front, lensed; anterior eyespots medial to
origins of lateral antennae. Nuchal organs on posterior part. Tentacular
segment slightly narrower than following segment; tentacular cirri 2 pairs,
on short cirrophores, similar to dorsal cirri of following segments. Trans-
verse rows of cilia on ventrum and on dorsal and ventral borders of para-
podial lobes on setigerous segments; cilia apparently not extending across
dorsum. Dorsal cirri without obvious regular arrangement of alternating
lengths, 2-3 mm long. Parapodial lobes (Fig. 4a) as long as body width, with
6-8 falcigerous compound setae in anterior segments and 4—5 in posterior
segments. Ventral cirri attached near middle of ventral margins of parapo-
dial lobes, extending about to tips. Shafts of compound setae (Fig. 4b—e)
with pronounced bifurcation on tips, with transverse row of serrations below
tips. Upper compound setae with stout shafts; lower compound setae with
shafts similar in diameter to those in middle of bundle and not as thick as
upper shafts; blades of compound setae stoutest in upper parts, longest in
middle, shortest in lower parts of bundle; tips of blades tridentate, with
principal tooth bifurcate in plane perpendicular to blade (Fig. 4b, c). Soli-
tary, slender, bidentate, superior simple setae on posterior setigers of 2
small, complete specimens; principal tooth longer than secondary, strongly
falcate. Two acicula in anterior parapodia (Fig. 4g); solitary aciculum in
posterior parapodia.

Pharynx usually in first 7 setigers in adults, proventriculus in setigers 8-
12 or 14; pharynx and proventriculus in more anterior segments of smaller
specimens. Pharynx slightly less than 114 times longer than proventriculus;
tip armed with large, middorsal tooth and ventral arc of 7 smaller teeth (Fig.
4h); rim otherwise smooth in figured specimen but appearing indistinctly
scalloped in holotype (however, pharynx of latter somewhat contorted, not
allowing satisfactory illustration). Proventriculus with about 33 rows of mus-
cle cells including 10 small rows anterior to chitinous ring.

Remarks.—Differences among species of Dioplosyllis were recently tab-
ulated by Mueller and Fauchald (1976). D. octodentata is closely allied with
D. cirrosa Gidholm (1962:253—255) from the Atlantic coast of France. D.
octodentata differs from D. cirrosa in the following characters. The pharynx
of D. octodentata has a ventral arc of 7 small teeth on the edge while the
pharynx of D. cirrosa has a ventral arc of 5 small teeth with tips removed
from the edge. Anterior eyespots are present on the prostomium of D.
octodentata and absent on D. cirrosa. The length of the pharynx compared
with the length of the proventriculus of D. octodentata is 1.5:1 while the
pharynx of D. cirrosa is almost twice as long as the proventriculus. The
proventriculus of D. octodentata has about 33 rows of muscle cells while
that of D. cirrosa has 45. Shafts of compound setae of D. octodentata are

1090 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distally much more deeply notched than those of D. cirrosa, and the 3 teeth
on tips of blades are much larger.

Specimens of D. octodentata may be easily misidentified as Eusyllis la-
melligera Marion and Bobretzky, 1875, since both species have long dorsal
cirri and a pharynx with a denticulate margin. [Pettibone (1963:120—122)
referred both Syllis fragilis Webster (1879:217—220, pl. 4, figs. 42, 43) and
Eusyllis tenera Verrill (1882:368) to E. lamelligera.| However, palps of E.
lamelligera are fused for about a third of their dorsal length, and the details
of the setae of the 2 species are greatly different.

Etymology.—The specific name, derived from the Latin adjectives, octo,
meaning eight, and dentata, meaning toothed, refers to the number of pha-
ryngeal teeth.

Exogone Orsted, 1845
Exogone dispar (Webster, 1879)

Paedophylax dispar Webster, 1879:223, pl. 4, fig. 49, pl. 5, figs. 50-55.

Exogone dispar.—Pettibone, 1963:130, fig. 35d (synonymy) [in part; not
Paedophylax longicirris Webster and Benedict].—Taylor, 1971:201-—204
[in part].—Hall and Saloman, 1975:12 [list; in part].—Day, 1973:33, 34 [in
part; not E. clavator Ehlers and E. uniformis Hartman].—Westheide,
1974: 106-109, figs. 48, 49 [in part; not Hartman and Fauchald, 1971, mix-
ture of species; not Kohn and Lloyd, 1973].—Gardiner, 1976:132, fig. 11f-
1 [in part].

Material examined.—4 specimens on slides, from Webster’s private col-
lection identified as Paedophylax dispar, now deposited at Smithsonian In-
stitution, without locality data but probably type-specimens from Virginia
(USNM 27560). NEW JERSEY: Great Egg Harbor; numerous specimens
(USNM 495, 498; as Paedophylax dispar by Webster). NORTH CARO-
LINA: Cape Lookout; S. Gardiner and H. Wilson, cols. (USNM 52918,
52919). Off Beaufort, J. H. Day, col., 16 specimens (USNM 51072). FLOR-
IDA: Lower Tampa Bay, 27°36'56’N, 82°41'05”"W, 9 m, J. Taylor and C.
Saloman cols., 25 Oct. 1963; 79 specimens (USNM 60478).

Description.—Long, slender, at most about 5 mm long, with up to 40
segments. Median antenna long, club-shaped, extending about to tips of
palps; lateral antennae papilliform; 4 eyes. Proventriculus cylindrical, short-
er than pharynx, with 17—20 rows of muscle cells, in 2—3 segments beginning
in setiger 4 when pharynx inverted. Dorsal cirri on all setigers. Obscure
oblate, subdermal gland below dorsal cirrus of each parapodium. Setae in-
cluding superior and inferior simple setae, compound spinigers and falcigers.
Superior simple setae of middle segments strongly bent near tips; tips point-
ed, smooth or irregularly dentate on outer edges below tips, without spines
or aristae. Spinigers similar throughout body, with shafts slightly enlarged

VOLUME 93, NUMBER 4 1091

near tips and spinous; blades shorter on posterior segments. Falcigers with
bidentate blades, primary tooth much smaller than secondary, serrate on
anterior segments, shorter, without serrations on posterior segments. Infe-
rior simple setae on posterior parapodia, with bidentate tips, secondary
tooth smaller than primary. Pygidium with 2 cirri. Sexually mature speci-
mens with natatory setae beginning on setiger 14 or 15; eggs and sperm
often found in more anterior segments, e.g., setiger 10.

Remarks.—The synonymy of the species is complicated. Exogone lon-
giceps (Verrill, 1879) and E. dispar were described at about the same time.
Verrill was aware of Webster’s species and differentiated E. longiceps from
E. dispar on rather questionable characters. Types of E. longiceps have
apparently been lost, but it appears very similar if not identical with E.
dispar (Pettibone, 1963). Pettibone (1954, 1963) also included E. longicirris
(Webster and Benedict) in synonymy with E. dispar; E. longicirris is a
different species. Day (1973) included E. clavator Ehlers from South Africa
and E. uniformis Hartman from California in synonymy with E. dispar.
However, according to Day (1967) spinigerous compound setae are absent
from posterior segments of E. clavator; they are present on posterior seg-
ments of E. dispar; and according to Banse (1972), E. uniformis has a long
proventriculus ‘‘with 26 or 27 rows of muscular columns’’ extending through
7-8 setigers, a short median antenna (Banse, 1972:201, fig. 5e) and setae
which are similar to E. lourei Berkeley and Berkeley.

Specimens reported as E. dispar from Alaska by Pettibone (1954) appar-
ently included 2 species. One of the species had superior simple setae with
an arista and is possibly E. longicirris; the other species may be E. lourei,
which she included in her synonymy. The same type of aristate superior
simple seta was described by Kohn and Lloyd (1973) for specimens from
Easter Island referred to E. dispar.

In Atlantic areas, specimens reported as E. dispar by Hartman (1965) and
Hartman and Fauchald (1971) are apparently a mixture of species and should
be re-examined. Specimens reported as E. dispar by Taylor (1971) from
Tampa Bay, Florida, include both E. dispar and E. arenosa, n. sp. About
half of the specimens from Bogue Sound and Banks Channel, North Caro-
lina (USNM 52916, 52917), reported as E. dispar by Gardiner (1976), are
similar to E. arenosa, n. sp., in having enlarged shafts of spinigers on setiger
2, but the shafts differ in shape from those of E. arenosa. The specimens
are similar to E. dispar in having superior simple setae without spines on
the tips and the proventriculus of mature specimens with about 20 rows of

muscle cells. All of Gardiner’s (1976) specimens from Cape Lookout

:

-(USNM 52918, 52919) are typical examples of E. dispar.

I can confirm the presence of the species on the east coast of North
America from Massachusetts to Florida and the Gulf of Mexico and from
the Galapagos Islands in the eastern Pacific Ocean.

1092 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Exogone longicirris (Webster and Benedict, 1887)
Fig. 5a—f

Paedophylax longicirris Webster and Benedict, 1887:722, pl. 3, figs. 46-50.
Exogone dispar.—Pettibone, 1954:259; 1963:130—131 [in part; not Paedo-
phylax dispar Webster].

Material examined.—MAINE: Eastport; 4 syntypes (USNM 439).

Description.—Slender, up to about 3 mm long and 0.3 mm wide. Median
antenna long, cylindrical, extending almost to tips of palps, lateral antennae
about % as long; 4 eyes. Proventriculus cylindrical, %4 as long as pharynx
with about 11 large and 5 small rows of muscle cells; in length of 3 segments
beginning in setiger 4 or 5; pharynx light-colored, about 5 segments long.
Dorsal cirri on all setigers. Setae including superior and inferior simple
setae, compound spinigers and falcigers. Superior simple setae on middle
and posterior parapodia (Fig. 5a, b), with 2 subequal teeth on tips, with
arista beginning below tips on concave edge and extending beyond. Spini-
gers (Webster and Benedict, 1887:pl. 3, fig. 48) similar throughout body,
shorter posteriorly. Blades of falcigers (Fig. 5c, d) with similar primary and
secondary teeth, with arista or hood beginning at bases and extending to
near tips. Inferior simple setae on posterior parapodia (Fig. 5e), similar to
superior simple setae but with shorter aristae and sharper teeth. Acicula
(Fig. 5f) solitary, with nobbed tips. Pygidium with 3 anal cirri.

Remarks.—Exogone longicirris differs from E. dispar (Webster) in having |
bidentate superior and inferior simple setae which are also aristate and in |
having compound falcigers with hooded blades. I cannot determine from |
examination of the types that compound spinigers are present on posterior |
setigers. There are no posterior segments on the syntypes I examined, and |
most setae of middle segments are broken.

Exogone lourei Berkeley and Berkeley, 1938

Exogone lourei Berkeley and Berkeley, 1938:44, figs. 6-12; 1948:79, fig. |
117.—Berkeley, 1967:1055.—Pettibone, 1967:5.—Banse and Hobson, |
1968:16, fig. 4d, e—Banse, 1972:200-202, figs. Sa—d; 1974:58, figs. 14h-j. |

Exogone uniformis Hartman, 1961:73, 74 [in part; pl. 6, fig. 1, pl. 7, fig. |
1 and 8 specimens from type-locality, but not holotype (AHF Poly 0170) |
or paratypes (AHF Poly 0171), fide Banse (1972)]; 1968:427, 428 [in part; |
unnumbered figures of anterior and posterior ends].

Material examined.—BRITISH COLUMBIA: False Narrows; holotype |
(USNM 32895). WASHINGTON: Puget Sound, 47°41'33’N, 122°24'18’W, |
23 m (16-36), Sta. 1 of Banse and Hobson (1968); 27 Feb. 1963; 1 specimen |
(USNM 36538). |

Description.—Slender, up to about 8 mm long with about 50 segments. |

VOLUME 93, NUMBER 4

200 um

NS
S a Sr Lot
(<= ey
Ke ——=—
= CT Ws | EESEEE ae SS 5
> Ss SS
Cs nal aa thee . ets aa
_—— or EET aes
= Sf Bess
= ag emia saa
A) ERP RD SS
S| U ‘ ot
= aa “ a
a F p+
Spi a a \\ Ses
SS eee
Cle ae
iss Gs, ,
= _ Be
a ;
qe

1093

Game

win G|

Fig. 5. Exogone longicirris: a, b, Superior simple setae; c, d, Compound falcigers; e, In-
ferior simple seta; f, Aciculum. Exogone arenosa: g, Anterior end, relaxed specimen; h,
Same, contracted specimen (FSBC I 23522); i, Posterior end, ventral view; j, Inferior simple

seta (FSBC I 23516).

1094 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Median antenna long, nearly reaching to anterior ends of palps; lateral an-
tennae short, ovoid; 4 eyes. Proventriculus cylindrical, shorter than phar-
ynx, with about 20 rows of muscle cells on mature specimens, in length of
3 or 4 segments beginning in setiger 3 or 4; pharynx dark red, with tip rolled
backwards when everted. Dorsal cirri on all setigers. Setae including su-
perior and inferior simple setae, compound spinigers and falcigers. Superior
simple setae beginning on anterior setigers, strongly bent near tips; tips
pointed; irregularly dentate on outer edges below tips but without spines or
aristae (Banse and Hobson, 1968:fig. 4d). Spinigers of setiger 2 enlarged,
with large, triangular process below tip of shafts and tongue-shaped process
on one side of tip, with blades shorter than on setiger 1 or 3 (Banse, 1972:fig.
5b). Spinigers continuing to posterior end but gradually shorter. Falcigers
(Banse and Hobson, 1968:fig. 4e) bidentate, with secondary tooth much
larger than primary, serrate on edge on anterior setigers, slightly shorter
and without serrations posteriorly. Inferior simple setae on middle and pos-
terior segments, sigmoid, tips bidentate with primary tooth much smaller
than secondary. Pygidium with 2 anal cirri. Natatory setae, sperm, 2 ova or
embryos per segment beginning on about setiger 17 and extending to about
setiger 35.

Exogone uniformis Hartman, 1961

Exogone uniformis Hartman, 1961:73, 74 [in part, not pl. 6, fig. 1, pl. 7, fig.
1 and 8 specimens from type-locality, fide Banse, 1972]; 1968:427, 428 [in
part, not unnumbered figures of anterior and posterior ends].—Banse,
1972:202, 203, fig. Se [diagnosis and additional description based on ex-
amination of holotype (AHF Poly 0170) and 3 paratypes (AHF Poly
0171)].

Remarks.—Both Hartman’s (1961) description and material of E. unifor-
mis included E. lourei Berkeley and Berkeley; however, the 2 species differ
as diagnosed by Banse (1972). Also see remarks under E. dispar and E.
arenosa.

Exogone arenosa, new species
Figs. 5g-j, 6

Exogone dispar.—Taylor, 1971:201—204.—Hall and Saloman, 1975:12 [list] |
[in part, not Paedophylax dispar Webster, 1879].

Material examined.—FLORIDA: Hutchinson Island Sta. V, 27°22.9'N, |
80°13.9'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher, |
col., Jan. 1972; USNM 60442), 54 paratypes (USNM 60443; AHF Poly 1308, |
1309; ZMH P-16389; FSBC I 23503-23511). Sta. II, 27°21.6’N, 80°13.2'W, |
about 11 m, coarse calcareous sand; 17 paratypes (ZMH P-16390; FSBC I |

VOLUME 93, NUMBER 4 1095

23512-23516). Sta. [V, 27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous
sand; 10 paratypes (FSBC I 23517-23523). Palm Beach, 0.5—0.75 mi off
Breakers Hotel, ‘‘Breakers Reef,’ 26°42.8’N, 80°01.2’W, 23.7 m; J. W.
Smith et al., cols., Mar. 1976; 1 paratype (FSBC I 23524). Tampa Bay,
27°36'56"N, 82°41’05’”W, 9 m, sand; J. Taylor and C. Saloman, cols., 25 Oct.
1963; 170 paratypes (USNM 60444).

Description.—Tube mucoid, covered to some extent with sand. Body
without color markings; eyes light to dark red, rarely black. Well-preserved,
sexually mature specimens mostly 5—6 mm long, about 0.25 mm wide, about
50 setigerous segments. Prostomium (Fig. 5g, h) about twice as wide as
median length; anteromedian part forming rounded, obtuse angle; lateral
sides rounded; posterior side often covered by fold of tentacular segment.
Two pairs of lensed eyes on posterior part in about middle and in contact
on each side; anterior pair slightly larger, more lateral. Long, fusiform me-
dian antenna originating about middle of prostomium, extending anteriorly
to near tips of palps; papilliform lateral antennae originating between base
of median antenna and anterior pair of eyes; all antennae on slight promi-
nence of prostomium. Palps together slightly wider than prostomium, slight-
ly longer than median prostomial length, with anterior notch and median
dorsal furrow. Tentacular segment slightly wider than prostomium, about
half length of following segments, with pair of small, ovoid tentacular cirri
on anterolateral margins. Small, pyriform or papilliform dorsal cirri on all
setigers of almost all specimens; few juvenile specimens (FSBC I 23525)
with dorsal cirri absent from setiger 2; cirri well above parapodia, usually
Shorter than parapodial lobes. Parapodial lobes stout, truncate, distally bi-
labiate. Ventral cirri slightly flattened, smaller than but otherwise similar to
dorsal cirri, originating on ventral median parts of parapodial lobes. Solitary,
superior simple setae (Fig. 6b, c) with long, thin secondary spine near tips,
beginning on first setiger of juveniles, on more posterior setigers of larger
specimens (setiger 1-15), not as thick as shafts of compound setae on an-
terior segments, becoming much stouter on posterior segments. Compound
setae (Fig. 6a, d—g) both spinigerous and falcigerous. Spinigers 1, occasion-
ally 2, in upper parts of bundles. Spinigers of setiger 2 (Fig. 6a, d) with stout
shafts having single, large, stout, triangular process below tips on same side
as pectinate edges of blades and tooth on one side of tips; blades 30-43 um
long. Shafts of spinigers of remaining segments (Fig. 6e) without triangular
process. Spinigerous blades of setiger 3 always longer than those of setiger
2, 38-49 um long; blades 26-30 um long at posterior end. Falcigerous blades
bidentate, with secondary tooth much stouter than primary, numbering 6—
7 on anterior parapodia, 3—4 on median parapodia, 2—3 on posterior para-
podia, about 10 um long on anterior segments with about 6 long serrations
below secondary tooth, shorter posteriorly, 6 wm or less in length and
smooth below secondary tooth. Solitary, bidentate, inferior simple setae

1096 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

15 ym
5
g
f
: is
i
h
———————————_ — ID ym
15 pm 15 um

Fig. 6. Exogone arenosa: a, Spiniger with stout, triangular process on shaft, setiger 2; b,
Superior simple seta, setiger 10; c, Same, posterior setiger; d-g (FSBC I 23516): d, Spiniger
with stout, triangular process on shaft, setiger 2; e, Spiniger without stout triangular process
on shaft, setiger 3; f, Falciger, setiger 2; g, Falciger, setiger 49; h, Acicula, setiger 3; i, Acic-
ulum, posterior setiger.

(Fig. 5j) on last 16 setigers of large specimen, on posterior S—7 segments of
most specimens. Three acicula in anterior parapodia; solitary, much stouter
one in posterior parapodia (Fig. 6h, i). Pygidium (Fig. 51) rounded, with pair
of anal cirri about as long as last 3 setigerous segments. Sexually mature
males with natatory setae beginning on setiger 19-20 for maximum of 22
segments; sperm mostly beginning on segment anterior to that on which
natatory setae begin, extending to 1-2 segments anterior to segment having
posteriormost natatory setae. External embryos 2 per segment, attached to
median side of ventral cirri of some females beginning on setiger 19-20,
continuing for 15-19 segments; natatory setae absent on females with ex-
ternal embryos, occasionally found on more immature specimens. Approx-
imately last 10 segments lacking sex products or natatory setae.

VOLUME 93, NUMBER 4 1097

Pharynx relatively thin walled, with large, anterior middorsal tooth; an-
terior end surrounded by 10 soft papillae; wall surrounded by brownish
glands. Location of pharynx depending upon state of contraction, usually
in first S setigerous segments of relaxed specimens, extending from anterior
of palps to posterior part of setiger 3 of contracted specimens. Proventric-
ulus about equal in length to pharynx, cylindrical, slightly greater than 3
times longer than diameter, with 25-28 rows of muscle cells with anterior
6 rows small in sexually mature adults, with fewer rows of muscle cells,
e.g., 12 large and 6 small rows in juveniles. Posterior part of proventriculus
usually extending to anterior part of setiger 9 with well-defined ventricle in
same.

Remarks.—Exogone arenosa differs from E. dispar Webster in having
shafts of spinigers of setiger 2 enlarged with a large triangular process near
the tips, in having a longer proventriculus and in having superior simple
setae with a spine on the tips. E. arenosa is very similar to E. lourei Berke-
ley and Berkeley from the northeast Pacific; compound setae of the 2 species
appear identical. However, superior simple setae of E. arenosa differ from
those of E. lourei in having a well-defined spine on the tips, and the pro-
ventriculus of E. arenosa is longer with up to 28 rows of muscle cells while
that of E. lourei has about 20 rows of muscle cells. The compound setae
and proventriculus of E. uniformis Hartman, 1961, are similar to those of
E. arenosa. E. uniformis has a shorter median antenna and superior simple
setae apparently do not have a spine on the tips (cf. Banse, 1972:203).

Etymology.—The specific name is derived from Latin and refers to the
sandy tube.

Exogone atlantica, new species
Fig. 7

Material examined.—FLORIDA: Eastern Gulf of Mexico off Egmont
Key, Project Hourglass Sta. C, 27°37'N, 83°28’W, 37 m, in scleractinian
Siderastrea radians (Pallas, 1766); R/V Hernan Cortez, B. Presley, col., 13
Dec. 1966; holotype (USNM 60345). Same, Project Hourglass Sta. B,
27°37'N, 83°07'W, 18 m; 3 Apr. 1967; 2 paratypes (FSBC I 23526). Same,
off Sanibel Island, Project Hourglass Sta. L, 26°24'N, 83°22’W, 55 m; 15
Nov. 1967; 1 paratype (USNM 60346). Hutchinson Island Sta. II, 27°21.6’N,
80°13.2’W, about 11 m, coarse calcareous sand; 5 paratypes (AHF Poly 1310;
ZMH P-13191; FSBC I 23527-23529).

Description.—Pharynx reddish brown, surrounded by thin, brownish
glands. Maximum length 3.2 mm, 38 setigerous segments; body anteriorly
tapered for first few segments, gradually tapered posteriorly. Prostomium
(Fig. 7a) longer than wide. Three pairs of eyes; anterior pair small, appar-
ently lensed, on margin anterior to lateral antennae, remaining 2 pairs larger,

1098 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

15 um

15 pm

150 ym

Fig. 7. Exogone atlantica: a, Anterior end of holotype, left lateral antenna partially ob-
scuring lateroanterior, lensed eye; b, Posterior end, ventral view; c, Parapodium, anterior
setiger, posterior view (FSBC I 23527); d-f, Superior simple setae: d, Setiger 12; e, Middle
setiger; f, Posterior setiger; g, Spiniger, anterior setiger (FSBC I 23527); h, Upper and lower
falcigers of same; i, Inferior simple seta.

lensed, on posterior part of prostomial lobe, arranged as flattened trapezoid
open to front, partially covered by fold of tentacular segment. Three minute
antennae between anterior pair of large eyes. Palps slightly narrower and
about as long as prostomial width, subtriangular, with small anterior notch
and dorsal furrow. Tentacular segment visible dorsally, anteriorly forming
fold covering posterior part of prostomium; length similar to following seg-
ments; width intermediate between those of prostomium and setiger 1. Sin-
gle pair of minute tentacular cirri smaller than antennae and dorsal cirri on

VOLUME 93, NUMBER 4 1099

lateral sides. Dorsal cirri absent from setiger 2, short, somewhat oval in
outline on other segments, originating well above parapodial lobes. Ventral
cirri about same length but more slender than dorsal cirri, originating on
median ventral edges of parapodial lobes, often longer on posterior than on
anterior segments. Solitary, superior simple setae (Fig. 7d-f) on all para-
podia, with acute tips, coarsely serrate near tips with 2—S pointed teeth on
edge. Upper compound setae (Fig. 7g) solitary spinigers, occurring only on
anterior half of body on 2 specimens, on all but posterior 4-10 setigers of
others; blades of spinigers long, thin, tips rounded or slightly hooked, finely
serrate on edge, about half as long on posterior as in anterior segments;
shafts thin, with distal part acutely tipped, broad at socket of blade. Falci-
gers (Fig. 7h) 3-4 in each parapodium; shafts stouter than those of spinigers;
all blades similar, delicate, slightly falcate, unidentate, almost as wide and
less than twice as long as shaft width, with up to 4 coarse serrations on
edge. Solitary, inferior simple setae on posterior segments (Fig. 7i), with
acute tips, with 1, occasionally 2, pointed secondary teeth on concave side
near tips. Acicula solitary; tips slightly enlarged, bent forward. Pygidium
(Fig. 7b) with pair of long, slender anal cirri. One somewhat damaged sex-
ually mature specimen with sex products in setigers 11-28; gravid segments
with bundles of about 10 short, very fine, natatory setae between parapodia
and dorsal cirri.

Pharynx long, extending posteriorly to setiger 4-6. Middorsal tooth rel-
atively large, anterior; anterior end surrounded by about 10 soft lobes. Pro-
ventriculus narrow, cylindrical, in setigers 4-6 to 7-8, with 17-20 rows of
muscle cells, anterior 5-6 small; prominent ventricle in setiger 8 or 9. Rel-
ative lengths of pharynx and proventriculus 1:0.8.

Remarks.—Exogone atlantica is similar to E. microtentaculata West-
heide (1974: 121-123, figs. 5la—d, 56) from the Galapagos Islands, but differs
in the relative lengths of the pharynx and proventriculus, in shapes of su-
perior and inferior simple setae and possibly in the shape of acicular tips.

Etymology.—The specific name refers to the type-locality of this species
as opposed to that of the closely related tropical Pacific species, E. micro-
tentaculata Westheide.

Odontosyllis Claparede,. 1863
Odontosyllis longigulata, new species
Fig. 8

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6'N,
80°13.2’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., May 1972; USNM 60445), 13 paratypes (USNM 60447; ZMH P-16392;
FSBC I 23530-23536). Sta. IV, 27°20.7’N, 80°12.8’W, about 11 m, coarse
calcareous sand; 13 paratypes (USNM 60446; AHF Poly 1311; FSBC I
23537-23544).

1100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

f | /9

15 ym
Fig. 8. Odontosyllis longigulata: a, Anterior end, dorsal view (FSBC I 23532); b, Posterior
end of holotype, dorsal view; c, Parapodium, slightly turned, anterior view; d-j (AHF Poly
1311): d, Upper falciger, setiger 10; e, Lower falciger of same; f, Falcigers, posterior setigers;
g, Superior simple seta of same; h, Inferior simple seta of same; i, Aciculum, setiger 10; j,
Same, posterior setiger; k, Anterior border of pharynx, dorsal view (FSBC I 23532).

VOLUME 93, NUMBER 4 1101

Description.—Body without color markings; pharynx often dark reddish
brown. Maximum length about 6.5 mm with maximum of about 50 segments.
Prostomial lobe oval, about twice as wide as long, posteriorly notched. Two
pairs of lensed eyes in trapezoidal, almost rectangular, arrangement, ante-
rior pair larger. Three short, similar, club-shaped to fusiform antennae,
about half as long as prostomial width; median originating from about middle
of prostomial lobe; laterals originating on anterolateral margin between an-
terior pair of eyes; all generally directed upward. Ciliated nuchal organs
posterior to eyes on curved lateral parts of prostomium. Palps basally fused,
free tips usually bent ventrally. Tentacular segment visible dorsally only as
short, anteriorly rounded, nuchal fold (Fig. 8a). Tentacular cirri generally
directed ventrally. Dorsal tentacular cirri slightly longer than ventral; ven-
tral tentacular cirri slightly longer than antennae. Setigerous segments
strongly arched dorsally, flattened ventrally. Dorsal cirri club shaped to
fusiform on anterior segments, fusiform posteriorly, about twice as long as
antennae on setiger 1; short, 12 to 4 body width without parapodia behind
first few segments. Parapodial lobes (Fig. 8c) relatively slender. Ventral cirri
originating near tips of parapodial lobes, appearing slightly compressed,
extending laterally about to tips; ventral margins forming line about 30° from
aciculum. Only bidentate compound setae on anterior parapodia, joined by
solitary superior and inferior simple setae on posterior parapodia. Superior
simple setae (Fig. 8g) smooth, slender, slightly curved, unidentate. Blades
of upper compound setae of anterior parapodia (Fig. 8d) with tips strongly
falcate, with large secondary tooth below primary; blade edges pectinate,
with about 12 long serrations. Distal ends of shafts of upper compound setae
of anterior parapodia strongly serrate. Blades of lower compound setae of
anterior parapodia shorter, stouter than upper blades, with primary teeth
not as falcate, and secondary teeth not as stout, with about 6, short serra-
tions on blade edges. Shafts of lower compound setae of anterior parapodia
(Fig. 8e) slightly stouter than dorsal; distal ends bifid, with about 5 small
serrations below tips. All compound setae of posterior segments similar;
blades with long, pointed, only slightly falcate primary tooth, secondary
tooth short, triangular, otherwise smooth. Inferior simple setae (Fig. 8h)
slender, half as thick as shafts of compound setae; tips bidentate, similar in
shape but smaller than blade tips of compound setae. Acicula (Fig. 81, j)
solitary, about twice as stout in posterior parapodia, slightly enlarged near
tips, truncate. Pygidium (Fig. 8b) with long, helically curved anal cirri;
length of cirri of 1 specimen equal to that of posterior 10-12 segments.

Pharynx long, thin walled, in setigers 2—7 to S—9 (S—6 segments), with
long dorsal projection at entrance; trepan (Fig. 8k) with ventral row of about
6 teeth and 2 lateral plates. Proventriculus long, in setigers 8-9 to 15 (7-8
segments), with 60-70 rows of muscle cells. Length ratio of pharynx to
proventriculus 1:1.8.

1102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—Odontosyllis longigulata is superficially similar to O. unde-
cimdonta Imajima and Hartman (1964:114, pl. 26, figs. h, i, pl. 27, figs. a—
e) from Japan in having a short, rounded nuchal fold, short antennae and
dorsal cirri, bidentate compound setae, superior and inferior simple setae,
and blunt-tipped acicula. O. undecimdonta is a larger species, 20 mm long
and 2 mm wide with 120 setigerous segments, the nuchal fold is larger, the
pharynx has 11 teeth, and parapodia are much stouter than those of O.
longigulata.

Etymology.—The specific name, derived from Latin, refers to the long
pharynx.

Parapionosyllis Fauvel, 1923
Parapionosyllis longicirrata (Webster and Benedict, 1884)
Fig. 9

Sphaerosyllis longicirrata Webster and Benedict, 1884:715, 716, pl. 8, figs.
95-100.

Pionosyllis manca Treadwell, 1931:1, 2, fig. 1.

Parapionosyllis longicirrata.—Pettibone, 1963:132, fig. 35e-f.—Day,
1973:32.—Gardiner, 1976:133, fig. 1lo-r.

Material examined.—MASSACHUSETTS: Orleans, Cape Cod; Petti-
bone, col., 25 Aug. 1954; 2 specimens (USNM 27522). Provincetown, Cape
Cod, muddy sand; Pettibone, col., 29 Aug. 1954; 1 specimen (USNM 32519).
VIRGINIA: Off Cape Henry, 38 m, bottom net, USFC Fish Hawk Sta.
8835, 1 paratype of Pionosyllis manca Treadwell (USNM 19600). NORTH
CAROLINA: Off Beaufort, 34°34’N, 76°25’W, 20 m, sand and broken shell;
4+ specimens (USNM 51070). FLORIDA: Hutchinson Island Sta. I,
27°21.3'N, 80°14.1’W, about 8 m, very fine to fine quartose sand; 1 specimen
(FSBC I 20795). Sta. II, 27°21.6’N, 80°13.2'W, about 11 m, coarse calcar-
eous sand; 10 specimens (FSBC I 20612-20621). Sta. III, 27°22.0'N,
80°12.4’W, about 7 m, medium calcareous sand; 21 specimens (FSBC I
20622-20633). Sta. 1V, 27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous
sand; 15 specimens (USNM 54517; FSBC I 20634-20645). Sta. V, 27°22.9'N,
80°13.9’W, about 11 m, coarse calcareous sand; 35 specimens (USNM
54516, 54518; FSBC I 20646-20657, 20659-20661). Palm Beach, 0.5—0.75 mi
off Breakers Hotel, *‘Breakers Reef,’ 26°42.8’N, 80°01.2’W, 23.7 m, in
galleries of fossil coral; J. W. Smith et al., cols., 3 March 1976; 1 specimen
(FSBC I 20662). Tampa Bay; J. Taylor, col., 1963; 4 specimens (USNM
54170). Same, 27°36'56’N, 82°41'05”W, 9 m, sand; J. Taylor and C. Saloman,
cols., 25 Oct. 1963; 17 specimens (FSBC I 17926). Same, 27°36'38’N,
82°43'29"W, 11 m, sand; J. Taylor and C. Saloman, cols., 29 Oct. 1963; 1
specimen (FSBC I 17422).

Description.—Body without color markings; pharynx often surrounded

VOLUME 93, NUMBER 4 1103

100 yum

oO
re
S

ee

m laid k

15 ym
Fig. 9. Parapionosyllis longicirrata: a-j, Florida specimens: a, Anterior end of juvenile,
dorsal view (FSBC I 20662); b, Anterior parapodium, dorsal view; c-f, Anterior setiger: c,
Upper falciger; d, Lower falciger; e, Aciculum; f, Superior simple seta; g—j, Posterior setiger:
g, Falciger; h, Aciculum; i, Superior simple seta; j, Inferior simple seta; k-m, Massachusetts
specimen (USNM 27522): k, Superior simple seta, middle segment; 1, Same, posterior segment;

m, Inferior simple seta of same.

1104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by brownish glands; paired, light brown glands often prominent above dorsal
cirri and occasionally on palps. Some juveniles lacking eyes on prostomium;
others with 3 pairs of eyes, single pair of anterior eyespots, 2 pairs of lensed
eyes on posterior half; eyespots mostly absent and posterior 2 pairs greatly
enlarged on sexually mature specimens. Subdermal glands, similar to para-
podial glands, visible near anterior ends of palps of few specimens. Dorsal
cirri on all setigerous segments. Dorsum not papillate, but each segment
with 2 subdermal, dorsal glands on each side with openings above dorsal
cirri (Fig. 9a, b). Superior simple setae (Fig. 9f, j) of anterior region of
moderate size but greatly enlarged in posterior region. Compound setae 5—
10 per parapodium (Figs. 9c, d, g); all finely serrate on blade edge; blades
unidentate, dorsally long, shorter ventrally in anterior region, all about same
length in posterior region; shafts with long tips smooth and pointed, collar
surrounding socket of hinge, in face view, relatively long, thin, broader than
remainder, easily broken. Inferior simple setae on posterior segments (Fig.
9j), about as stout as corresponding superior simple setae, strongly hooked
near tips, smooth. Acicula (Fig. 9e, h) solitary, with widened, circular tips,
relatively slender in anterior region, similar in shape but much stouter in
posterior region. Some mature males and females with natatory setae be-
ginning on setiger 11 or 12. Males with sperm from setiger 10 or 11 to near
posterior end; females with 2 internal eggs or external embryos per segment
beginning on setiger 11 or 12, extending to 5-10 segments from posterior
end; embryos apparently in subspherical cases attached medially to ventral
cirri.

Pharynx slightly longer than proventriculus, mostly found from anterior
prostomial margin to posterior border of setiger 2; anterior margin smooth,
surrounded by 10 soft lobes. Proventriculus found in 2-4 segments depend-
ing on contraction, mostly beginning at anterior border of setiger 3, cylin-
drical, about twice as long as wide, with about 15 transverse rows of muscle
cells, anterior 6 small.

Remarks.—Florida specimens are in agreement with Massachusetts spec-
imens identified by Pettibone (USNM 27522, 32519), except that superior
simple setae, although similar, are more strongly serrate, and inferior simple
setae are more strongly hooked (Fig. 9k—m). Setae of the paratype of Pion-
osyllis manca Treadwell appear identical with Florida specimens; however,
antennae were missing from the specimen. Pettibone (1963) stated that P.
longicirrata females had one egg per segment and embryos with up to 5
setigers attached ventrally to the body wall. Florida specimens have 2 eggs
per segment, and external embryos, which occurred on 3 Florida specimens,
were not highly developed. Florida, North Carolina and Massachusetts
specimens have 2 subdermal glands above parapodia of each setigerous
segment, but these could not be definitely confirmed on the paratype of
Pionosyllis manca from Virginia. The figure of an upper simple seta illus-

VOLUME 93, NUMBER 4 1105

trated from North Carolina specimens by Gardiner (1976:134, fig. 11p) is
incorrect; it possibly was drawn from a damaged shaft of a compound seta.

Pionosyllis Malmgren, 1867
Pionosyllis gesae, new species
Fig. 10

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6’N,
80°13.2’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher ef
al., cols., Mar. 1976; USNM 60456), 9 paratypes (USNM 60457-60459;
ZMH P-16393; FSBC I 23545-23548). Sta. III, 27°22.0’N, 80°12.4'W, about
7 m, medium calcareous sand; | paratype (FSBC I 23551). Sta. IV,
27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 4 paratypes
(USNM 60460; ZMH P-13691; FSBC I 23549, 23550). Sta. V, 27°22.9'N,
80°13.9’W, about 11 m, coarse calcareous sand; 1 paratype (FSBC I 23552).

Description.—Body without color markings. Slender; longest specimen
5.0 mm long (range, 2.9-5.0, n = 10), posteriorly incomplete, with 50 setig-
erous segments; maximum number of setigerous segments 53 (34—53 for all
8 apparently complete specimens, 49-53 for 4 complete specimens). Prosto-
mium (Fig. 10a, b) generally oval, slightly wider than long, composed of 3
indistinct lobes; anterior lobe between lateral antennae and in front of medial
antenna; lateral lobes comprising lateral sides of prostomium beginning an-
teriorly inside lateral antennae, ending posteriorly near midline. Some spec-
imens with 3 pairs of eyes; 2 large pairs in arc on posterior third of prosto-
mium in line with origin of median antenna; small anterior pair medially
adjacent to origins of lateral antennae; only anterior pair of eyespots visible
on some specimens. Median antenna originating slightly posterior to mid-
length of prostomium. Lateral antennae originating slightly posterolateral to
anterior eyespots near anterior border of prostomium. Antennae long, slen-
der, 3-4 times longer than prostomial width, considerably thicker than dor-
sal cirri. Palps flattened, anteriorly rounded, about as long as prostomial
length, only slightly fused, if at all, at base. Band of cilia on each side of
prostomium on anterior border of lateral lobes; ciliated nuchal organs on
posterior part separated by posterior extension of prostomium. Tentacular
segment occasionally similar in length to following segment, usually con-
tracting to less than half that length; transverse band of cilia on dorsal side.
Dorsal and ventral tentacular cirri similar to lateral antennae. Dorsal cirri
absent from setiger 2; alternately long and very short on other segments;
long dorsal cirri originating on short cirrophores, about 114 times body width
without parapodia; short cirri not as thick as long cirri, extending only about
half distance to tips of parapodial lobes, apparently originating directly from
body wall. Parapodial lobes (Fig. 10c, d) long, subcylindrical, truncate, well
separated, with row of 4—S tufts of cilia on dorsal and ventral sides; rows

1106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

[OO ym

&

FITS ARIZ

apne os eel
15 um

PS eS er
15 um

Fig. 10. Pionosyllis gesae: a, Anterior end, dorsal view (ZMH P-16393); b, Same (ZMH P-
13691); c, Parapodium with long dorsal cirrus (missing), posterior view; d, Parapodium with
short dorsal cirrus, posterior view; e, Superior simple seta, anterior setiger; f, Same, posterior
setiger; g-i, Compound falcigers, middle setiger: g, Upper; h, Middle; i, Lower; j, Aciculum
of same; k, Anterior end, showing proboscis, ventral view (FSBC I 23549).

VOLUME 93, NUMBER 4 1107

of tufts apparently extending to dorsum in middle of each segment. Ventral
cirri originating ventroposteriorly near tips of parapodia, subulate, extend-
ing slightly past parapodial tips. All parapodia with 3 compound falcigers
and 1 superior simple seta; simple setae (Fig. 10e, f) straight, fine tipped,
finely serrate near tips on anterior setigers, entirely smooth on middle and
posterior setigers. Blades of falcigers with serrate margins, slightly falcate,
bidentate with primary tooth larger. Blades of upper falcigers of anterior
and middle segments (Fig. 10g) with about 8 long, fine teeth below secondary
tooth, blade edges oriented ventrally. Two ventral falcigers (Fig. 10h, i) with
blades having 6—7 or more short triangular teeth below secondary tooth,
with blade edges oriented dorsally. Dentition of all falcigers in posterior
parapodia similar to that of ventral 2 of anterior parapodia. Shaft tips of
falcigers quadrilobate with pair of pointed lobes and pair of rounded lobes.
Acicula solitary, parallel sided except for tips; tips slightly enlarged, bent
anteriorly. Pygidium with pair of long anal cirri similar to longest dorsal
cirri. Most sexually mature specimens with sexual products in about 20
segments beginning with setiger 9, products occasionally beginning in setiger
8 or 10, setiger 12 on one specimen. None with natatory setae.

Pharynx brownish, extending posteriorly to setiger 4 when inverted, with
smooth anterior rim surrounded by 10 soft papillae (Fig. 10k). Median dorsal
tooth anterior, trilobed; lateral lobes small; median lobe much longer and
wider. Proventriculus slightly barrel-shaped, in setigers 5—7, with chitinous
ring possibly in position indicated by dashed lines (Fig. 10b) but not obvious,
with about 23 very irregular muscle cell rings.

Remarks.—Pionosyllis gesae is very similar to Eusyllis heterocirrata
Hartmann-Schroder (1959:118-121, figs. 64-66), from El Salvador and east-
erm Africa. According to Dr. Hartmann-Schréder (personal communica-
tion), E. heterocirrata has all falcigers with similar, relatively coarse ser-
rations, and ciliation is not visible. Further, she stated that she could not
definitely determine that the pharynx of E. heterocirrata has a denticulate
margin; if the pharyngeal margin of the latter species proves to be smooth,
it should be referred to Pionosyllis. Additionally, E. heterocirrata has 26—
30 segments and is 1.8—3.2 mm long including specimens from Tanzania and
Natal, southeastern Africa. Thus, in addition to a possible difference in the
pharynx and the presence of cilia, specimens of P. gesae are larger, have
a greater number of segments and have blades of upper falcigers of anterior
and middle segments with long, fine serrations. Pionosyllis gesae also shows
some similarity to Dioplosyllis Gidholm, 1962, particularly in the shape of
the prostomium and dorsal cirri and in having rows of cilia across the pro-
stomium, peristomium and setigerous segments. However, the ventral arc
of small teeth in the pharynx of Dioplosyllis is not present, and the chitinous
ring in the proventriculus, if present, is indistinct.

1108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology.—The species is named in honor of Dr. Gesa Hartmann-Schro-
der, whose advice made possible description of this species and others.

Pionosyllis uraga Imajima, 1966

Pionosyllis uraga Imajima, 1966a:114, 116, fig. 37.—Banse and Hobson,
1968:16, 17, fig. 4f, g.

Pionosyllis cf. uraga.—Day, 1973:33, fig. 4k—m.

Pionosyllis sp.—Gardiner, 1976:137.

Material examined.—NORTH CAROLINA: off Beaufort, 34°19'N,
75°56'W, 130 m; J. H. Day, col., 1965; 1 specimen (USNM 51071). FLOR-
IDA: Hutchinson Island Sta. II, 27°21.6’N, 80°13.2’W, about 11 m, coarse
calcareous sand; 3 specimens (USNM 54521; FSBC I 20663). Sta. IV,
27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 2 specimens
(USNM 54519; FSBC I 20664). Sta. V, 27°22.9'N, 80°13.9’W, about 11 m,
coarse calcareous sand; 10 specimens (USNM 54520; FSBC I 20665-20671).

Remarks.—Pionosyllis uraga was described from Japan and has also been
reported from Puget Sound, Washington (Banse and Hobson, 1968). North
Carolina and Hutchinson Island specimens are in agreement with Imajima’s
description, except that the middorsal tooth is near the midpoint rather than
near the anterior margin of the pharynx. Compound setae have fine ser-
rations on blade edges and 3-4 rows of serrations on shaft tips. Acicula
also appear to agree. All specimens from Florida and North Carolina are
anterior pieces only, as was also the case with Imajima’s specimens.

Plakosyllis Hartmann-Schroéder, 1956
Plakosyllis quadrioculata, new species
Figs. 11, 12

Material examined.—FLORIDA: Hutchinson Island Sta. V, 27°22.9’N,
80°13.9'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Sep. 1972; USNM 60219), 6 paratypes (USNM 54507; ZMH P-16394;
FSBC I 23553, 23554). Sta. II, 27°21.6’N, 80°13.2’W, about 11 m, coarse
calcareous sand; 14 paratypes (USNM 54505, 54506, 60220; AHF Poly 1312,
1313; FSBC I 23555-23558). Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m,
coarse calcareous sand; 3 paratypes (FSBC I 23559, 23560).

Description.—Body without color markings. Maximum length 3.0 mm,
width 0.3 mm; mature specimens with 30-55 setigers. Body strongly flat-
tened, ribbon-like, gradually broadening for first 7-8 segments, parallel sid-
ed except for few segments near posterior end, cuticle thick on prostomium
and body but not on cirri (Fig. 11b). Prostomium (Fig. 1la—c) about twice
wider than long, trapezoidal, widest near anterior margin, with pair of ven-
trolateral lobes on anterior margin and much larger, broader middle lobe;

VOLUME 93, NUMBER 4 1109

Fig. 11. Plakosyllis quadrioculata: a, Anterior end of holotype, dorsal view; b, Same, of
small specimen (FSBC I 23555); c, Same, ventral view; d, Posterior end of holotype, dorsal
view; e, Same (USNM 60220); f, Anterior end of everted pharynx, ventral view (USNM 60220).

1110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wrt OO|

\

SN

\o ys

15 um
aa a g\\ f| | el| d

Fig. 12. Plakosyllis quadrioculata (a-g FSBC I 23554): a, Parapodium of midregion, an-
terior view; b, Same, posterior view; c, Upper compound seta, anterior setiger (blade turned);
d, Same, posterior setiger; e, f, Lower compound setae, anterior setiger; g, Inferior simple
seta; h, Aciculum, dorsal view (USNM 54506); i, Same, lateral view.

median antenna spherical, attached on anterior border of middle lobe; lateral
antennae spherical, slightly smaller than median, attached on anterior bor-
ders of ventrolateral lobes; eyes lensed, 2 pairs, smaller pair dorsolateral
near posterior border, larger pair ventral, lateral to bases of palps, closer
together and more anterior than dorsal pair; palps originating near middle
of ventral side, extending to near anterior border, oval in outline, apparently
completely separated to bases; ciliary rows on ventrolateral lobes beginning
in front of ventral eyes and extending to dorsal side of lobes above lateral
antennae; ciliated nuchal organs ventrolateral, between prostomium and
tentacular segment. Tentacular segment shorter than following segment,
with 2 pairs of tentacular cirri; dorsal pair subequal to lateral antennae,
slightly larger than dorsal cirri; ventral pair originating ventrally and not
visible from dorsum, smaller than dorsal cirri. Setigerous segments (Fig.
12a, b) similar throughout, arched dorsally, straight or slightly concave ven-
trally, 8-10 times wider than long in mature specimens (Fig. 11a), relatively
narrower in immature specimens (Fig. 11b). Dorsal cirri spherical to oblong,
with single article, without visible sensory hairs but with small bumps in-
dicating they were originally present, attached on much broader, stouter

VOLUME 93, NUMBER 4 1111

cirrophores, tips extending laterally farther than setal lobes. Setal lobes
acutely pointed. Ventral cirri fused with lower parts of setal lobes for most
of their length, with unfused, conical tips extending almost to tips of setal
lobes on posterior sides. Setae compound falcigers with short unidentate
blades (Fig. 12d-f) on anterior and middle parapodia joined by slender,
slightly hooked inferior simple setae on posterior setigers (Fig. 12g); com-
pound setae 8—12, similar on all parapodia, of 2 types; 3-5 upper compound
setae having blades 9-10 um long, proximally with about 6 long, slender
serrations, with moderately hooked tips, shafts with 4—S long spines on edge
of distal end; 5—7 lower compound setae with smooth blades about 7 um
long, shafts slightly stouter than shafts of upper setae, lightly serrate on
distal end. Acicula (Fig. 12h, i) solitary, stout, slightly broadened near tips
in dorsal view, ending in long, acute tips, often emergent or confined in
pointed lobe on upper part of setal lobes. Dorsum of each segment of mature
specimens with transverse row of subcuticular glands containing granular
material; glands fewer and apparently empty in immature specimen (Fig.
11b). Pygidium (Fig. 11d, e) somewhat flattened, medially divided, with pair
of lateroposterior anal cirri similar but larger than dorsal cirri of posterior
segments; cirri well separated from remainder of pygidium by well defined
constrictions; constrictions becoming obscure on fully developed sexual sto-
lons. Sexual reproduction by posterior budding of stolons; stolons beginning
on setigers 23-29, completely filled with sexual products except for 2-3
posterior segments, 2 with pair of large ventrolateral eyes on anterior sto-
loniferous segment; single female stolon with long natatory setae between
dorsal cirri and setal lobes; female stolons with about 6 crowded, isohedral
eggs per segment.

Pharynx (Fig. 11f) longer than proventriculus, slender, thick walled, lo-
cated in setigers 1-5 or 6 of mature specimens when inverted, with trepan
of 10 small, widely spaced, pointed teeth in addition to larger middorsal
tooth, anterior border surrounded by 10 soft lobes. Proventriculus cylindri-
cal to barrel-shaped, with about 14 rows of muscle cells with anterior 4—5
rows small, about 3 segments long, usually in setigers 7-9. Ratio of phar-
ynx:proventriculus lengths about 3:2; both found in more anterior segments
of immature specimens.

Remarks.—Plakosyllis quadrioculata differs from P. brevipes Hartmann-
Schroder (1956:87-89, figs. 1-5) in having 2 pairs of eyes, one dorsal and
one ventral, shorter ventral cirri, compound setae having upper blades with
very long teeth throughout the body and shafts with very long, slender
serrations on the distal end, inferior simple setae, and distinctly articulated
anal cirri on specimens which are not sexually modified; also, P. quadri-
oculata lacks superior simple setae.

Type-specimens of P. brevipes were reported from Naples and Banyuls-
sur-Mer in the Mediterranean Sea. Weinstein (1961) described a pair of small

1112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ventral eyes on the prostomium of additional specimens of P. brevipes from
Banyuls-sur-Mer in addition to the 2 dorsal pairs described by Hartmann-
Schroder but did not report superior simple setae.

Gidholm (1962:250-252, fig. 1) reported 2 pairs of ventral eyes and a single
dorsal pair on the prostomium of specimens referred to P. brevipes from
Roscoff on the Atlantic coast of France. Setae of Gidholm’s specimens also
appear to differ from those described by Weinstein, and setae of posterior
setigers described by Weinstein appear identical with setae originally de-
scribed for the species by Hartmann-Schroder (1956:89, fig. 4). These dif-
ferences suggest that Gidholm’s specimens are referable to another species.

Plakosyllis quadrioculata differs from P. americana Hartman (1961:75,
76, pl. 33, fig. 1), which has a prostomium with 4 dorsal eyes and no ventral
eyes, about 100 segments maximally and compound setae with entirely
smooth shafts.

Hartmann-Schroder (1956) originally placed the genus in the subfamily
Exogoninae because of reduction in antennae, palps, and cirri and because
of the pronounced flattened condition of the body. Weinstein (1961) sug-
gested that the genus was related to Trypanosyllis Claparede of the Syllinae
and Eurysyllis Ehlers, 1864, which Fauvel (1923) placed in the Syllinae, but
which Fauchald (1977) placed in the Exogoninae. Gidholm (1962) referred
Plakosyllis to Eurysyllis. Eurysyllis species have large globular papillae on
the dorsum; this and other differences and similarities were noted by Gid-
holm.

The only significant difference of Plakosyllis from the small Trypanosyllis
species described herein is in the reduction of antennae, tentacular cirri,
and dorsal cirri to a single article from the multiarticulate condition, and it
is suggested that the genus be referred to the Syllinae along with the closely
related Eurysyllis. In this regard, Sphaerodoridium guilbaulti Rullier
(1974:33-35, fig. 2), based on a single specimen from the Atlantic coast of
the United States, appears to be a Eurysyllis.

Etymology.—The specific name is derived from the Latin prefix quadri,
meaning four, and oculata, having eyes, referring to the number of eyes.

Sphaerosyllis Claparede, 1863

Remarks.—Previous reports of this genus from eastern North America,
the Bahamas, and Cuba have been included under 10 specific names. S.
brevifrons Webster and Benedict, 1884, and S. longicauda Webster and
Benedict, 1887, have been referred to S. erinaceus Claparede, 1863 (Petti-
bone, 1963), although it appears that both are distinct from one another and
from S. erinaceus. S. hystrix Claparede was recorded by Pettibone (1963)
from South Norwalk, Connecticut; the specimens, first reported by Webster
and Benedict (1884) as S. brevifrons, are referred herein to S. taylori, n.

VOLUME 93, NUMBER 4 1113

sp. S. renaudae Hartmann-Schroder, 1958, was described from a single
anterior fragment from Bimini, The Bahamas. A solitary specimen reported
as S. pirifera [not Claparede, 1868] from North Carolina by Day (1973) and
Gardiner (1976) is referred herein to S. glandulata, n. sp. Specimens from
North Carolina reported as S. erinaceus by Gardiner (1976) are referred to
S. longicauda. S. centroamericana Hartmann-Schroder, 1959, was recently
reported from Cuba by Hartmann-Schroder (1979). An additional new
species, S. labyrinthophila, recently described by Gardiner and Wilson
(1979) from North Carolina, was also collected at Hutchinson Island.

Sphaerosyllis erinaceus and S. hystrix were also reported from Delaware
by Kinner and Maurer (1978), and S. hystrix by Maurer et al. (1976); I have
not examined their specimens and confirmed their records. However, S.
hystrix and S. erinaceus are European species, and I have no evidence that
they occur in the northwestern Atlantic from numerous specimens of this
genus that I have examined. S. hystrix specimens reported by Kinner and
Maurer (1978) and Maurer et al. (1976) are probably S. taylori, n. sp. Spec-
imens of S. erinaceus (a doubtful species in my opinion) reported by Kinner
and Maurer (1978) are probably S. longicauda. S. hystrix and S. erinaceus
are therefore not included in the key.

Sphaerosyllis longicirrata Webster and Benedict was discussed above as
a member of the genus Parapionosyllis Fauvel, 1923. Another species, S.
fortuita Webster, 1879, was described from a single Virginia specimen. The
specimen was apparently never deposited in the U.S. National Museum of
Natural History where most of Webster’s types are found. I cannot defi-
nitely decide from the original description that the species is a member of
Sphaerosyllis Claparede.

As far as I can determine, species names noted above include only 7
Sphaerosyllis species, 2 of which are newly described. An additional 8 new
species are described herein, increasing the number of known species in the
genus from eastern North America to 13.

Characters of systematic importance at the species level include: number
and position of prostomial eyes; degree and character of fusion between the
prostomium and tentacular segment; presence or absence of nuchal organs;
relative sizes and shapes of antennae, tentacular cirri, dorsal cirri, and anal
cirri; presence or absence of dorsal cirri on the second setigerous segment;
fine detail of the setae; changes in setae within parapodia and along the
body; shape of acicula and changes in acicular size along the body; arrange-
ment, number, and possibly length of glandular papillae; presence or ab-
sence of segmental glands opening above dorsal cirri and the contents there-
of; specific segments in which sexual products and natatory setae occur;
number of embryos per segment and position of incubated embryos, either
dorsal or ventral; segmental positions and length-width ratios of the pharynx
and proventriculus; size and position within the pharynx of the pharyngeal

1114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tooth; arrangement and number of rows of muscle cells of the proventric-
ulus; and possibly the length and width of the body and the number of
segments. Additionally, I have described what appear to be dorsal and lat-
eral lobes of the brain extending into the first and second setigers. The
structures were discovered in cleared, unstained specimens, and their ex-
istence was tentatively confirmed by histological examination. The struc-
tures are contiguous with the dorsal parts of the brain in the prostomial
lobe, and histological examination indicates them to be surrounded by a
membrane continuous with that surrounding the brain. The specimens, how-
ever, were poorly preserved for critical histological examination. The struc-
tures may be of systematic importance as indicated in the species descrip-
tions.

Key to Sphaerosyllis Species from Eastern North America

NEWS NAD SE Ml caren. oePa cu ah ages is. 6 Bc: aieasuane wor sds @> Se ak cs OER S. renaudae
= SSE VES TECSCME NG: cre soc cveeuls cScuecs, Siepeuie\ suede sieges «ogo sakes ont: ee eee 2
FED KLM Ne en en MME TUES S's 6 00 0 6 0 3
Bed STK CVC Si See gcuin ls: Ge Regs 5 laudcl oct Weng, 40s abc0h op oh op cel Eee 8
3. Eyes on anterior and posterior prostomial margins; acicula slender,
Stralgches pointe Ga Aor secs case cuke ie sere S. brevidentata, n. sp.
— Eyes on posterior half of prostomium; acicula stout, with tips bent
forward at about right angles... ......3 + senses | oe eee 4
4. Dorsal parapodial glands absent .............. S. piriferopsis, n. sp.
— Dorsal parapodial glands beginning on setiger 4 or5 ............ 5
5» Parapodial glands filled with rods ..s... .. <0: 0 «sc. ss + s06 sopeene 6
— Parapodial glands filled with spheres or granules................ 7

6. Superior simple setae and compound setae in middle segments ...
Fe Nee es ee ee Res TPA HR TE SS S. taylori, n. sp.

— Superior simple setae and simple acicular setae in middle segments
SE NARA USE Le OA Ma Re We eC eee) OI RNR ea RTs S. aciculata, n. sp.

7. Median antenna originating near posterior prostomial margin;

blades of upper compound setae of anterior segments about 12 um
TOM aires ps retea ayse eaa foes ans oe occas ots ee oma ee S. glandulata, n. sp.

— All antennae originating on anterior prostomial margin; blades of

upper compound setae of anterior segments about 40 um long
LE ee ete Sa re ee Cree Han ne S. magnidentata, n. sp.

8. Two pairs of eyes on prostomium in flattened, trapezoidal arrange-

ment open posteriorly and third pair near posterior border of ten-

tacular segment; anterior pair large ............ S. centroamericana
— All eyes on prostomium; anterior pair small ................... 9
9. Blades of compound setae bidentate ............ S. bilobata, n. sp.

— Blades of compound setae unidentate .......................-- 10

VOLUME 93, NUMBER 4 1115

10. Pygidium with midventral anal cirrus; integumental papillae both

longzandsShont s.r: eee eet ernie Gli Sacse gene ales S. longicauda
— Midventral anal cirrus absent; all integumental papillae similar in
SUZSM (STORE) Rearotes epee od (onsen oi tat intne oaks Wcee hese atari |. We. anche a 11

11. Paired, semicircular nuchal folds covering posterior eyes; blades of
upper compound setae of anterior segments about 30 um long

n'd OG: Wics NO Unio cckcgateacipe: 6c). 070) ot Al et nea? eS Ea EN S. labyrinthophila
— Paired nuchal folds absent; blades of compound setae of anterior

SecmentsraboutwlO pmelongs a2. o5.t lence. = oat mae ree ie aie 12
12. Prostomium and tentacular segments fused; anal cirri slender;

length to 1.5 mm, with 17 setigers ................. S. riseri, N. sp.
— Prostomium and tentacular segments distinct; anal cirri bulbous;

greater than 2.5 mm long, with 27 setigers ............ S. brevifrons

Sphaerosyllis aciculata, new species
Figs. 13, 14

Material examined.—FLORIDA: Hutchinson Island Sta. IV, 27°20.7'N,
80°12.8’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Nov. 1971; USNM 60221), 1 paratype (USNM 60222). Sta. I,
27°21.3'N, 80°14.1’W, about 8 m, very fine to fine quartose sand; 1 paratype
(FSBC I 23561). Tampa Bay, 27°36'20’N, 82°40'49”"W, 8 m, sand; J. Taylor
and C. Saloman, cols., 25 Oct. 1963; 1 paratype (FSBC I 23562). Tarpon
Springs, Anclote Anchorage, 1 mi W of Bailey’s Bluff, 4-5 m, shelly sand;
R. Ernest, col., Nov. 1973; 1 paratype (FSBC I 23563). Same, 28°11.8'N,
82°47.6’W, 2.5 m, sand with Syringodium cover; J. Studt and R. Ernest,
cols., 5 Dec. 1975; 1 paratype (ZMH P-16395).

Description.—Parapodial glands brown to light yellowish brown. Maxi-
mum length 2.2 mm; width about 0.1 mm excluding parapodia; maximum
of 21 setigerous segments. Prostomium clearly fused with tentacular seg-
ment (Fig. 13a, b), with latter surrounding posterior and lateral sides of
prostomium as fold covering posterior part of prostomial lobe. Median an-
tenna originating on anterior edge of fold, possibly shorter than lateral an-
tennae. Lateral antennae originating on anterolateral margins of prostomium
on short, anterior projections (distinct in contracted specimen). Anterior
margin of prostomium slightly concave. Two pairs of lensed eyes on pos-
terior half, in flattened, rectangular arrangement; anterior pair slightly larg-
er. Palps long when not turned under or contracted; anterior fourth not
fused; anterior notch and dorsal median furrow evident in relaxed specimen.
Tentacular cirri originating dorsally on lateral projections of tentacular seg-
ment on about same line as origins of lateral antennae (Fig. 13a); cirri di-
rected upward on short projections on relaxed specimen (Fig. 13b), directed
anteromedially around front of prostomium on contracted specimen. Glan-

1116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 13. Sphaerosyllis aciculata: a, Anterior end of contracted specimen, dorsal view
(USNM 60222); b, Same, holotype; c, Posterior end, dorsal view.

dular papillae on tentacular segment below and above tentacular cirri. Dor-
sal lobes of brain (Fig. 13b) extending posteriorly to middle of setiger 2;
lateral lobes extending to near posterior edge of setiger 1. Dorsum of each
segment with 8-10 small glandular papillae. Single specimen with dorsal
cirrus on one parapodium of setiger 2; dorsal cirri absent on setiger 2 of
remaining specimens; cirri relatively small, not extending past tips of para-
podial lobes of other segments, spherical bases and cylindrical tips of about
equal length. Parapodial lobes blunt with single anterior and single posterior
apical papillae; third papilla sometimes evident on anterior edge about half
distance from base to tip. Ventral cirri short, cylindrical. Solitary, superior
simple setae (Fig. 14a—c) on all parapodia, relatively broad in lateral view.
Setigers 1-8 with 4 short-bladed compound setae (Fig. 14d, e); blades of

VOLUME 93, NUMBER 4 1117

eas 5

i en Seer cae al
15 pm

Fig. 14. Sphaerosyllis aciculata: a, Superior simple seta, setiger 6, lateral view; b, Same,
dorsal view; c, Same, posterior setiger; d, Upper falciger, setiger 6; e, Lower falciger of same;
f, Upper simple acicular seta, posterior setiger; g, Lower simple acicular seta of same; h,
Inferior simple seta of same; i, Aciculum, setiger 6; j, Same, posterior setiger.

uppermost seta finely serrate. Following segments with compound setae
replaced with about 3 slightly enlarged, simple, acicular setae (Fig. 14f, g)
homologous with shafts of anterior compound setae; upper acicular setae
more highly modified, showing little resemblance to shafts; tips of lower-
most acicular setae strongly similar to tips of shafts of anterior compound
setae. Solitary, inferior simple setae (Fig. 14h) on posterior parapodia.
Acicula (Fig. 141, j) solitary, stout in anterior region, about twice as stout
in posterior region, with tips slightly enlarged and bent forward at almost
right angle. Parapodial glands (Fig. 13b) containing rods, beginning with
setiger 4, extending to posterior end; glands opening on dorsum above dorsal
cirri; one gland empty, another partly empty on one specimen. Pygidium
(Fig. 13c) with pair of long, stout, anal cirri, generally similar to dorsal cirri;
in addition, pygidium with 10 long papillae visible in dorsal view. One female
with internal eggs in setigers 7-15.

Pharynx very narrow, with moderately thick walls, surrounded by rela-
tively large but not prominently colored glandular region. Relatively large
middorsal tooth anterior, about as wide as anterior opening. Pharynx located
in setigers 1 and 2 of relaxed specimen. Proventriculus shorter than pharynx,
only slightly longer than wide, extending through about 1!2 segments in
setigers 3 and 4. Muscle cells arranged in 13-14 irregular transverse rows,
anterior 4—5 small. Relative lengths of pharynx to proventriculus 5:4.

Remarks.—Sphaerosyllis aciculata differs from other species in the genus
in having compound setae replaced entirely by simple, acicular setae in
middle and posterior segments.

1118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology.—The specific name is derived from Latin and refers to the
aciculated setae.

Sphaerosyllis bilobata, new species
Figs. 15, 16

Material examined.—FLORIDA: Hutchinson Island Sta. IV, 27°20.7'N,
80°12.8’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Nov. 1971; USNM 60465), 5 paratypes (USNM 60466, 60470; FSBC
I 23565-23567). Sta. II, 27°21.6’N, 80°13.2’W, about 11 m, coarse calcareous
sand; 4 paratypes (FSBC I 23568-23571). Sta. V, 27°22.9'N, 80°13.9'’W,
about 11 m, coarse calcareous sand; 10 paratypes (USNM 60467-60469;
FSBC I 23572-23575).

Description.—Body without color markings; eyes reddish. Maximum
length 1.9 mm, about 0.1 mm wide without parapodia, 26 setigers. Prosto-
mium (Figs. 15a, 16a) wider than long, anteriorly convex, almost semicir-
cular; posterior border nearly straight; sides rounded. Lateral antennae orig-
inating on anterolateral borders of prostomium lateral to anterior eyespots;
about half length of median. Median antenna originating from posterior bor-
der of prostomium on short lobe separate from tentacular segment and as-
sociated lobes; length about equal to median prostomia length. Three pairs
of eyes; pair of eyespots on anterior margin at about middle of each side;
2 pairs of lensed eyes on posterior half, with anterolateral pair much larger
on sexually mature specimens (Figs. 15a, 16a). Dorsal lobes of brain ex-
tending to posterior border of setiger 1; lateral lobes extending well into
setiger 2. Palps anteriorly rounded, much shorter than prostomium, ap-
pearing completely fused dorsally except for median furrow. Tentacular
segment distinct dorsally, about half as long as following segment, inter-
mediate in width between narrower prostomium and wider setiger 1; ten-
tacular cirri originating lateral to posterior pair of lensed eyes, about half
size of lateral antennae and dorsal cirri; pair of eyelid-like nuchal folds on
anterior margin of tentacular segment covering ciliated nuchal organs be-
tween prostomium and tentacular segment and posterior 2 pairs of eyes
(Figs. 15a, 16a), beginning medially near midline, extending around lateral
margins of prostomium, ending above tentacular cirri; no glandular papillae.
Dorsal cirri absent from setiger 2, replaced by glandular papillae. Dorsum
of each segment otherwise with about 4 short papillae. Dorsal cirri of re-
Maining segments with spherical bases and cylindrical tips, shorter than
setigerous lobes; spherical parts about equal in length to cylindrical parts.
Setigerous lobes (Fig. 15d) conical, acutely pointed; 2 small papillae usually
visible, one on anterior side about half distance from base to tip, other on
posterior side near tip. Ventral cirri cylindrical, less than half length of
setigerous lobes. Solitary, superior simple setae (Fig. 15j) on all parapodia,

VOLUME 93, NUMBER 4 1119

lOO yum

Ga 2)
/ ee
ea Baie ae
J
e J g
15 um

Fig. 15. Sphaerosyllis bilobata: a, Anterior end, dorsal view (USNM 60469); b, Same,
ventral view (USNM 60467); c, Posterior end, dorsal view (USNM 60469); d, Posterior para-
podium, posterior view; e-—h, Compound setae, anterior setiger: e, Upper; f, Upper middle; g,
Lower middle; h, Lower; i-k, Posterior setiger: i, Middle compound seta; j, Superior simple
seta; k, Inferior simple seta; 1, Aciculum, anterior setiger.

1120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 16. Sphaerosyllis bilobata, paratype (USNM 60466): a, Anterior end, slightly turned
dorsal view, showing cutaway of eyelid-like fold on right side beginning laterally with extreme
lateral margin of fold; b, Proventriculus, dorsal view, showing arrangement of muscle cells.

VOLUME 93, NUMBER 4 1121

slender, smooth, about as long as shafts of compound setae. About 5 com-
pound setae (Fig. 15e—h) on anterior parapodia, 4 on posterior parapodia,
similar; blades bidentate with similar primary and secondary teeth; upper
blades about 20 um long, with fine serrations on basal *%4; lower blades
smooth, about 10 um long. Inferior simple setae on posterior parapodia
(Fig. 15k), more slender, shorter than superior simple setae. Acicula (Fig.
15 1) solitary, slender; each with subdistal bulb and conical tip. Pygidium
short, subcylindrical, with few glandular papillae; anal cirri stouter than
dorsal cirri, 3-4 times longer. Embryos 2 per segment, found dorsally on
setigers 9-19 of some sexually mature females; females mostly with internal
eggs. Sperm in setigers 9 to 21—24 in males. Natatory setae on one specimen
beginning with setiger 9.

Pharynx surrounded by thick, brownish glands or lobes, usually extending
to posterior margin of setiger 2, with anterior end narrower than remainder;
chitinous wall relatively thick, narrow, slightly shorter than length of pro-
ventriculus; middorsal tooth anterior, of average size; anterior end sur-
rounded by soft lobes. Proventriculus barrel-shaped, in setigers 3-5; 18-19
transverse rows of muscle cells, anterior 5 small; cells in dorsal view also
arranged in 2 opposite diagonal planes (Fig. 16b).

Remarks.—Sphaerosyllis bilobata is very similar to S. erinaceus biden-
tata Hartmann-Schroder (1974b: 134, figs. 116-119) from West Africa [=S.
bidentata]. It differs in the following characters: shape of tentacular folds
covering posterior part of the prostomium; relative sizes of median and
lateral antennae and tentacular and dorsal cirri; relative lengths of upper
and lower compound setae; and lack of dentition on lower compound setae
and superior simple setae of S. bilobata.

Etymology.—The specific name is derived from the Latin prefix, bi-,
meaning double or twice, and the latinized Greek adjective, Jobata, meaning
lobed, and refers to the tentacular lobes.

Sphaerosyllis brevidentata, new species
Fig. 17

Material examined.—FLORIDA: Hutchinson Island Sta. V, 27°22.9'N,
80°13.9'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Jan. 1973; USNM 60215), 1 paratype (FSBC I 23564). THE BAHA-
MAS: S portion of Bimini Lagoon, 25°43’N, 79°16’ W, in submerged plastic
sponges; A. Schoener, col., 1970-1971; 1 paratype (USNM 51547).

Description.—Body without color markings; pharynx surrounded by
brownish glands. Length 1.2 mm; width 0.13 mm without parapodia, 0.18
mm with parapodia; male with 17 setigers; female with 14 setigers. Prosto-
mium (Fig. 17a) oval, about twice as wide as long. Two pairs of lensed eyes;
anterior pair in position usually occupied by minute eyespots on Sphaero-

1122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Q00 O oe ne
Se ae, g
a IN e j : e f

‘< alee a
ah [5 yum

Fig. 17. Sphaerosyllis brevidentata: a, Anterior end, dorsal view (FSBC I 23564); b, Same,
holotype, ventral view; c, Superior simple seta, posterior setiger; d, Upper compound seta; e,
Lower compound seta; f, Inferior simple seta; g, Aciculum.

syllis spp. having 3 pairs of eyes, on anterior margin slightly medial to lateral
antennae; posterior pair small, located near posterolateral margin. Median
antenna missing; single lateral antenna on anterior margin, extending ante-
riorly about to tips of palps, pyriform, without usual cylindrical tip. Lobes
of brain very short; dorsal lobes extending to beginning of setiger 1, covered
by very thin integument; lateral lobes extending about half that distance.
Palps very short, papillate, completely fused except for median furrow.
Tentacular segment dorsally distinct, completely separated from prostomi-
um, unmodified, about % length of following segment in relaxed specimen

VOLUME 93, NUMBER 4 1123

but reduced by about half in another specimen, without tentacular folds
covering part of prostomium, with glandular papillae above tentacular cirri.
Dorsum with 4 papillae per segment; larger pair situated slightly in front of
dorsal cirri; smaller pair medial to larger pair. Dorsal cirri on all setigerous
segments, similar in size and shape to tentacular cirri, extending laterally
for about same distance as parapodial lobes, with globular bases indistinctly
separated from tips, often with 2 constrictions of cylindrical parts. Para-
podial lobes conical, with slightly rounded tips, with at least 1 papilla on
anterior margin short distance from tips, and another on posterior side near
tips. Ventral cirri short, cylindrical, attached to bases of parapodial lobes.
Solitary, superior simple setae (Fig. 17c) on all parapodia and solitary, in-
ferior simple setae (Fig. 17f) on posterior segments; both smooth, tips acute;
inferior simple setae strongly curved, not as stout as superior simple setae.
Four compound setae (Fig. 17d, e) with short blades on all parapodia; blades
slightly curved, unidentate, upper 2 serrate, lower 2 smooth. Solitary acic-
ulum (Fig. 17g) in each parapodium, slightly curved near tip; tip long, con-
ical. Pygidium hemispherical, with long, glandular papillae; anal cirri lost.
Mature male with sexual products in setigers 7-12; scars of external em-
bryos visible on female, 2 per segment, on dorsum of setigers 7-10.
Pharynx (Fig. 17a, b) thickly chitinized, beginning near level of anterior
pair of eyes, extending to posterior part of setiger 1, about 0.11 mm long,
with small anterior opening; middorsal tooth small, located %4 to 4 of dis-
tance from anterior end. Proventriculus barrel-shaped, in setigers 2, 3, and
part of 4; length about 0.13 mm; width about 0.05 mm. Muscle cells in dorsal
view arranged in 3 planes: prominent, transverse plane and 2 opposite di-
agonal planes, with 19-20 transverse rows, anterior 5 small.
Remarks.—Sphaerosyllis brevidentata shows some resemblance to S.
brevifrons Webster and Benedict (1884:714, 715, pl. 3, figs. 24-30) and S.
riseri, n. sp., but differs from both in having only 2 pairs of eyes, one pair
of which is located on the anterior prostomial margin and the other located
on the posterior prostomial margin. Type-specimens and others of S. brev-

_ifrons have a similar tentacular segment, antennae, and cirri to those of S.

brevidentata, but S. brevifrons is a much larger species. Antennae and cirri
of S. riseri, n. sp., have distinct spherical bases and cylindrical tips, with
the tips contracted into the bases; S. riseri also has a prostomium and
tentacular segment showing a great deal of fusion.

Etymology.—tThe specific name is derived from Latin, and refers to the
small pharyngeal tooth of this species.

Sphaerosyllis glandulata, new species
Figs. 18, 19

_Sphaerosyllis sp. Taylor, 1971:229-231, fig. 5g—j.—Hall and Saloman,

1975:12 [list].

1124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 18. Sphaerosyllis glandulata: a, Anterior end, dorsal view (USNM 60217); b, Same,
contracted specimen (AHF Poly 1315); c, Middle segment, dorsal view (not scaled).

Sphaerosyllis pirifera.—Day, 1973:34.—Gardiner, 1976:131, fig. 10p—r [not
Claparede, 1868].

Material examined.—FLORIDA: Hutchinson Island Sta. IV, 27°20.7'N,
80°12.8’'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Nov. 1972; USNM 60216), 14 paratypes (USNM 60218; ZMH P-16396;
FSBC I 23578-23585). Sta. II, 27°21.6’N, 80°13.2'W, about 11 m, coarse
calcareous sand; 9 paratypes (USNM 60217; AHF Poly 1314; ZMH P-16397;
FSBC I 23586-23590). Sta. V, 27°22.9'N, 80°13.9’W, about 11 m, coarse
calcareous sand; 17 paratypes (AHF Poly 1315; FSBC I 23591-23598). Tam-
pa Bay, 27°36’ 15’N, 82°43'22"W, 9 m, sand; J. Taylor and C. Saloman, cols.,
25 Oct. 1963; 3 paratypes (FSBC I 23599). Same; J. Taylor, col., 1963; 7

VOLUME 93, NUMBER 4 1125

I 5 ym

bata

Fig. 19. Sphaerosyllis glandulata: a, Posterior end, dorsal view (USNM 60217); b, Superior
simple setae, anterior setigers; c, Upper falciger, anterior setiger; d, Lower falciger of same;
e, Upper falciger, posterior setiger; f, Lower falciger of same; g, Superior simple setae, pos-
terior setigers, dorsal and lateral views; h, Inferior simple seta, posterior setiger; i, Aciculum,
anterior setiger; j, Same, posterior setiger.

15 ym

paratypes (USNM 45555). NORTH CAROLINA: off Beaufort, 20 m, sand
and broken shell; J. H. Day, col., 1965; 1 paratype (USNM 51074).
Description.—Body without color markings; pharynx surrounded by
light, brownish glands; parapodial glands yellow, obscure. Maximum length
7.0 mm, width 0.18 mm without parapodia, 45 setigers. Prostomium and
tentacular segment fused (Fig. 18a, b); tentacular segment forming fold cov-
ering posterior part of prostomium; median antenna originating from fold.
Two pairs of lensed eyes on posterior half, in flattened trapezoid arrange-
ment, anterior and wider pair usually slightly larger. Lateral antennae orig-
inating on anterolateral margins of prostomial lobe in front of anterior pair
of eyes, with bulbous bases and cylindrical tips, similar to median antenna;
antennae about 1% times lengths and widths of tentacular and dorsal cirri.
Glandular papillae absent from prostomium. Dorsal lobes of brain extending
into setiger 1; lateral lobes slightly longer. Palps together about as wide as

1126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

prostomium, extended or bent ventrally, about as long as antennae when
turned ventrally. Tentacular segment enclosing most of lateral margins of
prostomial lobe, with arc of few glandular papillae on dorsum. Tentacular
cirri smaller than antennae, originating on transverse line below anterior
pair of eyes. Several short, glandular papillae on dorsum and parapodial
lobes of each setiger in median segments (Fig. 18c). Dorsal cirri absent from
setiger 2; replaced by glandular papillae. Dorsal cirri of other segments
similar to antennae but smaller, not extending past tips of setigerous lobes,
with bulbous bases oval in outline, mostly longer than cylindrical tips. Se-
tigerous lobes stout, bluntly rounded. Ventral cirri slender, cylindrical, not
extending past tips of setigerous lobes. Solitary, superior simple setae (Fig.
19b, g) present from setiger 1 on all but largest specimens, beginning behind
first several setigers on latter, with slightly bent, slightly serrate tips on
anterior setigers, much stouter on posterior segments, with 4 stout teeth on
lower sides near tips. Four compound setae on anterior parapodia (Fig. 19c,
d); blades of upper slightly longer than lower (12-8 um); edges with long
serrations; 3, occasionally 4, compound setae on posterior parapodia (Fig.
19e, f); all blades about 8 wm long; upper ones with long serrations; shafts
of compound setae similar throughout body, stouter on posterior segments,
distal ends smooth. Solitary inferior simple setae (Fig. 19h) on posterior
segments, curved, as stout as superior simple setae. Acicula (Fig. 191, j)
solitary, with tips strongly bent anteriorly at slightly less than right angle in
anterior parapodia, stouter, similar in shape in posterior parapodia. Spher-
ical parapodial glands containing yellow, opaque, spherical granules always
on dorsal side above each parapodial lobe medial to dorsal cirri beginning
with setiger 4, with opening above dorsal cirri; glands mostly empty in one
specimen, with granular particles present in and around spherical vesicles.
Pygidium (Fig. 19a) hexagonal in outline with pair of long, anal cirri on
posterolateral margins; transverse row of about 6 relatively long, glandular
papillae dorsally in middle of lobe and 4 widely spaced terminal papillae.
Sexually mature males with sex products beginning in setiger 10-11 and
continuing to setiger 31 in one specimen; natatory setae on one male begin-
ning on setiger 11, with sperm in setiger 10 in same. Females with internal
eggs, 2 per segment, beginning mostly in setiger 10-11; eggs beginning in
largest female (7 mm long, with 44 setigers) by setiger 12 (possibly setiger
11), extending posteriorly to setiger 32; no females with external embryos
or natatory setae. Number of segments containing sexual products appar-
ently related to animal size with posterior 10-12 segments usually empty.

Pharynx long and narrow, with relatively thick walls (usually not strongly
bent or folded), mostly located in first 3 setigers and part of tentacular
segment; middorsal tooth anterior, relatively large. Proventriculus cylindri-
cal, located in most of length of setigers 4 and 5, with 13-14 irregular trans-
verse rows of muscle cells, anterior 4-5 rows small.

|

VOLUME 93, NUMBER 4 1127

Remarks.—I am not aware of descriptions of any Sphaerosyllis species
having parapodial glands containing spherical granules. In view of the con-
stancy of this character, I feel that the specimens represent an undescribed
species of this genus. The specimen from North Carolina reported as S.
pirifera by Day (1973) and Gardiner (1976) is S. glandulata. Note, however,
that parapodial glands are not readily observable except in cleared speci-
mens, and gland contents become coagulated in preserved specimens.

Etymology.—The specific name is derived from the Latin, glandulata,
meaning glanded, and refers to the parapodial glands.

Sphaerosyllis longicauda Webster and Benedict, 1887
Figs. 20, 21

Sphaerosyllis longicauda Webster and Benedict, 1887:720, pl. 3, figs. 35—
39:

Brania sp. Hartman, 1944:pl. 24, figs. 1, 2.

Sphaerosyllis erinaceus.—Pettibone, 1963:135, 136, fig. 35a [in part].—Gar-
diner, 1976:131, fig. 10s—v [not Claparéde, 1863].

Material examined.—MAINE: Eastport; syntypes on 3 slides (USNM
27557) [aicoholic syntypes in poor condition (USNM 400)]. NORTH CAR-
OLINA: Banks Channel, Wrightsville Beach, intertidal, on rocks; R. Fox,
col.; 11 specimens (USNM 52915). FLORIDA: Hutchinson Island Sta. II,
27°21.6’N, 80°13.2’W, about 11 m, coarse calcareous sand; 2 specimens
(FSBC I 20672, 20673). Sta. V, 27°22.9’N, 80°13.9'W, about 11 m, coarse
calcareous sand; 15 specimens (USNM 54525, 54526; FSBC I 20674-20681).

Description.—Body without color markings; integument covered with
sediment. Females of Florida specimens maximally 2.6 mm long, 0.2 mm
wide without parapodia, with 26 setigerous segments; males smaller, max-
imally 2.1 mm long, with 24 setigerous segments. Prostomium (Fig. 20a, b)
oval, about 3 times wider than long, broadly fused dorsally with tentacular
segment. Three pairs of lensed eyes; anterior eyes small, located in usual
position of eyespots, on anterior margin of prostomial lobe medial to lateral
antennae; posterior 2 pairs larger, in arc on posterior part of prostomial
lobe; arc either anteriorly concave or convex depending upon state of con-
traction of specimen; lateral pair of eyes larger. Lateral antennae originating
on anterolateral margins of prostomial lobe, with bulbous bases and cylin-
drical tips, often extending slightly past palps. Median antenna originating
from anterior emargination of tentacular fold covering posterior part of pro-
stomium, 14 times lengths of lateral antennae, occasionally extending far-
ther anteriorly. Palps short, dorsally fused, with wide, median, anterior
notch and dorsal, median furrow, without papillae. Dorsal lobes of brain
extending to anterior part of setiger 1; lateral lobes extending to about mid-
dle of same segment. Tentacular segment visible dorsally principally as fold

1128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

100 pm

Fig. 20. Sphaerosyllis longicauda: a, Anterior end, dorsal view (USNM 54525); b, Same,
relaxed specimen; c, Posterior end, dorsal view.

covering posterior half of prostomial lobe; tentacular cirri similar to lateral
antennae, originating below lateral eyes; few short papillae on tentacular
fold. Dorsum of each segment papillated with both long and short papillae
(Fig. 20a). Dorsal cirri absent from setiger 2, replaced by glandular papillae,
on remaining segments relatively slender, much longer than parapodial
lobes. Parapodial lobes stout, with blunt tips, with about 4 long, prominent

VOLUME 93, NUMBER 4 1129

b Cc d f
: “|

15 yum
Fig. 21. Sphaerosyllis longicauda: a, Superior simple seta, anterior setiger; b, Same, pos-
terior setiger; c, Upper compound seta, anterior setiger; d, Lower compound seta, setiger 10;
e, Upper compound seta, posterior setiger; f, Same; g, Inferior simple seta, posterior setiger;
h, Aciculum, anterior setiger; i, Same, posterior setiger.

glandular papillae; long, most prominent papilla on posterior tip; 1 each on
anterodorsal part of tip, on anterior margin about half distance from base
to tip, and on posterior margin near base. Ventral cirri short, cylindrical.
Solitary, superior simple setae (Fig. 21a, b) on all setigers, slightly curved
near tips, smooth on anterior segments, stouter and serrate on concave side
near tips on posterior segments. About 7 compound setae on anterior para-
podia (Fig. 21c, d), with unidentate blades; upper blades longest, 22 um,
with fine serrations from bases to near tips; lower blades about 12 um,
smooth. Five or 6 compound setae with similar, shorter blades, 12-15 um
long, on posterior parapodia (Fig. 21e, f); upper blades with fine serrations
or smooth; blades and shafts often stouter than on anterior segments. Distal
ends of shafts of compound setae with about 3 serrations. Solitary, small,
smooth, inferior simple setae (Fig. 21g) on posterior parapodia. Acicula
(Fig. 21h, 1) solitary, relatively slender, sharply anteriorly curved near tips,
with acute tips in anterior parapodia; much stouter in posterior parapodia
but similar in shape except tips not as pointed. Pygidium hemispherical,
with 2 rather long, stout, lateral anal cirri about twice length of dorsal cirri
and prominent, midventral projection or cirrus, with about 10 long papillae
visible dorsally. Sexually mature males with sexual products beginning in
setiger 8, extending to setigers 16-21. Females with 4 internal eggs or dorsal,
external embryos, beginning on setigers 8 or 9, extending to setigers 16-21;
single pair of embryos between parapodial lobes and dorsal cirri; second

1130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pair above dorsal cirri; one male with natatory setae beginning on seti-
ger 8.

Pharynx (Fig. 20a) strongly chitinized, basally surrounded by lightly col-
ored glands, located in tentacular segment and first 2 setigerous segments
of relaxed specimens, ending posteriorly in setiger 2 with anterior end ex-
tending past palps of contracted specimens, about 0.17 mm long. Middorsal
tooth large, about 4 distance from anterior end; anterior end surrounded
by stiff cylinder with about 10 papillae on margin. Proventriculus barrel
shaped, located in setigers 3-5 of relaxed specimens, extending anteriorly
into setiger 2 in contracted specimens. Muscle cells in dorsal view arranged
in 3 planes: prominent transverse plane; 2 opposite, diagonal planes; 18-21
transverse rows; anterior 5 small.

Remarks.—Sphaerosyllis longicauda has been referred to S. erinaceus
Claparede, 1863 (Pettibone, 1963). However, it differs from S. erinaceus in
that the middorsal pharyngeal tooth is not at the anterior end of the pharynx,
and the pygidium has a ventrally originating median anal cirrus in addition
to the lateral pair. My specimens are in reasonably good agreement with
types of S. longicauda; however, examination of critical details of the type-
specimens was not possible because of poor quality of the slides, and new
material from the type-locality will possibly aid in confirming this determi-
nation. Specimens reported as S. erinaceus by Gardiner (1976; USNM
52915) are S. longicauda.

Sphaerosyllis erinaceus Claparéde (1863:45, 46, pl. 8, fig. 38) is based on
a very short description of a juvenile with figures of the prostomium and
pygidium only; type material has apparently been lost. It was redescribed
by Saint-Joseph (1887:207, 208, pl. 10, figs. 81-83) based on a single specimen
from near the type-locality. Saint-Joseph’s description is in agreement with
Claparede’s short description and must be accepted if the species is to stand.
However, paired tentacular lobes, that Saint-Joseph did not describe, have
been described on specimens from other areas and referred to S. erinaceus
(S. longicauda.—Eliason, 1920:11-13, not Webster and Benedict, 1887;
Hartmann-Schroder, 1971:168, synonymy), and Saint-Joseph’s specimen
should be re-examined to determine the presence of the lobes.

Sphaerosyllis longicauda was originally reported from New England and
is now known from North Carolina and Florida.

Sphaerosyllis magnidentata, new species
Fig. 22

Material examined.—FLORIDA: Monroe County, John Pennekamp
State Park, South Creek channel marker No. 2, 3.8 m, in sand, shell, and
coral rubble with Thalassia, Penicillus, and Halimeda cover; R. J. Helbling,
col., 13 Nov. 1975; holotype (USNM 60452), 2 paratypes (USNM 60453;
FSBC I 23600). Same; col., 1979; 4 paratypes (USNM 60454; FSBC I 23601).
THE BAHAMAS: S portion of Bimini Lagoon, 25°43’N, 79°16’W, in sub-

VOLUME 93, NUMBER 4 1131

100 um

15 ym
Fig. 22. Sphaerosyllis magnidentata: a—c, Holotype: a, Anterior end, dorsal view; b, Mid-
dle segment, dorsal view; c, Posterior end, dorsal view; d-i, Paratype (USNM 60453): d,
Superior simple seta, setiger 4; e, Same, posterior setiger; f, Upper compound seta, setiger 4;
g, Lower compound seta of same; h, Inferior simple seta, posterior setiger; i, Aciculum, setiger

4; j, Same, posterior setiger.

1132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

merged plastic sponges; A. Schoener, col., 1970-1971; 1 paratype (USNM
60455).

Description.—Body without color markings. Holotype 1.8 mm long, 0.25
mm wide including parapodia across proventriculus, 20 setigers; paratypes
with 18—21 setigers. Prostomium and tentacular segment almost completely
fused dorsally (Fig. 22a), separated only by obscure groove. Prostomium
about 3 times wider than long with 2 pairs of relatively small, lensed eyes
on posterior half in flattened trapezoidal arrangement open to front, anterior
pair slightly larger. All three antennae similar, each about half as long as pro-
stomial width, constricted at base, with globular base and cylindrical tip, all
originating near anterior prostomial margin. Palps dorsally smooth, with
fused part almost as wide, longer than prostomium; anterior divided parts
usually turned ventrally. Dorsal lobes of brain extending to about middle of
setiger 1; ventrolateral lobes slightly shorter. Tentacular segment about as
long as, slightly wider than, and laterally surrounding posterior half of pro-
stomium; tentacular cirri originating below eyes, about % lengths of anten-
nae, similar to dorsal cirri of following segments. Six glandular papillae in
transverse row above tentacular cirri. Dorsal cirri absent from setiger 2,
constricted at bases on other anterior setigers, with bulbous bases and cy-
lindrical tips, with elongate, nearly cylindrical bases and narrower, cylin-
drical tips on middle and posterior setigers; all extending to about tips of
parapodial lobes. Dorsal parapodial glands beginning on setiger 5, possibly
also on setiger 3, extending to posterior end; contents granular; dorsal open-
ing not seen. Setal lobes relatively stout, blunt. Ventral cirri cylindrical,
slender, not extending to tips of setal lobes. Solitary, superior simple setae
(Fig. 22d, e) emerging above acicula on all parapodia, curved dorsally, ser-
rate on ventral border near tips, longer and stouter on posterior setigers.
About 7 compound setae on anterior parapodia, 4 on posterior parapodia
(Fig. 22f, g); blades with unidentate, hooked tips; edges serrate to near tips;
serrations relatively long, large, closely spaced proximally, shorter and
more slender distally; upper blades long on anterior parapodia, gradually
shorter below (43 and 17 um on setiger 4, slightly shorter anteriorly); blades
similar on posterior parapodia but upper blades shorter (about 30 um).
Shafts of compound setae with 2 or 3 rows of serrations subdistally. Solitary,
inferior simple setae on posterior parapodia (Fig. 22h), with hooked, uni-
dentate tips, serrate near tips on concave margins. Acicula (Fig. 22i, j)
solitary, stout, with tips bent forward at about right angle, slightly stouter
in posterior parapodia. Glandular papillae on dorsum and parapodia of se-
tigerous segments as figured on anterior and middle segments (Fig. 22a, b).
Pygidium subspherical, with dorsal transverse row of 8 large papillae, 4
similar, terminal papillae surrounding anus; anal cirri similar in shape to
dorsal cirri of posterior segments but longer and stouter. No sexually mature
specimens seen.

VOLUME 93, NUMBER 4 1133

Pharynx (Fig. 22a) about 140 um long, thick walled, surrounded by thick
glands, extending from anterior margin of setiger 1 to near posterior margin
of setiger 3; middorsal tooth anterior, massive; soft papillae surrounding
anterior end possibly absent. Proventriculus short, subcylindrical, about 120
wm long, 105 wm wide, extending to about middle of setiger 5 in slightly
less than 2 segments, with about 14 irregular, transverse rows of muscle
cells, anterior 4 rows small.

Remarks.—Sphaerosyllis magnidentata resembles S. ovigera Langerhans
(1879:567, 568, pl. 32, fig. 23; Fauvel, 1923:302—304, fig. 116a—d) in the origin
of the median antenna, but differs in having smooth palps, long- and short-
bladed compound setae, and parapodial glands containing granular material,
not rods as in S. ovigera.

In addition to differences indicated in the key, S. magnidentata differs
from other Sphaerosyllis species reported herein with dorsal parapodial
glands and acicula bent forward at about a right angle, in having a massive
middorsal pharyngeal tooth and upper compound setae with very long
blades.

Etymology.—tThe specific name is derived from the Latin magna, mean-
ing large, and dentata, meaning toothed, and refers to the large pharyngeal
tooth.

Sphaerosyllis piriferopsis, new species
Figs. 23, 24

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6’N,
80°13.2'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., May 1972; USNM 60210), 18 paratypes (USNM 16398; ZMH P-60211;
FSBC I 23602-23608). Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse
calcareous sand; 21 paratypes (AHF Poly 1316; ZMH P-16399; FSBC I
23609-23617). Sta. V, 27°22.9’N, 80°13.9’W, about 11 m, coarse calcareous
sand; 6 paratypes (FSBC I 23618-23622). Monroe County, John Pennekamp
State Park, South Creek channel marker No. 2, 3.8 m, in sand, shell, and
coral rubble with Thalassia, Penicillus, and Halimeda cover; R. J. Helbling,
col., 13 Nov. 1975; 1 paratype (FSBC I 23622). THE BAHAMAS: S portion
of Bimini Lagoon, 25°43’N, 79°16’W, in submerged plastic sponges; A.
Schoener, col., 1970-1971; 2 paratypes (USNM 54326).

Description.—Body without color markings; pharynx surrounded by
brownish glands; dark brown eye pigment often in dorsal lobes of brain,
extending posteriorly into setiger 1. Maximum length 2.3 mm, width 0.12
mm without parapodia, 30 setigers. Prostomium and tentacular segment
fused (Fig. 23a, b), with tentacular segment dorsally forming fold covering
posterior part of prostomium; median antenna often originating from ante-
rior margin of fold. Prostomium about twice as wide as long, with 2 pairs

1134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wit OO}

{OO ym

f

[eer ree eon
1OO pm

Fig. 23. Sphaerosyllis piriferopsis: a, Anterior end, dorsal view, pharyngeal tooth darkened
(AHF Poly 1316); b, Same, holotype; c, Middle segment, dorsal view; d, Same (ZMH P-16398);
e, Posterior end, dorsal view; f, Same, ventral view.

of lensed eyes on posterior half arranged in flattened trapezoid open to front;
posterior pair often indistinct, possibly absent, with eye pigment in dorsal
lobes of brain; anterior pair slightly larger. Lateral antennae attached on
anterolateral margins, similar in shape but slightly larger than median an-
tenna, usually not extending past palps. All antennae with bulbous bases
and cylindrical tips; length of tips about equal to that of bases. Dorsal lobes

VOLUME 93, NUMBER 4 [135

———

aeiwaibieven nd

My Piss. Nueee
15 um

Fig. 24. Sphaerosyllis piriferopsis: a, Superior simple seta, anterior setiger (AHF Poly
1316); b, Same, posterior setiger; c, Upper compound seta, anterior setiger; d, Same; e, Shaft
of same; f, Lower compound seta, posterior setiger; g, Inferior simple seta, posterior setiger;
h, Aciculum, anterior setiger; i, Same, posterior setiger.

of brain extending to about middle of setiger 1; lateral lobes slightly longer.
Palps together slightly wider and about as long as prostomium, tips usually
turned ventrally, large papillae or bumps on dorsal side and smaller papillae
on tips. Tentacular segment visible dorsally although fused with prostomi-
um, enclosing lateral sides of prostomium, with several glandular papillae
dorsally. Tentacular cirri usually originating ventrolateral to eyes or more
anteriorly on contracted specimens, about as stout but shorter than lateral
antennae. Dorsal cirri absent from setiger 2, replaced by glandular papillae.
Dorsal cirri on other setigers similar to antennae but not as stout; tips ex-
tending to or slightly past parapodial tips. Parapodial lobes stout, bluntly
rounded. Ventral cirri slender, cylindrical, originating near parapodial bas-
es, not extending past tips. Small, spherical, glandular papillae on dorsum
and parapodia of median segments (Fig. 23c, d). Solitary superior simple
setae above acicula on all parapodia, serrate on edge near tips, slender on
anterior segments, stouter on posterior segments (Fig. 24a, b). About 5
compound setae on anterior parapodia (Fig. 24c—e), about 3 on posterior
parapodia; upper blades on anterior parapodia about twice as long (17 wm)
as lower ones, with long, thin serrations from bases to near tips; lower
blades smooth or nearly so; tips unidentate; blades of compound setae on
posterior parapodia all about same length as lower ones of anterior para-
podia but stouter, more strongly hooked (Fig. 24f). Shafts of compound
setae similar throughout body but stouter posteriorly, with few serrations
on distal ends. Solitary, inferior simple setae (Fig. 24g) on posterior para-
podia, about as stout as corresponding superior simple setae but more

1136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

strongly curved. Acicula (Fig. 24h, 1) solitary, relatively stout, with tips bent
anteriorly at right angle; posterior acicula only slightly stouter than anterior.
Dorsal parapodial glands always absent. Pygidial lobe (Fig. 23e, f) triangular
in outline, with pair of anal cirri similar but longer and thicker than dorsal
cirri, several relatively long, glandular papillae dorsally and ventrally be-
tween anal cirri and 4 terminal papillae. Sexually mature females with 2
external or internal embryos per segment; external embryos with 4-6 seti-
gers each, attached ventrally by pygidium near parapodial lobes beginning
on setiger 14 in some females, but on setiger 13 in one, continuing on 10
segments of one female; short natatory setae on 10 segments of one female,
beginning on setiger 13. Males with sperm in setigers 11-24 and natatory
setae on setigers 12—24; sperm in setiger 11 apparently originating in setiger 12.

Pharynx (Fig. 23a, b) long, narrow, thin walled, often bent from contrac-
tion in preserved specimens, usually in first 3 setigers, surrounded by prom-
inent glands; middorsal tooth relatively large, anterior. Proventriculus short-
er than pharynx, cylindrical, usually in setigers 4 and 5; 13-14 irregular,
transverse rows of muscle cells, anterior 4—5 small.

Remarks.—Sphaerosyllis piriferopsis shows similarity to S. pirifera Cla-
parede (1868:515, 516, pl. 14, fig. 2; Viguier, 1884:96-99, pl. 4, fig. 38, pl.
5, figs. 39-43; Fauvel, 1923:301, 302, fig. 115 1—p). However, the prostomium
and tentacular segment of S. piriferopsis are not completely fused; yellowish
‘‘glands’’ in setiger 1 are absent; glandular papillae are numerous but small,
and thus the body is not always encrusted with sand; compound setae are
fewer in number in each parapodium (3-5 vs. 6-8) and have shorter blades
(maximum length 50 um in S. pirifera according to Claparéde’s apparently
incorrectly scaled figure, and 23 wm according to Viguier’s figure vs. 17 wm
in §. piriferopsis); and sexual products and natatory setae begin in more
posterior segments of S. piriferopsis. All these differences appear to be
reliable taxonomic characters. Yellowish ‘‘glands’’ of S. pirifera may be
homologous with ‘‘lateral lobes of the brain’’ reported in S. piriferopsis.
Differences between Claparede’s and Viguier’s figures of S. pirifera, other
than length of blades of compound setae, are the presence or absence of
dorsal cirri on setiger 2 and the degree of fusion of the prostomium and
tentacular segment.

Further, several authors have stated their opinion that S. hystrix Cla-
paréde, 1863, and S. pirifera are synonymous, and that presence or absence
of parapodial glands containing rods from setiger 4 posteriad is an unreliable
specific character. This is absolutely contrary to my experience, and in my
opinion, S. pirifera and S. hystrix cannot be considered synonyms.

Etymology.—The specific name, derived from the previously described
species, pirifera, and the Greek suffix, -opsis, meaning likeness, refers to
the similarity between the two species.

VOLUME 93, NUMBER 4 1137

Sphaerosyllis riseri, new species
Fig. 25

Material examined. —FLORIDA: Monroe County, John Pennekamp
State Park, South Creek channel marker No. 2, 3.8 m, in sand, shell and
coral rubble, with Thalassia, Penicillus and Halimeda cover; R. J. Hel-
bling, col., 10 May 1976; holotype (USNM 60471). Hutchinson Island Sta.
V, 27°22.9’N, 80°13.9’W, about 11 m, coarse calcareous sand; 2 paratypes
(USNM 54451; FSBC I 23626). Sta. II, 27°21.6’N, 80°13.2'W, about 11 m,
coarse calcareous sand; 2 paratypes (USNM 54452). Sta. IV, 27°20.7’N,
80°12.8’W, about 11 m, coarse calcareous sand; 3 paratypes (USNM 54450;
FSBC I 23624, 23625).

Description.—Body without color markings; pharynx surrounded by
brownish glands; lateral lobes of brain yellowish brown; sperm brown; ova
or embryos light yellow. Minute; maximum length 1.5 mm, width 0.14 mm
without parapodia, 17 setigers. Prostomium fused dorsally with tentacular
segment (Fig. 25a, b); tentacular segment forming thin fold over posterior
part of prostomial lobe covering posterior pair of eyes, with median antenna
originating from anterior part of fold between posterior 2 pairs of eyes on
relaxed specimens or slightly in front of fold on contracted specimens. Three
pairs of eyes; pair of anterior eyespots adjacent to origins of lateral anten-
nae; 2 pairs of lensed eyes on posterior half above origins of tentacular cirri,
very close together laterally. Lateral antennae originating on anterior margin
of prostomium on line slightly medial to posterior 2 pairs of eyes, closer to
midline than lateral margins; antennae all of similar size, with bulbous bases
and cylindrical tips; tips contracted into bases and distally ciliated. Dorsal
lobes of brain wide, extending to posterior part of setiger 1; lateral lobes
slightly longer. Palps short, each broader than long, papillate dorsally. Ten-
tacular segment partially surrounding lateral margins of prostomial lobe;
tentacular cirri similar in shape but smaller than antennae and dorsal cirri,
Originating on anterior part of lateral extensions. Four glandular papillae per
segment on dorsum in 2 longitudinal rows plus 2 per segment occurring
singly in front of dorsal cirri (Fig. 25c); 12 on ventrum in 3 transverse rows
of 4 each. Dorsal cirri on all setigerous segments, similar to antennae but
slightly smaller on anterior setigers, slightly longer posteriorly. Parapodial
lobes short, pointed (Fig. 25a, b, e), triangular in dorsal view. Body of
sexually modified regions of females often narrower, resulting from extru-
sion of eggs from body cavity, with parapodial lobes appearing much longer
and more acutely pointed. Parapodia with anterior and posterior papillae
subdistally (Fig. 25c) with posterior one closer to tips. Ventral cirri shorter
than parapodial lobes, digitiform. Solitary, superior simple setae on all para-
podia (Fig. 25g), smooth on anterior segments, lightly serrate near tips
posteriorly. Five compound setae on anterior parapodia, 4 on posterior para-

1138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

podia; all blades 7-8 um long, unidentate, often with fine serrations on
blade edge of upper ones (Fig. 25f); blades slightly longer on first 2-3 seti-
gers. Solitary, curved, inferior simple setae on posterior parapodia (Fig. 25h).
Acicula (Fig. 25i) solitary, slender, slightly enlarged near tips; tips long,
thin. Pygidium with pair of anal cirri about 3 times longer than dorsal cirri
(Fig. 25d). Sexually mature specimens including 3 males and 2 females;
males with brownish sperm in setigers 7-16; females with 2 spherical em-
bryos per segment in setigers 7-13; embryos carried between parapodial
lobes and dorsal cirri when external (Fig. 25c). Natatory setae not seen.
Pharynx (Fig. 25a, b) in anterior 2 setigers of relaxed specimens, extend-
ing only to middle of setiger 2 in contracted specimens, thin walled, wide,
vase-shaped, with anterior opening narrower than widest part or anterior
end often flared when everted; middorsal tooth minute, originating about
0.6 of distance from posterior to anterior margins. Ratio of pharynx to pro-
ventriculus lengths 1:1.1. Pharynx often turned slightly ventrally, appearing
shorter in contracted specimens. Proventriculus barrel shaped, in total
length of 3 segments, middle of setiger 1 to setiger 5; muscle cells arranged
in 2 opposite diagonal planes in addition to more prominent transverse
plane; 17-18 transverse rows of muscle cells, anterior 5—6 rows small.
Remarks.—Sphaerosyllis riseri is very similar to S. tetralix Eliason
(1920: 13-15, fig. 4; Hartmann-Schroder, 1971:165—167, fig. 54a—c). S. riseri
is maximally 1.5 mm long with 17 setigers, while S. tetralix is maximally 4
mm long with 29 segments. Antennae and cirri of S. riseri have distal parts
ciliated on the tips and strongly contracted into the bases, while on S.
tetralix they not distally ciliated and are flask shaped, as typical of other
members of the genus, on anterior segments and cirriform on posterior
segments. Antennae of S. riseri are all similar, while the median antenna of
S. tetralix is larger than the lateral two. Eggs and sperm first begin in setiger
7 of mature specimens of S. riseri and in setiger 8 of S. tetralix, and S.
riseri has 2 eggs per segment vs. 4 for S. tetralix. The proventriculus of S.
riseri has about 18 rows of muscle cells while that of S. tetralix has about
20 rows. Papillation of the dorsum is identical for the 2 species; S. riseri
has about 12 small papillae per segment on the ventral side in regular ar-
rangement, while S. tetralix was reported to have diffusely arranged papillae
ventrally. Diffusely arranged papillae reported on the ventrum of S. tetralix

=>

Fig. 25. Sphaerosyllis riseri: a, Anterior end, dorsal view, pharyngeal tooth darkened
(USNM 54450); b, Same (USNM 54451); c, Middle segment, dorsal view (USNM 54450); d,
Posterior end, dorsal view (USNM 54451); e, Parapodium, ventral view (USNM 54452); f,
Compound setae, setiger 7; g, Superior simple seta; h, Inferior simple seta, posterior segment;
i, Acicula, dorsal and lateral views.

1139

VOLUME 93, NUMBER 4

100 ym

thi

vu |) \
ees

"yg

Ai WN

f = ii

15 ym

1140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

indicates an irregular arrangement, but the arrangement is probably regular
with numerous papillae and therefore difficult to determine.

Etymology.—The species is named in honor of Dr. Nathan W. Riser, who
assisted me in describing this species and others.

Sphaerosyllis taylori, new species
Fig. 26

Sphaerosyllis brevifrons.—Webster and Benedict, 1884:714, 715 [in part,
specimens from South Norwalk, Connecticut].

Sphaerosyllis hystrix.—Pettibone, 1963:136, 137, fig. 35g [not Claparéde,
1863].

Material examined.—FLORIDA: Hutchinson Island Sta. V, 27°22.9’N,
80°13.9'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Sep. 1972; USNM 60212), 33 paratypes (USNM 54522, 54524, 60213;
AHF Poly 1317; ZMH P-16400; FSBC I 23627-23637). Sta. II, 27°21.6’N,
80°13.2'W, about 11 m, coarse calcareous sand; 7 paratypes (USNM 54523;
FSBC I 23638-23640). Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse
calcareous sand; 10 paratypes (AHF Poly 1318; ZMH P-16401; FSBC I
23641-23645). Tampa Bay, 27°34'54’"N, 82°43'01”W, 7 m, sand; J. Taylor
and C. Saloman, cols., 10 Oct. 1963; 3 paratypes (USNM 60214; FSBC I
23646). CONNECTICUT: South Norwalk; paratypes (USNM 27556; spec-
imens on 3 slides originally identified by Webster as S. brevifrons Webster
and Benedict). MARYLAND: Chincoteague Bay in Zostera beds; R. Orth,
col., 29 Nov. 1970; 3 paratypes (USNM 44084).

Description.—Body without color markings; brownish glands surrounding
pharynx. Maximum length 2.4 mm, width 0.2 mm without parapodia, 24
setigers. Prostomium and tentacular segment almost completely fused dor-
sally (Fig. 26a, b). Two pairs of lensed eyes on posterior half, very close
together laterally, in nearly rectangular arrangement, anterior pair slightly
larger. Lateral antennae originating on anterolateral margin of prostomium
in front of eyes, with bulbous bases and cylindrical tips; lengths slightly less
than distance between right and left eyes. Median antenna originating be-
tween posterior pair of eyes of relaxed specimens, often attached to anterior

=

Fig. 26. Sphaerosyllis taylori: a, Anterior end, dorsal view (USNM 54524); b, Same, con-
tracted specimen (USNM 54522); c, Parapodium, middle segment, dorsal view (USNM 54524);
d, Posterior end of holotype, dorsal view; e, Superior simple seta, anterior setiger; f, Same,
posterior setiger; g, Upper compound seta, anterior setiger; h, Compound setae, posterior
setiger; i, Inferior simple seta, posterior setiger; j, Aciculum, anterior setiger; k, Same, pos-
terior setiger.

1141

VOLUME 93, NUMBER 4

1142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

part of tentacular fold on contracted specimens; antennae similar in length
or median slightly shorter. Dorsal and lateral lobes of brain extending to
anterior part of setiger 2 of relaxed specimens, extending only into setiger
1 of contracted specimens. Palps together about as wide as prostomium,
exceeding length of lateral antennae by about half in extended specimens,
with tips mostly bent ventrally. Tentacular segment completely enclosing
lateral margins of prostomium, with few, glandular papillae below tentacular
cirri and on dorsum; tentacular cirri originating on about same transverse
line as lateral antennae, similar to dorsal cirri, about as thick as antennae
but shorter. Dorsum of each segment with about 20 long, glandular papillae
in characteristic arrangement (Fig. 26c). Dorsal cirri absent from setiger 2,
replaced by glandular papillae; dorsal cirri of other setigers with bulbous
bases and cylindrical tips often extending past tips of parapodial lobes.
Parapodial lobes stout, blunt; each with 2 apical papillae, anterior one slight-
ly below tip on ventral side; posterior one nearer to tip, nearly dorsal in
position, 2 other papillae anteriorly and posteriorly below dorsal cirrus (Fig.
26c). Ventral cirri cylindrical, not extending past tips of parapodial lobes.
Solitary, smooth, slightly curved, superior simple setae on all parapodia
(Fig. 26e, f), slender anteriorly, much stouter posteriorly. Blades of com-
pound setae (Fig. 26g, h) all similar in size, 10-8 wm long dorsally to ven-
trally on anterior parapodia, similar in shape except upper ones of anterior
parapodia with about 6 long, fine serrations on edge; all unidentate. Shafts
of compound setae mostly stouter on posterior segments. Solitary, inferior
simple setae (Fig. 261) on posterior parapodia, slightly slenderer than cor-
responding superior simple setae. Acicula (Fig. 26), k) solitary, bent forward
near tips at about right angle; tips pointed; relatively slender in anterior
segments, about twice as stout in posterior segments. Spherical to oblate
parapodial glands containing needle-like rods; glands always above para-
podial lobes medial to dorsal cirri beginning on setiger 4, with dorsal opening
slightly in front of and medial to origins of cirri. Pygidium (Fig. 26d) short,
bluntly rounded, with pair of anal cirri similar in shape but about twice as
large as posterior dorsal cirri, with about 6 glandular papillae on dorsal side.
Sexually mature males with sex products beginning in setiger 6 or 7, ex-
tending to setiger 20, but usually to about setiger 15. Natatory setae on
some males beginning on segment following that with sex products, usually
setiger 7. Females with internal ova beginning in setiger 7 or 8, extending
to about setiger 15; none with externally attached embryos or natatory setae.
Lengths of pharynx and proventriculus (Fig. 26a, b) about equal. Pharynx
in setigers 1 and 2, surrounded by thick glands, often lobed about middle;
middorsal tooth anterior. Proventriculus cylindrical, not much longer than
wide, usually, if not always, within setigers 3 and 4, with 13-14 very irreg-
ular, transverse muscle cell rows, anterior 4—5 rows small.
Remarks.—Sphaerosyllis taylori differs from European descriptions of S.

VOLUME 93, NUMBER 4 1143

hystrix Claparede (1863:45, pl. 13, figs. 36, 37; Fauvel, 1923:301, fig. 115g,
h; not i, k; Hartmann-Schroder and Stripp, 1968:13, 14, fig. 6a—c; Hart-
mann-Schroder, 1971:67, fig. 54d-f; 1974a:197) in having a pharynx and
proventriculus of nearly equal length, compound setae with shorter blades
with fewer serrations and of similar length in each parapodium and through-
out the body, smooth, superior simple setae on all parapodia, and in being
smaller with fewer segments.

Claparéde’s figure (1863:fig. 36) of the anterior end of S. hystrix shows
the pharynx in almost 3 segments and the proventriculus in setigers 3 and
4. Fauvel’s figure (1923:fig. 115g), attributed to Claparede, shows the phar-
ynx occupying three segments and the proventriculus about two segments
long mostly in setigers 5 and 6. Figures of Hartmann-Schroder and Stripp
(1968) and Hartmann-Schroder (1971) show compound setae to be larger
with many more serrations than those of S. taylori, and superior simple
setae with a denticulate ventral margin. Claparede’s figure (1863:fig. 36)
shows only compound setae on anterior parapodia; Fauvel’s figure (1923:fig.
115h), attributed to Claparéde, shows only compound setae on a sexually
modified parapodium. Fauvel (1923) and Hartmann-Schroder (1971) stated
that superior simple setae were present on middle and posterior segments;
Hartmann-Schroder (1971) stated that blades of compound setae were mod-
erately long to short.

There are obvious problems with other European descriptions of S. hys-
trix. Saint-Joseph (1887:204—207) reported 2 larval forms with 2 modes of
ventral incubation of embryos, and embryos first occurring on different
segments on specimens from Dinard, France. The description of McIntosh
(1908: 156-159, pl. 59, figs. 3, 4, 8; pl. 70, fig. 1; 1910:pl. 79, figs. 11-13)
included specimens with both dorsal and ventral incubation of embryos, and
is so confused that it can be ignored. Southern (1914:19, 20) reported females
from Ireland with both 2 and 4 eggs per segment in setigers 8-19 and males
with natatory setae beginning on setigers 10 or 11. It appears certain that
the accounts of Saint-Joseph, McIntosh and Southern each referred to more
than one species, and their specimens should be reexamined.

Etymology.—It is a pleasure to name this species in honor of Dr. John L.
Taylor, who assisted in the Hutchinson Island study, and has been of much
help to me by providing specimens and advice.

Streptosyllis Webster and Benedict, 1884
Streptosyllis pettiboneae, new species
Figs. 27, 28

Material examined.—FLORIDA: Hutchinson Island Sta. III, 27°22.0’N,
80°12.4’W, about 7 m, medium calcareous sand; holotype (R. Gallagher,
col., Sep. 1971; USNM 60448), 7 paratypes (USNM 60450; AHF Poly 1319;

1144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 27. Streptosyllis pettiboneae: a, Anterior end, dorsal view; b, Same; c, Anterior end,
ventral view; d-f, Parapodia (FSBC I 23648): d, Setiger 1, anterior view; e, Setiger 3, posterior
view; f, Setiger 20, anterior view.

VOLUME 93, NUMBER 4 1145

ieee

= a

Fi wit c cer ny me
1S um

Fig. 28. Streptosyllis pettiboneae: a, Superior simple seta, anterior setiger; b, Same, middle
setiger, lateral view; c, Same, ventral view; d, Upper compound seta, anterior setiger; e, Lower
compound seta, anterior setiger; f-i, Compound setae, middle segments: f, Lateral view (of
blade); g, Dorsal view (of blade); h, Lateroventral view of blade; i, Shaft.

ZMH P-16402; FSBC I 23647-23649). Sta. II, 27°21.6’N, 80°13.2’W, about
11 m, coarse calcareous sand; 1 paratype (USNM 60449). Sta. IV,
27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 2 paratypes
(FSBC I 23650, 23651). Tampa Bay, 27°36'56"N, 82°41’05”W, 9 m, sand; J.
Taylor and C. Saloman, cols., 25 Oct. 1963; 3 paratypes (USNM 60451).

Description.—Pharynx uncolored to reddish brown or brown. Largest
specimen slightly greater than 4.0 mm long, width 0.9 mm without parapodia
across proventriculus, 48 setigers. Prostomium (Fig. 27a, b) pentagonal;
width less than twice length; posterior border slightly convex. Mature spec-
imens with 3 pairs of lensed eyes; anterior pair small, lateroposterior to
origins of lateral antennae; 2 posterior pairs larger, arranged in flattened
trapezoid open to front; interior pair on posterior margin. Antennae smooth;
median antenna originating between posterior 2 pairs of eyes; origins of
lateral antennae near anterior margin of prostomium at middle of each side,
anteromedial to anterior pair of eyespots; shapes and lengths of antennae
variable; shape cylindrical to clavate, with median usually longer than pro-
stomial width, longer and stouter than lateral antennae. Palps (Fig. 27b, c)
composed of 2 distinct parts, pair of cushion-shaped ventral lobes and pair
of filiform lobes emanating from middle of cushion-shaped lobes; cushion-
shaped lobes rarely visible from dorsum; filiform lobes relatively long, often
observable below anterior pair of eyes when pharynx everted. Rows of cilia
on prostomium in front of anterior pair of larger eyes and on posterolateral

1146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

margins. Prostomial integument light brown, appearing glandular. Tentac-
ular segment with same integumental consistency as prostomium; length
depending on state of eversion of pharynx and state of contraction of spec-
imen, short when pharynx everted, always visible dorsally. Tentacular cirri
not articulate, directed slightly anterolaterally; dorsal tentacular cirri from
slightly longer to almost twice as long as ventral tentacular cirri. Dorsal cirri
(Fig. 27a) of anterior 5 setigers not articulate, similar to dorsal tentacular
cirri and lateral antennae, with short cirrophores; becoming slightly longer,
almost as long as body width after setiger 5; articulate on setiger 6; smooth
or only wrinkled on setigers 7 and 8; thereafter alternately articulate and
smooth or only wrinkled to posterior end; articulate dorsal cirri with 9-10
articles; each article containing 2 large cells of granular material; smooth
dorsal cirri long, clavate to cylindrical, diameter slightly less than, and
lengths equal to articulated dorsal cirri. Parapodial lobes (Fig. 27d-f) blunt,
much stouter on setigers 2—S than on other setigers. Ventral cirri attached
near lower compound setae, about as long as parapodial lobes, slightly con-
stricted at bases, with stout bases and long, slender tips. Solitary, superior
simple setae on all parapodia (Fig. 28a—c) short, stout, without hyaline
hoods, abruptly bent near tips, serrate on ventral edge near tips on setigers
1-5; after setiger 5 longer, more slender, serrate on ventral borders near
tips, with short, distally bifid, hyaline hoods. Compound setae of first 5
setigers (Fig. 28d, e) including 1-3 upper, long-bladed falcigers, 10-12 lower,
stout, short-bladed falcigers; long blades serrate on edge, with slender, bi-
dentate tips; shafts of long-bladed falcigers with longest part smooth, short-
est part trilobed; short falcigerous blades stout, bidentate, with 2 serrations
below secondary tooth; shafts of short-bladed falcigers similar to those of
long-bladed ones but stouter. Compound setae behind setiger 5 (Fig. 28f-i)
relatively long falcigers; blades flattened, serrate on edge; tips unequally
bifid in plane perpendicular to narrow axis of blades; bifid tips in ventral
view appearing hooded or encapsulated; shafts distally with 1 long and 3
short, pointed lobes. Inferior simple setae absent. Acicula solitary, greatly
enlarged and emergent in setigers 2—S, with nobbed tips. Patches of cilia
medial to dorsal and ventral cirri, apparently also continuing onto dorsum
and ventrum of each segment. Pygidium triangular in outline, with short,
ventrally originating median cirrus; one specimen with single long, slightly
thicker lateral cirrus about as long as last 8 segments. Sexually mature
specimens with sexual products beginning in setigers 13-14, extending to
near posterior end, often ‘‘spilling over’’ from originating segments into
adjacent anterior or posterior segments, appearing similar in all specimens.
Natatory setae, when present, either long or short (probably indicating fe-
males and males), usually beginning on segment after appearance of internal
sexual products, usually on all segments containing sexual products. Soli-
tary, slender notoacicula in parapodia with natatory setae.

VOLUME 93, NUMBER 4 1147

Pharynx (Fig. 27a, b) broad, much shorter than proventriculus, often bent,
extending from mouth to anterior part of setiger 3 when inverted, extending
to about anterior part of setiger 1 when everted; 10 soft papillae around
anterior end; chitinous part often everted past soft papillae by as much as
half total length; middorsal tooth absent. Proventriculus long, barrel-shaped,
in 6 segments, setigers 3-8 or 2—7, with about 55 rows of muscle cells.

Remarks.—Streptosyllis pettiboneae is similar to S. websteri Southern
(1914:26-28, pl. 2, fig. 3) and differs in the following characters: longer
compound setae of setigers 1-5 have bidentate blades; shafts of compound
setae behind setiger 5 are not serrate near the tips; blades of compound
setae after setiger 5 appear unidentate, but tips are covered by hyaline hoods
which are distally bifid; hoods of superior simple setae are concave on tips;
some dorsal cirri behind setiger 5, in alternate arrangement, are distinctly
articulate; and sexual modification first appears on a more posterior seg-
ment, setigers 13-14 vs. 11 for S. websteri.

Etymology.—The species is named in honor of Dr. Marian H. Pettibone
in an attempt to express my sincere appreciation for all the help she has
provided me during the last several years.

Syllides Orsted, 1845
Syllides bansei, new species
Figs. 29, 30

Material examined.—FLORIDA: Hutchinson Island Sta. IV, 27°20.7'N,
80°12.8’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Nov. 1972; USNM 60440), 9 paratypes (AHF Poly 1320; ZMH P-16403;
FSBC I 23653-23659). Sta. II, 27°21.6'N, 80°13.2’W, about 11 m, coarse
calcareous sand; 3 paratypes (AHF Poly 1321; ZMH P-16404; FSBC I
23660). Sta. V, 27°22.9'N, 80°13.9’W, about 11 m, coarse calcareous sand;
1 paratype (AHF Poly 1322). Tarpon Springs, Anclote Anchorage,
28°12.6'N, 82°47.4’W, 2.5 m, sand with Syringodium cover; J. Studt and R.
Ernest, cols., 17 Jan. 1975; 12 paratypes (USNM 60441). Same, 28°12.6'N,
82°46.7'W, 1.5 m, sand with Halodule cover; 17 Jan. 1975; 1 paratype
(FSBC I 23661). Same, 28°12.5'N, 82°46.5’W, 0.5 m, sand; 5 Dec. 1975; 1
paratype (FSBC I 23662).

Description.—Interior body (sexual?) contents brown after first several
segments; pharynx surrounded by brown glands. Maximum length 2.5 mm,
30 setigerous segments. Prostomium (Fig. 29a—d) about 2% times wider than
long; anterior margin projecting as obtuse angle; lateral margins rounded;
posterior margin straight or concave. Two pairs of large, lensed eyes on
posterior half; anterior pair larger, on lateral margins; smaller pair near
posterior margin in about middle of each side; pair of eyespots almost al-
ways on anterior margin on line between 2 larger pairs, slightly lateral to

1148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Og ee

200 ym

Fig. 29. Syllides bansei: a, Anterior end, dorsal view; b, Same; c, Anterior end, ventral
view; d, Anterior end, dorsal view; e, Parapodium, setiger 2, anterior view; f, Same, middle
setiger, posterior view.

origins of lateral antennae. Lateral antennae originating on anterior margin
at about middle of each side, about as long as prostomium plus palps; me-
dian antenna originating between posterior pair of eyes, 112 to 2 times longer
than lateral antennae. Antennae, tentacular cirri, and dorsal cirri of first 2

VOLUME 93, NUMBER 4 1149

=a

2)

Aan

ae,

A.

f
:
j
a b g h
i

20 ym
Fig. 30. Syllides bansei: a, Superior simple seta, anterior setiger; b, Same, middle setiger;
c, Tip of same, dorsal view; d-i, Compound setae: d, Upper, anterior setiger; e, Middle of
same; f, Lower of same; g, Upper, middle setiger; h, Middle of same; i, Lower of same; j,
Aciculum, anterior setiger.

setigerous segments smooth or only wrinkled. Palps in dorsal view not ap-
pearing medially fused; each subtriangular, slightly shorter than length of
prostomium; anterior edges often ventrally turned; lateral edges always ven-
trally turned; pair of small digitiform appendages ventroposteriorly (Fig.
29c). Tentacular segment shorter than following segment, definitely sepa-
rated by grooves from prostomium and setiger 1. Dorsal tentacular cirri
similar to median antenna and dorsal cirri of setigers 1 and 2; ventral ten-
tacular cirri similar to lateral antennae. Dorsal cirri after first 2 setigers
articulate, maximally with 20 articles; basal parts smooth or pseudoarticu-
lated; articles composed of 2 oval cells containing material of mostly uniform
texture and grey-green color. Parapodial lobes (Fig. 29e, f) well extended
from body, almost half as long as body width; tips rounded in dorsal view,
obliquely truncate laterally at greater or lesser angle to ventrum. Ventral
cirri Originating from about middle of lower margins of parapodial lobes;
anterior 5 pairs clavate, short, stouter than following; more posterior ventral
cirri longer, more filiform. Solitary, superior simple setae on all setigers

1150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Fig. 30a, b), slightly curved toward dorsum, lightly serrate on convex edge
for about half emergent length; tips covered by slightly expanded, evenly
rounded, hyaline hoods; setae shorter, stouter than following, abruptly ta-
pering in width only near tips on first 5 setigers, gradually tapering toward
tips more posteriorly. Compound setae similar in shape and number in all
but last few parapodia, 7—9 in anterior parapodia, 5—7 in posterior parapodia,
longer on middle parapodia than on setigers 1-5. Blades of compound setae
26-65 um long, of 3 basic types (Fig. 30d-i), all bidentate; upper blades
usually solitary, often absent, long, lightly serrate on basal half; middle
blades, 2-3, intermediate in length, with single, basal spur, otherwise
smooth (very fine serrations may be present above spur); lower blades, 3—
5, similar to upper blades but shorter than upper and middle types. Shafts
of compound setae similar throughout body; distal end serrate in side view,
pointed; short part of hinge broader than remainder of shaft in plane view.
Inferior simple setae absent. Acicula (Fig. 30j) (neuroacicula) solitary, sim-
ilar in size and shape throughout body, relatively slender, gradually nar-
rowed to slightly enlarged tips; tips truncate or slightly concave, appearing
ragged. Slender notoacicula in sexually modified segments of adults begin-
ning in parapodia of setigers 8 or 9, extending to 2-3 segments from pygidi-
um. Pygidium with paired lateral anal cirri apparently lost; several speci-
mens with slender, short median anal cirrus. Single sexually mature
specimen with 3-5 very short natatory setae on parapodia of setigers 11-
29; no specimens with internal eggs or external embryos; sexual products
in segments with notoacicula.

Ratio of pharynx to proventriculus lengths about 2:3; pharynx (Fig. 29a,
b, d) thickly chitinized, slightly less than twice long as wide, without mid-
dorsal tooth, surrounded anteriorly by about 10 semicircular, soft papillae
proximal to smooth anterior end of chitinous part when completely everted.
Proventriculus barrel-shaped, about 12 times longer than wide, occupying
3-S5Y2 segments depending on size of specimen and state of eversion of
pharynx, usually ending near posterior part of setiger 5, with 35 transverse
rows of muscle cells, also arranged in 2 opposite, diagonal planes (Fig. 26a).

Remarks.—The genus Syllides Orsted, 1845, was recently reviewed by
Banse (1971). S. bansei is similar to S. benedicti Banse (1971:1478, 1479,
fig. 6), differing in the following characters: smaller size (2.5 mm vs. 6.5
mm); smaller maximum number of segments (30 vs. 70); smaller length to
width ratio of proventriculus (1.5:1 vs. 3:1); more anterior position of pro-
ventriculus (ending at posterior part of setiger 5 vs. about setiger 10 for S.
benedicti); shape of tips of superior simple setae; fine detail of blades of
upper and lower compound setae (finely toothed in S. bansei vs. smooth);
and possibly in shape of tips of shafts of compound setae.

Etymology.—The species is named in honor of Dr. Karl Banse, whose
review of the genus facilitated the description of this species.

VOLUME 93, NUMBER 4 1151

Syllides floridanus, new species
Figs. 31, 32; Table 1

Material examined.—FLORIDA: Hutchinson Island Sta. V, 27°22.9’N,
80°13.9’'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Nov. 1971; USNM 60438), 2 paratypes (USNM 60439). Sta. II,
27°21.6'N, 80°13.2'W, about 11 m, coarse calcareous sand; 1 paratype
(FSBC I 23663).

Description.—Articulate dorsal cirri brownish yellow; middle and poste-
rior segments often dark yellow, possibly from sexual products. Dimen-
sions: see Table 1. Prostomium (Fig. 31a, b) about twice as wide as long,
pentagonal, anterior margin forming obtuse angle, lateral margins almost
Straight, posterior margin straight or convex. Three pairs of lensed eyes;
posterior 2 pairs on posterior half of prostomial lobe, in trapezoidal arrange-
ment, with posterior ones about equidistant between origin of median an-
tenna and lateral pair of eyes; lateral pair near middle of lateral margins;
anterior pair small, on anterior margin on line between 2 posterior pairs,
often damaged. Median antenna originating between lateral pair of eyes, all
missing. Lateral antennae originating between median antenna and anterior
pair of eyes, slightly nearer to anterior eyes than median antenna; length
about 0.18 mm, equal to prostomial width or slightly less. Palps (Fig. 3la—
c) short, triangular, not appearing medially fused; pair of digitiform ap-
pendages ventrally on lateroposterior parts. Lateroanterior prostomial mar-
gins ciliated. Texture of dorsal surface of prostomium glandular. Tentacular
segment slightly shorter than following segment; dorsal tentacular cirri
about twice longer than lateral antennae (0.24:0.13 mm); ventral tentacular
cirri shorter than dorsal; texture of integument glandular, same as that of
prostomium. Dorsal cirri of setigers 1 and 2 smooth or only wrinkled; long
on setiger | (0.5 mm), much shorter (0.3 mm) on setiger 2. Articulated dorsal
cirri beginning with setiger 3 (Fig. 31b), continuing to posterior end, with
maximum of 20 articles; articles distinct from proximal parts, each com-
posed of 2 cells containing numerous particles showing granular appearance.
Parapodial lobes (Fig. 31d, e) long, cylindrical; dorsal anterior lobe on end;
rows of cilia above and below apparently also extending across dorsum of
each segment. Ventral cirri originating from about middle of ventral border
of parapodia, extending laterally to parapodial tips, subulate. Solitary, su-
perior simple setae (Fig. 32a, b) on all parapodia, long, slender, fine tipped,
distally serrate, basally rounded; serrate parts flattened; tips slightly
hooked, capitate. Compound setae (Fig. 32c-i) long falcigers, 9-10 in each
parapodium; blades bidentate, distal teeth similar in size, primary tooth
strongly hooked, secondary tooth pointed; upper blades 1-2 per parapodium
(often absent), serrate on proximal third, 80-100 um long, with basal ser-
rations only slightly enlarged; blades below upper ones with about 3 long,

1152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 31. Syllides floridanus: a, Anterior end, dorsal view; b, Same, holotype; c, Same,
ventral view; d, Parapodium, setiger 10, anterior view; e, Same, setiger 22, posterior view.

basal spines more or less parallel to blade edge proximally, with smaller
serrations distally on basal third, 2—3 per parapodium, 64-81 wm long; about
2 shorter blades below basally spined blades, 48-58 um long, with edges
similar to those of upper blades; lower blades 24-34 um long, finely serrate

VOLUME 93, NUMBER 4 1153

eee

O um

Fig. 32. Syllides floridanus: a, Superior simple seta, setiger 10; b, Tip of same (magnified);
c, Upper compound seta, setiger 10; d, Base of blade of same, setiger 30; e, Upper middle
compound seta, setiger 10; f, Same, setiger 20; g, Bases of blades of same, setiger 30; h, Lower
middle compound seta, setiger 20; i, Lower compound seta of same; j, Aciculum, setiger 10
(b, d and g not scaled).

on edge. Shafts similar throughout; distal ends pointed, weakly to strongly
serrate; short parts of hinge wider than width of remainder of shaft. Inferior
simple setae absent. Neuroacicula (Fig. 32j) solitary; tips slightly enlarged,
blunt. Notoacicula (Fig. 3le, f) slender, beginning at least by setiger 10.
Pygidium with all lateral anal cirri lost, with short, slender median cirrus.
No sexually mature specimens with natatory setae, ova, or external em-
bryos or internal sexual products observed; all probably first appearing in
segments with notoacicula.

Pharynx surrounded by brown glands, with about 10 reduced, soft papillae
surrounding anterior end; middorsal tooth absent. Proventriculus in 4—5
segments, barrel-shaped; about 42 transverse rows of muscle cells in dorsal
view, also arranged in 2 diagonal planes.

Remarks.—Syllides floridanus is similar to specimens from the northeast

1154

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Morphometric characters of Syllides japonica from Japan, the northeastern Pacific

and New England, S. j. edentatus from the Galapagos Islands and S. floridanus.

S. japonica SS. floridanus
S. japonicus S. japonicus edentata N. sp.,
S. japonicus Banse, Banse, Westheide, Hutchinson
Imajima, Northeast New Galapagos Island,
Character Japan Pacific England Islands Florida
Length (mm) 6-7 Incomplete 4 3.3 3.5
Width (without parapodia, mm) 0.36 0.38 0.5 0.3 0.3
No. setigerous segments 42-46 Incomplete 34 32 38
Pharyngeal papillae Present Absent Present ? Present
(reduced)
Pharynx:proventriculus, length i 1:1.5 1:1.4 ? 1:1.5
ratio
Proventriculus, width:length ? 1:1.75 1:2.1 # 1:1.6
ratio
Muscle cell rows u i 50 45-50 42
Maximum number articles 30 y % 13 20
(dorsal cirri)
Natatory setae (notoacicula) 2 10 10 9 10
from setiger
Superior simple seta, tip Blunt Fine Recurved, Fine Fine,
bidentate capitate
Superior simple seta, length 100 250 150 150 150
(um)
Blades of compound setae:
Dorsalmost, proximal teeth Long Long Long Fine Stiff
spines spines spines hairs hairs
and stiff
hairs
Dorsalmost, length (um) 84 125 150 65 80-100
Ventralmost, length (um) 34 25-30 30-40? ai 24-34
Ventralmost:dorsalmost 12255 1185) 1:3.8 1:2.4 1:3
length ratio
Basally spined, length (um) 45 (-84) ? 80 (—150) 47 64-81
No. compound setae (setiger 7 (2) 20 (4) 17 (4) 15 (ant) 10 (4)
number)
No. compound setae (setiger 8 (10) 10 (25) 10 (mid) 10 (13) 9 (21)

number)

Pacific (USNM 45264) and New England (USNM 10080, 33152) referred to
S. japonicus Imajima, 1966, by Banse (1971:1477, 1478, fig. 5) and to S. /.
dentata Westheide (1974:81-83, figs. 36e, 37) from the Galapagos Islands.
[One of the New England specimens (USNM 10080), from Cape Cod Bay,
was originally identified by Vernill as S. setosus Verrill (1882:369); the latter
may be one of Verrill’s types but was not considered to be such by Banse.]
S. japonicus Imajima from Japan (Imajima, 1966a:112, 114, fig. 36) has a
pharynx with a subterminal, middorsal tooth and perhaps should not be a

VOLUME 93, NUMBER 4 1155

member of the genus. Banse’s specimens and those of S.j. edentatus do not
have a pharyngeal tooth and appear to be typical members of the genus.
Banse’s northeast Pacific specimens lack soft papillae surrounding the an-
terior end of the pharynx, but such papillae are present in New England
specimens. The integument of the tentacular segment and first setigerous
segment of northeast Pacific specimens is glandular and similar to that of
the prostomium, while in New England specimens only the integument of
the tentacular segment is similar to that of the prostomium. Eyes are faint
in the epitokous specimen and absent in the nonepitokous specimens Banse
reported from New England, but well developed on the Pacific specimens.
Superior simple setae of New England specimens are sturdy and flattened
in the serrated area and have a recurved tip which under interference mi-
croscopy is bidentate with lower cusp small; superior simple setae of north-
eastern Pacific specimens are very thin and finely pointed (Dr. N. W. Riser,
personal communication). Superior simple setae of S. floridanus are stout
at the base and very fine tipped, with tips entire, slightly hooked and capitate
or hooded. Additionally, large basal spines of certain blades of compound
setae of New England specimens are curved toward the opposite edges of
blades, while on S. floridanus such are curved parallel to the toothed edges
of the blades. Other differences are noted in Table |. Such differences seem
to indicate 5 distinct species in specimens noted above, 4 of which are
typical members of Syllides. Certainly, S.j. edentatus should be elevated to
specific rank. I am unable at this time to describe the New England and
northeast Pacific specimens.

Trypanosyllis Claparede, 1864
Trypanosyllis coeliaca Claparéde, 1868
Figs. 33, 34

Trypanosyllis coeliaca Claparéde, 1868:513, 514, pl. 13, fig. 3.—Fauvel,
1923:270, fig. 101f-h.—Cognetti, 1957:27-29, text-fig. 5a, pl. 1, fig. 5.—
Hartmann-Schroder, 1977:85, figs. 23, 24.

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6'N,
80°13.2’W, about 11 m, coarse calcareous sand; 7 specimens (USNM 54537;
FSBC I 20682-20686). Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse
calcareous sand; 9 specimens (FSBC I 20687-20692). Sta. V, 27°22.9'N,
80°13.9’W, about 11 m, coarse calcareous sand; 3 specimens (USNM 54538;
FSBC I 20693, 20694).

Description.—Body without color markings; eyes dark reddish brown.
Flattened, ribbonlike; largest specimen (posteriorly incomplete) 6.0 mm
long, 0.65 mm wide across proventriculus, 73 setigers; complete specimen
5.5 mm long, 0.6 mm wide, 79 setigers. Prostomium (Fig. 33a, b) about

| twice as wide as long; dorsal outline rectangular to trapezoidal; anterior

1156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ew
uit OO|

ry
)

\)
Ons
AIT)
iY
SY

400 um

O
EA
CY e

(7
|
at

a)
108G88

Fig. 33. Trypanosyllis coeliaca: a, Anterior end, dorsal view; b, Same, ventral view (FSBC
I 20689); c, Posterior end of same, ventral view; d, Parapodium, anterior setiger, anterior view
(cirrus missing); e, Same, posterior setiger; f, Anterior border of pharynx, dorsal view (FSBC |

I 20689); g, Same (USNM 54538).

margin nearly straight, with rounded corners, mostly wider than posterior
margin; posterior margin convex, with slight median concavity; 5 lobes dor-
sally, consisting of 2 triangular, anterolateral lobes, triangular median an-
terior lobe, and 2 posterior lobes with triangular anterior extensions between

VOLUME 93, NUMBER 4 1157

15 um

Fig. 34. Trypanosyllis coeliaca: a, Upper compound seta, anterior setiger; b, Lower com-
pound seta of same; c, Upper compound seta, posterior setiger; d, Lower compound seta of
same; e, Inferior simple seta of same; f, Acicula, anterior setiger.

anterolateral and anteromedial lobes. Lateral antennae and anterolateral
pair of dorsal eyes on anterolateral lobes; median antenna originating on
anterior margin of anteromedial lobe; posterior pair of dorsal eyes on an-
terolateral margins of posterior lobes. Ciliated nuchal organs extending
across posterior margin of prostomium; ciliated areas also on anterior mar-
gin of medial lobe. Dorsal eyes in trapezoidal arrangement open to front,
anterolateral pair larger; third pair of eyes on ventral side of prostomium
posterior to origin of lateral antennae and palps, generally situated on each
side between 2 dorsal pairs. Median antenna with 13 articles, about twice
length of prostomium, extending for greater than twice length of palps;
lateral antennae slightly shorter, with 10-11 articles, originating ventrally
on short ceratophores. Palps originating from anteroventral prostomial mar-
gin, shorter than prostomium, appearing superficially articulated, constrict-
ed and completely separated at bases in ventral view, ending in round tips,
often anteriorly divergent. Tentacular segment dorsally reduced to % length
of following segment. Dorsal tentacular cirri originating on short cirro-
phores, with 10-11 articles, slightly longer than lateral antennae and shorter
than dorsal cirri of setiger 1. Ventral tentacular cirri *%4 length of dorsal
tentacular cirri, with 7-8 articles. Setigerous segments 7-8 times wider than
long. Dorsal cirri with short cirrophores, with 15-16 articles on setiger 1,
thereafter with about 10 articles on anterior setigers, long and short in mid-
dle segments with about 8 and 12 articles. Parapodial lobes (Fig. 33d, e)
flattened, convex posteriorly, more convex anteriorly, with tips acutely
pointed, slightly acuminate in dorsal view. Ventral cirri originating below
emergence of setae, flattened, with round tips, often extending past tips of
setal lobes of anterior segments, shorter posteriorly. Compound setae (Fig.

1158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

34a—d) 9-10 in each parapodium; blades bidentate, with relatively long ser-
rations on edge, upper blades longer (18-13 um), with more serrations than
lower blades on anterior segments, only slightly longer posteriorly (14-12
yum). Shafts of compound setae with few short serrations subdistally. Soli-
tary, slender, bidentate, inferior simple setae (Fig. 34e) on posterior para-
podia. Acicula (Fig. 34f) 2 in each parapodium of anterior setigers, 1 in
posterior parapodia; smaller one of anterior parapodia slightly bent near
tips; tips of larger acicula stout, emergent, obliquely acute; acicula of pos-
terior setigers similar to larger, anterior ones. Pygidium (Fig. 33c) composed
of 3 lobes, posteriorly divided; anal cirri long, stout, with 12-13 articles.
Sexual reproduction by stolons; stolons formed by posterior budding; one
with dorsal and ventral pairs of eyes, 27 setigerous segments long.

Pharynx (Fig. 33a, f, g) relatively wide, long, in setigers 2—9 or 3-10,
slightly more than 1% times length of proventriculus; trepan consisting of
10 small, acutely pointed teeth and larger, middorsal tooth; anterior end
surrounded by 10 prominent, soft lobes; posterior part surrounded by light
brown glands. Proventriculus (Fig. 33a) short, thick, opaque in transmitted
light in glycerin, in about 4 segments; length to width ratio 3:2; about 20
transverse rows of muscle cells also arranged in 2 opposite, diagonal arcs;
rows of muscle cells not obviously divided by median line. Ventricle oc-
cupying about 2 segments, medially divided.

Remarks.—Hutchinson Island specimens are in agreement with the orig-
inal description and other European descriptions of T. coeliaca. The phar-
ynx of Hutchinson Island specimens appears similar to that described for
T. fertilis Verrill (1900:616, 617) from Bermuda, but many other characters
are not in agreement.

Trypanosyllis coeliaca was described from the Gulf of Naples. It was
previously known from the Mediterranean Sea, the eastern North Atlantic
and adjacent seas (Hartmann-Schroder, 1977), Florida (Rullier, 1974), and
the Solomon Islands (Gibbs, 1971). Gibbs (1971:142) gave no description for
the single specimen he reported, and I have not examined it. Rullier
(1974:29, 30) reported 2 very young individuals, 2 mm long with 20 setigers,
and I cannot confirm from his short description that Hutchinson Island
Specimens are the same species as the specimens he reported.

Trypanosyllis inglei, new species
Fig. 35

Material examined.—FLORIDA: Hutchinson Island Sta. IV, 27°20.7'N,
80°12.8’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Sep. 1972; USNM 60223); 12 paratypes (USNM 60224; AHF Poly 1323;
ZMH P-16405; FSBC I 23664-23672). Sta. II, 27°21.6’N, 80°13.2'W, about
11 m, coarse calcareous sand; 13 paratypes (USNM 60225; AHF Poly 1324;

VOLUME 93, NUMBER 4 1159

Fig. 35. Trypanosyllis inglei: a, Anterior end, dorsal view; b, Posterior end, dorsal view;
c-e, Setae, setigers 1-4: c, Incompletely fused hook; d, Completely fused hook; e, Stout
compound seta; f-—h, Compound setae, setiger 8: f, Upper; g, Middle; h, Lower; i, Aciculum,
anterior setiger; j, Same, posterior setiger; k, Anterior border of pharynx, ventral view (c-j
not scaled).

ZMH P-16406; FSBC I 23673-23680). Sta. V, 27°22.9'N, 80°13.9’W, about
11 m, coarse calcareous sand; 2 paratypes (FSBC I 23681, 23682).
Description.—Without color markings. Maximum length 8.5 mm, width
across proventriculus 0.6 mm, 100 segments; flattened, ribbonlike. Prosto-
mium (Fig. 35a) generally bilobed, about twice as wide as long, reniform,

1160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

posterior margin convex, lateral margins rounded, anterior margin straight
or concave. Single pair of lensed eyes near anterior margin at about middle
of each side. Median and lateral antennae originating near anterior margin
on short ceratophores, with about 11 articles; lateral antennae originating
above lateral edges of palps; distal articles longer. Palps shorter than mid-
prostomial length, together occupying about half total width of anterior pro-
stomial margin, completely separated basally, short, oval in outline. Ten-
tacular segment distinct dorsally, about half length of following segments,
appearing biannulate; dorsal tentacular cirri similar to antennae; ventral
tentacular cirri about half length of dorsal tentacular cirri, with about 8
articles. Width of setigerous segments 5-7 times length, appearing biannu-
late; integument very thick. Dorsal cirri of setiger 1 similar to antennae and
dorsal tentacular cirri; short on setigers 2 and 3, with 2-3 articles; longer
on setigers 4 and 6, with 3-4 articles; short on setigers 5, 7, and 8, with 1-
2 articles; alternately long and short thereafter, with 2 and 4 articles ante-
riorly and 1 and 2 articles posteriorly; all originating on short, distinct cir-
rophores. Setigerous lobes flattened, acutely pointed, more convex on an-
terior border than posterior. Ventral cirri of anterior parapodia small, oval,
flattened, constricted at base, originating near tips of parapodial lobes on
posterior side; broader, longer, originating obliquely on posterior sides with
more lateroanterior origins, extending laterally to about tips of parapodial
lobes, with nipple-shaped tips on posterior parapodia. Parapodia of setigers
1-4 with about 3 stout, simple, unidentate hooks formed from fusion of
blades and shafts of compound setae (Fig. 35c, d) and mostly single, stout,
compound setae with unidentate blades (Fig. 35e); hooks as stout as or
stouter than acicula; stoutest hook in each parapodium completely fused,
other hooks not completely fused. Setae beginning on setiger 5 compound
falcigers (Fig. 35f—h) with short, hooked, unidentate blades; dorsal blades
with long serrations; ventral blades mostly smooth; blades otherwise similar
in size and shape throughout body, length about 10 um. Solitary, inferior
simple setae on posterior parapodia, about as stout as shafts of compound
setae, slightly curved, unidentate. Acicula (Fig. 35i, j) solitary in all para-
podia, stout, with rounded tips, not emergent, confined within acute dor-
soposterior lobes. Pygidium (Fig. 35b) composed of 2 flattened, divided
lobes, with pair of anal cirri similar to dorsal cirri of midregion, with length
of each cirrus not exceeding total width of pygidium.

Pharynx (Fig. 35k) long, slender, often somewhat bent from contraction
during fixation, normally in setigers 4-13 when inverted; trepan with 10
widely spaced, acute teeth; 10 soft, rounded lobes encircling distal end.
Proventriculus long, narrow, usually in setigers 13-21 (about 9 segments),
maximum length 1.0 mm, width 0.16 mm, with 50-57 irregular transverse
rows of muscle cells.

Remarks.—Trypanosyllis inglei cannot be assigned to the subgenera

|
}

VOLUME 93, NUMBER 4 1161

erected by Imajima and Hartman (1964) since both compound setae and
simple setae formed from fusion of shafts and blades of compound setae are
present; I know of no described species having both compound unidentate
setae and simple setae of this type. However, the species appears to be
more aligned with the subgenus 7rypanobia than with Trypanosyllis or Try-
panedenta, since a middorsal pharyngeal tooth is absent, some simple setae
formed from fusion of shafts and blades are present, and blades of com-
pound setae are unidentate.

Etymology.—The species is named in honor of Robert M. Ingle, previous
Director of the Florida Department of Natural Resources Marine Research
Laboratory, who was instrumental in initiating the project in which this
species was collected.

Trypanosyllis parvidentata, new species
Fig. 36

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6’N,
80°13.2'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Sep. 1972; USNM 54540). Sta. IV, 27°20.7’N, 80°12.8'W, about 11 m,
coarse calcareous sand; 2 paratypes (USNM 54541; FSBC I 23683). Sta. V,
27°22.9'N, 80°13.9'W, about 11 m, coarse calcareous sand; 1 paratype
(FSBC I 23684).

Description.—Body without color markings; eyes dark red. All specimens
posteriorly incomplete; largest specimen 10 mm long, width 0.45 mm across
proventriculus, gradually widening for anterior half of body, 90 segments,
flattened, ribbonlike. Prostomium (Fig. 36a) wider than long; anterior mar-
gin straight; lateral margins rounded; posterior margin convex, with median
concavity; 5 lobes dorsally including pair of anterolateral lobes, anterome-
dial lobe, and pair of posterior lobes; ciliated nuchal organ covering posterior
width. Eyes 2 pairs, well separated, in trapezoidal arrangement open to
front; anterior pair larger, lensed, on anterolateral lobes; posterior pair pos-
sibly not lensed, on posterior lobes; ventral eyes apparently absent. Median

_ antenna originating from anteromedial lobe; lateral antennae originating

from anterolateral lobes; all on short ceratophores, with 8-10 articles. Palps
about as long as prostomial lobe without nuchal organ, narrow at bases,
well separated, originating from anteroventral margin, bent ventrally in
specimen figured (Fig. 36a). Tentacular segment relatively short dorsally;
dorsal tentacular cirri originating on short cirrophores, similar to antennae;
ventral tentacular cirri about half as long as dorsal, with about 5 articles.
Anterior setigerous segments about 6 times wider than long. Dorsal cirri
originating on short cirrophores, with about 15 articles on setiger 1, 10-13
articles on setiger 2, with about 10 articles on following few anterior setigers;
dorsal cirri beginning at about setiger 5, alternately long and short, with 15

1162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

b

a
Ao

SDA 1%

= a)
a Stoo? eaiconnn aaa
)
£ Se
O k)

h

ea Ay errant
lOO yum

(iar: Se
15 jum

Fig. 36. Trypanosyllis parvidentata: a, Anterior end, dorsal view [composite figure of ho-
lotype and paratype (FSBC I 23684)]; b, Parapodium, anterior setiger, somewhat oblique an-
terior view (cirrus missing); c, Same, posterior setiger; d, Upper compound seta, anterior
setiger; e, Same, posterior setiger; f, Lower compound seta of same; g, Aciculum, anterior
setiger; h, Anterior border of pharynx of holotype, dorsal view.

VOLUME 93, NUMBER 4 1163

and 10 articles. Parapodial lobes (Fig. 36b, c) flattened, convex anteriorly
and posteriorly; tips acuminate. Ventral cirri originating from near tips of
parapodial lobes, stout, superficially appearing articulate, with somewhat
ventrally pointing apices. Compound setae (Fig. 36d—f) 8-9 in each para-
podium; blades falcigerous, bidentate, similar in anterior and posterior seg-
ments but possibly more strongly hooked and with primary and secondary
teeth more widely separated in posterior segments, about 16 um long dor-
sally, about 11 ~m long ventrally; edges with numerous fine serrations from
bases to secondary teeth; shafts serrate subdistally. Inferior simple setae
not observed. Acicula (Fig. 36g) solitary in all parapodia, with slightly bev-
eled, rounded tips, stouter in posterior segments. Pygidium not observed.

Pharynx (Fig. 36a, h) occupying length of setigers 2-8 or 3-9, often strong-
ly bent from contraction, with thin walls; trepan with 10 very small, obscure
teeth and large middorsal tooth; 10 soft lobes surrounding anterior end.
Proventriculus (Fig. 36a) 3-4 segments long (setigers 9-11 to 12-13), about
twice longer than wide (150 um X 290 um, 170 um x 340 um), with 18
transverse, medially divided muscle cell rows; ventricle about 2 segments
long, also medially divided.

Remarks.—Trypanosyllis parvidentata differs from other members of the
genus in having a trepan of 10 minute teeth in addition to a large, middorsal
tooth. Additionally, the species differs from other Trypanosyllis species
reported herein in the length and number of articles of antennae and cirri,
and the fine detail of the compound setae.

Etymology.—The species name is derived from Latin and refers to the
minute teeth on the trepan.

Trypanosyllis savagei, new species
Fig. 37

Material examined.—FLORIDA: Hutchinson Island Sta. II, 27°21.6’N,
80°13.2’W, about 11 m, coarse calcareous sand; holotype (R. Gallagher,
col., Nov. 1971; USNM 60461), 2 paratypes (FSBC I 23685). Sta. V,
27°22.9'N, 80°13.9’W, about 11 m, coarse calcareous sand; 4 paratypes
(USNM 60462-60464; FSBC I 23686).

Description.—Body without color markings; eyes light reddish orange.
Length less than 5 mm; width 0.4 mm at proventriculus; flattened, ribbon-
like, maximum 80 segments. Prostomium (Fig. 37a) oval in outline, wider
than long, with concave posterior indentation; 5 lobes dorsally including
small, anteromedial lobe, pair of small, anterolateral lobes, and pair of large,
posterolateral lobes; latter with acute extensions between anteromedial and
anterolateral lobes. Two pairs of dorsal eyes close together on opposite
sides of midline; anterior pair slightly more widely separated, possibly sit-
uated on anterolateral lobes and covered by posterolateral lobes; both dorsal

1164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wit OO!

h g f

oe Lia =a.)
40 ym IS jum

Fig. 37. Trypanosyllis savagei: a, Anterior end, dorsal view; b, Middle segments, including
anterior end of developing reproductive stolon, dorsal view; c, Posterior end, dorsal view; d,
Anterior parapodium, slightly oblique anterior view; e, Same, posterior parapodium; f, Upper
compound seta, anterior setiger; g, Compound seta, posterior setiger; h, Inferior simple seta,
posterior setiger; i, Anterior border of pharynx, dorsal view.

pairs near lateral margins; smaller, ventral pair of eyes slightly medial to 2
dorsal pairs. Median antenna originating from raised anteromedial lobe, with
about 8 articles; lateral antennae originating on anterior margin on short
ceratophores, extending for about same distance and similar to median;
distal articles about 2 times longer than others. Palps small, about equal in
length to half prostomial length, completely separated at bases. Ciliated
nuchal organs on posterior margin of prostomium and tufts of cilia on an-

VOLUME 93, NUMBER 4 1165

terior margin. Tentacular segment dorsally distinct but reduced to less than
half length of following segment, laterally surrounding posterior half of pro-
stomium. Tentacular cirri originating on short cirrophores; dorsal tentacular
cirri similar to antennae; ventral tentacular cirri more slender, about half as
long as dorsal, with about 6 articles. Mouth opening on ventral posterior
border of prostomial lobe. Dorsal cirri of setiger 1 similar to dorsal tentac-
ular cirri and antennae; length equal to about half body width, with 5-6
articles on other anterior segments, with 3—4 or fewer articles on segments
behind proventriculus; all originating on distinct, short cirrophores. Para-
podial lobes (Fig. 37d, e) flattened; tips usually acutely pointed; anterior
and posterior borders convex; posterior borders more convex than anterior
ones. Ventral cirri originating from about middle of ventral margins of para-
podial lobes about half distance from bases to tips, projecting laterally for
about same distance or for slightly greater distance than parapodial lobes;
cirri stout, pyriform in outline; tips rounded. Blades of all compound setae
similar (Fig. 37f, g), about 12 um long, slender, falcigerous and bidentate,
with primary tooth more prominent than secondary; blade edges pectinate
to secondary tooth, with longer serrations on upper blades. Shafts smooth.
Solitary, inferior simple setae on posterior parapodia, slender, curved, bi-
dentate. Acicula solitary, stout, acute, often emergent. Pygidium (Fig. 37c)
composed of 2 flattened posterior lobes and median, anterior lobe, with pair
of anal cirri originating from outer posterior corners on ventral side; each
cirrus with about 3 articles. Sexual reproduction by stolons formed by pos-
terior budding; stolon beginning on setiger 45 in specimen figured (Fig. 37b),
containing sexual products in all but last few segments.

Pharynx long, narrow, mostly in setigers 2—9; trepan of 10 almost equi-
triangular, small teeth and larger middorsal tooth; anterior margin surround-
ed by 10 soft lobes (Fig. 371). Proventriculus (Fig. 37a) slightly greater than
half pharyngeal length, cylindrical, greater than twice longer than wide (220
um long, 90 wm wide), occupying 3-4 segments (setigers 10-14), with about
17 medially divided, transverse rows of muscle cells. Ventricle in figured
specimen convoluted, medially divided.

Remarks.—Trypanosyllis savagei is similar to specimens of T. coeliaca
Claparede from Hutchinson Island but differs in several important charac-
ters. Although mature specimens of both species have about the same num-
ber of segments, 7. savagei is much smaller and slenderer, and characters
such as number of setae per parapodium, number of articles on antennae
and cirri, and number of acicula in anterior parapodia are reduced as com-
pared with T. coeliaca. The principal difference between the two species
is in the shape and character of the proventriculus. The proventriculus of
T. coeliaca is very stout and opaque in glycerin with transmitted light (see
Cognetti, 1957:pl. 1, fig. 5), while the proventriculus of 7. savagei is slender
(greater than 2 times longer than wide) and translucent in glycerin.

1166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology.—The species is named in honor of Thomas Savage, formerly
of the Florida Department of Natural Resources Marine Research Labora-
tory.

Dentatisyllis, new genus

Type-species.—Syllis carolinae Day, 1973.

Diagnosis.—Body long, cylindrical, with numerous segments. Prostomi-
um with 3 antennae; tentacular segment with 2 pairs of cirri, all articulated.
Palps free at base. Eversible pharynx with trepan of 10 teeth and large
middorsal tooth; anterior margin surrounded by 10 soft lobes. Nuchal organs
absent. Dorsal cirri articulate; parapodial lobes conical; ventral cirri pres-
ent. Setae compound falcigers joined by superior and inferior simple setae
on posterior segments.

Remarks.—Dentatisyllis has the same relationship to Typosyllis Langer-
hans, 1879, as Geminosyllis Imajima, 1966, has to Haplosyllis Langerhans,
1879. Both Typosyllis and Dentatisyllis have numerous cylindrical segments
with conical parapodial lobes, and similar antennae, cirri, palps and setae.
Dentatisyllis differs from Typosyllis in having an eversible pharynx with
several teeth on the margin while the latter has a pharynx with a smooth
anterior margin. Dentatisyllis differs from Trypanosyllis Claparede, 1864,
in that the latter has numerous flattened segments and flattened parapodial
lobes.

Etymology.—The generic name is formed from the Latin dentata, meaning
toothed, and the stem generic name, Syllis. Gender: feminine.

Dentatisyllis carolinae, new combination
Fig. 38

Syllis (Typosyllis) regulata carolinae Day, 1973:30, fig. 4a—f.—Gardiner,
1976:141, fig. 12x-z, 13a.

Material examined.—NORTH CAROLINA: off Beaufort, 34°34’N,
76°25'W, 20 m, sand and broken shell; J. H. Day, col., 19 May 1965; ho-
lotype (USNM 43146), 4 paratypes (USNM 43147). SOUTH CAROLINA:
32°13'00"N, 79°52'02”W, 24 m, 17 Feb. 1977; 1 specimen (VIMS). GEOR-
GIA: 31°11'59’N, 81°08’00"W, 11 m; R. Bertelsen, col.; 1 specimen (USNM
60472). FLORIDA: Hutchinson Island Sta. I, 27°21.3'N, 80°14.1’W, about
8 m, very fine to fine quartose sand; 1 specimen (FSBC I 20695). Sta. II,
27°21.6'N, 80°13.2'W, about 11 m, coarse calcareous sand; 90 specimens
(USNM 54535; FSBC I 4543, 20696-20729). Sta. III, 27°22.0’N, 80°12.4'W,
about 7 m, medium calcareous sand; 2 specimens (FSBC I 20731, 20732).
Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 134
specimens (FSBC I 20733-20767). Sta. V, 27°22.9'N, 80°13.9’W, about 11

VOLUME 93, NUMBER 4 1167

wt OOg

Fig. 38. Dentatisyllis carolinae: a-e, Setae, posterior setigers (FSBC I 20779): a, Superior

simple; b, Small, upper compound; c, Large, upper compound; d, Lower compound; e, Inferior
simple; f-h, Anterior border of pharynx: f, Paratype, lateral view, everted; g, Small specimen,

ventral view (FSBC I 20767); h, Large, mature specimen (USNM 60474).

1168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

m, coarse calcareous sand; 85 specimens (USNM 54534, 60473, 60474; VIMS;
FSBC I 20768-20783, 20785-20794). Johns Pass, Tampa Bay area, Pinellas
County; B. Howton and R. Dalton, cols., 12 July 1978; 1 specimen (FSBC
I 20840). Northeastern Gulf of Mexico, 29°20’00.4"N, 84°44'02.3”W, 30 m,
coarse calcareous sand; U.S. Dept. Interior, Bureau of Land Management
Miss.-Ala.-Fla. (MAFLA) Study, Transect IV, Sta. 2423; R/V Columbus
Iselin, July 1976; 1 specimen (USNM 55831).

Additional description.—Body slightly tapered for anterior third, broadest
in middle, tapered for most of posterior third. Antennae and dorsal cirri of
anterior segments relatively slender; dorsal cirri of middle segments much
stouter and longer. Prostomium of juveniles with pair of eyespots near an-
terior margin in addition to 2 larger pairs. Median antenna with about 35
articles; lateral antennae originating posterior to anterior border on short
ceratophores, with about 25 articles. Dorsal tentacular cirri similar to me-
dian antenna; ventral tentacular cirri similar to lateral antennae, with about
22 articles. Dorsal cirri of setiger 1 much longer than median antenna, with
up to 55 articles; setiger 2 dorsal cirri similar to lateral antennae, with 27
articles; setiger 3 dorsal cirri similar to median antenna; dorsal cirri of se-
tigers 4, 6, and 9 similar to cirri of setiger 1; dorsal cirri with about 30
articles on setigers 5, 7, and 8; thereafter alternately short and long to pos-
terior end, with about 30 articles on short cirri and about 55 articles on long
cirri of middle segments. Superior simple setae truncate but distinctly bi-
lobed or bifid, with few serrations on ventral borders (Fig. 38a). Falcigers
of anterior setigers numerous, with slender shafts and blades. Upper falci-
gers of posterior setigers with slender shafts and blades; blades as long as
upper ones of anterior parapodia on some mature specimens but mostly
shorter and similar in length to lower blades (Fig. 38b, c); lower falcigers
of posterior setigers with stout shafts and blades (Fig. 38d). Inferior simple
setae with tips similar to tips of blades of lower compound setae of posterior
segments, with several stiff hairs below secondary tooth (Fig. 38e). Up to
5 acicula in anterior parapodia, 3 in middle parapodia, reduced to 1-2 in
posterior parapodia. Pygidium with short, slender, smooth, ventrally origi-
nating anal cirrus in addition to 2 ventrolaterally originating articulated ones.

Pharynx (Fig. 38f—h) longer than proventriculus, thin walled, especially
on anterior margin, light brown, usually located in setigers 1—2 to 9-11, in
10-11 segments when inverted, with trepan of 10, somewhat irregular teeth
on anterior chitinous margin and large middorsal tooth near anterior margin
with tip not extending to trepan; anterior end surrounded by 10 soft lobes
in apposition to teeth of trepan. Proventriculus usually located in setigers
10-12 to 14-17, in 5-8 segments when pharynx not everted, subcylindrical;
length: width ratios 3:1 to 5:1.

Remarks.—Compound setae of D. carolinae are remarkably similar to
those of Typosyllis regulata Imajima, 1966, from Japan; antennae and cirri

VOLUME 93, NUMBER 4 1169

are similar but shorter on 7. regulata. However, the anterior border of the
pharynx of T. regulata is smooth (Imajima, 1966b:290, fig. 64b). Therefore,
the 2 species cannot be placed in the same genus, and D. carolinae is
certainly not a subspecies of T. regulata as it was originally described (Day,
1973).

The length and number of articles of antennae and cirri described herein
appear to be typical for syllid species with long, articulated dorsal cirri of
alternate lengths. The arrangement may become obscured when antennae
or cirri have been broken off and are regenerating, resulting in their being
shorter, with fewer articles. Small specimens of the species also have an-
tennae and cirri with fewer articles.

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, and M. L. Hollinger. 1977. Nearshore marine ecology at Hutchinson Island, Florida,

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Gardiner, S. L. 1976. Errant polychaete annelids from North Carolina.—J. Elisha Mitchell

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, and W. H. Wilson, Jr. 1979. New records of polychaete annelids from North Carolina

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description de deux nouvelles especes.—Cah. Biol. Mar. 3:249-260.

Hall, J. R., and C. H. Saloman. 1975. Distribution and abundance of macroinvertebrate
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Hartman, O. 1942. A review of the types of polychaetous annelids at the Peabody Museum

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VOLUME 93, NUMBER 4 1171

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Florida Department of Natural Resources, Marine Research Laboratory,
100 Eighth Avenue, S.E., St. Petersburg, Florida 33701.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1173-1176

BARBIDRILUS PAUCISETUS, NEW GENUS,
NEW SPECIES (OLIGOCHAETA: ENCHYTRAEIDAEB),
FROM EASTERN NORTH AMERICA

Michael S. Loden and Steven M. Locy

Abstract.—Barbidrilus paucisetus, anew genus and species of freshwater
enchytraeid oligochaete was collected from the Apalachicola River, Florida.
Specimens identical in setal morphology and placement have been collected
from North Carolina and Ohio. The new species is unusual in that the setae
are present only in the ventral setal bundles of segments II and III. Straight,
anteriorly directed setae, elongate spermathecal ducts, and compact penial
bulbs are characteristic.

Investigations of North American Enchytraeidae have lagged consider-
ably behind the work on this group by specialists in Europe, Asia, and South
America. Likely synonyms of several of the species, lost type-specimens,
and species transferred to other genera have hindered identifications of ter-
restrial and freshwater species and resulted in few North American investi-
gators willing to study enchytraeids. Tynen’s (1975) checklist of species
known to occur in North America has provided a much-needed impetus for
additional work on this group of oligochaetes.

We believe this to be the first twentieth-century description of a possibly
endemic North American genus.

Although most descriptions of enchytraeid species have been based on
living material, the specimens we examined were, unfortunately, received
in a preserved state. This was of some disadvantage in distinguishing such
structures as the nephridia, but because of the large numbers of specimens
received, we were able to observe most of the other taxonomically impor-
tant features of the specimens. Some of the specimens had been stained
with a red dye which facilitated observations of internal structures.

Barbidrilus, new genus

99 66

Etymology.—‘‘barba,’’ L. ‘‘beard’’; “‘drilos, worm ’’; in reference to
the anterior ventral placement of elongate setae.

Diagnosis.—Setae long, straight, without nodulus, restricted to anterior
ventral bundles. Head pore at 0/I. Sorsal pores absent. Esophageal-intestinal
transition gradual, no esophageal or intestinal diverticula. Dorsal vessel aris-
ing anterior to clitellum. Three pairs of primary septal glands; secondary

septal glands absent. Sperm sac present. Sperm ducts long, narrow; penial

1174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Br

= E22.
PSI GES
DITA ory

As
SGN RS
A ait SORE.
eG Vitenasgesaae

Eris

aio
Cererirey tt

Crh -y :

5
a
By
2
ee
-
a
ee
4
ac
AN
=
=
Gg
=
io
Cy
=
=

E
3 J “ei
~ Ay ui 2"
oF > woe FX
A)
K/ a Hee
4 me
Bist
148
hia
yea ¢
el
H
a
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B
A
aor A AEH an _
ee IAS pes an ah ps ee
EEE = Soe pg PE

D

Fig. 1. Barbidrilus paucisetus. A, seta; B, lateral aspect of anterior end; C, male duct; D,
semidiagrammatic dorsal aspect of anterior end. Br, brain; SG, septal gland; SpA, spermathecal

ampulla; SpD, spermathecal duct.

VOLUME 93, NUMBER 4 1175

bulbs compact. Ovisac present. Spermathecal ampullae sac-like, not com-
municating with esophagus; duct long, without associated glands.
Type-species.—The following new species:

Barbidrilus paucisetus, new species
Fig. 1

Etymology.—‘‘paucus, L. ‘‘few’’; referring to the reduction in setal
number.

Holotype. —NMNH 60006. Florida: Apalachicola River. River Mile 87.5.
15 December 1978. B. Pruitt, Jr.

Paratypes.—Same location and date, 10 specimens (NMNH 60007-
60016). Florida: Apalachicola River, River Mile 67.6, 14 December 1978, B.
Pruitt, Jr., 9 specimens (LSU 3070).

Description.—Length 5-9 mm; width 0.08—0.14 mm. Up to 50 segments.
Prostomium rounded, conical, as long as width at peristomial junction. Head
pore indistinct, at 0/I; dorsal pores absent. Setae 2—3 per bundle, only in
ventral bundles of II and III. Setae long, straight, without nodulus, directed
anteroventrally; tips with rounded bifurcations. Setal follicles absent from
other segments. Brain as long as wide, incised posteriorly. Esophageal-in-
testinal transition gradual, no intestinal or esophageal diverticula. Chlor-
ogogue cells sparce, beginning in V. Primary septal glands paired, in V, VI,
and VII, varied in pattern, but not united dorsally or ventrally. First 4
segments aseptate. Nephridia present, but characters unobservable in fixed
specimens. Peptonephridia present, paired. Clitellum extending XI/XII—XII/
XIII with large, closely associated gland cells.

Sperm funnels cylindrical, as long as wide, collar of same diameter as
funnel. Sperm duct long, narrow, extending posteriorly as far as XIV. Penial
bulb compact, bulbous. Spermathecal pores lateral in V. Spermathecal ducts
long, without glands, extending posteriorly to VII or VIII. Spermathecal
ampullae thin-walled, sac-like, elongate, free in body cavity, located as far
posteriorly as septum IX/X. Spermatozoa in lumen of ampulla, not orga-
nized into aggregations.

Discussion.—Barbidrilus paucisetus is unique among the Oligochaeta in
the placement, relative shape, and orientation of the setae.

A reduction in setae such as occurs in B. paucisetus is found in two other
enchytraeid genera. In Marionina achaeta Lasserre, 1964, setae are absent
from all segments; in M. preclitellochaeta Nielsen and Christensen, 1963,
setae are present only in the ventral bundles of segments II through VI.
Setae are completely absent in all known species of Achaeta.

Barbidrilus paucisetus differs from Marionina species because of several
features. Spermathecae in species of Marionina are generally spherical to
ovoid; ampullae usually communicate with the esophagus, and the ducts are

1176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mostly confined to segment V with glands associated with either the sper-
mathecal pores or ducts. In B. paucisetus the spermathecal ampullae are
elongate, sac-like, and free from the gut. There are no gland cells associated
with either the ducts or the spermathecal pores, and the spermathecae may
extend posteriorly to segment IX.

It is perhaps to species of Achaeta that B. paucisetus is most closely
related. Several species of Achaeta (e.g. A. danica Nielsen and Christen-
sen, 1959) have elongate spermathecal ducts. The dorsal vessel of all species
of Achaeta arises in the anteclitellar region, and all lack setae. While most
species of Achaeta have setal follicles in the position of the missing setal
bundles, others (e.g. A. camerani (Cognetti, 1889), A. iridescens Christof-
fersen, 1979, A. littoralis Lasserre, 1967) share with B. paucisetus a lack
of them.

Differences between Achaeta and Barbidrilus include the esophageal-in-
testinal transition (abrupt in Achaeta, gradual in Barbidrilus), presence of
an ovisac in Barbidrilus and an absence in Achaeta, location of the head
pore (0 in Achaeta, O/I in Barbidrilus), and the shape of the penial bulb
(subdivided to form gland-like structures in Achaeta, compact and pear-
shaped in Barbidrilus).

Barbidrilus paucisetus is known only from freshwater habitats; it has been
collected from rivers at locations where fine silt overlies sand. While the
type-locality is Florida, specimens with setae that were identical in shape
and position to those of B. paucisetus have been collected from the South
River, North Carolina, and from Sandusky Bay, Ohio.

Acknowledgments

We wish to thank Buford Pruitt, Jr. and his colleagues at Water and Air
Research, Inc. for bringing the specimens to our attention. Acknowledg-
ment is due to Joseph Boda of the Mobile District, U.S. Army Corps of
Engineers. Specimens were collected during work performed under COE
Contract Number DACWO0O1-78-C-0101.

Literature Cited

Tynen, M. J. 1975. A checklist and bibliography of the North American Enchytraeidae (An-
nelida: Oligochaeta).—Syllogeus No. 9, 14 pp.

Department of Zoology, Louisiana State University, Baton Rouge, Lou-
isiana 70803.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1177-1183

THE TADPOLE OF THE COSTA RICAN
FRINGE-LIMBED TREE-FROG,
HYLA FIMBRIMEMBRA

Jay M. Savage

Abstract.—A single large, long-tailed, tuberculate, and generalized hylid
tadpole from the Isla Bonita-Cinchona area of the Cordillera Central of
Costa Rica cannot be associated with any of the 93 tadpoles of the family
Hylidae described from Mexico and Central America. Consideration of ge-
ography and tadpole characteristics typical of the species groups for the 29
hylid species from the region, whose tadpoles are unknown, leads to asso-
ciation of the larva with the fringe-limbed flying frog, Hyla fimbrimembra.
The tadpole is the first to be described for the 5 species of flying frogs that
comprise the miliaria group.

In 1963 Norman J. Scott and I collected a single large tadpole from the
La Cinchona area of the Cordillera Central of Costa Rica made famous by
Edward H. Taylor as the type locality of many amphibians and reptiles (see
Savage, 1974). The tadpole was peculiar in the combination of small labial
disk, very long tail (77% of total length) and peculiarly tuberculate flanks.
The understanding of the characteristics of tadpoles of most Costa Rican
anurans was minimal at that time and it was not possible to hazard a guess
at our example’s identity.

Fortunately, the intervening years have seen the knowledge of larval
stages for Costa Rican anurans increase several fold and tadpoles for most
species, except the many forms in the genera Bufo and Centrolenella, are
now known. In the course of preparing a review of the tadpoles of Costa
Rican frogs and toads, I have again turned to an attempt at associating the
Cinchona example (Fig. 1) with a named adult population.

The example in question was collected at Costa Rica: Provincia de Ala-
juela: Canton Alajuela: Cinchona (1,360 m) by Jay M. Savage and Norman
J. Scott on 26 September 1963, between 8:30 and 10:00 p.m. It is number
7015 in the CRE collection at the University of Southern California. Its
salient features are summarized below, using the terminology of Duellman
(1970:33—43), where applicable.

General Features.—Stage 38 (system of Gosner, 1960). Body robust,
width somewhat greater than depth. Nostrils directed anteriolaterally, eyes
directed laterally. Anus dextral; spiracle sinistral, located on side of body
below level of eye, about % of distance from eye to posterior margin of

1178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

[oxpeezrcae oe ae
8mm

Fig. 1. Tadpole of Hyla fimbrimembra. Upper, oral disk; lower, lateral view.

VOLUME 93, NUMBER 4 1179

body. Lateral area of posterior portion of body heavily tuberculate. Caudal
fin extending onto body as a low ridge that reaches almost to level of spi-
racle; dorsal portion about 4 depth of caudal musculature, ventral portion
reduced; fin rounded at end.

Oral disk small, anterioventral, directed ventrally. Disk complete, not
indented laterally; disk bordered laterally by 2-3 rows of small papillae,
below by 1 row; oral papillae incomplete across upper labium. Upper beak
a narrow arch, lower beak less well-developed, both with small serrations.
Denticles in 2/3 rows, row immediately above upper beak interrupted me-
dially, uppermost row and row just below lower beak longest, about equal
in length, others only slightly shorter.

Coloration.—Dorsum and sides of head uniform brown, gradually grading
into dirty white venter; flanks lightly pigmented but appear black because
black parietal peritoneum visible through skin. Hindlimb rudiment strongly
barred with black above. Tail musculature light pinkish tan; caudal fins and
overlying skin transparent with distinct network of mottled black pigment.

Measurements (mm).—Standard length (body length) 17; tail length 57;
total length 74; tail length 77% of total length.

An Attempt at Identification

At the present time Costa Rica has a known anuran fauna of approxi-
mately 120 species. Thirty-three of these forms, all members of the genus
Eleutherodactylus, are either known to have direct development without a
tadpole stage or are presumed to share this feature in common, as far as
known with all approximately 300 members of the genus. The remaining
species are allocated to 8 frog families as follows, with the denominator
indicating the number of Costa Rican species in the family, and the numer-
ator the number for which tadpoles have been described: Rhinophrynidae
(1/1); Microhylidae (1/3); Leptodactylidae (6/6); Bufonidae (5/14); Hylidae
(34/39); Dendrobatidae (5/7); Centrolenidae (7/13); and Ranidae (4/4).

The characteristics of the Cinchona example do not match the description
of any known tadpole from Costa Rica or from adjacent areas of Central
America. It must therefore represent a known species for which the tadpole
is not known or an unknown species. If one follows a process of elimination
based on the available data, the number of feasible alternative possibilities
can be substantially reduced. In the following comparisons, features for the
unnamed tadpole are indicated in parentheses.

There is little likelihood that the mystery tadpole represents an unde-
scribed species of either the Rhinophrynidae or Microhylidae. All known
members of these families have tadpoles that lack beaks and denticles and
have either two spiracles or a single ventrally located one (beaks and den-
ticles present; a single sinistral spiracle).

1180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Members of the family Ranidae are also excluded from consideration
since the tadpoles of all known Central American species have been de-
scribed and have either the oral disk indented laterally (Pipiens group) or
high numbers of denticle rows 3/4 to 6/4 (complete oral disk and denticle
rows 2/3).

The tadpoles of members of the genera Atelopus and Bufo (family Bu-
fonidae) are distinctive and cannot be confused with the present specimen.
Atelopus has a well-developed ventral disk (no ventral disk) and Bufo has
the oral papillae incomplete on upper and lower labia (complete on lower
labium).

The Centrolenidae may also be eliminated from consideration since all
known tadpoles of this family have a median anus, very short denticle row
segments just above the mouth that are separated from one another by a
huge gap above the beak and are pale pink to bright red (in life) with minimal
dark pigment (dextral anus, only a narrow gap in long denticle row segments
just above beak, and with considerable dark pigmentation).

Members of the family Leptodactylidae known from lower Central Amer-
ica that have free-living larvae may also be eliminated from consideration.
Leptodactylus has a median anus (dextral); Physalaemus has the oral disk
indented laterally (oral disk complete) and Pleurodema has a median anus
(dextral).

Members of the family Dendrobatidae show a considerable diversity of
larval features (Savage, 1968; Silverstone, 1975, 1976). In known Phyllo-
bates and most Colostethus the oral disk is indented laterally (oral disk
complete). In most Dendrobates the anus is median (dextral) but in those
with a dextral anus the oral disk is indented (oral disk complete). In the
Costa Rican form Colostethus nubicola which has a non-indented umbelli-
form oral disk (complete but non-umbelliform oral disk) the anus is dextral
and denticle rows are absent (2/3 denticle rows). The tadpole of Dendrobates
pumilio of Costa Rica also has a complete oral disk but has a median anus
(dextral) and 0—1/2—2 rows of denticles (2/3 rows of denticles).

The mystery tadpole has been compared directly to tadpoles of all lower
Central American species for which tadpoles are known in the 7 families
discussed above. There can be little doubt that it is distinct from all of them
and almost certainly does not represent an undescribed species or any
known species for which the tadpole remains unknown at this time. This
comparison eliminates from serious consideration as the possible congeners
of the tadpole representatives of all the lower Central American genera of
these families except for the monotypic bufonid Crepidophryne. No tadpole
is known for this genus which seems to be a specialized derivative from
Bufo (Savage and Kluge, 1961). Most likely, if the species has a tadpole
stage, it would agree with Bufo, which uniformly has the oral disk indented
laterally and the oral papillae broadly interrupted across both the upper and

VOLUME 93, NUMBER 4 1181

lower labia (oral disk complete; complete series of papillae across lower
labia). In addition, Crepidophryne is known from only a few localities in the
Cordillera de Talamanca (Savage, 1972), the closest some 75 km southeast
of the collecting site for the unknown larvae.

The unidentified tadpole agrees in basic features with generalized tadpoles
of species in the large family Hylidae, the only family not eliminated from
consideration in the above review. These basic features include: complete
oral disk, dextral anus, sinistral spiracle located laterally, denticles 2/3, and
oral papillae incomplete across upper and complete across lower labium.
Among the hylids of lower Central America, 5 members of the genus Hyla
(angustilineata, elaeochroa, loquax, pseudopuma and staufferi) and 4 mem-
bers of the genus Smilisca (baudinii, phaeota, puma and sila) resemble the
Cinchona tadpole in these basic features. Direct comparison of comparable
stages of tadpoles of these 9 forms to the unnamed larva clearly indicates
that it cannot be associated with any of them. It may most easily be sepa-
rated from H. elaeochroa and H. staufferi by the shape of the body and fin
form. In the latter 2 species the body is deep, with the depth much greater
than width and the tail fins are deep, with upper and lower portions much
deeper than tail musculature, and taper to a point (body ovoid, depth and
width about equal; depth of upper and lower portions of caudal fin less than
depth of tail musculature and tip of fin rounded). H. loquax, S. baudinii and
S. phaeota most obviously differ from the Cinchona tadpole in having
smooth bodies (tuberculate). H. angustilineata, H. pseudopuma and S. sila
do not have the dorsal portion of the fin extending onto the body (fin ex-
tending onto body).

The above review indicates that the mystery tadpole represents a species
of the tree-frog family Hylidae. An analysis of the characteristics of the
larvae of this family from Mexico and Central America, almost all of which
are so ably described and illustrated by Duellman (1970), confirms this view.
Of 122 hylid species now recognized from the region, the tadpoles of 29
remain undescribed. Most of the species (20) for which tadpole data are
lacking occur in the highlands of southern Mexico and Guatemala (15) or
are inhabitants (5) of sites in western Mexico (one of these forms reaches
Guatemala and El Salvador). All of these species are found at sites far
removed geographically and ecologically from the Cordillera Central of Cos-
ta Rica.

Of the remaining 9 forms without known tadpoles, all from lower Central
America, three species, Agalychnis calcarifer and A. litodryas and Phyl-
lomedusa venusta, belong to genera whose tadpoles are so distinctive as to
eliminate them from consideration. Tadpoles for these genera uniformly
have the spiracle located ventrally although sinistral to the midventral region
and have a pointed tail fin (lateral spiracle and rounded tail fin). One form,
Hyla pugnax of the Pacific lowlands in western Panama and northern Co-

1182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lombia (Kluge, 1979) belongs to a species group in which all known tadpoles
have 2/4 denticle rows.

The elimination of the 4 mentioned species from consideration leaves 4
forms, Hyla fimbrimembra, H. lythrodes, H. miliaria and H. xanthosticta,
all known from Costa Rica, and H. thysanota, known only from extreme
eastern Panama, as candidates for association with the tadpole. H. lythrodes
and xanthostica, each known from a single male holotype, are unlikely to
be conspecific with the tadpole. All of the allies of these two forms (Savage,
1967; Duellman, 1970) have larvae with oral papillae complete across the
upper labium (incomplete). Direct comparison of the Cinchona tadpole with
known tadpoles of these 7 allied forms convinces me that they are not
related. Additional differences include: H. rufioculis and uranochroa, 2/2
denticle rows (2/3); H. legleri, 2/5 denticle rows (2/3); H. rivularis, pictipes
and tica, 4-7 rows of papillae on lower labium (1); and H. debilis, body
depressed (ovoid). H. lythrodes is most similar to rufioculis and xanthostica
to pictipes in adult features.

The analysis leaves only the Costa Rica and Panama members of the
Hyla miliaria group (Duellman, 1970) (imbrimembra, miliaria, and Thy-
sanota) as probable adult representatives of the mystery tadpole. Signifi-
cantly, the tadpole of no member of this distinctive group of large frogs has
ever been described. The group as currently understood contains 5 species:
echinata of southern Mexico; fimbrimembra of Costa Rica; miliaria of Nic-
aragua, Costa Rica, Panama and Colombia; thysanota of eastern Panama;
and valancifer of southeastern Mexico and Guatemala. Only the 2 Costa
Rican species seem reasonable candidates for association with the Cinchona
tadpole.

Of the 2 forms, it is clear that the tadpole almost certainly is H. fimbri-
membra. This species is known from only 2 specimens (Duellman, 1970) an
adult female (the holotype) and a juvenile male (the holotype of the name
H. richardtaylori). Both examples were collected in the Isla Bonita—Cin-
chona area (Savage, 1974), a few kilometers apart, by Edward H. Taylor
(1948). The tadpole subject of this paper was collected virtually at the type
locality of fimbrimembra. Hyla miliaria, on the other hand, is known only
from one locality, at 600 m elevation in Costa Rica.

Adult members of the miliaria group are very large (known to reach 106
mm in male miliaria) frogs with immense hands and feet that are extensively
webbed. In addition the outer margins of the hands, forearms, and feet have
extensive fleshy fringes. Known adult males have the prepollex modified as
a projecting spine (miliaria), a spadelike plate (valancifer), or clump of
spines (echinata). Duellman (1970) has described and illustrated the use of
the hands, feet, webs and fringes in the flight, or more accurately glide, in
an example of miliaria. All known juveniles of the group have a tuberculate
dorsum and the unique and extensively tuberculate body of the tadpole here

VOLUME 93, NUMBER 4 1183

associated with H. fimbrimembra is another confirmation of the correctness
of this allocation. I have seen no other hylid tadpole from Mesoamerica
having this condition.

Certainly adults of the miliaria stock are among the rarest and most bi-
zarre of New World tree-frogs. The tadpole of H. fimbrimembra is quite
ordinary and shares basic character combinations with many other hylids.
This situation offers support for Starrett’s Rule which states that the most
specialized and uniquely modified tadpoles develop into ordinary frogs
whereas the most bizarre and distinctive frogs have ordinary, generalized
tadpoles.

Acknowledgments

I wish to thank Norman J. Scott, the John Simon Guggenheim Memorial
Foundation, and the Organization for Tropical Studies (OTS) for aid in the
fieldwork that made this paper possible. Frances Runyan prepared the tad-
pole drawings and I greatly appreciate her contribution to the effectiveness
of the account.

Finally, I wish especially to acknowledge the advice on, and insight into,
tadpole biology provided over the past 20 some years by my colleague, the
discoverer of Starrett’s Rule, Dr. P. H. ‘“‘Holly”’ Starrett.

Literature Cited

Duellman, W. E. 1970. The hylid frogs of Middle America.—Monogr. Mus. Nat. Hist. Univ.
Kansas 1(2 vols.):xii—754.

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae.—Herpetolo-
gica 16(3):183-190.

Kluge, A. G. 1979. The gladiator frogs of Middle America and Colombia—a reevaluation of
their systematics (Anura: Hylidae).—Occ. Pap. Mus. Zool. U. Mich. 688:1-24.
Savage, J. M. 1967. A new red-eyed tree-frog (family Hylidae) from Costa Rica, with a review

of the Hyla uranochroa group.—Bull. So. Calif. Acad. Sci. 67(1):1-20.

. 1968. The dendrobatid frogs of Central America.—Copeia 1968(4):745-776.

. 1972. The systematic status of Bufo simus O. Schmidt with description of a new toad

from western Panama.—J. Herp. 6(1):25-33.

. 1974. Type localities for species of amphibians and reptiles described from Costa

Rica.—Rev. Biol. Trop. 22(1):71-122.

, and A. G. Kluge. 1961. Rediscovery of the strange Costa Rica toad, Crepidius epi-
oticus Cope.—Rev. Biol. Trop. 9(1):39-S51.

Silverstone, P. A. 1975. A revision of the poison-arrow frogs of the genus Dendrobates

Wagler.—Sci. Bull. Nat. Hist. Mus. Los Angeles Co. 21:1-—S55.

. 1976. A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra

(Family Dendrobatidae).—Sci. Bull. Nat. Hist. Mus. Los Angeles Co. 27:1-S3.

Taylor, E. H. 1948. Two new hylid frogs from Costa Rica.—Copeia 1948(4):233-238.

Department of Biological Sciences and Allan Hancock Foundation, Uni-
versity of Southern California, Los Angeles, California 90007.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1184-1203
COMMON INSHORE BARNACLE CYPRIDS OF THE
OREGONIAN FAUNAL PROVINCE
(CRUSTACEA: CIRRIPEDIA)!

Jon D. Standing

Abstract.—Seven species of barnacle cyprids from field collections are
illustrated, described, and compared. Cyprids described in detail are Polli-
cipes polymerus, Chthamalus dalli, Balanus glandula, Balanus nubilus?,
and Semibalanus cariosus. Previous descriptions of the cyprids of Balanus
crenatus and Balanus improvisus are amplified. Ecological notes and a key
to the species are also given.

The sessile stages of barnacles, their systematics and biology, have re-

ceived considerable study for more than a century. Yet our knowledge of !

barnacle larvae has advanced slowly, despite their prominence in inshore
waters, and their importance in unifying the four orders of Cirripedia. This
situation has prevailed for two main reasons. First, larvae have been re-
garded historically as mere developmental stages in the life history of the
all-important adult. Fortunately, many biologists now view them as func-
tional, independent animals having important ecological roles, and deserving
of more detailed study.

Second, barnacle larvae, especially cyprids, have proved difficult to dis-
tinguish and to identify (Pyefinch, 1948; Costlow and Bookhout, 1958;
Moyse, 1961; Crisp, 1962; Barker, 1976; Lang, 1979). Specimens collected

from plankton samples are not easily linked to known adults, and laboratory-

reared cyprids frequently differ from their ‘‘wild’’ counterparts in size and
shape (Freiberger and Cologer, 1966; Lang, 1979). Workers dealing with
only one or two species have usually lacked the comparative material nec-
essary for good descriptions, and most studies on barnacle larvae have
focused on the naupliar stages rather than on the cyprids. Thus, illustrations
and measurements of cyprids are fairly common in the literature, but full
descriptions are rare.

This paper describes seven species of cyprids from the Oregonian Prov-
ince. These are Pollicipes polymerus, Chthamalus dalli, Balanus glandula,
Balanus crenatus, Balanus improvisus, Balanus nubilus?, and Semibalanus
cariosus. Although some of these species have been illustrated or briefly

1 Part of a doctoral dissertation to be submitted to the Department of Zoology, University
of California, Berkeley.

VOLUME 93, NUMBER 4 1185

described previously, they are included here either because earlier accounts
do not compare them to other Oregonian forms, or because this study has
revealed characters not observed before. Also included here are notes on
identification, distribution, and abundance, and a key to the species.

Materials and Methods

Unless otherwise noted, cyprids were collected in Bodega Harbor, Cali-
fornia, from April 1975 to November 1978. Bodega Harbor is a small, par-
tially closed embayment located about 100 km north-northwest of San Fran-
cisco. It has coastal water conditions and considerable tidal exchange. The
collection site was the U.S. Coast Guard wharf, near the mouth of the
harbor. Of the cyprids examined from Bodega Harbor, 99.93% were refer-
able to the species described here (excepting Balanus improvisus and Sem-
ibalanus cariosus); 0.07% were of uncertain identity.

Samples were taken with a gasoline-driven pump, fitted with an intake
hose, and filtered through a 158 wm mesh plankton net. In 1976 and 1977,
cyprids were collected at three depths: 30 cm above the bottom (depth 3-
5 m, depending on tidal level), 15 cm below the surface, and midway be-
tween the surface and bottom. Volumes sampled at each depth were 1.0 m°.
Samples were collected halfway between the predicted times of lower low
water and higher high water levels on flood tides. Sampling frequency was
about every eight days. Cyprid densities are based on samples collected
from October 1976 to November 1977. Adult distributions are modified from
Newman (1979a).

Identification of cyprids was accomplished in several different ways.
First, some identifications were based primarily on laboratory rearing: either
cyprids were raised from embryos taken from identified adults, or field-
collected cyprids were induced to settle and grow into identifiable juveniles.
Second, spatial and temporal coincidences of large numbers of cyprids and
newly settled juveniles were frequently observed in the field. The cyprids
were later identified by following cohorts of juveniles through time until
they became identifiable. Third, some cyprids could be identified by com-
paring them to illustrations or descriptions in the literature, or by comparing
them to cyprids identified in other parts of the world. Finally, considerable
information on the local and geographical distributions of adults was avail-
able. However, caution must be exercised in inferring cyprid identifications
from large-scale distributions of adults. Identifications based on several dif-
ferent kinds of evidence are likely to be most reliable.

Cyprids were studied with dissecting and compound microscopes for de-
scriptive characters. Special attention was directed towards taxonomic fea-
tures not requiring high power microscopy and microdissection. These fea-
tures were carapace shape, size, and sculpturing, together with special

1186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON |

pigmentation patterns. Shape and size characters were initially observed by |
carefully aligning the body in side and dorsal views. This alignment was
accomplished by sticking the animal to a dish with a tiny dab of silicone |
grease, and then orienting it so that the body was as horizontal as possible |
in dorsal view, and so that the compound eyes were superimposed in side
view. Size measurements were made with a dissecting microscope fitted |
with an ocular micrometer and were representative of all cyprids collected
in 1977, unless otherwise indicated. The measurements taken were length—
from anterior to posterior carapace margins, depth—greatest distance be-
tween dorsal and ventral carapace margins, and width (breadth)—greatest
distance between one side and the other. Special pigmentation characters,
as referred to here, are those that appear to differ from general ground color
and from color that may be associated with specific organs. These characters
were observed on freshly killed cyprids, as well as formalin-preserved ones,
because pigmentation characters sometimes fade in preservatives. Obser-
vation of sculpturing characters required the high dry magnification of a
compound microscope. The drawings were made with the aid of a camera
lucida. 3

Voucher specimens of the species described here have been placed in the
National Museum of Natural History (USNM 181245—181251).

Subclass Cirripedia
Order Thoracica
Suborder Lepadomorpha
Family Scalpellidae
Subfamily Pollicipinae Zevina, 1978
Pollicipes polymerus Sowerby, 1833 (=Mitella polymerus)
Fig. 1

Distribution.—Adults: lower midtidal and low intertidal zones of wave-
swept shores; on rocks and hard-shelled animals. Cyprids: throughout water
column, mean density 10 m3; mainly in fall and winter. British Columbia
south to Cape San Lucas, Baja California.

Identification.—(1) Cyprids reared in the laboratory from embryos, using
the methods of Lewis (1975). (2) Juveniles and adults present on floating
pier adjacent to Bodega Harbor collection site. (3) Cyprid illustrations in
Broch (1922) and Lewis (1975), although Lewis’s drawing does not resemble
P. polymerus. Naupliar stages described and figured in Lewis (1975).

Diagnosis.—Small, mean length 494 um. Translucent when fresh. Rela-
tively translucent when preserved, with light orange cement glands pos-
teroventral to compound eyes. No special pigmentation. Carapace profile
relatively high in side view; anterior end broadly rounded; posterior end
narrowly rounded; anterodorsal margin broadly curved; posterodorsal mar-

VOLUME 93, NUMBER 4 1187

Table 1.—Sizes of barnacle cyprids in wm. Depth is the measurement between the dorsal
and ventral carapace margins; width, the measurement between sides. Data are expressed as

means; ranges are given in parentheses.

Species Collection site Length Depth Width n

Pollicipes polymerus Bodega Harbor, 494 256 177. 50
California (420-520) (220-260) (160-200)

Chthamalus dalli Bodega Harbor, 529 270 230 50
California (440-580) (240-300) (200-260)

Balanus glandula Bodega Harbor, 728 388 309 50
California (640-780) (320-440) (240-360)

Balanus crenatus Bodega Harbor, 852 374 328 50
California (700-960) (300-440) (260-400)

Balanus crenatus Point Barrow, 1,001 473 450 67
Alaska (900-1,060) (400-520) (380-520)

Balanus improvisus San Francisco Bay, 584 304 285 25)
California (540-610) (270-340) (250-320)

Balanus nubilus? Bodega Harbor, 932 431 345 50
California (800-1,000) (360-480) (280-400)

Semibalanus cariosus San Juan Island, 1,111 499 493 21
Washington (960-1,200) (420-560) (440-560)

gin curved, breaking about two-thirds of the way towards posterior end;
ventral margin broadly curved anterior to, and relatively straight posterior
to, compound eyes. Carapace profile narrowly fusiform in dorsal view, mean
width 177 wm; both ends broadly rounded. Entire carapace sculptured with
uniform, rounded, regularly-spaced papillae (about 4.5 um in diameter and
1.5 um high) and occasional spine-like processes (about 7 wm high).

Size variation.—Mean length 494 um, range 420 to 520 wm at Bodega
Harbor, California (Table 1).

Remarks.—This cyprid is the smallest one discussed here, although its
size overlaps broadly with that of Chthamalus dalli. It differs from the
cyprids of C. dalli, Balanus improvisus, and most other species in having
a broadly rounded anterior end in side view, a break in the posterodorsal
margin, a narrow carapace profile in dorsal view, and a carapace sculptured
with small papillae. The greatest depth of the carapace is at about the level
of the compound eyes, rather than considerably behind this level as in the
other species. The cyprid of P. polymerus is similar to the somewhat larger
cyprid of B. improvisus in having a break in the posterodorsal margin, but
differs in having the carapace sculptured with papillae rather than pits. The
latter two species are also spatially separated, co-occurring mainly around
the mouths of some enclosed bays.

1188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

500 jum

Figs. 1-4. Lateral (a) and dorsal (b) views of barnacle cyprids. 1, Pollicipes polymerus; 2,
Chthamalus dalli; 3, Balanus improvisus; 4, Balanus glandula. Insets show carapace sculp-
turing; pigmented areas are stippled. Straight arrows indicate some diagnostic characters used
inthe key. |

Suborder Balanomorpha
Superfamily Chthamaloidea
Family Chthamalidae
Subfamily Chthamalinae
Chthamalus dalli Pilsbry, 1916
Fig. 2

Distribution.—Adults: intertidal zone of exposed and protected shores; |
on rocks, pier pilings, and hard-shelled animals. Cyprids: surface and mid- |
depths mainly, mean density 11 m~*; mainly in summer and fall. Northern
Japan and Alaska south to San Diego, California.

VOLUME 93, NUMBER 4 1189

Identification.—(1) Juveniles reared in the laboratory from field-collected
cyprids. (2) Adults present on pilings adjacent to collection site. Naupliar
stages have not been described or figured.

Diagnosis. —Small, mean length 529 um. Translucent when fresh, with
paired, dark areas located about halfway between compound eyes and pos-
terior end. Relatively translucent when preserved, but more opaque ante-
riorly; with prominent light brown cement glands posteroventral to com-
pound eyes. No special pigmentation. Carapace profile relatively high in
side view and peaked dorsally at midlength; anterior end narrowly rounded
to angular; posterior end narrowly rounded; anterodorsal margin slightly
curved; posterodorsal margin evenly curved; ventral margin curved anterior
to, and slightly curved posterior to, the compound eyes. Carapace profile
in dorsal view broadly fusiform, with both ends rounded. Carapace smooth,
unsculptured.

Size variation.—Mean length 529 um, range 440 to 580 um at Bodega
Harbor, California (Table 1).

Remarks.—This cyprid is most likely to be confused with those of P.
polymerus and B. improvisus, which it overlaps in size. Unlike P. poly-
merus, the carapace profile peaks well behind the compound eyes in side
view and is relatively wide in dorsal view. The carapace is glassy smooth,
lacking the sculpturing of both P. polymerus and B. improvisus cyprids. In
side view the cyprid of C. dalli also differs from that of B. improvisus in
having a more angular anterior end and an evenly curved posterodorsal
margin with no break. C. dalli is also probably similar to Chthamalus fissus,
but the cyprid of the latter species is undescribed at present.

Superfamily Balanoidea
Family Balanidae
Subfamily Balaninae Newman, 1979b
Balanus improvisus Darwin, 1854
Fig. 3

Distribution.—Adults: low intertidal zone and subtidal areas in estuaries
and some enclosed bays; particularly tolerant of brackish water; on rocks,
pilings, and hard-shelled animals. Cyprids: mainly low in water column in
estuaries and some enclosed bays (Bousfield, 1955; Mohammad, 1961); sum-
mer, also spring and fall (Weiss, 1948; Jones and Crisp, 1954; Blom, 1965;
Sneli, 1972). Introduced into the North Pacific; Columbia River, Oregon
south to the Salinas River, California; occasionally in harbors south of Point
Conception; Ecuador, Japan, Australia, North and South Atlantic.

Identification.—(1) Presence of large numbers of cyprids' and adults in
low salinity (3.4%) waters in San Francisco Bay estuarine system (Newman,
1967; personal observation). (2) Comparison with laboratory-reared B. im-

1190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

provisus cyprids from Rhode Island. (3) Cyprid illustrations and descriptions
in Buchholz (1951), Doochin (1951), and Jones and Crisp (1954); photograph
in Freiberger and Cologer (1966). Naupliar stages described and figured in
Buchholz (1951), Jones and Crisp (1954), and Lang (1979).

Material examined.—San Francisco Bay, California (October 1978).

Diagnosis.—Small, mean length 584 um. Relatively opaque when pre-
served, especially at anterior end. No special pigmentation in preserved
specimens. Carapace profile relatively high in side view; anterior end round-
ed; posterior end narrowly rounded; anterodorsal margin curved; postero-
dorsal margin curved, breaking just in front of the posterior end; ventral
margin curved anterior to, and slightly curved posterior to, the compound
eyes. Extreme posteroventral margin of carapace finely crenulated. Cara-
pace profile wide in dorsal view, mean width 285 u~m; with both ends round-
ed. Carapace sculptured with rounded pits (about 4 wm in diameter).

Size variation.—Mean length 584 um, range 540 to 610 um in San Fran-
cisco Bay, California (Table 1); mean length 587 um in Biscayne Bay, Flor-
ida (Doochin, 1951), 523 ~m on the east coast of England (Jones and Crisp,
1954), 514 wm in the Netherlands (de Wolf, 1973).

Shape variation.—Sometimes the anterior end is narrowly rounded in side
view, and both ends are broadly rounded to truncate in dorsal view.

Remarks.—This cyprid is most similar to that of C. dalli but could also
be mistaken for those of B. glandula or P. polymerus. It differs from the
cyprid of C. dalli in having a more rounded anterior end in side view, a
posterodorsal margin that breaks just in front of the posterior end, and a
carapace sculptured with small pits and fine crenulations, the latter along
the posteroventral margin. The sizes of B. improvisus cyprids do not gen-
erally overlap those of P. polymerus or B. glandula, and the sculpturing of
B. improvisus cyprids differs from the papillae of the latter two species. In
addition, unlike P. polymerus, the carapace peaks well behind the com-
pound eyes in side view and is considerably wider in dorsal view. Unlike
B. glandula, the anterior end is more evenly rounded in side view and less
truncate in dorsal view. The break in the posterodorsal margin also sepa-
rates the cyprid of B. improvisus from that of B. glandula. Finally, B. im-
provisus cyprids are not likely to be found along the outer coast but do
occur in the upper reaches of estuaries. Therefore, they co-occur with the
cyprids of C. dalli and B. glandula mainly in enclosed bays and in the
mouths of estuaries.

Balanus glandula Darwin, 1854
Fig. 4

Distribution.—Adults: upper and sometimes lower midtidal zone of ex-
posed and protected shores; on rocks, pier pilings, and hard-shelled animals.

VOLUME 93, NUMBER 4 1191

Cyprids: mainly high in water column; mean density 28 m-, 1,436 m~? on
26 March 1977; mainly spring, also summer and fall. Aleutian Islands south
to San Quintin Bay, Baja California.

Identification.—(1) Coincidence of large numbers of cyprids and juveniles
at collection site. (2) Adults present on pilings and floating pier adjacent to
collection site. (3) Photograph of cyprid in Strathmann and Branscomb
(1979). Naupliar stages have not been described or figured.

Diagnosis.—Medium-sized, mean length 728 um. Translucent when
fresh. Chartreuse pigmentation observable only in fresh specimens; con-
sisting of large (5—6 times the size of the compound eye) paired, irregularly
shaped areas located in posterior third of body; several small, unpaired
spots posterodorsal to compound eyes; and sometimes a pair of large (2-3
times size of compound eye) paired and fused areas anterodorsal to com-
pound eyes. Golden (or rarely brown) when preserved, with light brown
cement glands ventral to compound eyes. Carapace profile high in side view,
mean depth 388 um; both ends narrowly rounded; anterodorsal margin
slightly curved; posterodorsal margin evenly curved; ventral margin curved
both in front of and behind compound eyes, becoming relatively straight
posteriorly. Carapace profile in dorsal view fusiform, with anterior end
somewhat truncate and posterior end narrowly rounded; median raphe con-
tinuous from anterior end to posterior end. When viewed from the anterior
end, raphe is seen as a seam where sides of carapace meet at an acute angle
along dorsal margin. Entire carapace sculptured with elliptic papillae (about
7 wm long and 2 wm wide).

Size variation.—Mean length 728 um, range 640 to 780 um at Bodega
Harbor, California (Table 1); mean length 790 um, range 670 to 860 um on
San Juan Island, Washington (Strathmann and Branscomb, 1979).

Remarks.—This cyprid is medium-sized, ordinarily overlapping in size
only with Balanus crenatus. In side view it differs from the cyprid of B.
crenatus and most other species in having a high carapace profile and a
ventral margin curved well behind the compound eyes. Other distinguishing
features include an anterior end somewhat truncate in dorsal view, a dorsum
acutely angled rather than rounded when viewed from the anterior end, and
a carapace sculptured with papillae. Also distinctive are the chartreuse-
pigmented areas in fresh specimens and the golden (or rarely brown) ground
color in preserved ones. Most other species are more or less white when
preserved. Unlike Balanus nubilus? and B. improvisus cyprids, the cara-
pace of B. glandula, in side view, has a narrowly rounded anterior end and
an evenly curved posterodorsal margin. Although considerably larger than
P. polymerus, the cyprid of B. glandula is similar in having the carapace
sculptured with papillae. However, the papillae are elliptic rather than
rounded.

192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Balanus crenatus Brugiére, 1789
Fig. 5

Distribution.—Adults: low intertidal zone and especially subtidal areas to
depths of 182 m; exposed and protected shores; on rocks, pier pilings, hard-
shelled animals, and sometimes seaweeds. Cyprids: mainly low in the water
column; mean density 92 m~, 5,909 m~ on 26 March 1977; summer, fall,
and especially spring. Northern Japan and Alaska south to Santa Barbara,
California; North Atlantic.

Identification.—(1) Juveniles reared in the laboratory from field-collected
cyprids. (2) Coincidence of large numbers of cyprids and juveniles at col-
lection site. (3) Adults present on pilings and floating pier adjacent to col-
lection site. (4) Cyprid descriptions and illustrations in Herz (1933) and
Pyefinch (1948). Naupliar stages described and figured in Herz (1933) and
Pyefinch (1948, 1949).

Material examined.—Mostly from Bodega Harbor, California; some from
Point Barrow, Alaska (August, 1976).

Diagnosis.—Medium-sized, mean length 852 um. Translucent when
fresh. Relatively opaque (white) when preserved, especially at anterior end.
A single pair of black pigment spots, somewhat smaller than compound
eyes, located just posterior to midlength, and persisting after preservation.
Carapace profile low in side view, mean depth 374 um; anterior end nar-
rowly angular; posterior end narrowly rounded; anterodorsal margin only
slightly curved; posterodorsal margin evenly curved; ventral margin curved
anterior to, and relatively straight posterior to, compound eyes. Carapace
profile in dorsal view fusiform, with both ends narrowly rounded. Carapace
smooth, unsculptured.

Size variation.—Quite variable, mean length 852 um, range 700 to 960
jm at Bodega Harbor, California (Table 1); mean length 1,001 um, range
900 to 1,060 um at Point Barrow, Alaska (Table 1); mean length 946 um,
range 734 to 1,102 um in Scotland (Barnes, 1953); mean length 643 um in
the Netherlands (de Wolf, 1973).

Remarks.—This cyprid is most similar to that of B. nubilus?, a slightly
larger form. It differs in having a narrowly angular anterior end in side view
and an evenly curved posterodorsal margin. The cyprids of B. glandula and
Semibalanus cariosus also overlap B. crenatus in size. However, the cyprid
of B. crenatus has the ventral margin relatively straight behind the com-
pound eyes and a smooth carapace, unlike B. glandula, and the anterior
end is narrowly rounded in dorsal view, unlike both of these other species.
It also differs from the cyprid of S. cariosus in lacking a break in the pos-
terodorsal margin and in having a carapace profile of medium width in dorsal
view. The cyprid of B. crenatus is distinguished from all of the above-
mentioned species by having a low carapace profile in side view and a pair
of black pigment spots just posterior to midlength.

|

VOLUME 93, NUMBER 4 1193

5a
5b
Ve
i
Wa
6a
6b

500 ym

Figs. 5-6. Lateral (a) and dorsal (b) views of barnacle cyprids. 5, Balanus crenatus; 6,
Balanus nubilus?. Pigment spots are stippled. Arrows indicate some diagnostic characters used
in the key.

Balanus nubilus? Darwin, 1854
Fig. 6

Distribution.—Adults: low intertidal zone and subtidal areas to depths of
90 m; exposed and protected shores; on rocks, pier pilings, and hard-shelled
animals. Cyprids: mainly low in water column; mean density 6 m~°; spring,
summer, and fall, especially summer. Southern Alaska south to La Jolla,
California.

Identification.—(1) Juveniles reared in the laboratory from field-collected
cyprids. (Although identification of the juveniles was not completely certain,
the presence of large numbers of this cyprid as far north as Crescent City,

1194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

California suggests that the correct identification is B. nubilus rather than
the more southern form Balanus aquila.) (2) Adults present on pilings and
floating pier adjacent to collecting site. Naupliar stages described and fig-
ured in Barnes and Barnes (1959).

Materials examined.—Mostly from Bodega Harbor, California; some
from Moss Landing, California and Crescent City, California.

Diagnosis.—Medium-sized, mean length 932 um. Translucent when
fresh. Relatively translucent when preserved, although more opaque ante-
riorly and posteriorly; with light orange cement glands posteroventral to
compound eyes. No special pigmentation. Carapace profile relatively low
in side view; anterior end rounded; posterior end narrowly rounded; antero-
dorsal margin curved; posterodorsal margin curved, breaking slightly about
three-fourths of the way towards posterior end; ventral margin curved an-
terior to, and nearly straight posterior to, compound eyes. Carapace profile
in dorsal view fusiform, with both ends narrowly rounded. Carapace
smooth, unsculptured.

Size variation.—Mean length 932 wm, range 800 to 1,000 um at Bodega
Harbor, California (Table 1).

Remarks.—This cyprid is most similar to that of B. crenatus. It differs,
in side view, in having a break in the posterodorsal margin and a more
rounded anterior end. This latter character also distinguishes the cyprid of
B. nubilus? from that of S. cariosus, which overlap each other in size. In
addition the cyprid of B. nubilus? has a relatively narrow width and a nar-
rowly rounded anterior end in dorsal view, unlike S. cariosus. All of these
characters, together with the smooth carapace of B. nubilus?, separate it
from the cyprid of B. glandula, a smaller species.

Family Archaeobalanidae Newman and Ross, 1976
Subfamily Semibalaninae Newman and Ross, 1976
Semibalanus cariosus (Pallas, 1788) (=Balanus cariosus)
Fig. 7

Distribution.—Adults: lower midtidal and sometimes low intertidal zones
of wave-exposed shores; on rocks and hard-shelled animals. Cyprids: prob-
ably low in water column; mainly spring and summer (Connell, 1970; Day-
ton, 1971; Strathmann and Branscomb, 1979). Japan, Bering Sea, and Alas-
ka south to Morro Bay, California.

Identification.—(1) Coincidence of large numbers of cyprids and juveniles
(R. R. Strathmann, pers. comm.). (2) Photograph of cyprid in Strathmann
and Brarscomb (1979). Naupliar stages have not been described or figured.

Material examined.—San Juan Island, Washington (May 1976).

Diagnosis.—Large, mean length 1,111 wm. Very opaque when preserved,
except for translucent area along dorsum. No special pigmentation in pre-

———_ > ——

VOLUME 93, NUMBER 4 1195

ke ~

500 um

Fig. 7. Lateral (a) and dorsal (b) views of Semibalanus cariosus. Arrows indicate some
diagnostic characters used in the key.

served specimens. Carapace profile relatively low in side view; anterior end
broadly angular; posterior end narrowly rounded; anterodorsal margin
curved, breaking somewhat just behind anterior end; posterodorsal margin
curved, breaking prominently just in front of posterior end; ventral margin
curved anterior to, and slightly curved posterior to, compound eyes. Car-
apace profile wide in dorsal view, mean width 493 ym; anterior end truncate;
posterior end narrowly rounded. Compound eyes small and deeply inset.
Carapace smooth, unsculptured.

Size variation.—Mean length 1,111 um, range 960 to 1,200 um at San
Juan Island, Washington (Table 1); a larger sample (N = 80) from San Juan
Island had a mean length of 1,140 um, range 980 to 1,240 um (Strathmann
and Branscomb, 1979).

Remarks.—This cyprid is the largest one considered here, although its
size overlaps with that of B. nubilus? and B. crenatus. It differs from the
former species in having a broadly angular anterior end in side view and in
having a wide carapace profile, a truncate anterior end, and relatively small,
deeply inset compound eyes, all visible in dorsal view. These characters,

1196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

together with the breaks in the anterodorsal and posterodorsal margins of
S. cariosus, serve to distinguish it from the cyprid of B. crenatus.

Discussion

Over 50 species of thoracican cyprids have been previously illustrated or
described, mostly from the North Atlantic, India, and Australasia (cf. Nils-
son-Cantell, 1921, 1978; Lang, 1979). The present paper describes seven
species of cyprids from the Oregonian Province of the temperate Northeast
Pacific region. They include most of the intertidal barnacles characteristic
of this province, along with some of the more subtidal forms. The ranges
of most of these species extend into the adjoining Aleutian and Californian
Provinces, where still other species occur. Nevertheless, the descriptions
given here will account for most of the intertidal species encountered in the
Aleutian and Californian Provinces; they will be less useful for the subtidal
species.

The taxonomic features studied here were size, shape, special pigmen-
tation, and sculpturing. Size is usually expressed in the descriptive literature
as carapace length; carapace depth appears occasionally, width rarely. Cy-
prids range in length from less than 100 wm (Bocquet-Védrine, 1961) to
more than 2,200 um (Standing, personal observation). The very smallest
cyprids, in general, are rhizocephalans, while lepadids and some scalpellids
tend to be very large (Nilsson-Cantell, 1921; Zevina, 1971). Other scalpellids
are considerably smaller. For example, Calantica spinosa and Pollicipes
polymerus have lengths of 850 um and 494 wm, respectively (Batham, 1946b;
the present study). The trends in the Balanomorpha are as follows: chtham-
aloidean cyprids are small (less than 625 um), but not as small as rhizoce-
phalans; coronuloideans are mostly small to medium sized; and balanoi-
deans range from small to large, although Chirona hameri is very large,
averaging 1,454 wm (Crisp, 1962). The smallest and largest balanoidean cy-
prids observed in the present study were Balanus improvisus and Semibal-
anus cariosus, respectively (Table 1).

Cyprid size is quite variable, even within a single species. In the labora-
tory, size varies inversely with rearing temperature (Lang, 1979), and in the
field, size often varies directly with latitude (Pyefinch, 1948; Crisp, 1962).
This size-latitude phenomenon, probably involving both temperature and
food supply, also applies to barnacle nauplii and embryos (Barnes and
Barnes, 1965), as well as many other marine animals. The present study
provides additional evidence of the size-latitude phenomenon. Balanus
crenatus cyprids from Point Barrow, Alaska, were larger than those from
Bodega Harbor, California (Table 1), and B. glandula cyprids from San
Juan Island, Washington (Strathmann and Branscomb, 1979) were larger
than those from Bodega Harbor.

————————

VOLUME 93, NUMBER 4 1197

Some species of cyprids in the present study overlapped considerably in
size (Table 1). For this reason, size is best considered a taxonomic feature
of secondary importance. However, it is useful for initially distinguishing
between groups of species and for helping to recognize species after initial
identification. Carapace width was a good character for separating Pollicipes
polymerus and Semibalanus cariosus cyprids from other species of similar
length.

Carapace shape characters, unlike size, proved to be particularly impor-
tant for discriminating between species. Therefore, shape characters have
been described in detail and illustrated carefully, operations not always
accomplished in previous studies. Still, the existing literature does suggest
the variation to be expected in shape characters, and some general trends
in cyprid shapes.

Side views of shape characters are especially revealing. The anterior ends
of cyprids vary from pointed or angular through rounded to truncate.
Anterodorsal margins are usually inflated to various extents but can be
depressed. Pollicipes polymerus has a broadly rounded anterior end and a
broadly curved anterodorsal margin, resembling Calantica (formerly Polli-
cipes) spinosa but not Capitulum (formerly Pollicipes) mitella (Yasugi,
1937; Batham, 1946b). Also similar in these respects are some other scal-
pellids and most lepadids (Darwin, 1851; Aurivillius, 1894; Nilsson-Cantell,
1921; Broch, 1931; Zevina, 1971). Chthamalus dalli has an angular anterior
end similar to that of some other chthamalids (Barker, 1976; Karande and
Thomas, 1976; Lang, 1979), but it is unclear whether or not this character
is representative of the group. The break in the anterodorsal margin of
Semibalanus cariosus cyprids is a character different from any other illus-
trated in the literature.

Posterior ends of cyprids range from sharply pointed through narrowly
rounded to broadly rounded; posterodorsal margins are usually inflated to
varying degrees but can be depressed. Some species of cyprids have angular
(i.e., “‘broken’’) posterodorsal margins: Pollicipes polymerus, Balanus im-
provisus, B. nubilus?, and Semibalanus cariosus in the present study. The
literature suggests that this character is more common in lepadomorphs
(Aurivillius, 1894; Broch, 1931; Anderson, 1965; Standing, personal obser-
vations) than in balanomorphs (Buchholz, 1951; Moyse, 1961), but it is also
likely that the character has frequently been overlooked in previous studies.
Another interesting character on the posterior end is the finely crenulated
margin of Balanus improvisus, a character similar to that present in the
acrothoracican Trypetesa nassaroides (Turquier, 1967). This character is
reported here for the first time in B. improvisus from San Francisco Bay
(and from Rhode Island), even though the cyprid has been illustrated many
times before. It is not yet clear whether this character is representative of

1198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the species and previously overlooked, or present in only some populations.
The existing illustrations of this species suggest that other characters may
be quite variable as well.

Ventral margins also differ between species. They may be straight through
curved to recurved. Most species have convexly curved ventral margins.
However, Pollicipes polymerus cyprids and those of some other lepado-
morphs have relatively straight ventral margins (Darwin, 1851; Aurivillius,
1894; Broch, 1931; Batham, 1946b; Zevina, 1971).

Dorsal views of cyprids show fewer shape characters than side views.
Nevertheless, some of these characters have taxonomic value. Particularly
interesting are anterior ends, which can vary from angular through rounded
to truncate. Most species have rounded anterior ends, but those of Balanus
glandula and Semibalanus cariosus are nearly truncate. The curvature of
the sides and the shape of the posterior end also vary in cyprids, but these
characters appear to have less taxonomic value than some of the other
characters, at least in balanomorphs.

Carapace sculpturing characters are infrequently referred to in the de-
scriptive literature. The present study suggests that they may be more wide-
spread than is generally believed, but often overlooked. At present a number
of lepadomorphs are known to have parallel carinae along the length of the
carapace (Darwin, 1851; Broch, 1931; Batham, 1946a, b; Zevina, 1971;
Standing, unpublished observations), and ‘‘brickwork’’ sculpturing has been
described for bla cumingi (Karande, 1974). In addition, Semibalanus bal-
anoides has tiny pits (Walker and Lee, 1976), which appear to be similar to
those observed for the first time here on Balanus improvisus from San
Francisco Bay (but not on specimens from Rhode Island). However, the
papillae described here for Pollicipes polymerus and Balanus glandula cy-
prids are different from any sculpturing previously described, including the
papillae that occur on the ascothoracican Ulophysema oresundense (Bratt-
strom, 1948) and the ‘‘peg-plates’’ present on some acrothoracicans (Tom-
linson, 1969). Some acrothoracican and rhizocephalan cyprids also have the
carapace ornamented with large spines, but these are very different from
the small spine-like processes seen on Pollicipes polymerus. Clearly, much
additional work is needed to fully assess the importance of carapace sculp-
turing and ornamentation to cyprid taxonomy. SEM studies, for example,
might reveal ultramicroscopic sculpturing in species that appear smooth
under the light microscope.

Special pigmentation characters, like sculpturing characters, are not often
mentioned in the literature, probably because most species lack them. The
positions of pigment spots or areas seem to vary widely in different species,
but in general, cyprids with special pigmentation can be conveniently divid-
ed into two color classes: those having bright yellow or orange areas of
pigmentation (Batham, 1946b; Walley, 1969; Lang, 1976) and others having

VOLUME 93, NUMBER 4 1199

dark purple or black pigmentation (Pillai, 1958; Barker, 1976). In the present
study, chartreuse areas were observed in fresh Balanus glandula cyprids,
and black spots were present on B. crenatus. The pigmentation in B. glan-
dula cyprids appears to be similar to the ‘‘yellow cells’’ found in Semiba-
lanus balanoides (Walley, 1969); the spots in B. crenatus cyprids may be
most similar to the ‘‘black patches’’ described for Balanus amphitrite am-
phitrite (Pillai, 1958), and also present on B. improvisus from Rhode Island
(Standing, personal observation). The special pigmentation observed in the
cyprids of B. glandula and B. crenatus has not been previously reported,
although both species have been briefly described before.

Distributional information can also be useful in larval identification, but
it must be interpreted carefully. Cyprids are likely to be found outside the
geographical ranges of adults only occasionally, and inshore species are not
often collected far out to sea, at least in the Oregonian Province. A few
species may be found in special situations where other species are not to be
expected. Balanus improvisus cyprids, for example, occur in very low sa-
linities that probably cannot be tolerated by other species. Although they
have been collected in bays and estuaries on this coast, they were not taken
at Bodega Harbor and are probably not ordinarily present along the outer
coast. Another cyprid not collected at Bodega Harbor was Semibalanus
cariosus, although adults are present in the area. The adults of this species
become uncommon south of San Francisco, and recruitment may be infre-
quent in central California. Vertical distributions in the water column and
seasonal abundances of cyprids also differ, but identifications should not be
based on this information because of considerable overlap between species.
The ecological data collected in the present study will be presented in great-
er detail elsewhere.

In conclusion, the present work has shown considerable differences be-
tween some species of cyprids present in the Oregonian Province. Appli-
cation of microdissection and electron microscope techniques will undoubt-
edly reveal further variation. Even now, the differences separating the
cyprids of Balanus improvisus, B. glandula, and B. crenatus may be greater
than those known to separate the juveniles or adults of these species. Fur-
ther study of cyprids, like nauplii (Lang, 1979), should enable us to view
the life histories and phylogenies of cirripedes in new and revealing ways.

Key to Cyprids of the Oregonian Province

[FeSimiall tenechless:tmamOl 5 00 cscs ras eeu oc ies opel a leyscn, ue ei siers) soaue = pa

— Medium to large, length greater than 625 um................... 5

2. Anterior end narrowly rounded to angular in side view; posterodor-
Sal’mareinvevenly curved (Bigs. 2a; 4a)iracci dee 25 ee cee Soe e 3

1200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

— Anterior end broadly rounded to rounded in side view; posterodorsal
margin with break in curve (Figs. la, 3a) ....................-- 4
3. Distinct golden (rarely brown) ground color when preserved, char-
treuse pigmented areas when fresh; carapace surface dull under low
magnification, sculptured with papillae under high magnification
steal hcdsirans een iis at cate atta ela a nai os Balanus glandula? (Fig. 4)
— Relatively translucent when preserved, no special pigmentation
when fresh; carapace surface shiny under low magnification, smooth
under high magnification .................. Chthamalus dalli (Fig. 2)
4. Carapace depth greatest about one-third of the way back from an-
terior end; carapace breadth narrow in dorsal view, about one-third
length; mainly outer coast.............. Pollicipes polymerus (Fig. 1)
— Carapace depth greatest about halfway back from anterior end; car-
apace breadth wide in dorsal view, about one-half length; bays and

estuaniesn yedaes hei teen ees ! Balanus improvisus (Fig. 3)

5. Anterior end narrowly angular to narrowly rounded in side view;
posterodorsal margin evenly curved (Figs. 4a, Sa) ............... 6

— Anterior end broadly angular to rounded in side view; posterodorsal
margin with break in curve (Figs. 6a, 7a) ............00 cece eeues ir |

6. One pair of black pigment spots just posterior to midlength when
fresh; white ground color when preserved, with pigment spots per- |
sisting; carapace surface shiny under low magnification, smooth un- |
der high magnification ................... Balanus crenatus (Fig. 5) |

— Several chartreuse pigmented areas when fresh; golden (rarely |
brown) ground color when preserved, with no special pigmentation :
persisting; carapace surface dull under low magnification, sculp-
tured with papillae under high magnification Balanus glandula (Fig. 4)

7. Anterior end truncate, posterior end narrowly rounded in dorsal
view; carapace breadth wide in dorsal view, about one-half length;
compound eyes small, eye capsules less than 68 wm .............

Kd A pda fh tenet muni a bury eaielieais ae awt ola ulin OM Semibalanus cariosus (Fig. 7)

— Anterior and posterior ends both narrowly rounded in dorsal view;
carapace breadth relatively narrow in dorsal view, about one-third
length; compound eyes large, eye capsules more than 68 um .....

yh Asante .ee eke ati cath eenet od eter tinea Balanus nubilus? (Fig. 6) |

Acknowledgments

I am grateful to William Newman and Ralph Smith for reading and com-
menting on the manuscript. Cadet Hand also read the manuscript and en-

2 Only rarely less than 625 um in length.

VOLUME 93, NUMBER 4 1201

couraged the study. Richard Strathmann and James Carlton provided the
material from San Juan Island and Point Barrow, respectively. Anne Hutch-
inson and William Lang gave me Balanus improvisus cyprids from San
Francisco Bay and Rhode Island, respectively. Lorraine Andrade typed the
manuscript. I thank all of these people.

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1202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Freiberger, A., and C. P. Cologer. 1966. Rearing acorn barnacle cyprids in the laboratory for
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VOLUME 93, NUMBER 4 1203

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Bodega Marine Laboratory, University of California, Bodega Bay, Cali-
fornia 94923.

Present address: Duke University Marine Laboratory, Beaufort, North
Carolina 28516.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1204-1219

A NEW SPECIES AND A NEW GENUS OF CERVINITDAE
(COPEPODA: HARPACTICOIDA) FROM THE
BEAUFORT SEA, WITH A REVISION
OF THE FAMILY!

Paul A. Montagna

Abstract.—Cervinia unisetosa n. sp. from the bathyal zone of the Beau-
fort Sea (Arctic Ocean) is unique in the Cervinia in that it possesses a
reduced fifth leg, moderate length caudal rami, and only one seta on the
bases of the mandible and the maxilliped. The discovery of the male of
Pseudocervinia magna (Smirnov, 1946) proves that the species should be
redesignated as a Cervinia as originally described. Expansicervinia glacieria
n. gen. & n. sp. from the deep-sea of the same area proves to be unique in
the Cerviniidae in three expansions; the ventral margins of second thoracic
segment; the basal segment of the antennule; and the terminal endopodite
of the second leg. The genus Stratiopontotes Soyer, 1970 is synonymized
with Ameliotes Por, 1969. Keys to the genus Cervinia and the family Cer-
viniidae are presented.

Benthic infaunal studies off the northern coast of Alaska conducted by
the Oregon State University Marine Benthic Ecology Group have yielded
many harpacticoid copepods of the family Cerviniidae Sars, Lang (Monta-
gna and Carey, 1978). Two species, Cervinia langi Montagna, 1979 and Pseu-
docervinia magna (Smirnov, 1946) have been discussed in detail elsewhere
(Montagna, 1979). More recent sampling has yielded enough specimens of
a new species of Cervinia and a new genus to warrant description.

Cervinia unisetosa n. sp. is most closely related to Cervinia tenuicauda
Brodskaya, 1963 in that the P; is much reduced, a character previously
unique in the genus. However, C. unisetosa differs from C. tenuicauda in
the length of the CR and details of the Md and Mxp.

The discovery and analysis of the male of P. magna requires that this
species be reinstated to its original designation of Cervinia magna. Descrip-
tions of the new species and the males of C. magna are presented with a
revised key to the genus.

Expansicervinia glacieria n. gen. & n. sp. is a member of the subfamily
Cerviniinae Brodskaya, 1963 and is unique in the subfamily in that the P,—

1 Contribution No. 368 from the Belle W. Baruch Institute for Marine Biology and Coastal
Research, USC.

VOLUME 93, NUMBER 4 1205

P, terminal exopodites have only two outer spines; the antennules have 8
segments and possess lateral expanded knobs on the basal segments; there
iS no rostrum; and the ventral margin of the second thoracic segment is
greatly enlarged.

The family Cerviniidae has not been revised since Brodskaya (1963).
Since then, two new genera have been added; however, as will be discussed,
Stratiopontotes Soyer, 1970 is a synonym of Ameliotes Por, 1969. In the
present text a new genus is added, and Pseudocervinia Brodskaya, 1963 is
synonymized with Cervinia Norman, 1878. A discussion of the taxonomic
decisions and a key to the subfamilies and genera is presented.

All figures were made with the aid of a camera lucida. The nomenclature
and descriptive terminology are adopted from Lang (1948, 1965) and Coull
(1977). The following abbreviations are used throughout the text, figures
and tables: R = rostrum, A, = antennule, A, = antennae, Md = mandible,
Mx! = maxillula, Mx = maxilla, Mxp = maxilliped, P,-P, = legs 1-6,
exp. = exopodite, end. = endopodite, benp. = baseoendopodite, GF =
genital field, and CR = caudal ramus. Body length measurements are from
the base of the R to the base of the CR, excluding both. CR L/W (=length
to width ratio) is measured from the inner proximal edge to the inner distal
edge for length, and the outer proximal edge for width.

Family Cerviniidae Sars, 1903; Lang, 1948
Subfamily Cerviniinae Brodskaya, 1963
Genus Cervinia Norman, 1878
Cervinia unisetosa, new species
Figs. 1-2

Cervinia sp. B. Montagna and Carey, 1978:119.

Material.—31 2? 2. Holotype, 1 2 USNM 180117 (United States National
Museum of Natural History). Paratypes 16 2 USNM 180118.

Type-locality.—Bathyal zone of the Beaufort Sea off the northern coast
of Alaska, USA (71°45.1'N, 150°35.0’W) 2,325 m, described as station 6 in
Bilyard and Carey (1979).

Description.—Female: Based on nonovigerous female 1.63 mm long.
Body typical Cervinia shape (Fig. 1), cephalothorax broadened anteriorly,
segment bearing P, distinct. Body tapers greatly through metasome, but to
lesser extent in urosome. CR divergent, longer than last 2 segments, L/W =
7.5. R triangular, fused to cephalothorax. A, extending beyond A,.

A, (Fig. 1): 7-segmented with aesthetasc on segment-3. 5 setae on terminal
segment.

A, (Fig. 1): Allobasis with 2 setae, exp. 4-segmented with 2.1.1.3 setae
respectively. Terminal segment with 3 proximal, 6 distal broad spines.

1206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Wu

_0.5 mm_,
HABITUS

Fig. 1 Cervinia unisetosa °.

VOLUME 93, NUMBER 4 1207

Fig. 2. Cervinia unisetosa °.

Md (Fig. 2): Precoxa with bidentate pars incisiva, unidentate lacina mob-
ilis, 4 additional spines, 1 seta. Coxa-basis with | inner seta. End. 1-seg-
mented with 3 inner, 5 terminal setae. Exp. 4-segmented with 1.1.1.2 setae
respectively.

Mxl (Fig. 2): Arthrite of precoxa with 9 distal spines, 4 surface setae
arranged in opposing pairs on opposite surfaces. Coxa with 6 inner setae.
Basis and end. fused with 12 setae. Exp. reduced with 3 setae.

Mx (Fig. 2): Syncoxa with 3 endites, with 2.3.2 setae respectively. Basis

1208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with 2 claws, 1 seta. End. 3-segmented with 1 spine, 1 seta; 2 spines; 3
spines, 2 setae respectively.

Mxp (Fig. 2): Not prehensile, basis with 3 claw-like spines, 1 seta asso-
ciated with innermost. End. 3-segmented with 1 claw, 1 seta; 1 seta; 2 claws,
2 setae respectively.

P, (Fig. 1), P,—P, (Fig. 2): Each ramus of all legs 3-segmented. Setation
as figured and listed below:

Exopod Endopod
P, 1.1.123 1.1.221
12, 1.1.223 e227
P, 1.1.223 e232
P e224 2221

_

P, (Fig. 1): Basis and exp. fused into broad plate which is indistinctly
separated. No inner expansions of benp., or outer seta. Exp. part much
reduced with 3 distal setae.

P, & GF (Fig. 1): Located medioventrally. P, with 3 terminal setae located
just lateral to GF. GF medial with sclerotized posterior curving ridge. Gen-
ital pore triangular but indistinct.

Male.—Unknown.

Variability.—3 specimens were dissected; all were identical except for
the following: the CR L/W varied between 7.1 and 7.9, and 1 specimen had
only 1 setae on the middle segment of the P, endopodite instead of the
normal 2.

Etymology.—The specific name unisetosa (L. 'unus' = one + L. ’seta’ =
bristle) refers to the single seta on the coxa-basis of the Md and in associ-
ation with the 3 claws of the Mxp.

Discussion.—Cervinia unisetosa is most nearly related to C. tenuicauda
based on the identical structure of the P; which is unique in the Cerviniidae
but common to both. However, the CR of C. unisetosa is only half as long
as that of C. tenuicauda, the longest in the genus. Cervinia unisetosa is
unique in the genus in 2 characters: there is only one seta on the coxa-basis
of the mandible, and there is only one seta associated with the 3 claws of
the maxilliped basis.

In a previous study of Cervinia I concluded that the setation of the swim-
ming legs is a conservative character in this genus (Montagna, 1979). How-
ever, the setation of the mandible and the maxilliped are unique among all
the species of the genus. Taxonomic decisions in the Harpacticoida usually
are not based on the details of the mouthparts. Because of their small size,
the details may be difficult to ascertain, particularly for earlier workers who
did not have the advantage of modern research microscopes. However, I
feel an exception must be made for Cervinia because of their unusually

VOLUME 93, NUMBER 4 1209

large size. The mouthparts are as large as the swimming legs (Fig. 2), and
well figured in even the earliest descriptions of the genus.

Cervinia magna Smirnov, 1946
Figs. 3-4

Pseudocervinia magna (Smirnov).—Brodskaya, 1963:1801.

Material.—9 gravid 2° 2, 492 2 2, 205 copepodites, and 75 36, collected
from the continental slope off Pitt Point, Alaska, USA, described as stations
30-34 by Montagna & Carey (1978), depth 25-100 m. Specimens of the
males have been deposited in USNM 180119.

Description.—Male: Body 1.20 mm long. Typical Cervinia male; cepha-
lothorax broadened anteriorly, R slightly broader than 2, A, with many
aesthetascs, first segment distinct, body tapered throughout, urosome with
many rows of spinules (Fig. 3). Sexual dimorphism pronounced; all char-
acters are dimorphic except A,, P,—P, exp. and Cr, which are female-like.

A, (Fig. 3): 8-segmented, asthetascs on segments 4 and 6, 2 on segment 8.

Md (Fig. 3): Precoxa greatly reduced, pars incisiva and lacina unidentate.
Coxa-basis, end. and exp. as in °.

Mxl (Fig. 3): Greatly reduced, arthrite with 6 spines, coxa with 5 setae.
Basis with 2 inner, end. represented by 6 terminal setae, exp. represented
by 2 outer setae.

Mx (Fig. 3): Basis claw transformed into shape resembling ‘bottle opener,’
with proximal seta, 1 seta on surface. End. 3-segmented with 1.2.3 setae.

Mxp (Fig. 3): Basis reduced (as large as 3 end. segments combined) with
3 claws; innermost transformed, widened and prehensile; middle claw with
1 seta. End. 3-segmented 2.1.4 setae and spines respectively.

P,—P, (Fig. 3): All rami 3-segmented, setation as listed below. Only end.
figured, P, and P, terminal end. with transformed spines.

Exopod Endopod
P, fall23 e221
P, e223 1.2.221
P, e228 1.2.321
P 1.1.223 te 2221

_

P, (Fig. 3): Setation and shape as ¢ except terminal seta greatly enlarged
and proximal seta reduced.

P, (Fig. 3): Lateral with 3 setae.

Copepodite: The last copepodite stage of C. magna resembles an adult
in that the urosome has 5 segments (Fig. 4). Just one stage younger speci-
mens have 4 urosomal segments with the last two segments combined. All
the appendages of the cephalothorax are the same as the adult °, as are the

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1210

WI ALN

HUM A ty

Md, Mxl,Mx, Mxp

Cervinia magna co.

Fig. 3.

VOLUME 93, NUMBER 4 1211

0.2 mm
VENTRAL

Fig. 4. Cervinia magna copepodite.

P, and P,. But P, and P, endopodites vary as figured (Fig. 4) with the setal
formulae being 1.521 and 1.321 respectively.

Discussion.—In my previous study of this species, I corrected several
character misdiagnoses of the female (Montagna, 1979). These character
traits were significant because they were the basis of a generic description
by Brodskaya (1963). If C. magna had possessed opposed CR as previously
thought, it would have been in the subfamily Cerviniopsinae, thus requiring
a distinct genus. However, since the CR are divergent, C. magna belongs
in the Cerviniinae (Montagna, 1979). Only one character in the genus de-
scription of Pseudocervinia remained unique; that the male had a 2-seg-
mented end. on the first leg. On that basis I previously decided that it would
be premature to declare Pseudocervinia synonymous with Cervinia since
I had not yet seen the male of the species. Though Smirnov (1946) did not
figure the male, he described the P, end. as 2-segmented, and the P,—P, end.
as 3-segmented with the setation as described above.

Considering the highly dimorphic nature of the males, I was not sure that
I had the male of C. magna. In fact, at first I thought the copepodite (Fig.
4) was the male because it was identical to the female except for the P., P,
and P, endopodite setation. Most convincing was that the urosome was
composed of five segments as in adult males. But in all 11 of these morphs
I never found spermatophores, as I did in all 75 of the true males. Further,
dissection of the morphs which undoubtedly were juveniles (i.e., with four
urosomal segments, and unfused genital segments) showed the exact struc-
ture of P,—P; found in the juvenile of Fig. 4. In true juveniles (4-segmented

1212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

urosome) the last 2 segments were always fused, whereas they are distinct
in the figured juvenile.

The evidence that the figured male described here is the male of C. magna
is very strong. Ecological evidence: In the samples that yielded the material,
781 cerviniids were found in three stations, all within 10 miles. No other
cerviniid was found in these samples, and no other is known to occur in the
sample areas (Montagna and Carey, 1978). Males were most abundant in
the fall, the only time gravid females were found. Morphological evidence:
In my previous (Montagna, 1979) and present study (see discussion of C.
unisetosa), I came to the conclusion that in the genus Cervinia the mouth-
parts and particularly the mandibular palp (coxa-basis, end., and exp.) and
the Mxp basis setation were the most conservative and taxonomically im-
portant features in distinguishing the species. The male is consistent with
this hypothesis. Though all legs and mouthparts are dimorphic, only the
palp of the mandible and setation of the maxilliped are exactly as in the
female of C. magna (the female is figured in Montagna, 1979).

Given that the male described here is the male of C. magna, consistent
with the definition of Cervinia (Lang, 1948) and inconsistent with the defi-
nition of Pseudocervinia, C. magna is restored its original designation and
Pseudocervinia 1s a synonym to Cervinia.

Key to the Species of Cervinia

Based on females. Cervinia brevipes Brodskaya, 1963, cannot be placed
in the key since its swimming leg morphology is unknown; further, it may
not be a Cervinia at all, since it has no rostrum as do all others in the genus.
Consult Table 1 for C. brevipes distinguishing characteristics.

1. P,—P, with 2-segmented end.—‘‘synarthra”’ group ............... 2
— P,P, with 3-segmented end.—‘‘bradyi’’ group .................. 5
Dork, With-3-segmented,.end vse 0/s. 28.6 hyn Aincctnds (eran: ee ee 3
— WP withi2-segmentedvendi wy. “emer ans o4- slo. C. magna Smirnov
3. P,-P, end. terminal segment with 6, 7, 6 setae and spines respec-
tively 2.0600. 6.80. tends Reheat akaluitenew See ee Eee 4
— P,—P, end. terminal segment with 7, 8, 7 setae and spines respec-
tivelVnwr ers ods. ideo AS ee tank al cee eee C. langi Montagna
4. CR with long lateral-proximal seta ................... C. pilosa Lang
— CR with short lateral-proximal seta ................ C. synarthra Sars
5. CR longer than last urosome segment (L/W = 5) ................ 6
— CR as long as last urosome segment (L/W = 4) ... C. bradyi Norman
6. Basis of Mxp twice as long as end. segments; P; exp. with 3 seta.. 7

Basis of Mxp as long as 3 end. segments; P, exp. with 2 setae ....
EDL OOO UIE SEEN, Sle ty « Qh Don. SH aewaES C. tenuiseta Brodskaya

VOLUME 93, NUMBER 4 1213

Table 1.—Salient morphological characters of the species of Cervinia (se. = number of
setae, sp. = number of spines, sg. = number of segments).

Endopod setal formulae’ CR Md
Coxa- Exo-
P, Prox- basis pod
se.- =n imal = se.- sg.- Mxp
Species P, P, P, P, s? L/W LUS® setae _ sp. se. basis®
bradyi
INeormaniy) 1et22 12227 12321) e222 1s=e%, 4 1 Ss —- — —
brevipes
Brodskaya 1.1.221 — — — 3-e 60h 7 Zz A 1-2 4-4? 1.1.0
langi

Montagna 1.1.221 1.421 5211 1.421 5-CunG Z S pa lesny 93-5). p ieZal

magna
Smirnov 1.321 1.421 1.421 1221 3-e 64 1 = 22215 3-5), 4051-0

pilosa
Lang 122 16321 1.421 1.321 3 2 L—- — =

synarthra
Sars Ile le 32 1.421 13211 3-Cys 7) 2 Sir 3- Ome? Ne lel

tenuicauda
Brodskayanloi-221- 122-221) ik2-321) 162-221) 3 eal8 S55) eAV 2 4-4 led

tenuiseta
Brodskaya tele221 91-2.221 422.321 122:221 2-e _12:5¢ 4 L 40 46 1.2.1

unisetosa
Nn. sp. R22 el eS Al at 8 Sia aS 1-0 4-5 1.0.0

1 For all species exp. formula is P, = 1.1.123, P,-P, = 1.1.223.

2 s = shape, either reduced (r) or elongate (e).

3 >n LUS = as long as or longer than n Last Urosome Segments.

4L, S, A = long, short, or absent; short includes reduced.

5 n.n.n. = number of se. associated with (inner, middle, outer) claws, respectively.

7. CR as long as last 4 urosome segments (L/W = 12) ..............
dfarta Mond Shs A aPaae eA eres hice, fhe CIS LESTER Boy | C. tenuicauda Brodskaya
— CR as long as last 2 urosome segments (L/W = 8) ...............

Expansicervinia, new genus

The genus Expansicervinia is proposed for E. glacieria, and definition
preliminary, based on the type species described herein. The genus has two
features which are unique in all the described Cerviniidae. (1) The ventral
expansion of the second metasomal segment (Fig. 5), with an associated
broadening of the entire P, end. (2) An outer knoblike expansion on the first

1214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A, segment. The genus belongs to the subfamily Cerviniinae (by virtue of
the divergent CR) and is unique in the subfamily by having only 2 outer
spines on the terminal P,—P, exopodites, no rostrum, and an 8-segmented
A,. P; is minute and laterally located; there is an outer expansion of the
basis and a very long outer seta.

Expansicervinia glacieria, new species
Figs. 5-6

Cervinia sp. A Montagna and Carey, 1978:119.

Material.—3 2°? 2°. Holotype 1 2 dissected on slide USNM 180115. Para-
type 1 2 USNM 180116.

Type-locality.—Deep-sea floor off the Arctic coast of Alaska, USA, de-
scribed as station 49 (72°58'N, 146°29’W, 3,576 m) and station 50 (72°42’N,
143°40’W, 3,386 m) in Montagna and Carey (1978).

Description.—Female: Based on nonovigerous ¢ 1.1 mm long (Fig. 5).
Body Cervinia-like, broadened anteriorly and tapering throughout. R ab-
sent. Segment bearing P, distinct, segment bearing P, ventrally expanded.
CR divergent, as long as entire urosome. Lateral excrescences on first 2
urosomal segments (Fig. 5).

A, (Fig. 6): 8-segmented, basal segment broadened with outer node.
Aesthetasc on segments 3 and 4.

A, (Fig. 6): Allobasis with 2 setae. Terminal segment bears 3 inner and
7 terminal setae. Exp. 4-segmented with 2.1.1.3 setae respectively.

Md (Fig. 6): Precoxa with bidentate pars incisiva, tridentate lacina mob-
ilis, and 8 associated spines and a seta. Coxa-basis with 4 long setae. Exp.
4-segmented with 2.1.1.2 setae. End. 1-segmented with 3 inner and 6 ter-
minal setae.

Mxl (Fig. 6): Arthrite with 8 spines, 4 surface setae arranged as opposing
pairs. Coxa-basis with 5 inner and 1 outer setae. End. with 10 terminal
setae. Exp. represented by 2 setae.

Mx (Fig. 6): Syncoxa with 4 endites, with 5.3.3.2 setae respectively. Basis
with 2 claws, 3 setae. End. 2-segmented with 2.4 setae respectively.

Mxp (Fig. 6): Not prehensile. Basis as long as remaining segments, with
3 claws and 1.2.1 setae. 3 end. segments with | claw, 1 seta; 3 setae; 2 claws,
2 setae.

P,-P, (Fig. 5): All rami 3-segmented as figured, setation also listed below:

Exopod Endopod
P, 1.1.023 te leazil
P, e222 1e2221
P, e222 1.2.221
P 1.1.222 1.1.121

PS

1216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Expansicervinia glacieria °.

P, (Fig. 5): Located laterally (see habitus). No inner expansion of benp.,
but outer part expanded with 1 long seta. Exp. very small, 2 terminal setae.

P, & GF (Fig. 5): GF located medio-ventrally, genital pore circular, prox- |

imal and distal sclerotized ridges. P, lateral to GF with 3 terminal seta.
CR (Fig. 5): L/W = 12, 1 short proximal-lateral seta, 1 middorsal seta,

2 distal setae. Terminally with 2 principal caudal setae.
Variability.—None noticed in 3 specimens examined.

VOLUME 93, NUMBER 4 1217

Male.—Unknown.

Etymology.—The generic name Expansicervinia (L. ‘expansus’ = ex-
pand) refers to a Cervinia-like species with expanded characters; the first
segment of the A,, the ventral part of the second thoracic segment, and the
terminal segment of the P, end. The gender is feminine. The specific name
glacieria is in honor of the USCGC Glacier, from which the species was
collected, and without which the 4 years of sampling in the Beaufort Sea
would not have been possible.

Discussion.—Expansicervinia most closely resembles Cervinia in general
body shape, P, thoracic segment being distinct, shape and setation of the
CR, form and setation of the A,, segmentation of the swimming legs, and
general form of the mouthparts. Both these genera are distinct from others
in the subfamily by these same characteristics; hence they are probably
close phylogenetically. However, Expansicervinia is distinct from Cervinia
in that A, is 8-segmented with the basal segment enlarged (7 segments in
Cervinia), R is absent, P, lacks an inner seta on the terminal exp., P, seg-
ment is ventrally enlarged, P,-P, have only 2 outer spines on terminal ex-
opodites (opposed to 3 in Cervinia), and there is an outer expansion of P;
basis.

The expansion of the P, segment is unique in all the Harpacticoida and
its function may be peculiar to deep-sea life. Though this character is un-
known in other deep-sea harpacticoids, only further research could test this
hypothesis. The morphological result of the expansion is that the animal
can never straighten; thus the anterior end of the body must always be
oblique to the sediment. Lengthened body parts in other harpacticoids usu-
ally are associated with epipelic life; the expanded segment would further
lengthen the body and is probably an adaption toward this existence. As-
sociated with the second segment is the peculiar shape of the P, endopod,
which is much longer than the exopod, outwardly broadened, and possesses
broad, thick spines. Thus the entire second segment is specialized, perhaps
for some form of epipelic existence in the deep-sea.

Subfamily Cerviniopsinae Brodskaya, 1963
Ameliotes Por, 1969

Stratiopontotes Soyer, 1970:379.

Discussion.—As pointed out by Bodin (1979), the two genera Ameliotes
and Stratiopontotes are very similar to one another. In fact, the generic
descriptions are alike. Priority in this case is obfuscated by the publication
date of the journal Vie et Milieu; though the cover says 1969, it was printed
and published in 1970. Following Corliss (1979) I assign priority to Por’s
description (1969) over Soyer (1970). The latter’s species should now be
referred to as Ameliotes mediterraneus (Soyer, 1970).

1218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Key to Subfamilies and Genera of Cerviniidae

IC Redivergeenty sei cic ace ee subfam. Cerviniinae Brodskaya 2
— CR opposed (parallel) ........ subfam. Cerviniopsinae Brodskaya 6
2. EXPs Pi =P. With 3. SELMENES «A acnine ols Bo, SE ae oe ee 3
— Exp. P,-P, with 1 or 2 segments ................ Cerviniella Smirnov
3. CR at least as long as last urosomal segment; A, 7-8 segmented .. 4
— CR at most half as long as urosomal segment; A, 6-segment-

SG. sags Sete eres oral eat cv sca GEE cate oaks Sea Paracerviniella Brokskaya
4.,A, 7-segmented; with. rostrum :.. 5,002.0 6 sAverdic vies: } atieyenskes oe ee 5
— A, 8-segmented; rostrum absent; ventral expansion of segment bear-

Ue ag CRM eae) ie ae AACE eRe, yee. ml Expansicervinia, n. gen.
SCR Of Cqualslenethsy 4 ce sihoc cases cares cee oe Cervinia Norman
— | Right CRelongernthanleft 2. eee asus ele Eucanuella T. Scott
6. Prosome segments lacking or with reduced lateral ornamental ex-

CHESCHICES Feat eee Pi aes coe hoc sigeren oe ek SieR mous fous a, MeO 7

— Prosome with lateral or dorsal ornamentation ...................
> eins oul pe Reade Reusiteps aceeydeauny Nan Berio «ape cen cana Pontostratiotes Brady

Te PGeS2SC OMENS a cis ow Ae weve ok bv ee Sdn de ee 8
1) AGuOsSCOMENLEG 635 8.006% ah Jenna 4s Lpanaticyigt> ok e ee Cerviniopsis Sars
8. P, end. 3-segmented without apophysis ................0.eeeeeee 9

=, Pend: Z2-segmented with, apoplhysiS a2i-,..-c:10<i- «5 nner
Ee Te oak ae, OE EAN Ne ae eet Hemicervinia (1. C. Thompson)

97 Ends Of Mxi3-segmented. pce eae cued scat oes wees eee Ameliotes Por

— End. of Mxl reduced and fused to basis ... Herdmaniopsis Brodskaya

Acknowledgments

I am grateful to the Oregon State University benthic group, headed by A.
G. Carey, Jr., for their help and support during the collection and processing
of the samples which yielded the specimens reported on here. I also ac-
knowledge B. C. Coull for enthusiastic encouragement and for reading ear-
lier drafts of this manuscript.

Literature Cited

Bilyard, G. R., and A. G. Carey, Jr. 1979. Distribution of western Beaufort Sea polychaetous
annelids.—Mar. Biol. 54:329-339.

Bodin, P. 1979. Catalogue des nouveaux Copepodes Harpacticoides marins.—Université de
Bretagne Occidentale, Brest Cédex, France, 228 pp.

Brodskaya, V. A. 1963. [A survey of the family Cerviniidae (Crustacea, Copepoda). ]—Zool.
Zhur. 42:1785—1803. (In Russian with English summary.)

Corliss, J. O. 1979. Taxonomic importance of true publication date.—Trans. Amer. Micros.
Soc. 99:571.

Coull, B. C. 1977. Marine flora and fauna of the northeastern United States. Copepoda:
Harpacticoida.—NOAA Tech. Rep. NMFS Circ. 399, 48 pp.

VOLUME 93, NUMBER 4 1219

Lang, K. 1948. Monographie der Harpacticiden, I & II.—Hakan Ohlsson, Lund, 1682 pp.

. 1965. Copepoda Harpacticoidea from the Californian Pacific coast.—Kungl. Svenska

Vetensk. Handl., Fjarde Serien 10(2): 1-560, pls. 1-6.

Montagna, P. A. 1979. Cervinia langi n. sp. and Pseudocervinia magna (Copepoda: Harpac-

ticoida) from the Beaufort Sea (Alaska, USA).—Trans. Amer. Micros. Soc. 98:77-88.

, and A. G. Carey, Jr. 1978. Distributional notes on Harpacticoida (Crustacea: Cope-

poda) collected from the Beaufort Sea (Arctic Ocean).—Astarte 11:117-122.

Por, F. D. 1969. Deep-sea Cerviniidae (Copepoda: Harpacticoida) from the western Indian
Ocean, collected with R/V Anton Brunn in 1964.—Smithsonian Contrib. Zool. 29: 1-60.

Smirnov, S. S. 1946. [New species of Copepoda Harpacticoida from the northern Arctic
Ocean.|—Trudy Dreif Exped. Glaseumor Ledkol. Par ‘“‘Sedov.”’ 3:231—263. (In Russian
with English summary.)

Soyer, J. 1970. Contribution a l'étude des Copépodes Harpacticoides de Méditerranée. I.
Cerviniidae Sars, Lang.—Vie et Milieu 20(2B) [year 1969]:367-386.

Belle W. Baruch Institute for Marine Biology and Coastal Research and
Department of Biology, University of South Carolina, Columbia, South
Carolina 29208.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1220-1222

TWO NEW RECORDS OF THE CARIBBEAN
MARINE TUBIFICID KAKETIO INERI
RIGHI AND KANNER (OLIGOCHAETA)

Christer Erséeus

Abstract.—Kaketio ineri Righi and Kanner, 1979, originally described
from Bonaire in the southern Caribbean Sea, is reported from subtidal sands
in Bermuda and southern Florida. The genus Kaketio Righi and Kanner,
1979, is supposed to be closely related to Thalassodrilides Brinkhurst and
Baker, 1979.

In a recent taxonomic account of some marine oligochaetes from the
Caribbean Sea, Righi and Kanner (1979) established a new genus and species
of tubificid from Bonaire, Kaketio ineri. Two small lots of marine oligo-
chaetes from Bermuda and Florida, received for identification by me,
proved to belong to this species. The four worms, two from each area, were
Stained in paracarmine and mounted whole in Canada balsam. The genital
region of one of the specimens from Florida was cut into two halves before
mounting. Although being scanty, the new material adds valuable infor-
mation as to the geographical distribution of K. ineri.

Kaketio ineri Righi and Kanner, 1979
Kaketio ineri Righi and Kanner, 1979:55-62, figs. 39-47.

Type-material and type-locality.—See Righi and Kanner (1979).

Material examined.—United States National Museum of Natural History
Cat. No. 60130, 2 specimens from Whale-Bone Bay, Bermuda, subtidal
sand, shallow water, coll. 14 Aug. 1975, by M. L. Jones; USNM 60131, 2
specimens from Buttonwood Sound, Key Largo (Monroe Co.), S of Miami,
Florida, muddy sand with algae, 1-2 m depth, coll. May 1977, by R. Rehrer
(University of Miami).

Remarks.—K. ineri was very carefully described and depicted by Righi
and Kanner (1979). My new specimens conform well to their description in
all fundamental characters, although the two specimens from Bermuda are
only partially mature. The single specimen (from Florida) that is fully grown
posteriorly is 15.5 mm long and consists of 74 segments. This is slightly
different from the only complete specimen examined by Righi and Kanner
(25 mm, 52 segments).

One of the spermathecae in one of the worms from Florida contains the

VOLUME 93, NUMBER 4 1221

a ps ss
Qe >

Fig. 1. Map of the Caribbean area showing geographical records of Kaketio ineri Righi and
Kanner. Triangle = type-locality; circles = new records.

strange hyaline structure with 6 pairs of denticles which was described by
Righi and Kanner (cf. their fig. 46).

Habitat.—Marine shallow-water, muddy sands.

Geographical distribution.—Known from Bermuda, Florida (new rec-
ords) and Bonaire (Fig. 1).

Discussion.—The new material extends the known geographical distri-
bution of K. ineri (Fig. 1); it can be expected to occur throughout the
Caribbean.

According to Righi and Kanner, the male efferent ducts of Kaketio, su-
perficially at least, resemble those of Aulodrilus Bretscher, 1899. However,
in my opinion, Kaketio appears more closely related to Thalassodrilides
Brinkhurst and Baker, 1979. [Synonym: Curacaodrilus Righi and Kanner,
1979. This genus name was published the same year as Thalassodrilides,
but the publication was distributed later than that of Brinkhurst and Baker.].
Both Kaketio and Thalassodrilides are characterized by their possession of

1222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

relatively wide vasa deferentia entering more or less cylindrical atria, which
bear discrete, but not stalked, prostate glands, and which terminate into
glandular atrial ducts before ending in folded, eversible pseudopenes that
are enclosed in large muscular sacs. It was not mentioned in the original
definition of Thalassodrilides, but all species of that genus have a barrel-
shaped, thick-walled and dilated, portion of the intestine with a plexus of
fine blood vessels in segment IX (Erséus, in preparation; cf. Righi and Kan-
ner, 1979:54). This occurs also in K. ineri (cf. Righi and Kanner, 1979:57),
but is otherwise not common in the Tubificidae. The principal distinguishing
characters of the two genera are found in the morphology of their atria. In
Kaketio, each atrium is divided into two parallel structures: a proper atrium,
which is thin-walled and narrow, and a glandular and compact body (termed
‘‘atrial diverticulum’’ by Righi and Kanner), which covers the ventrolateral
surface of the proper atrium, and to the full length of which the prostate
glands are attached. In Thalassodrilides, this ‘‘diverticulum’’ is absent, and
the attachment of the prostates is not as broad as in Kaketio.

Acknowledgments

Dr. M. L. Jones (Smithsonian Institution, Washington, D.C.) and Dr. G.
H. Darcy (NOAA, National Marine Fisheries Service, Miami, Florida) kind-
ly provided me with the material.

Literature Cited

Brinkhurst, R. O., and H. R. Baker. 1979. A review of the marine Tubificidae (Oligochaeta)
of North America.—Canadian J. Zool. 57:1553-1569.

Righi, G., and E. Kanner. 1979. Marine Oligochaeta (Tubificidae and Enchytraeidae) from the
Caribbean Sea.—Stud. Fauna Curacao and Caribbean Is. 58:44-68.

Swedish Museum of Natural History, Stockholm, and Department of Zo-
ology, University of Gothenburg, Box 25059, S-400 31 Goteborg, Sweden.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1223-1229

A NEW SPECIES OF THE MARINE AMPHIPOD GENUS
GAMMAROPSIS FROM THE SOUTHEASTERN
UNITED STATES (PHOTIDAE)!

Walter G. Nelson

Abstract.—A new species of the amphipod genus Gammaropsis, G. suth-
erlandi, encountered during ecological studies of the amphipods of the es-
tuaries of North Carolina, is described here. The new species appears
closely related to Gammaropsis maculata (Johnston) 1827.

During ecological investigations of the amphipod crustaceans associated
with beds of eelgrass (Zostera marina) in the sounds near Beaufort, N.C.
(Nelson, 1979a, b), specimens of the marine photid genus Gammaropsis
were collected. The material proved to be a species new to science and
constitutes an addition to the list of amphipod species found in the shallow
estuarine waters of North Carolina (Fox and Bynum, 1975; Bynum and Fox,
1977; Nelson, 1979c) as well as to the amphipod fauna of the southeastern
United States.

Gammaropsis sutherlandi, new species
Figs. 1-3

Eurystheus maculatus.—Pearse and Williams, 1951.
Eurystheus erythopthalmus.—Pearse and Williams, 1951.
Eurystheus maculatus. —McClosky, 1970?

Gammaropsis maculata.—Fox and Bynum, 1975?
Gammaropsis maculata.—Fox, 1978?

Description.—Male.—8.5 mm. Eye oval, on acutely rounded produced
cephalic lobes. Head equals width of 1.5 pereonites. Antenna 1, 42% of
total body length, 103% of antenna 2; ratio of articles 1:2:3—1:1.4:.93, ven-
tral margin with long setae; flagellum with 16 articles, 92% of peduncle
length; accessory flagellum with 7 articles. Antenna 2, 41% of total body
length, ratio of articles 3:4:5—1:2.4:2.2; flagellum 57% of peduncle length,
with 13 articles.

Gnathopod 1, article 2 with posterodistal group of setae. Gnathopod 2,
ratio of articles 5:6—1:1.7; article 2 with a few medium setae on anterior

1 Contribution from Harbor Branch Foundation, Inc.

1224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Gammaropsis sutherlandi, male, 8.5 mm.

edge; article 5 with 9-10 clusters of setae on posterior margin; article 6 with
7 clusters of setae on posterior margin behind palm and 7 clusters on palm,
4 rows of setae on anterior margin, palm not defined by angular projection,
smooth curve from hind margin into concave palm, 2 small teeth projecting
from curve of palm dividing palm roughly into thirds; setae on articles 5 and
6 longer than width of article 6.

Epimeron 2 with small tooth on posterodistal corner. Epimeron 3 with
strong tooth on posterodistal corner.

Uropod 1, peduncle with distoventral edge produced into upturned spur
4 length of rami; inner ramus slightly longer than outer; peduncle equal to
outer ramus in length, outer margin with 8 spines, inner margin with 9 small
spines and 1 large distal spine; outer ramus with 6 outer and 4 inner marginal
spines; inner ramus with 7 outer and 3 inner marginal spines. Uropod 2,
peduncle with 3 spines on outer margin, | distal spine on inner margin; inner
ramus longer than outer; outer ramus equals length of peduncle; outer ramus
with 5 outer and 4 inner marginal spines; inner ramus with 6 outer and 3

VOLUME 93, NUMBER 4 1225

yee

Fig. 2. Gammaropsis sutherlandi, male, 8.5 mm: A, Antenna 1; B, Antenna 2; C, Gnathopod
2; D, Gnathopod 1; E, Detail of palm, gnathopod 2, setae removed; F, Detail of palm, gnathopod
2, setae present; G, Uropod 3; H, Uropod 2; I, Uropod 1; J, Pereopod 3; K, Pereopod 4; L,
Telson.

1226 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ly
—
~ SS

INSSK

G

6

J L

\ ee

Fig. 3. Gammaropsis sutherlandi, female, 8.5 mm: A, Gnathopod 1, scale = 1 mm; B,
Gnathopod 2, scale = 1 mm; C, Mandible; D, Maxilla 1; E, Maxilliped; F, Maxilla 2; G, Detail
of palm, gnathopod 2; H, Pereopod 5; I, Pereopod 6; J, Pereopod 7; K, Lower lip, male 8.5
mm; L, Epimera 2 and 3.

VOLUME 93, NUMBER 4 1227

inner marginal spines. Uropod 3, peduncle with 1 small dorsomedial spine
and 3 flattened dorsomedial spines at junction with rami, 1 large distal spine
on inner margin; outer ramus with 3 outer and 0 inner marginal spines; inner
ramus with 3 inner and 3 outer marginal spines. Telson moderately incised
with stout medial spines.

Female.—8.5 mm. Gnathopod 1, like male except article 5 lacks medial
line of setae. Gnathopod 2, much less massive than in male; article 2 with
a few setae on anterior margin; article 6, 2 times length of article 5, palm
oblique with 2 small acute projections and 1 spine inserted medially near
first projection, palm defined only by presence of second spine—not by an
angular projection.

Material examined.—Types: Holotype male, 7.7 mm, Frying Pan Shoal,
5 Mar. 1935, USNM 173231. Allotype female, 6.7 mm, Frying Pan Shoal,
5 Mar. 1935, USNM 173232. Paratype series, Beaufort, N.C., USNM
173233, including paratype male, 8.5 mm, and paratype female, 8.5 mm,
figured herein. Duke University Marine Laboratory, Beaufort, N.C. from
fouling tiles, 14 Apr. 1974, 1 ovigerous female, | immature; 19 Mar. 1975,
2 females, 1 male; 20 May 1975, 1 female, 1 immature. Shackleford Jetty,
Bogue Sound, N.C. from algae, 20 Jan. 1976, 1 ovigerous female; 17 Mar.
1976, 1 male, 1 immature. Material from Duke University Marine Labora-
tory Reference Museum collections, from dredge samples collected by R.V.
Eastward off the coast of N.C., 19 Apr. 1965, 1 female, 34°34.5’N,
76°25.5’'W, 20 m; Eastward station 17069, 1 male, 2 females; Eastward
station 3645, 13 Jan. 1966, 1 male.

Material from USNM collections: From North Carolina.—Off Bogue In-
let, 19 July 1915, 2 females, 2 immatures, No. 8286; Shackleford Banks, 12
Sept. 1928, 5 females, USNM 102997; Frying Pan Shoal, 5 Mar. 1935, 2
males, 5 females, 1 immature, USNM 133318; Frying Pan Shoal, 2 Apr.
1935, 1 ovigerous female, USNM 134066; Black Rocks off New River, 19
June 1949, 4 males, 5 females, USNM 183351; Black Rocks off New River,
19 June 1949, 1 female, USNM 183864. From South Carolina.—Albatross
Sta. 20037, 12 Dec. 1919, 1 ovigerous female, USNM 64746; east of Cape
Romain, 1 July 1935, 3 females, USNM 135202; off Little River Inlet, 17
Aug. 1949, 3 males, 2 females, USNM 183864. From Dry Tortugas, Flori-
da.—Tortugas, 1 female, Sta. No. 33-31; south of Tortugas, 31 July 1930,
1 male, 11 females, Sta. No. 38-30; south of Tortugas, 23 July 1932, 1 male,
Sta. No. 59-32; Tortugas (Haul 210 by W. L. Schmitt), 10 June 1925, (2 lots)
1 female and 1 male, 2 females and 3 immatures, USNM 93400; Loggerhead
Key, 7 Aug. 1931, 5 males, 3 females, 3 immatures, USNM 115488; Mis-
cellaneous, probably Tortugas, 17 June 1932, 2 ovigerous females, Sta. No.
3-32; south of Tortugas, 4 Aug. 1931, 1 female; Fish Hawk Sta. No. 8499,
5 males, 19 females; Fish Hawk Sta. No. 8895, 1 ovigerous female.

1228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution.—Cape Hatteras, south side, south to the Dry Tortugas of
Florida, in depths from shallow subtidal to 300 m.

Relationships.—Gammaropsis sutherlandi appears to be closely related
to Gammaropsis maculata (Johnston) 1827 but is distinct from it in several
features. The comparisons below are based on G. maculata (=G. erythop-
thalma figured by Sars, 1894, pl. 198) material from Norway in the collection
of the Zoological Museum of the University of Oslo. The confused synon-
omy of G. maculata has been discussed by Krapp-Schickel and Myers
(1979). The flagellum of antenna 1 is equal to 70-80% of the length of the
peduncle in G. maculata and 92% in G. sutherlandi, with the flagellum
being composed of 13-14 articles in males of G. maculata and 16-18 in G.
sutherlandi.

The posterodistal corner of article 2 of gnathopod 1 possesses a group of
setae in males of G. sutherlandi and lacks it in G. maculata. On gnathopod
2, setae are found on the anterior margin of article 2 of both males and
females of G. sutherlandi and are lacking in G. maculata. Article 5 of male
gnathopod 2 possesses 6 groups of setae in G. maculata and 9-10 in G.
sutherlandi. Article 6 of male gnathopod 2 possesses 9 groups of setae on
the posterior margin in G. maculata and 14 in G. sutherlandi. The palm of
male gnathopod 2 is defined by a small distally pointing angular projection
in G. maculata and is not defined by any angular projection in G. suther-
landi. The setae on articles 5 and 6 of male gnathopod 2 are less than the
width of article 6 in G. maculata and equal to or greater than the width of
article 6 in G. sutherlandi. Both species possess 2 toothlike projections on
the palm. Although the degree of development of these teeth varies in G.
sutherlandi, maximum development is considerably less than that for G.
maculata, although Krapp-Schickel and Myers (1979) suggest the palmar
teeth of G. maculata become obsolete in old males.

Etymology.—This species is named for Dr. John P. Sutherland in honor
of his contributions to the study of marine fouling communities and the field
of marine ecology.

Ecology.—This species appears quite generalized in its epifaunal habitat.
It has been collected from the shallow subtidal from algae on rock jetties,
from the fouling community, and rarely, from eelgrass (Zostera marina)
beds in the estuary near Beaufort, N.C. It has also been collected from such
substrates as stones, the backs of spider crabs, ceramic fouling plates, and
was also common in the stomach contents of sea bass taken from approx-
imately 15 m depth on an artificial reef structure off Wilmington, N.C. In
the region of Beaufort, N.C. available data indicate a breeding season of
from at least January through July, and this species may well breed year
round.

VOLUME 93, NUMBER 4 1229

Acknowledgments

I would like to thank the following for providing material for this study:
Dr. J. L. Barnard, USNM; Dr. Wim Vader, University of Tr@mso; Dr. M.
E. Christiansen, University of Oslo; Dr. W. Kirby-Smith, Duke University.
Dr. E. L. Bousfield has provided information and encouragement through-
out this study, and both he and Dr. J. L. Barnard kindly reviewed the
manuscript. M. A. Nelson inked the plates. Preparation of the manuscript
was made possible by a Harbor Branch Institution Postdoctoral Fellowship.

Literature Cited

Bynum, K. H., and R. S. Fox. 1977. New and noteworthy amphipod crustaceans from North
Carolina, U.S.A.—Chesapeake Sci. 18:1-33.
Fox, R.S. 1978. Amphipoda. In R. Zingmark, Ed., An annotated checklist of the biota of the
coastal zone of South Carolina.—University of South Carolina Press, Columbia, 364 pp.
and K. H. Bynum. 1975. The amphipod crustaceans of North Carolina estuarine
waters.—Chesapeake Sci. 16:233-237.

Krapp-Schickel, G., and A. A. Myers. 1979. The Mediterranean species of Gammaropsis
Liljeborg (Crustacea, Amphipoda).—Boll. Mus. Civ. St. Nat. Verona 6:441—467.
McClosky, L. R. 1970. The dynamics of the community associated with a marine scleractinian

coral.—Int. Rev. Ges. Hydrobiol. 55:13-81.
Nelson, W. G. 1979a. Experimental studies of selective predation on amphipods: Conse-
quences for amphipod distribution and abundance.—J. Exp. Mar. Biol. Ecol. 38:225—
245.
. 1979b. The analysis of structural pattern in an eelgrass (Zostera marina) amphipod
community.—J. Exp. Mar. Biol. Ecol. 39:231-264.
. 1979c. Additions to the amphipod crustaceans of North Carolina. —Estuaries 2:65.
Pearse, A.S., and L. G. Williams. 1951. The biota of the reef off the Carolinas.—J. Elisha
Mitch. Sci. Soc. 67:133-161.

Harbor Branch Institution, Inc., RR 1, Box 196-A, Ft. Pierce, Florida
33450 (present address: Institute of Marine Biology, University of Bergen,
N-5065 Blomsterdalen, Norway).

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1230-1238

CESTODE PARASITES OF SOME
VENEZUELAN STINGRAYS

Monte A. Mayes and Daniel R. Brooks

Abstract.—Forty-seven stingrays representing 8 species collected in Ven-
ezuelan coastal waters were examined for cestode parasites. Specimens of
18 tetraphyllidean, lecanicephalidean, and trypanorhynchan cestode species
were collected, 3 described as new. Rhinebothrium margaritense sp. n.
from Dasyatis guttata and D. americana most closely resembles R. spini-
cephalum and R. tetralobatum but differs by having 3-6 rather than only
2 testes per proglottid and a different number of bothridial loculi. Rhodo-
bothrium paucitesticulare sp. n. from Rhinoptera bonasus possesses fewer
than half as many testes per proglottid as any other known member of the
genus. Dioecotaenia campbelli sp. n. differs from D. cancellata by having
24 rather than 21 bothridial loculi and smaller eggs and embryos. Other
collected species include: Acanthobothrium electricolum in Narcine bras-
iliensis, A. fogeli in Gymnura micrura, Disculiceps sp. and Acanthoboth-
rium tortum in Aetobatus narinari, Tylocephalum sp. and Rhinoptericola
megacantha in Rhinoptera bonasus, Rhinebothrium magniphallum and
Parachristianella cf. monomegacantha in Himantura schmardae, Acan-
thobothrium americanum, Phyllobothrium centrurum. Rhinebothrium cor-
ymbum and Rhodobothrium pulvinatum in Dasyatis americana, and Acan-
thobothroides thorsoni, Acanthobothrium tasajerasi, A. urotrygoni,
Rhinebothrium magniphallum, and Rhodobothrium pulvinatum in Dasyatis
guttata. All represent new locality records, and those species occurring in
Dasyatis guttata are reported from that host for the first time.

Little is known about the helminth fauna of elasmobranchs living along
the Venezuelan coast. Troncy (1969) described a nematode, Echinocephalus
diazi, from Lake Maracaibo and listed the freshwater stingray Potamotry-
gon hystrix as host. Later, Diaz-Ungria (1973) corrected the host identifi-
cation to the euryhaline dasyatid stingray Himantura schmardae (Werner).
Deardorff, Brooks, and Thorson (in press) discovered E. diazi in H.
schmardae from Colombia, thus confirming Diaz-Ungria’s statement. We
found no other reports of helminths infecting Venezuelan coastal stingrays.

This paper reports 18 species of tetraphyllidean, lecanicephalidean and
trypanorhynchan cestodes collected by the first author during examinations
of 47 stingrays representing 8 species. Stingrays were collected during the
summer from the Gulf of Venezuela and from Lake Maracaibo (1977 and
1978) and from Isla de Margarita (1978). Table 1 summarizes our findings.

VOLUME 93, NUMBER 4 1231

Helminths were removed from hosts, examined alive when possible, fixed
with AFA or 10% formalin and stored in 70% ethanol. Most specimens were
stained with Mayer’s hematoxylin and mounted in Canada balsam for study
as whole mounts. Serial cross sections, cut at 8 wm and stained with he-
matoxylin-easin, were used to confirm some aspects of proglottid morphol-
ogy. All measurements are in um unless otherwise stated; figures were
drawn with the aid of a drawing tube. Representative specimens of all
species have been deposited in the University of Nebraska State Museum,
Division of Parasitology, Harold W. Manter Laboratory.

Rhinebothrium margaritense, sp. nov.
Figs. 1-2

Description (based on 15 specimens).—Strobila up to 5.7 mm long, cras-
pedote, apolytic, composed of 75-100 proglottids. Scolex up to 1.86 mm
wide, composed of 4 pedicellated, elongate, bilobed, septate bothridia.
Bothridia 744—1,209 long by 232-279 wide; divided longitudinally by median
septum, horizontally by hingelike constriction between lobes; posterior lobe
divided horizontally by 12-13 septa forming 26-28 loculi; anterior lobe di-
vided horizontally by 12 septa forming 26 loculi plus terminal loculus at tip;
total number of.loculi per bothridium 53 or 55. Pedicels 418-558 long. Ce-
phalic peduncle short, aspinose, 46-93 long. Immature proglottids wider
than long. Mature proglottids 120—456 long by 182-240 wide, length greater
than width only in last 6-10 proglottids if at all. Genital pore 48-52% (* =
50%, n = 50) of proglottid length from anterior end. Testes 3-6 (¢ = 4,n =
68) in number, 20-60 in diameter. Cirrus sac elongate, 84-120 long by 36—-
72 wide, containing spined eversible cirrus. Genital atrium shallow, simple.
Vagina anterior to cirrus sac, sphincter present. Ovary X-shaped in cross
section, bilobed in frontal view, lobate, poral lobe extending anteriorly to
level of genital pore; 65-156 long by 50-165 wide at isthmus. Vitellaria
follicular; follicles 3-7 in diameter, extending nearly entire length of pro-
glottid. Gravid proglottids not collected.

Hosts.—Dasyatis guttata (Bloch and Schneider) (type); Dasyatis ameri-
cana Hildebrand and Schroeder.

Site of infection.—Spiral valve.

Locality.—Isla de Margarita, near Robledal, Venezuela.

Holotype.—USNM Helm. Coll. No. 75715. Paratypes: USNM Helm.
Coll. No. 75716; Univ. Nebraska State Museum No. 21036.

Etymology.—This species is named for the island from which it was col-
lected.

Remarks.—Rhinebothrium margaritense is a member of a monophyletic
species-group within Rhinebothrium characterized by being small worms
with relatively to markedly long bothridial pedicels, more than 25 proglottids

1232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Host-parasite list for cestodes collected in Venezuelan coastal waters during 1978
and 1979. All previously known species are reported from Venezuela for the first time. * = new

host record.
Host

Narcine brasiliensis
(Olfers)

Rhinobatos percellens
(Walbaum)

Aetobatus narinari
(Euphrasen)

Gymnura micrura
(Bloch and
Schneider)

Rhinoptera bonasus
(Mitchill)

Himantura schmardae
(Werner)

Dasyatis americana
Hildebrand and
Schroeder

Dasyatis guttata
(Bloch and
Schneider)

Cestode species

Acanthobothrium electricolum
Brooks and Mayes, 1978

Negative

Acanthobothrium tortum Linton,
1890
Disculiceps sp.

Acanthobothrium fogeli
Goldstein, 1964

Rhinoptericola megacantha
Campbell and Carvajal, 1975

Rhodobothrium paucitesticulare
sp. n.

Dioecotaenia campbelli sp. n.

Tylocephalum sp.

Parachristianella cf.
monomegacantha

Rhinebothrium magniphallum
Brooks, 1977

Phyllobothrium centrurum
Southwell, 1925

Rhodobothrium pulvinatum
Linton, 1889

Rhinebothrium corymbum
Campbell, 1975

Rhinebothrium margaritense sp.
n.

Acanthobothrium americanum
Campbell, 1969

Rhodobothrium pulvinatum*
Linton, 1889

Rhinebothrium magniphallum*

Rhinebothrium margaritense sp.
n.

Locality and incidence

Robledal, Isla de
Margarita (1/3)

Isla de Margarita (2)

Gulf of Venezuela,
Caimare Chico (1/1)
same

Isla de Margarita (1/2)

Gulf of Venezuela,
Caimare Chico (1/9)

same

same
same

Boca Canonero, Bahia de
Tablazo, Lake
Maracaibo (1/2)

same

Isla de Margarita (3/3)

same (2/3)
same (1/3)
same (2/3)

same (2/3)

Gueria, Gulf of Paria (1/3)

Mouth of Lake Maracaibo
(1/9)

El Guano, S. of Rio Apon,
Lake Maracaibo (1/5)

Robledal, Isla de
Margarita (1/2)

VOLUME 93, NUMBER 4 1233

Table 1.—Continued.

Host Cestode species Locality and incidence
Acanthobothroides thorsoni* Robledal, Isla de
Brooks, 1977 Margarita (1/2)
Acanthobothrium urotrygoni* Gulf of Venezuela,
Brooks and Mayes, 1980 Caimare Chico (1/5)

Mouth of Lake Maracaibo
(1/9)

Robledal, Isla de
Margarita (1/2)

Acanthobothrium tasajerasi Mouth of Lake Maracaibo
Brooks, 1977 (1/9)

per strobila, craspedote proglottids which are wider than long except for
terminal proglottids, bilobed bothridia with a single median septum and at
least 32 loculi, and an average of fewer than 10 testes per proglottid. The
genealogical relationships of that species-group’s members have been pre-
sented elsewhere (Brooks, Mayes, and Thorson, in press). The new species
most closely resembles R. spinicephalum Campbell, 1969 and R. tetralo-
batum Brooks, 1977 both of which possess 2 testes per proglottid rather
than 3-6 (< = 4) exhibited by R. margaritense. Rhinebothrium spinice-
phalum further differs from the new species by possessing 32—34 bothridial
loculi rather than 53 or 55 as exhibited by R. margaritense; R. tetralobatum
possesses 48—54 loculi. Both R. spinicephalum and R. margaritense possess
compact lobate ovaries, differing from the fragmented 4-part ovaries of spec-
imens of R. tetralobatum.

Rhodobothrium paucitesticulare, sp. nov.
Figs. 3-5

Description (based on 7 specimens).—Tetraphyllidea, Phyllobothriidae,
Rhodobothrium Linton, 1890 as emended by Campbell and Carvajal, 1979.
Strobila craspedote, serrate posteriorly, maximum dimensions 20-31 mm
long by 7.4-9.0 mm wide. Scolex 1.0-1.9 mm wide, composed of 4 pedi-
cellated bothridia. Bothridia 465-930 long by 465-930 wide, trumpet-shaped
when relaxed, adherent surfaces convex, traversed by numerous convolu-
tions forming irregular pattern. Bothridial faces round or subtriangular in
cross section, margins ruffled. Pedicels approximately 250 long. Neck up to
1.86 mm long by 700 wide. Number of segments 400-600. Immature pro-
glottids wider than long, markedly protandric. Mature proglottids squared,

1234

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ee
Cy ><)
DALY

ene”
oto Mg

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20 4s oe

of
ro

e
Se
ve

Decszeorszen
cts

@
Cee
eo
oe

s
2.
)
Ste

2 Po;
°
00%,

Figs. 1-5. Rhinebothrium margaritense: 1, Scolex; 2, Mature proglottid. Rhodobothrium
paucitesticulare: 3, Scolex; 4, Immature proglottid; 5, Mature proglottid.

372-651 long by 372-605 wide. Genital pore alternating irregularly in ante-
rior 48-52% ( = 50%, n = 25) of proglottid; genital atrium well-developed.
Cirrus sac 186-279 long by 93-140 wide, containing spined eversible cirrus.
Testes 40-80 in number (x = 50, n = 100) in immature proglottids; few se-
nescent testes present or testes lacking in mature proglottids. Testes sub-
spherical, 24-60 in diameter. Vagina anterior to cirrus sac, sphincter pres-

VOLUME 93, NUMBER 4 1235

Figs. 6-11. Dioecotaenia campbelli: 6, Scolex; 7, Early mature female proglottid; 8, Mature
female proglottid; 9, Gravid female proglottid; 10, Maturing male proglottid; 11, Fully mature
male proglottid ready for hypodermic insemination following detachment.

ent. Ovary follicular, bilobed in frontal view, X-shaped in cross section;
lobes expanded anteriorly in maturing proglottids, 167-353 long by 373-465
at isthmus. Vitellaria forming lateral bands extending nearly entire length
of proglottid; follicles 24-120 in diameter. Uterus saccate, preovarian in
mature proglottids. Gravid proglottids not collected.

Host.—Rhinoptera bonasus (Mitchill).

Site of infection.—Spiral valve.

Locality.—Gulf of Venezuela, Caimare Chico, Venezuela.

Holotype.—USNM Helm. Coll. No. 75717. Paratypes: USNM Helm.
Coll. No. 75718; Univ. Nebraska State Museum No. 21034.

Etymology.—tThe specific name refers to the presence in this species of
only about half as many testes per proglottid as are present in members of
the other known species.

Remarks.—Campbell and Carvajal (1979) reviewed Rhodobothrium Lin-
ton, 1889, recognizing it as the senior synonym of Inermiphyllidium Riser,

1236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1955 and Sphaerobothrium Euzet, 1959. They considered R. pulvinatum
Linton, 1889, R. lubeti (Euzet, 1959) Campbell and Carvajal, 1979, and R.
mesodesmatum (Bahamonde and Lopez, 1962) Campbell and Carvajal, 1979
members of Rhodobothrium and R. brachyascum (Riser, 1955) Campbell
and Carvajal, 1979 a provisional member of the genus. The new species
differs from the above by exhibiting approximately half as many testes per
proglottid, 40-80 with a mean of 50 vs. 117-149, 120-160, and 150-210,
respectively for the 3 species listed above. Rhodobothrium paucitesticulare
further differs by being more markedly protandric. No evidence beyond
ovarian anlagen indicates female genitalia in proglottids containing fully de-
veloped testes. The testes senesce markedly in proglottids containing de-
veloping ovaries and other female genitalia.

Such an observed ontogenetic feature is of potential interest when hosts
for Rhodobothrium species are compared. Rhodobothrium lubeti and R.
mesodesmatum infect Myliobatis spp., R. pulvinatum infects members of
Dasyatis, and the new species infects Rhinoptera bonasus. Rhinoptera bon-
asus already is known to host the only dioecious tetraphyllidean cestode
(we describe another species from the same host next), raising the possibility
that the presence of a markedly protandric species of tetraphyllidean in R.
bonasus relates more to a more general phenomenon involving some aspect
of the host’s biology.

Dioecotaenia Schmidt, 1969

Schmidt (1969) redescribed Rhinebothrium cancellatum Linton, 1890 and
reported it to be a dioecious cestode, the only known such tetraphyllidean.
His generic diagnosis was so extensive that a second new species which we
describe herein could not be accommodated within the genus unless we
emended Schmidt’s (1969) diagnosis slightly, as follows:

Emended diagnosis.—Tetraphyllidea, Dioecotaeniidae. Sexes completely
separate. Sexual dimorphism apparent. Scolex with 4 bothridia on short
peduncles, each divided into loculi interpretable as 3 longitudinal rows of
equal numbers of loculi or a center row of loculi surrounded by a circle of
marginal loculi. Myzorhynchus, accessory suckers, and hooks lacking.
Neck present. External segmentation feeble, proglottids acraspedote. Os-
moregulatory canals consisting of 6 major medullary trunks, of which the
most lateral are highly ramified and anastomose with irregular ducts leading
to lateral margins. Cortex thin. Muscle bindles feeble.

Testes medullary, in 2 layers, arranged in a semicircle or circle on all
sides of cirrus sac. Genital pores lateral, alternating irregularly. Cirrus sac
large, containing spined eversible cirrus and internal seminal vesicle. Cirrus
long, armed at base with hooks possessing bifid roots. External seminal
vesicle lacking.

VOLUME 93, NUMBER 4 1237

Ovary bilobed, transversely elongate, medullary. Oviduct short. Vagina
medullary, convoluted; vaginal pore lacking. Seminal receptacle embedded
in substance of one ovarian lobe, irregularly alternating sides. Vitellaria
compact, surrounding posterolateral margins of both ovarian lobes. Uterus
bilobed, saccate, preovarian. Uterine pore preformed, medioventral. Em-
bryo with simple outer membrane. Sperm transfer by hypodermic impreg-
nation. Medullary continuous dorsomedian sheath containing injected cirri
present in female. Parasites of elasmobranchs. Type-species: D. cancellata
(Linton, 1890) Schmidt, 1969. Other species:

Dioecotaenia campbelli, sp. nov.
Figs. 6-11

Description (based on 15 specimens).—With characters of the genus given
above.

Males (7 specimens): Strobila up to 26 mm long by 1.21—1.73 mm wide.
Scolex 962-979 long by 1,270-—1,490 wide. Bothridia 790-—1,023 long by 651-
698 wide, possessing 24 loculi arranged either as 3 longitudinal rows of 8
loculi or as a median row of 6 loculi surrounded by 18 marginal loculi. Neck
approximately 5 mm long. Strobila with up to 200 proglottids. Testes 60-90
in number (x = 80, n = 20), 19-36 in diameter. Testes atrophy markedly in
posteriormost proglottids as internal seminal vesicle and cirrus sac enlarge.
Cirrus sac curved posteriorly, more markedly in older proglottids; sac 279-
623 long by 47-139 wide. Genital atrium prominent, capable of protruding
as suckerlike papilla, surrounded by darkly-staining parenchymal cells.

Female (8 specimens): Strobila up to 50 mm long by up to 1.9 mm wide.
Scolex 962-974 long by 1,270-1,490 wide. Bothridia and neck as in male.
Strobila with up to 200 proglottids. Ovary 139-279 long by 465-605 wide.
Seminal receptacle 93-139 in diameter. Uterus first a transverse sac, be-
coming bilobed when gravid, with thin ventral isthmus. Uterine pore near
posterior end of proglottid. Eggs 45-65 in diameter, oncospheres 36-48 in
diameter.

Host.—Rhinoptera bonasus.

Site of infection.—Spiral valve.

Locality.—Gulf of Venezuela, Caimare Chico, Venezuela.

Holotype. —USNM Helm. Coll. No. 75719 (male). Allotype: USNM
Helm. Coll. No. 75720 (female). Paratypes: USNM Helm. Coll. No. 75721;
Univ. Nebraska State Museum No. 21033.

Etymology.—This species is named for Dr. Ronald A. Campbell, South-
eastern Massachusetts University, in recognition of his contributions to the
systematics of cestodes infecting elasmobranchs.

Remarks.—Dioecotaenia campbelli closely resembles D. cancellata, the
only consistent anatomical differences being number of bothridial loculi (24

1238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in the new species vs. 21 in D. cancellata) and size of eggs and embryos
(36-48 um in diameter eggs vs. 45-65 um in diameter eggs; 19-26 um in
diameter oncospheres vs. about 35 um in diameter oncospheres). Because
both species inhabit the same host species and occur in adjacent geograph-
ical areas, it seems likely that they are sister-species. Thus, their close
morphological similarity does not necessarily indicate any degree of mor-
phological conservatism in Dioecotaenia.

Literature Cited

Brooks, D. R., M. A. Mayes, and T. B. Thorson. In press. Systematic review of cestode
infecting freshwater stingrays (Chondrichthyes: Potamotrygonidae) including four new
species from Venezuela.—Proc. Helm. Soc. Wash.

Campbell, R. A., and J. Carvajal G. 1979. Synonymy of the phyllobothriid genera Rhodo-
bothrium Linton, 1889, Inermiphyllidium Riser, 1955, and Sphaerobothrium Euzet, 1959
(Cestoda: Tetraphyllidea).—Proc. Helm. Soc. Wash. 46:88-97.

Deardorff, T. L., D. R. Brooks, and T. B. Thorson. In press. Two species of Echinocephalus
(Nematoda: Gnathostomidae) from Neotropical stingrays.—J. Parasitol.

Diaz-Ungria, C. 1973. Helmintos endoparasitos de Venezuela.—Cien. Vet., Maracaibo 3:37-
242.

Schmidt, G. D. 1969. Dioecotaenia cancellata (Linton, 1890) gen. et comb. n., a dioecious
cestode (Tetraphyllidea) from the Cow-Nosed Ray, Rhinoptera bonasus (Mitchill), in
Chesapeake Bay, with the proposal of a new family, Dioecotaeniidae.—J. Parasitol.
$5:271-275.

Troncy, P. M. 1969. Description de deux nouvelles especes de nematodes parasites de pois-
sons.—Bull. Mus. Natn. Hist. Nat. 2nd series 41:598—605.

(MAM) Environmental Sciences Research, 1702 Building, Dow Chemical
USA, Midland, Michigan 48640; (DRB) Department of Pathology, National
Zoological Park, Smithsonian Institution, Washington, D.C. 20008 (present
address: Department of Zoology, University of British Columbia, 2075 Wes-
brook Mall, Vancouver, British Columbia, Canada V6T 1W5S).

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1239-1252

CESTODE PARASITES IN MYLIOBATIS GOODEI
GARMAN (MYLIOBATIFORMES:
MYLIOBATIDAE) FROM RIO DE LA PLATA,
URUGUAY, WITH A SUMMARY OF CESTODES
COLLECTED FROM SOUTH AMERICAN
ELASMOBRANCHS DURING 1975-1979

Daniel R. Brooks, Monte A. Mayes, and Thomas B. Thorson

Abstract.—Specimens of 7 cestode species, 4 described as new, were
collected from spiral valves of 4 Myliobatis goodei captured in the La Plata
estuary near Montevideo, Uruguay. Discobothrium arrhynchum sp. n. dif-
fers from all other members of the genus by lacking a myzorhynchus. It
most closely resembles D. myliobatidis by having relatively large suckers,
markedly craspedote immature proglottids, an average of 21 testes per pro-
glottid, and a vagina extending laterally rather than medially to the testes.
Caulobothrium is recognized as a valid genus but Rhabdotobothrium is
considered a junior synonym. Two species-groups within Caulobothrium
are recognized, and a new species in each one is described. Caulobothrium
uruguayense sp. n. most clearly resembles C. tetrascaphium by having
more than 100 proglottids per strobila, craspedote proglottids, preovarian
testes, more than 100 testes per proglottid, a very long cephalic peduncle,
and genital pores in the anterior 14 of the proglottid. It differs by having 14
or 15 rather than 25 bothridial loculi and recurved rather than straight cirrus
sacs. Caulobothrium ostrowskiae sp. n. most closely resembles C. myliob-
atidis, C. opisthorchis and C. multorchidum by exhibiting postovarian
testes. The new species differs from C. opisthorchis by having fewer testes
and by lacking vitelline follicles encircling the postovarian testes; it differs
from C. multorchidum by having elongate rather than broad flaplike bo-
thridia; and it differs from C. myliobatidis by having fewer bothridial loculi.
Rhabdotobothrium dollfusi and R. anterophallum become Caulobothrium
dollfusi and C. anterophallum. Phyllobothrium myliobatidis sp. n. differs
from P. auricula, which it most closely resembles, by having longer and
thinner bothridial pedicels and much smaller cirrus sacs. Contracted spec-
imens of Phyllobothrium sp. and immature specimens of 2 species of Acan-
thobothrium are briefly described and discussed. A table listing hosts ex-
amined and one listing cestodes collected during study of South American
elasmobranch parasites from 1975-1979 are included.

1240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

From 1975-1979 the authors collected helminth parasites from a total of
117 euryhaline stingrays captured in various parts of northern and eastern
South America. This report represents the last in a series of taxonomic
papers reporting our findings. Herein we report helminths parasitizing My-
liobatis goodei Garman (Myliobatiformes: Myliobatidae) from Rio de la
Plata, Uruguay and present a list of hosts examined and parasites collected
as a result of our study. Specimens representing 7 species of cestodes were
collected from the spiral valves of 4 M. goodei. All appear to represent new
species, but the condition of our material permits description of only 4.

Spiral valves were removed from hosts and placed in ice-water for one
hour. They were then slit longitudinally and immersed in 10% formalin for
transportation to the laboratory. Spiral valves were systematically dissected
and examined for helminth parasites; those present were removed and
stored in 70% ethanol. Most collected specimens were stained with Mayer’s
hematoxylin and mounted in Canada balsam for study as whole mounts.
However, some specimens were serially cross-sectioned, cut at 8 wm and
stained with hematoxylin-eosin, to confirm certain aspects of proglottid
morphology. All figures were drawn with the aid of a drawing tube; mea-
surements are in wm unless otherwise stated.

Discobothrium arrhynchum, sp. nov.
Figs. 1-2

Description (based on 30 specimens).—Strobila craspedote, apolytic, as-
pinose, up to 3,350 long, composed of 43-48 proglottids. Scolex 177-186
long by 233-326 wide, composed of 4 suckers each with lateral flap partially
enclosing suctorial opening; suckers and scolex spinose. Suckers 132-216
long by 120-256 wide. Apical organ or myzorhynchus lacking. Cephalic
peduncle lacking. Neck extremely short, not measured. Immature proglot-
tids wider than long, markedly craspedote. Mature terminal proglottids 408—
672 long by 192-312 wide. Testes 29-72 in diameter, 18-28 (x = 21, n = SO)
in number, 3-4 (x = 3.5) preporally, 5—9 (« = 6.8) postporally, 9-16 (« =
10.7) antiporally. Cirrus sac in anterior % of proglottid, 36-79 long by 12-
48 wide, containing unspined eversible cirrus. Genital pore 25-32% (x =
27.5%) of proglottid length from anterior end. Genital atrium shallow, sim-
ple. Vagina opening anteriorly to cirrus sac, extending posteriorly lateral to
postporal testicular field, reaching near posterior !/s of proglottid. Ovary in
posterior '/s of proglottid, bialate, 36-96 long by 36-108 wide at isthmus,
anterior to posteriormost extent of vagina. Mehlis’ gland prominent, ootype
immediately posterior to ovarian isthmus. Vitellaria follicular, follicles ex-
tending nearly entire length of proglottid, S—17 in diameter.

_ Host.—Mbyliobatis goodei Garman (Myliobatiformes: Myliobatidae).

VOLUME 93, NUMBER 4

Table 1.—South American elasmobranchs examined by the authors for parasitic helminths

during 1975-1979.

1241

Host species Locality Number Year(s)
Narcine brasiliensis Cartagena, Colombia 17 1976
Isla de Margarita, Venezuela 3 1978
Urolophus jamaicensis Cartagena, Colombia 5) 1976
Urotrygon venezuelae Cienaga Grande, near Santa 1 1975
Marta, Colombia
Cartagena, Colombia 16 1976
Aetobatis narinari Cartagena, Colombia 2 1976
Gulf of Venezuela, Venezuela 1 1978
Rhinoptera bonasus Gulf of Venezuela, Venezuela 9 1977
Rhinobatus percellens Cartagena, Colombia 1 1976
Isla de Margarita, Venezuela 2 1978
Gymnura micrura Isla de Margarita, Venezuela 2) 1978
Dasyatis americana Cartagena, Colombia 1 1976
Isla de Margarita, Venezuela 3 1978
Dasyatis guttata Cienaga Grande, near Santa 10 1975-1976
Marta, Colombia
Cartagena, Colombia 1 1976
Lake Maracaibo, Venezuela Vp) 1977-1978
Isla de Margarita, Venezuela 3 1978
Himantura schmardae Cienaga Grande, near Santa 12 1975-1976
Marta, Colombia
Lake Maracaibo, Venezuela 2 1977
Myliobatis goodei La Plata, Uruguay 4 1979
Site of infection.—Spiral valve. :

Locality.—Rio de la Plata estuary, near Montevideo, Uruguay.

Holotype. —USNM Helm. Coll. No. 75722. Paratypes: USNM Helm.
Coll. No. 75723; Univ. Nebraska State Museum No. 21003.

Discobothrium arrhynchum differs from all other members of the genus
by lacking a myzorhynchus. It most closely resembles D. myliobatidis Dai-
ley and Mudry, 1968 and D. japonicum Yamaguti, 1934 by having relatively
large suckers and D. myliobatidis by exhibiting markedly craspedote im-
mature proglottids, an average of 21 testes per proglottid, and a vagina
extending posteriorly lateral to the testes rather than medial to them. By
possessing 18-28 testes per proglottid, the new species differs markedly
from D. japonicum, which has 6.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1242
3
O}
oF
OC;
O}
O}
er;
i
2

Figs. 1-6. Discobothrium arrhynchum: 1, Scolex; 2, Mature proglottid. Caulobothrium
uruguayense: 3, Scolex; 4, Mature proglottid. Caulobothrium ostrowskiae: 5, Scolex; 6, Mature

proglottid.

Caulobothrium Baer, 1948

Two recognized genera, Caulobothrium Baer, 1948 and Rhabdotoboth-
rium Euzet, 1953, possess scolices comprising 4 pedicellated septate both-
ridia without marginal loculi and no myzorhynchus along with proglottids

VOLUME 93, NUMBER 4 1243

exhibiting postvaginal testes. The validity of the genera as natural (mono-
phyletic) groups has been questioned, most recently by Appy and Dailey
(1977). Appy and Dailey concluded that the species included in Cauloboth-
rium and Rhabdotobothrium represented a group distinct from those in-
cluded in Rhinebothrium Linton, 1890 whose species exhibit only prevaginal
testes. However, they did not agree that the distinctions between Caulo-
bothrium and Rhabdotobothrium, the presence or absence of a cephalic
peduncle, respectively, constituted valid grounds for generic distinction be-
cause species assigned to both Caulobothrium and Rhinebothrium possess
peduncles of varying lengths. We examined type-specimens of the following
species of Caulobothrium and Rhabdotobothrium as well as published de-
scriptions of all 10 previously-known species and specimens of 2 new
species described in this paper: Caulobothrium myliobatidis Carvajal, 1977
(USNM Helm. Coll. No. 74143), C. anacolum Brooks, 1977 (73969, 73970),
C. multorchidum (Young, 1954) Appy and Dailey, 1977 (45976, 74598), C.
opisthorchis Riser, 1955 (37415), C. tetrascaphium Riser, 1955 (37414), C.
longicolle (Linton, 1890) Baer, 1948 (7663, 34959, 35940, 36008), and Rhab-
dotobothrium anterophallum Campbell, 1977 (73203-4). We discovered 2
distinct groups regardless of peduncle length. One group of species is char-
acterized by 40 or fewer proglottids per strobila, acraspedote proglottids,
and fewer than 100 testes per proglottid. Members of that group include C.
opisthorchis, C. multorchidis, C. myliobatidis, C. anacolum, and one of the
new species described herein. Of those, only C. anacolum lacks postovarian
testes, a trait unique among rhinebothriine cestodes to the other 4 species.
The second group, containing C. tetrascaphium, C. longicolle, C. insignis
(Southwell, 1911) Baer, 1948, C. tobijei (Yamaguti, 1934) Baer, 1948, Rhab-
dotobothrium dollfusi Euzet, 1953, R. anterophallum, and the second new
species to be described, is characterized by more than 100 proglottids per
strobila, craspedote proglottids, preovarian testes, and more than 100 testes
per proglottid. Of those species, the 2 placed in Rhabdotobothrium lack any
cephalic peduncle whereas the other species all possess very long cephalic
peduncles. Members of the first group of species all exhibit moderate-length
peduncles. Two interpretations seem possible; first, all the above species
represent a monophyletic group with 2 divergent lineages or, secondly, each
of the 2 groups is derived independently from a different Rhinebothrium
species-group. In neither case would Rhabdotobothrium be logically con-
sidered a valid genus unless each of the 2 groups of Caulobothrium were
also accorded generic status. If the second possibility is true, suggesting
that the presence of postvaginal testes is either a homoplastic or plesio-
morphic trait, species assigned to Rhinebothrium would also have to be
segregated into various generic groupings. Pending a phylogenetic analysis
of all rhinebothriine groups, we retain Caulobothrium and consider Rhab-
dotobothrium a junior subjective synonym. Rhabdotobothrium dollfusi be-

1244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

comes Caulobothrium dollfusi (Euzet, 1953) comb. n. and R. anterophallum
becomes C. anterophallum (Campbell, 1977) comb. n.

Caulobothrium uruguayense, sp. nov.
Figs. 3-4.

Description (based on 30 specimens).—Strobila craspedote, apolytic, up
to 30 mm long, composed of 100-150 proglottids. Scolex with 4 pedicellated
bothridia, 825—1,100 long by 825-1,100 wide; pedicels 75-160 long. Both-
ridia elongate, 930-1,302 long by 232-418 wide, divided horizontally by 13
or 14 septa forming 14 or 15 total loculi. Cephalic peduncle long, aspinose,
1,860—2,418 long. Immature proglottids wider than long. Mature proglottids
465-651 long by 391-512 wide. Testes in anterior 4/s of proglottid, 12—26 in
diameter, 136-223 (¢ = 185, n = SO) in number, 29-50 (% = 44) preporally,
36-58 (x = 52) postporally, 67-103 (x = 89) antiporally. Cirrus sac in an-
terior 4% of proglottid, elongate, posterior end recurved, 72-384 long by 19—
36 wide, containing spined eversible cirrus. Genital atrium shallow, simple.
Genital pore 32-39% (x = 35%) of proglottid length from anterior end. Va-
gina anterior to cirrus sac, vaginal sphincter weakly developed. Ovary fol-
licular, bialate, X-shaped in cross section, in posterior '/s of proglottid, 186-
279 long by 326-419 wide at isthmus. Vitellaria follicular, follicles 7-10 in
diameter.

Host.—Mpyliobatis uruguayensis.

Site of infection.—Spiral valve.

Locality.—Rio de la Plata estuary, Uruguay.

Holotype. —USNM Helm. Coll. No. 75724. Paratypes: USNM Helm.
Coll. No. 75725; Univ. Nebraska State Museum No. 21002.

Etymology.—This species is named after the country in which its host
was collected.

Caulobothrium uruguayense possesses a very long cephalic peduncle,
thus differing from C. dollfusi and C. anterophallum which lack peduncles.
By possessing genital pores in the anterior 4 of the proglottid, C. uru-
guayense resembles C. tetrascaphium; all other species exhibit genital pores
at mid-proglottid. The new species differs from C. tetrascaphium by pos-
sessing 14 or 15 bothridial loculi rather than 25 and by exhibiting cirrus sacs
which are curved anteriorly at their posterior ends rather than extending in
a direct line or slightly posterior from the genital pore.

Caulobothrium ostrowskiae, sp. nov.
Figs. 5-6

Description (based on 15 specimens).—Strobila acraspedote, apolytic,
composed of 16-20 proglottids, up to 15 mm long. Scolex with 4 pedicellated
bothridia, 550-825 long by 550-825 wide; pedicels 93-112 long. Bothridia

VOLUME 93, NUMBER 4 1245

elongate, 744-791 long by 279-326 wide, divided longitudinally by single
median septum, horizontally by 17-19 septa forming 2 parallel rows of 36,
38, or 40 loculi plus terminal loculus at anterior tip of proglottid; total num-
ber of loculi 37, 39, or 41 (mode = 41). Cephalic peduncle aspinose, 232-
373 long. Immature proglottids wider than long. Mature proglottids 893-
1,256 long by 167-233 wide. Testes extending nearly entire length of pro-
glottid, 19-26 in diameter, 41-63 (< = 52, n = 25) in number, 6-8 (x = 7)
preporally, 13-24 (< = 19) postporally, 21-33 ( = 26) antiporally. Cirrus
sac elongate, 72-132 long by 12-17 wide, containing spined eversible cirrus.
Genital atrium shallow, simple. Genital pore 25—33% (* = 29%) of proglottid
length from anterior end. Vagina anterior to cirrus sac, vaginal sphincter
weakly-developed. Ovary in posterior %3 of proglottid, follicular, bialate,
X-shaped in cross section, 36—96 long by 24—60 wide at isthmus. Postovarian
testes present. Vitellaria follicular, extending nearly entire length of pro-
glottid; folicles 12-17 in diameter.

Host.—Myliobatis goodei.

Site of infection.—Spiral valve.

Locality.—Rio de la Plata estuary, near Montevideo, Uruguay.

Holotype. —USNM Helm. Coll. No. 75726. Paratypes: USNM Helm.
Coll. No. 75727; Univ. Nebraska State Museum No. 21004.

Etymology.—This species is named for Dr. Margarita Ostrowski de Nu-
nez, who reported the first known elasmobranch cestodes from the Rio de la
Plata region.

Caulobothrium ostrowskiae resembles C. myliobatidis, C. opisthorchis,
and C. multorchidum by exhibiting postovarian testes. Caulobothrium op-
isthorchis differs by having 78-90 rather than 41-63 testes per proglottid
and vitelline follicles which encircle the postovarian testes. Caulobothrium
multorchidum possesses flaplike rather than elongate bothridia which are
markedly different from those of the new species. Finally, C. ostrowskiae
differs from C. myliobatidis by having 37-41 rather than 54—58 bothridial
loculi. :

Phyllobothrium Van Beneden, 1850

The genus Phyllobothrium contains a number of species infecting a va-
riety of elasmobranchs and possessing a variety of scolex morphologies
(Williams, 1968). The most simplified, and presumably plesiomorphic, mor-
photype consists of 4 flaplike bothridia lacking a muscular rim and therefore
lacking a distinct shape. Some species possessing such bothridia have apical
suckers surmounting each bothridium and some do not; likewise, some
species possess marginal loculi, such as P. centrurum Southwell, 1925 or
even marginal loculi and horizontal septa across the face of the bothridia,
producing medial loculi, as in Phyllobothrium kingae Schmidt, 1978. The

1246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

genus almost certainly represents a polyphyletic group whose phylogenetic
relationships are virtually unknown. Williams (1968) suggested that mem-
bers of the genus be catalogued and identified according to their hosts.
However, in this study we collected specimens of 2 species of Phylloboth-
rium, a large species with the archetypa scolex morphology and a second,
smaller species possessing amorphous bothridia and marginal loculi. There-
fore, for convenience and better consistency we prefer to catalogue Phyl-
lobothrium species according to general scolex morphology, recognizing
that only a thorough phylogenetic analysis will produce an adequate clas-
sification of the species involved.

Phyllobothrium myliobatidis, sp. nov.
Figs. 7-9

Description (based on 4 specimens).—Strobila acraspedote, apolytic,
composed of 50-75 proglottids, up to 30 mm long. Scolex up to 2.5 mm
wide, composed of 4 pedicellated bothridia with marginal loculi. Pedicels
419-512 long. Bothridia 651—-1,395 wide, amorphous, with single row of 83-
90 marginal! loculi. Cephalic peduncle aspinose, 1,023—1,302 long. Immature
proglottids wider than long. Mature proglottids 465-2,418 long by 251-474
wide. Testes in anterior %4 of proglottid, 24-72 in diameter, in single field
of 122-150 in number. Cirrus sac in anterior 4 of proglottid, elongate, 168—
240 long by 84-168 wide, containing spined eversible cirrus. Genital atrium
shallow, simple. Genital pore 15-27% of proglottid length from anterior end.
Vagina anterior to cirrus sac, vaginal sphincter present. Ovary in posterior
1/. of proglottid, H-shaped with posterior lobes expanding posteriorly as
proglottid matures; ovarian lobes X-shaped in cross section, follicular. Vitel-
laria follicular; follicles 7-24 in diameter, extending entire length of pro-
glottid, becoming more extensive as proglottid matures. Terminal proglottids
possessing vitelline fields extending into medial portion of proglottid dorsally
and ventrally.

Host.—Myliobatis goodei.

Site of infection.—Spiral valve.

Locality.—Rio de la Plata estuary, Uruguay.

Holotype.—USNM Helm. Coll. No. 75728. Paratypes: USNM Helm.
Coll. No. 75729.

Phyllobothrium myliobatidis most closely resembles P. auricula Bene-
den, 1858 by having acraspedote proglottids with genital pores in the ante-
rior 4% and 122-150 testes, band-like vitellaria which proliferate as the pro-
glottid matures, and by having similar ovarian and bothridial morphology
(marginal loculi with no transverse septa present on bothridia). Euzet’s
(1959) illustration of P. auricula shows an apical sucker on each bothridium

VOLUME 93, NUMBER 4

@00%cee
Cecceore

1247

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= =
Gb =
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RY

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Figs. 7-9. Phyllobothrium myliobatidis: 7, Scolex; 8, Mature proglottid; 9, Gravid pro-

glottid.

which appears more like a strongly developed marginal loculus than a true
sucker. The new species exhibits a similar structure in some specimens.
Phyllobothrium myliobatidis differs from P. auricula primarily by exhibiting
longer and thinner bothridial pedicels, and by having cirrus sacs 168-240
wm long by 84-168 ~m wide whereas those in P. auricula measure up to
400 wm long by 200 wm wide. Phyllobothrium auricula parasitizes Dasyatis
pastinaca in French coastal waters while P. myliobatidis occurs in Myliob-

atis goodei from Uruguayan coastal waters.

1248 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Phyllobothrium sp.

Host.—Mpyliobatis goodei.

Site of infection.—Spiral valve.

Locality.—Rio de la Plata estuary Uruguay.

Specimens deposited.—Univ. Nebraska State Museum No. 21001.

Ten specimens of a species of Phyllobothrium possessing amorphous
bothridia lacking apical suckers or marginal loculi infected one M. goodei.
The specimens were too contracted for adequate identification or descrip-
tion, but we have deposited all specimens in hope that they will aid future
workers.

Acanthobothrium sp.

Host.—Myliobatis goodei.

Site of infection.—Spiral valve.

Locality.—Rio de la Plata estuary, Uruguay.

Specimens deposited.—Univ. Nebraska State Museum No. 21000.

A single intact contracted specimen of a large species of Acanthobothrium
occurred in one M. goodei. The species, apparently undescribed, belongs
to a group of species characterized by large size, anapolysis, long aspinose
cephalic peduncles, expanded necks, sessile and relatively broad bothridia,
large and robust bothridial hooks, more than 100 proglottids per strobila,
flat follicular ovaries, more than 100 testes per proglottid, and relatively
square proglottids. Due to the contracted nature of the strobila in our single
specimen, we present only the following partial description:

Scolex 825 long by 1,017 wide, comprised of 4 sessile triloculate bothridia
each surmounted by apical pad and sucker, armed with pair of bifid hooks;
bothridia connected by velum at posterior ends. Bothridia 651-670 long by
409-418 wide; ratio of length to width 1:0.63. Ratio of locular lengths
1:0.44:0.32. Apical pad 326-335 in diameter; suckers 74-93 in diameter.
Bothridia hook prongs markedly dimorphic. Bothridial hook formula (mod-
ified from that of Euzet, 1956):

108 96-120 55-72
204-211
Neck expanded at insertion to scolex, aspinose. Cephalic peduncle 1,925

long, aspinose. More than 100 proglottids present. Mature proglottids ap-
parently wider than long to squared. Ovary follicular, with flat lobes.

|

VOLUME 93, NUMBER 4 1249

Acanthobothrium sp.

Host.—Myliobatis goodei.

Site of infection.—Spiral valve.

Locality.—Rio de la Plata estuary, Uruguay.

Specimen deposited.—Univ. Nebraska State Museum No. 20999.

We collected a single apparently very young specimen of Acanthoboth-
rium possessing 2 elongate immature proglottids. We present a description
of the scolex of this specimen:

Scolex 825 long by 1,237 wide, comprised of 4 triloculate bothridia free
at posterior end, each surmounted by apical pad and sucker, armed with
pair of bifid hooks. Bothridia 837-930 long by 604-698 wide; ratio of locular
lengths 1:0.75:0.90. Apical pad 192 in diameter, sucker 72 in diameter. Both-
ridial hook formula (modified from that of Euzet, 1956):

41-48 84-89 96-99
137-144

Neck expanded at insertion to scolex, spinose.

Ostrowski de Nunez (1971) reported 2 species of Acanthobothrium from
the Argentinean side of the La Plata estuary infecting Zapteryx brevirostris
(Miller and Henle) (Rhinobatiformes: Rhinobatidae). She collected a single
immature specimen, Acanthobothrium sp., possessing markedly dimorphic
bothridial hook prongs, and bothridial hook proportions, scolex, and stro-
bilar morphology similar to the first specimen listed above. However, both
hook prongs and thus the total hook length in our specimens are greater
(96-120 vs. 78 for the inner prong, 55—72 vs. 26-39 for the outer prong, and
204-211 vs. 156-169 for total hook length) than that reported by Ostrowski
de Nunez (1971). It is possible that both specimens represent the same
species. The second species reported by Ostrowski de Nunez, Acantho-
bothrium zapterycum, exhibits smaller scolex dimensions ‘and bothridial
hook sizes as adults than does the single immature specimen we collected.
Thus, at least 3 and possibly 4 different species of Acanthobothrium occur
in stingrays inhabiting the La Plata region, although only one has been
named.

Discussion

As mentioned in the introduction, this paper concludes a 5-year study of
the cestode parasites infecting marine elasmobranchs along the eastern and
northern coast of South America. Table 1 presents the hosts collected and

1250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Cestode species collected in South American marine elasmobranchs during 1975—

1979.

COLOMBIA
DASYATIS AMERICANA
Phyllobothrium cf. kingae Brooks and
Mayes, 1980
Polypocephalus medusius: Brooks and
Mayes, 1980
HIMANTURA SCHMARDAE

Acanthobothroides thorsoni Brooks,
1977
Acanthobothrium tasajerasi Brooks,
1977
Acanthobothrium himanturi Brooks,
1977
UROLOPHUS JAMAICENSIS

Acanthobothrium cartagenensis
Brooks and Mayes, 1980
Phyllobothrium cf. kingae Brooks and
Mayes, 1980
UROTRYGON VENEZUELAE
Acanthobothrium urotrygoni Brooks
and Mayes, 1980
AETOBATIS NARINARI
Acanthobothrium colombianum
Brooks and Mayes, 1980
NARCINE BRASILIENSIS

Acanthobothrium lintoni: Brooks and
Mayes, 1978

VENEZUELA
NARCINE BRASILIENSIS
Acanthobothrium electricolum: Mayes
and Brooks, 1980
RHINOPTERA BONASUS
Rhinoptericola megacantha: Mayes
and Brooks, 1980
Dioecotaenia campbelli Mayes and
Brooks, 1980
GYMNURA MICRURA
Acanthobothrium fogeli: Mayes and
Brooks, 1980
AETOBATUS NARINARI

Acanthobothrium tortum: Mayes and
Brooks, 1980

Lecanicephalum peltatum: Brooks and
Mayes, 1980

Rhinebothrium magniphallum Brooks,
1977

Rhinebothrium tetralobatum Brooks, 1977

Caulobothrium anacolum Brooks, 1977

Rhinebothrium magniphallum: Brooks
and Mayes, 1980

Rhinebothrium magniphallum: Brooks
and Mayes, 1980

Acanthobothrium electricolum Brooks and
Mayes, 1978

Rhodobothrium paucitesticulare Mayes
and Brooks, 1980

Tylocephalum sp. Mayes and Brooks,
1980

Disculiceps sp. Mayes and Brooks, 1981

VOLUME 93, NUMBER 4

1251

Table 2.—Continued.

HIMANTURA SCHMARDAE

Parachristianella cf.
monomegacantha Mayes and
Brooks, 1980

DASYATIS GUTTATA

Acanthobothroides thorsoni: Mayes
and Brooks, 1980

Acanthobothrium tasajerasi: Mayes
and Brooks, 1980

Acanthobothrium urotrygoni: Mayes
and Brooks, 1980

DASYATIS AMERICANA

Acanthobothrium americanum: Mayes
and Brooks, 1980

Phyllobothrium centrurum: Mayes and
Brooks, 1980

Rhodobothrium pulvinatum: Mayes
and Brooks, 1980

URUGUAY

MYLIOBATIS GOODEI

Discobothrium arrhynchum sp. n.
Caulobothrium uruguayense sp. n.
Caulobothrium ostrowskiae sp. n.

Rhinebothrium magniphallum: Mayes and
Brooks, 1980

Rhodobothrium pulvinatum: Mayes and
Brooks, 1980

Rhinebothrium magniphallum: Mayes and
Brooks, 1980

Rhinebothrium margaritense Mayes and
Brooks, 1980

Rhinebothrium corymbum: Mayes and
Brooks, 1980

Rhinebothrium margaritense Mayes and
Brooks, 1980

Phyllobothrium myliobatidis sp. n.
Phyllobothrium sp.
Acanthobothrium sp.

Acanthobothrium sp.

the collection localities, and Table 2 presents an annotated list of the ces-

todes we collected.

Literature Cited .

Appy, R., and M. D. Dailey. 1977. A new species of Rhinebothrium and redescription of
three rhinebothriate species from the Round Stingray, Urolophus halleri Cooper in
Southern California.—Bull. So. Calif. Acad. Sci. 76:116—-127.

Brooks, D. R. 1977. Six new species of tetraphyllidean cestodes, including a new genus, from
a marine stingray Himantura schmardae (Werner, 1904) from Columbia.—Proc. Helm.

Soc. Wash. 44:51-59.

and M. A. Mayes. 1978. Acanthobothrium electricolum sp. n. and A. lintoni Goldstein,

Henson, and Schlicht, 1969 (Cestoda: Tetraphyllidea) from Narcine brasiliensis (Olfers)
(Chondrichthyes: Torpedinidae) in Colombia.—J. Parasitol. 64:617-619.

Soc. Wash. 47:22-29.

. 1980. Cestodes in four species of euryhaline stingrays from Colombia.—Proc. Helm.

Euzet, L. 1959. Thése presentées a la Faculté des Sciences de Montpellier pour obtenir le
grade de Docteur es Sciences Naturelles: 1. Recherches sur les cestodes Tetraphyllides
de Selaciens des cétes de France.—Causse, Graille, and Castelnau, Montpellier, 263 pp.

1252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mayes, M. A., and D. R. Brooks. 1980. Cestode parasites of some Venezuelan stingrays.—
Proc. Biol. Soc. Wash. 93(4):1230-1238.

Ostrowski de Nunez, M. 1971. Estudios preliminares sobre la fauna parasitaria de algunos
elasmobranquios del litoral bonaerense, Mar del Plata, Argentina. I. Cestodes y tre-
matodes de Psammobatis microps (Gunther) y Zapteryx brevirostris (Muller y Henle).—
Physis 30:425—446.

Williams, H. H. 1968. The taxonomy, ecology, and host-specificity of some Phylobothriidae
(Cestoda: Tetraphyllidea), a critical revision of Phyllobothrium Beneden, 1849 and com-
ments on some allied genera.—Phil. Trans. R. Soc. London Ser. B, 253:231-307.

(DRB) Department of Pathology, National Zoological Park, Smithsonian
Institution, Washington, D.C. 20008 (present address: Department of Zo-
ology, University of British Columbia, 2075 Wesbrook Mall, Vancouver,
British Columbia, Canada V6T 1W5); (MAM) Environmental Sciences Re-
search, 1702 Building, Dow Chemical USA, Midland, Michigan 48640; (TBT)
School of Life Sciences, University of Nebraska—Lincoln, Lincoln, Ne-
braska 68588.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1253-1263

NOTES ON AXIOPSIS (AXIOPSIS) SERRATIFRONS
(A. MILNE EDWARDS) (CRUSTACEA:
DECAPODA: THALASSINIDEA)

Brian Kensley

Abstract.—Axiopsis (Axiopsis) serratifrons (A. Milne Edwards) is record-
ed for the first time from several localities in the Atlantic. The species is
redescribed, and observations on geographical distribution, morphological
variation, and behavior in captivity are provided. It is concluded that this
is a widely distributed tropical shallow water species.

The capture of several specimens of the common axiid shrimp inhabiting
the shallow back-reef area at Carrie Bow Cay, Belize, led to problems of
identification, which were complicated by examination of material from
Florida and Bermuda. That axiid taxonomy, especially the status of the
genera, is in flux (de Saint Laurent, 1979) did not help to resolve these
problems. It was felt that refiguring and describing this, the type-species of
the genus Axiopsis, and commenting on the range of variation, would cast
a little light on axiid taxonomy.

Family Axiidae
Axiopsis (Axiopsis) serratifrons (A. Milne Edwards)
Figs. 1-5

Axia serratifrons A. Milne Edwards, 1873:11, pl. 2, fig. 6.

Axiopsis serratifrons.—Borradaile, 1903:538.—Sendler, 1923:44, pl. 6,
fig. 10.

Axiopsis (Axiopsis) serratifrons.—de Man, 1925:72, pl. 6, fig. 12.

Axius serratifrons.—Rathbun, 1906:895.—Edmondson, 1923:27.

Axius spinipes de Man, 1888:464, pl. 19, fig. 6.—Zehntner, 1894:195.

Axiopsis spinipes.—Nobili, 1906:91.—Borradaile, 1903:538; 1910:262.

Axius affinis de Man, 1888:469, pl. 20, fig. 1.

Axiopsis affinis.—Borradaile, 1903:538; 1904:752.—Nobili, 1906:92.

Description.—Male. Carapace strongly sclerotised, with scattered shal-
low pits over entire surface posterior to cervical groove; rostrum triangular,
extending beyond eyes, margins dentate/spinose, continuous with spinose
lateral carina of gastric region of carapace; lateral carina not quite reaching
cervical groove posteriorly; spinose median carine starting at about mid-
length of rostrum and extending as far as lateral and submedian carinae;

1254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

eo OF I tg
Gy Ei 8 i
<é Vee A J
\ Up ; SE
TMK\\ ®)
l \
TSN HN )

Bi \

4h

Yun
foe ZH
C4 U

Fig. 1. Axiopsis (A.) serratifrons. Adult in lateral view.

latter starting in gastric region, spinose; numerous rounded and acute scat-
tered tubercles between median and submedian carinae; antennal spine very
short, subacute; anteroventral region of carapace broadly rounded; supra-
branchial groove faint.

Pleon segments with scattered shallow pits. Pleuron of pleonite 1 ventro-
laterally narrowed, with short distal marginal spine; pleuron of pleonite 2
broadly rectangular; pleura of pleonites 3—6 each with ventrolateral marginal
Spine.

Eyes extending to distal end of basal antennular segment; latter with short
distolateral spine; second and third segments subequal; flagella equal in
length. Antenna with outer peduncle spines flattened, elongate; inner spines
short; fifth segment about two-thirds length of fourth. Mandibular palp
3-segmented, second segment twice length of basal segment; third segment
longer and broader than second, with cluster of short curved spines on
rounded apex; cutting edge bluntly rounded; molar transverse, rounded,
with outer part hollowed. Maxilla 1 bilobed, both lobes distally broad, bear-
ing short spines and setae; slender sparsely setose exopod 2-segmented.
Maxilla 2, posterior lobe of scaphognathite with elongate modified seta
armed with tiny spinules. Maxilliped 1 with broad bilobed epipodite. Max-
illiped 2 terminal segment of endopod armed with short curved spines;

VOLUME 93, NUMBER 4 1255

fourth segment elongate, with elongate setae on median margin; epipodite
narrow, flattened, with attached podobranch. Maxilliped 3 with dactylus
and propodus setose; carpus with single distal spine; merus with 4 spines
on median margin increasing in length distally; ischium with 3 short spines
on median margin, triquetral, inner surface with row of about 20 spines on
raised ridge; basal segment with apically acute lobe on inner distal angle;
epipodite slender, elongate, with attached podobranch. Pereopod 1, smaller
cheliped, with fingers of chela shorter than palm, with ragged cutting edges;
palm parallel-sided; carpus shorter than propodal palm; merus with single
strong spine on upper margin, lower margin with 4 strong spines; ischium
with 4 spines on inner ventral margin; coxa with short mediodistal spine.
Larger cheliped, fingers only slightly shorter than palm; dactylus distally
strongly curved, with single strong triangular tooth proximally; propodal
finger with several tiny teeth proximally, becoming more irregular distally,
inner and outer propodal surfaces bearing low tubercles. Pereopod 2 fingers
slightly shorter than palm of chela, cutting edges straight, armed with short
spines; carpus equal in length to chela; merus and ischium each with 4
spines on ventral margin; coxa with hooked spine at posterodistal corner.
Pereopod 3 dactylus with 2 rows of short articulating ventral spines; pro-
podus with single row of about 6 short spines on outer ventral submargin,
distal spine strongest; carpus shorter than propodus, unarmed; merus with
single strong ventrodistal spine; coxa with hooked spine at posterodistal
corner. Dactylus of pereopod 4 with 2 rows of spines on outer surface;
propodus with 2 rows of spine clusters on outer surface, plus dense ven-
trodistal cluster of slender finely serrate spines; carpus shorter than pro-
podus, unarmed; merus with single ventrodistal spine; coxa with median
hooked spine; sternal plate with strong anteriorly-directed spine on some-
what flattened process on each side. Pereopod 5 shorter than pereopod 4,
dactylus with several ventral spines; grooming apparatus consisting of prox-
imal hollow in dactylus with row of peg-like scales on outer margin; hollow
accommodating ventrodistal spinose lobe of propodus, latter with several
simple and dentate spines, plus dense cluster of ventrodistal slender finely
serrate spines; carpus about half length of propodus; merus, ischium, basis,
and coxa unarmed; flattened articulating plate at outer proximal part of coxa
distally either with single spine and small papilla or (more often in Belize
material), bispinose. First pleopods absent. Endopod of pleopod 2 with slen-
der appendix interna bearing distal patch of short hooks; appendix mascu-
lina slightly shorter but broader than appendix interna, with 7 elongate sim-
ple distal setae. Outer uropodal ramus with row of 11 to 13 small spines
along transverse articulation in distal quarter, strong articulating spine on
outer margin at transverse articulation; outer surface with 2 strong rounded
ridges, outer ridge bearing 4 spines; outer margin with 4 distal spines; inner
uropodal ramus with 3 spines on outer (anterior) margin, single strong ridge

1256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. ] Y

j G

Uy

y
a Z
Z 2B

Set
Ee er ee ee

Fig. 2. Axiopsis (A.) serratifrons: a, Anterior carapace in dorsal view; b, Telson and right
uropod; c, Mandible, inner view; d, Mandible, outer view; e, Maxilla 1; f, Maxilliped 1; g,
Maxilla 2.

VOLUME 93, NUMBER 4 1257

Fig. 3. a, Axiopsis (A.) serratifrons: Maxilliped 2; b, Maxilliped 3; c, Pereopod 4, dactylus
and propodus; d, Maxilliped 3, inner view of ischium; e, Pleopod 1 °; f, Pereopod 5 coxal
plate, Gilbert Island specimen; g, Pereopod 5 coxal plate, Bermuda specimen; h, Pereopod 5
coxal plate, Carrie Bow Cay specimen. ‘

on outer surface bearing 5 spines. Telson with short median spine on distal
margin; latter almost straight, with 3 small submarginal spines laterally;
lateral margin with 2 strong spines; outer surface with 2 broadly rounded
ridges each ending in strong spine on each half; distal margins of both uro-
podal rami and telson bearing dense plumose setae.

Female.—Essentially similar to male, first pereopods more robust. Pleo-
pod | 2-segmented, slender, distal segment bearing marginal plumose setae.

1258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Axiopsis (A.) serratifrons, grooming apparatus of pereopod 5: a, Gilbert Island; b,
Zululand; c, Bermuda; d, Carrie Bow Cay; e, Florida.

Branchial formula.—
Maxilliped 2 epipodite + podobranch
Maxilliped 3 2 arthrobranchs, epipodite + podobranch
Pereopod 1 2 arthrobranchs, epipodite + podobranch
Pereopod 2 2 arthrobranchs, epipodite + podobranch
Pereopod 3 2 arthropods, epipodite + podobranch

Pereopod 4 2 arthrobranchs, epipodite

VOLUME 93, NUMBER 4 1259

a b Cc

Fig. 5. Axiopsis (A.) serratifrons, appendix masculina and appendix interna: a, Bermuda;
b, Gilbert Island; c, Carrie Bow Cay.

Color notes.—(Belize and Bermuda specimens.) Carapace, pleon, and
appendages chestnut-brown to almost black in males (becoming purple-blue
in alcohol), paler brown to olive green in females. Carapace with pale blotch
on posterior branchiostegite and on anteroventral region of cervical groove.
Pleura of pleonites 3 to 6 with pale anterior half; middorsal articulating areas
of pleonites pale. Finger and thumb of both first chelae orange-brown, rest
of cheliped dark.

Behavioral and biological notes.—The following observations were
made at Carrie Bow Cay, Belize, where A. (A.) serratifrons was common
in the back-reef area. The shrimps were confined to areas of coarse calcar-
eous sands mixed with dead coral rubble and pieces of pavement rock, in
water depths of 0.5-2.0 meters. An estimate of density was difficult, but
one burrow per 5 m? would not be unreasonable.

The burrows were nearly always situated in shallow depressions, with
two or three entrances visible between rubble pieces. The animals could
sometimes be seen at the main entrance of the burrow, with the antennae
exploring the surrounding area. Even with fish or conch bait, however, the
animals could not be induced to leave the burrows.

1260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

During March 1980, a mature male and female were captured and placed
in a narrow aquarium containing coarse sand and rubble, with circulating
sea water. Burrow construction and feeding behavior were observed. When
released onto the surface of the sand, rapid movements of the pleon pro-
duced a hollow which was soon enlarged by excavation. The first 2 pairs of
pereopods plus the third maxillipeds, together with the elongate setae on
their ventral margins, form an efficient basket capable of moving consid-
erable quantities of sand, which were dumped beyond the margin of the
excavation. Within 24 hours a branching burrow system had been excavat-
ed, with interlocking rubble pieces preventing sandfills. The rubble pieces
were manipulated with the first pair of chelipeds, and when placed in po-
sition, rapidly vibrated as if to ensure or improve solidity. Construction and
modification of the burrow continued throughout the 2 weeks of observa-
tions. During the day the shrimps never left the burrow, but at night roamed
about the surface a little distance from the burrow entrance.

Food in the form of lumps of fish or conch would be dragged into the
burrow, and held in place by the third maxillipeds while pieces were torn
off by the mandibles. On several occasions, after feeding for a short while,
the piece of food was crammed into a crevice and sand piled on top to
cover it completely. This food item would be excavated later, more eaten
from it, and reburied.

Ventilation of the burrow could sometimes be seen, with either the male
or female sitting in the lowest part of the burrow, with the pleon fully
extended and the pleopods beating rapidly.

Grooming was frequently performed, the entire body, especially the
clumps of setae on the carapace, and the antennular and antennal flagella
being combed by the grooming apparatus at the distal end of the fifth per-
eopods.

Distribution.—Pacific Ocean: Hawaii (type-locality); Palmyra Is.; Fan-
ning Is.; Gilbert Is.; Bikini Atoll; Samoa; Palau Is.; Noordwachter Is. (Java
Sea); Ambon Is., Obi Is., Damar Is., Lucipara Is., Kur Is., Roti Is. (Indo-
nesia). Indian Ocean: Maldive Is.; Chagos Archipelago; Obock (French So-
maliland), Red Sea; Zululand (South Africa). Atlantic Ocean: Belize; Flor-
ida; Bermuda.

Material examined.—Onotoa, Gilbert Islands, from tidepool; 2 Aug. 1951;
det. L. B. Holthuis ovig. 2, USNM 95559, CL 12.1 mm, TL 29.2 mm; 2°,
USNM 95561, CL 14.3 mm, TL 33.2 mm; 6, USNM 95561, CL 15.0 mm,
TL 35.0 mm. Narnu Is., Bikini Atoll, back reef in lagoon, 3 Apr. 1946; det.
L. B. Holthuis. 6, USNM 95560, CL 11.1 mm, TL 27.6 mm. Sodwana Bay,
Zululand, South Africa, offshore reef, 50 ft, 24-28 July 1976; °, South
African Museum, CL 8.8 mm, TL 22.5 mm; 2 juv., South African Mu-
seum, CL 6.9 mm, 5.5 mm, TL 17.4 mm, 13.4 mm. Carrie Bow Cay,
Belize, back reef coral rubble and coarse sand, 1-2 m, Mar. 1980; ovig.

VOLUME 93, NUMBER 4 1261

2, USNM 18905, CL 23.0 mm, TL 59.0 mm; 2°, USNM 18905, CL
17.8 mm (damaged); 4 6, USNM 18905, CL 19.5 mm, 19.0 mm, 15.6 mm,
11.8 mm, TL 50.5 mm (damaged), 39.1 mm, 29.3 mm. Carrie Bow Cay,
Belize, coral patches, 1 m, 16 May 1977; 2 juv., USNM 18908, CL 6.0 mm,
5.3 mm, TL 14.0 mm, 12.3 mm. Carrie Bow Cay, Belize, 14 May 1977; juv.,
USNM 18907, CL 4.2 mm, TL 10.2 mm. Pigeon Key, Monroe County,
Florida, under dead coral slab, 2 m, 29 May 1970; 6, USNM 18903, CL
23.0 mm, TL 62.0 mm. Pigeon Key, Monroe County, Florida, under dead
coral, 1.5—2 m, 20 Nov. 1969; 6, USNM 18904, CL 22.4 mm, TL 58.0 mm.
North Rocks, Bermuda, 10 m, 7 Oct. 1976; ¢, juv., CL 15.3 mm, 5.0 mm,
TL 40.0 mm, 12.9 mm.

Discussion.—The presence of a transverse suture on the outer uropodal
ramus, the rounded posterodorsal carapace, the rostrum continuous with
the gastric region, elongate antennal spines, and pigmented eyes, place the
present species in’ Axiopsis (Axiopsis), using de Man’s key (1925:1). This
definition may be expanded to include a triangular rostrum with dentate
margins, unequal first chelipeds with fingers shorter than palm, telson with
spinose dorsal keels, pleopod 1 absent in male, and pleopod 2 with rod-
shaped appendix interna and appendix masculina. The present species
agrees with all these features, and falls into that group of species having five
spinose carinae in the gastric region of the carapace. de Man’s key to the
species of Axiopsis leads to A. serratifrons (A. Milne Edwards), a species
which possesses numerous spines or tubercles between the median and
submedian carinae. Although widely distributed throughout the Indo-Pacif-
ic, this species has not previously been recorded from the Atlantic.

That A. serratifrons from the Indo-Pacific is a variable species, was noted
by de Man (1925) when he drew A. affinis and A. spinipes into the same
species. (Borradaile 1903, designated the former as the type-species of the
genus.) This variation, also seen in the Atlantic specimens, is to some degree
a function of age, the larger and smaller first chelae being more noticeably
different in older and fully mature specimens. Variation is also seen in sev-
eral other morphological features, in addition to the chelipeds. These are
summarized in Table 1.

The coxal plate of pereopod 5 varies from having the anterodistal corner
rounded, through rectangular, to a spinose condition. In the Belize sample
of 6 specimens, this plate is either bispinose or rectangular/spinose, with no
apparent correlation with age.

The ventrodistal grooming lobe of the propodus of pereopod 5 is armed
with a varying number of dentate and smooth spines, which also vary in
relative lengths.

The ratio of length of appendix masculina to appendix interna ranges from
0.6 to 0.9, but may be a function of maturity. The appendix masculina bears
a varying number of elongate terminal setae (6-10).

1262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparison of material of A. serratifrons from 5 localities.

Gilbert Is.,
Belize Bermuda Florida Bikini At. Zululand
Carapace carinae spination
1 (left) 19-22 20 20-21 18-22 17-19
2 12-14 13 13-14 12-14 10-11
3 (median) 20-24 23 23 14-22 15-17
4 12-14 13 13 12-15 10-12
5 (night) 20-24 20 19-21 17-21 17-19
Marginal spine on pleura 1, 3-6 1, 3-6 1, 3-6 1, 3-6 1, 3-6
Pereopod 1
Larger chela, length/width 2.1 1.6 1.9 ed 1.9
1.8 1.6
1.6 1.5
Smaller chela, length/width 2,52) 2.1 Dah Dall 223
oad) 1.9
2.1
Pereopod 5, coxal plate Rectangular Rectan- Rectan- Rounded _ Rectan-
to gular gular gular
bispinose
Pereopod 5, grooming lobe 9 5) 12 10 3
of propodus, spination
Appendix masculina/appendix 0.7 0.6 0.7 0.9 2
interna
Appendix masculina setation 7 6 9 10 2
terminal terminal terminal terminal

The palm length/width ratio of the first chelae varies considerably, with
the smaller chela being consistently more slender.

Variation in the degree of pitting of the carapace and rugosity of the
chelae can also be seen, but is difficult to describe other than in relative
terms.

From Table | it is difficult to justify splitting the present material either
into species or subspecies. The Atlantic population perhaps represents an
incursion from the Pacific which took place at a time when the Isthmus of
Panama was submerged. A record of the species from the eastern Pacific
would strengthen this surmise.

The species recorded by Sakai (1970) as Axiopsis (Axiopsis) sp. aff. ser-
ratifrons from Tsushima Is., Japan, is almost certainly not this species, as
both the acute anterior margin of the carapace, and the more spinose merus
and ischium of maxilliped 3 (as figured by Sakai) differ markedly from Milne
Edwards’ species.

VOLUME 93, NUMBER 4 1263

Acknowledgments

My sincere thanks are due to Dr. Richard Heard for the specimens from
Florida, Dr. Wolfgang Sterrer for the specimens from Bermuda, Mr. Antony
Rath for assistance with the capture of some of the Belize specimens and
with keeping the animals in aquaria, and to Drs. F. A. Chace, Jr., R. H.
Gore, and R. B. Manning for reading the manuscript and making many
useful comments.

Collecting trips to Belize by the author were financed in part from a grant
from the Exxon Corporation. This paper is contribution number 64 of the
Smithsonian Institution’s Investigations of Marine Shallow Water Ecosys-
tems Project, Dr. Klaus Ruetzler, Chief Coordinator.

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Rathbun, R. J. 1906. The Brachyura and Macrura of the Hawaiian Islands.—U.S. Fish Com-
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Sakai, K. 1970. A small collection of thalassinids from the waters around Tsushima Islands,
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the Seto Marine Biological Laboratory 18:37—47.

Sendler, A. 1923. Die Decapoden und Stomatopoden der Hanseatischen Stidsee Expedition.—
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Zehntner, L. 1894. Crustacés de |’ Archipel Malais.—Revue Suisse de Zoologie et Annales du
Musée d’ Histoire Naturelle de Genéve 2:135-214.

Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1264-1265

OBANDYELLA, A NEW NAME FOR AIRSUTELLA
BANDY, 1972 (FORAMINIFERIDA),
NON COOPER AND MUIR-WOOD,
1951 (BRACHIOPODA)

D. Haman, R. W. Huddleston, and J. P. Donahue

Hubner (1822:1-10) erected the genus Agrodiaetus to accommodate all
56 species of the European Lycaenidae (Insecta, Lepidoptera) known to
him at that time. The author did not, however, designate a type-species for
the genus. Hemming (1934:109) subsequently designated Papilio damon
Denis and Schiffermuller, 1775, as the type-species of Agrodiaetus Hubner,
1822 (see also Hemming, 1967:33). Tutt (April 1909a:154) established the
genus Hirsutina (type-species Papilio damon Denis and Schiffermuller,
1775) for representatives of West European butterflies. The same author in
the following month (Tutt, May 1909b:108) redundantly redescribed Hir-
sutina with the identical type-species designation. The selection of Papilio
damon Denis and Schiffermuller, 1775, by Hemming (1934:109) as the type
for Agrodiaetus Hubner (ICZN Art. 69a, v) consequently made Hirsutina
Tutt 1909 a junior objective synonym (ICZN Art. 61b).

Kirchner (1933:106) described a new Triassic brachiopod genus from Ger-
many which he named Hirsutina (type-species Spirifer? hirsutus Alberti,
1864). Despite the fact that Hirsutina Tutt, 1909, became a junior objective
synonym of Agrodiaetus Hubner, 1822 as a result of the action of Hemming
(1934) at the time Kirchner (1933) erected Hirsutina, the epithet was not
available for redesignation (ICZN Art. 17). Consequently, Hirsutina Kirch-
ner, 1933 at the time of its establishment became a junior homonym of
Hirsutina Tutt, 1909. This fact was recognized by Cooper and Muir-Wood
(1951:195), and they proposed the replacement name Hirsutella for the ge-
nus Hirsutina Kirchner, 1933 non Tutt, 1909.

Bandy (1972:310) in his discussion of the evolutionary lineage of turbor-
otaloids and globorotaloids distinguished three new subgenera of the genus
Globorotalia Cushman, 1927. These new planktic subgenera were Fohsella,
type-species Globorotalia (F.) praefohsi Blow and Banner, 1966; Menar-
della, type-species G. (M.) menardii (d’Orbigny, 1826); and Hirsutella,
type-species G. (H.) hirsuta (d’Orbigny, 1839). These subgenera have not
been widely recognized by foraminiferologists, a fact commented upon by
Stainforth et al. (1975:28, 32). Despite this lack of acceptance by the sci-
entific community, the epithet Hirsutella Bandy, 1972, is a junior homonym
of Hirsutella Cooper and Muir-Wood, 1951, and consequently requires re-

VOLUME 93, NUMBER 4 1265

jection and replacement under the Law of Homonymy (ICZN Arts. 52, 53,
56).

The name Obandyella is hereby proposed to replace the name Hirsutella
Bandy, 1972, non Hirsutella Cooper and Muir-Wood, 1951. The name is
constructed in honor of Orville Lee Bandy (1917-1973) for his pioneer and
semantic studies in micropaleontology.

Publication permission and assistance by Chevron Qil Field Research
Company is gratefully acknowledged.

Literature Cited

Bandy, O. L. 1972. Origin and development of Globorotalia (Turborotalia) pachyderma
(Ehrenberg).—Micropaleontology 18(3):294-318.

Cooper, G. A., and H. M. Muir-Wood. 1951. Brachiopod homonyms.—J. Wash. Acad. Sci.
41(6): 195-196.

Hemming, F. 1934. The generic names of Holartic butterflies: vol. 1, 1758—1863.—Brit. Mus.

(Nat. Hist.), London, viii + 184 pp.

. 1967. The generic names of the butterflies and their type-species (Lepidoptera: Rho-

palocera).—Bull. Brit. Mus. (Nat. Hist.), Supp. 9, 509 pp.

Hiibner, J. 1822. Systematisch-alphabetisches Verzeichniss aller bisher bey den Fiirbildungen
zur Sammlung europaischer Schmetterlinge angegebenen Gattungsbenennungen; mit
Vormerkung auch augsburgischer Gattungen.—Augsburg, vi + 81 pp.

Kirchner, H. 1933. Die Fossilien der Wtirzburger Trias, Brachiopoda.—Neues Jahrb. Min-
eral., Geol and Palaont., Biel. 71:88-138.

Stainforth, R. M. et al. 1975. Cenozoic planktonic foraminiferal zonation and characteristics
of index forms.—Univ. Kansas Paleo. Contribs., Art. 62 (Univ. Kansas), 425 pp.

Tutt, J. W. 1909a. Nat. Hist. British Butterflies (3):154.

1909b. The generic subdivision of the tribe Lycaenid Plebeiidi—Entomol. Rec.
21:107-108.

(DH and RWH) Chevron Oil Field Research Co., La Habra, California
90631; (JPD) Natural History Museum of Los Angeles County, Los Angeles,
California 90007.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1266-1276

A TABULATION OF THE CRUSTACEA
DECAPODA OF BERMUDA!

John C. Markham and John J. McDermott

Abstract.—tThe first list of decapod crustaceans of the northwestern At-
lantic oceanic island group of Bermuda compiled since 1922 is presented.
Based on published accounts and recent collections, the list includes 276
species in 48 families; of these, 69 species have not been recorded previously
from Bermuda. Only 6 species are endemic. An expanded and fully anno-
tated checklist of Bermudian decapods as well as considerations of the zoo-
geography of that fauna will be published later.

Currently in preparation is a field guide to the *‘Marine fauna and flora of
Bermuda’’ under the joint editorship of Wolfgang and Christiane Sterrer of
the Bermuda Biological Station for Research. It is intended that the field
guide, containing illustrations, brief descriptions, and ecological notes for
all of the common, endemic or otherwise noteworthy species of marine life
from Bermuda being prepared by leading specialists, will be published in
1981. A consequence of that project has been much more intensive collect-
ing in Bermuda than had been undertaken in many years. The present tab-
ulation of the Crustacea Decapoda draws in part on material gathered for
the field guide, but, unlike the latter, it is intended to be comprehensive in
its coverage and is not a manual for identification. It is essentially a very
abbreviated version of a report in preparation to be titled “‘An annotated
checklist of the Crustacea Decapoda of Bermuda,’’ which will constitute a
Special Publication of the Bermuda Biological Station. That checklist wiil
include all known citations of decapods in and near Bermuda along with a
complete catalog of specimens in the collection of the museum of the Ber-
muda Biological Station and notes on their taxonomy, ecology, breeding
periods, etc. Subsequent papers which we anticipate preparing will deal
with the zoogeography of certain species or higher taxa of decapods from
Bermuda and other aspects of their biology.

The systematic arrangement used for the higher taxa is mainly that of
Glaessner (1969). Families and genera of Caridea follow Holthuis (1955),
while families and genera in the Brachyura are ordered after Glaessner

1 Contribution No. 859, Bermuda Biological Station for Research, St. George’s West, Ber-
muda.

VOLUME 93, NUMBER 4 1267

(1969) and Rathbun (1918, 1925, 1930, 1937). Various authorities were con-
sulted for the arrangements of other groups. All specific names, insofar as
possible, have been corrected in light of current opinions.

Bermuda, an oceanic island mass lying some 1,000 km off the eastern
coast of North America, stands atop the Bermuda Rise, which extends
upward from abyssal depths. For purposes of this account, ‘‘Bermuda’’ is
that region from land to a depth of 2,750 m (1,500 fathoms) around the
islands of Bermuda. This is an irregularly shaped area very close to 4,000
km?, approximately within the latitudes from 32°00’ to 32°45’N and longi-
tudes from 64°30’ to 65°15'’W. It extends roughly northeast-southwest and
contains Bermuda and its platform, 2 shallow banks to the southwest (Chal-
lenger and Plantagenet [also called Argus] Banks) and a deep bank to the
northeast.

Although casual observations on Bermuda Decapoda were made from the
time of the first settlement of the islands at the start of the 17th century,
catalogs of the decapod fauna first appeared only rather recently. The ear-
liest published list of Bermuda decapods was that of Jones (1859), who cited
12 species assignable to eight families, with some remarks on diagnostic
characters and habitats in which they occurred. The names which he used
were those of close relatives in Europe and boreal North America, but his
accounts were good enough so that it is possible to refer them to known
Bermuda species. Godet (1960) mentioned 8 species, also by names of
species found elsewhere; because of inadequate characterizations, it is dif-
ficult to determine all of the species to which he was referring. Heilprin
(1888, 1889) made a carefully documented collection and strove to establish
the proper names for the 30 decapod species recorded. Hurdis (1897) ap-
parently copied the list of Jones (1859) directly with no attempts to add or
to correct names. Two major oceanographic expeditions, the Challenger
Expedition (reported by Miers, 1886; Bate, 1888; Henderson, 1888; sum-
marized by Cole, 1901) and the Plankton-Expedition (Ortmann, 1893) col-
lected decapods at and near Bermuda, but there was no effort to make the
collections comprehensive. Young (1900), without additional collecting,
compiled records from most of the authors listed above. He recorded 27
decapod species in 14 families from Bermuda, for which he presented di-
agnoses, Synonymies and numerous illustrations. Rankin (1900), drawing on
very well documented collections of his own plus other material available
to him and consulting most of the papers then published, recorded 57 deca-
pod species in 17 families from Bermuda, including several new records.
Finally, in what remain the standard references, Verrill (1908, 1922) drew
up comprehensive accounts of 119 species of Bermuda decapods in 28 fam-
ilies; he presented full descriptions complete with many good illustrations
for all of the species including several described as new or recorded for the
first time from Bermuda.

1268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Results

In the list of decapods presented below, the status of many species has
been indicated by three symbols. Those whose identification or whose ac-
tually documented occurrence in Bermuda is subject to doubt are indicated
by question marks (?); there are 10 such species. The 69 species recorded
from Bermuda for the first time are labeled with the symbol §. Those
species, 6 in number, which are evidently endemic to Bermuda, are denoted
with an asterisk (*). New species are also indicated by ‘‘n. sp.’’; all are
anomurans and represent perhaps 5 species.

Table 1, on page 1275 following the list, summarizes the decapod crusta-
ceans by infraorder after the removal of 11 questionable records.

CRUSTACEA DECAPODA OF BERMUDA

CLASs CRUSTACEA
SUBCLASS MALACOSTRACA
ORDER DECAPODA
SUBORDER DENDROBRANCHIATA
INFRAORDER PENAEIDEA
SUPERFAMILY PENAEOIDEA
FAMILY PENAEIDAE

Metapenaeopsis goodei (Smith) Penaeus brasiliensis Latreille
§M. hobbsi Pérez Farfante P. duorarum duorarum Burkenroad
M. smithi (Schmitt) Trachypenaeus constrictus (Stimpson)

Trackypenaeopsis mobilispinus (Rathbun)

FAMILY SOLENOCERIDAE
Hymenopenaeus laevis (Bate)

FAMILY SICYONIDAE

Sicyonia wheeleri Gurney

FAMILY ARISTEIDAE
Gennadas elegans (Smith)

SUPERFAMILY SERGESTOIDEA
FAMILY SERGESTIDAE
SUBFAMILY LUCIFERINAE

Lucifer faxoni Borradaile L. typus (H. Milne Edwards)

SUBFAMILY SERGESTINAE

Sergestes arachinpodus (Cocco)
. armatus (Kr@yer)

. atlanticus H. Milne Edwards
. cornutus (Kr@yer)

. crassus Hansen

. edwardsi Kr@yer

. grandis Sund

. hanseni (Ortmann)

. japonicus Bate

. kroyeri Bate

. pectinatus Sund

. robustus Smith

. Sargassi Ortmann
. splendens Hansen
. talismani Barnard
. tenuiremis Krgyer
S. vigilax Stimpson

ANHAARHARRHAR
ANRAHARHRARAURARADA

VOLUME 93, NUMBER 4 1269

SUBORDER PLEOCYEMATA
INFRAORDER STENOPODIDEA
FAMILY STENOPODIDAE

Stenopus hispidus (Olivier) S. scutellatus Rankin
§Microprosthema semilaeve (Von Martens)?

INFRAORDER CARIDEA
SUPERFAMILY OPLOPHOROIDEA
FAMILY OPLOPHORIDAE

Oplophorus spinicauda A. Milne Edwards M. marptocheles (Chace)
O. spinosus (Brullé) M. miccycla (Chace)
Acanthephyra acutifrons Bate M. mollis (Smith)

A. brevirostris Smith M. vesca (Smith)

A. curtirostris Wood-Mason Notostomus distirus Chace
A. eximia Smith N. perlatus Bate

*A. gracilipes Chace N. robustus Smith

A. pelagica (Risso) 2N. westergreni Faxon

A. purpurea A. Milne Edwards Ephyrina bifida Stephensen
2A. rostrata Bate E. hoskynii Wood-Mason
A. stylorostrata (Bate) Hymenodora gracilis Smith
*Meningodora compsa (Chace) Systellaspis braueri (Balss)

S. debilis (A. Milne Edwards)

SUPERFAMILY PASIPHAEOIDEA
FAMILY PASIPHAEIDAE

Pasiphaea hoplocerca Chace P. sulcatifrons Smith
P. loicerca Chace ?P. gilesii Wood-Mason
Parapasiphae macrodactyla Chace ?P. serrata Rathbun

Leptochela bermudensis Gurney

SUPERFAMILY BRESILIOIDEA
FAMILY BRESILIIDAE

Lucaya bigelowi Chace Discias atlanticus Gurney
D. serratirostris Lebour

FAMILY RHYNCHOCINETIDAE

Rhynchocinetes rigens Gordon

SUPERFAMILY PALAEMONOIDEA
FAMILY PALAEMONIDAE
SUBFAMILY PALAEMONINAE

Leander tenuicornis (Say) Palaemon northropi (Rankin)
Brachycarpus biunguiculatus (Lucas)

SUBFAMILY PONTONIINAE

Periclimenes americanus (Kingsley) §P. perlatus Boone

P. anthophilus Holthuis & Eibl-Eibesfeldt §Pontonia mexicana Guérin-Méneville
P. harringtoni Lebour Anchistioides antiguensis (Schmitt)
P. irridescens Lebour Typton tortugae McLendon

Periclimenaeus bermudensis (Armstrong) §Tuleariocaris neglecta Chace

1270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

FAMILY GNATHOPHYLLIDAE
Gnathophyllum americanum Guérin- §Gnathophylloides mineri Schmitt
Méneville

SUPERFAMILY ALPHEOIDEA
FAMILY ALPHEIDAE

Alpheopsis labis Chace A. normanni Kingsley
A. trigonus (Rathbun) §A. nuttingi (Schmitt)
§Neoalpheopsis hummelincki (Schmitt) A. paracrinitus Miers
§Automate gardineri Coutiére A. peasei (Armstrong)
Salmoneus ortmanni (Rankin) 8A. viridari (Armstrong)
§Alpheus armatus Rathbun A. websteri Kingsley

A. armillatus H. Milne Edwards ?Synalpheus brevicarpus Herrick
A. bahamensis Rankin S. fritzmuelleri Coutiére
A. beanii Verrill S. goodei Coutiere

A. bouvieri A. Milne Edwards S. hemphilli Coutiére
A. cylindricus Kingsley 2S. longicarpus (Herrick)
§A. floridanus Kingsley S. minus (Say)

A. formosus (Gibbes) S. townsendi Coutiére
2A. heterochaelis Say Synalpheus spp.

FAMILY HIPPOLYTIDAE

Trachycaris restrictus (A. Milne Edwards) §T. manningi Chace

Hippolyte coerulescens (Fabricius) Latreutes fucorum (Fabricius)
H. zostericola (Smith) Tozeuma carolinense Kingsley
§Thor amboinensis (de Man) §Lysmata grabhami (Gordon)
2T. floridanus Kingsley L. intermedia (Kingsley)

§L. moorei (Rathbun)

FAMILY PROCESSIDAE

Processa bermudensis (Rankin) *P. wheeleri Lebour

SUPERFAMILY PANDALOIDEA
FAMILY PANDALIDAE

Parapandalus richardi (Coutiére) P. martia (A. Milne Edwards)
§Plesionika edwardsii (Brandt) Plesionika sp.
FAMILY PHYSETOCARIDIDAE
Physetocaris microphthalma Chace
INFRAORDER ASTACIDEA

FAMILY NEPHROPIDAE
SUBFAMILY NEPHROPINAE

Nephropsis rosea Bate

INFRAORDER PALINURA
SUPERFAMILY PALINUROIDEA
FAMILY PALINURIDAE

Justitia longimanus (H. Milne Edwards) P. guttatus (Latreille)
Panulirus argus (Latreille) P. laevicauda (Latreille)

VOLUME 93, NUMBER 4 1271

FAMILY SYNAXIDAE

Palinurellus gundlachi Von Martens

FAMILY SCYLLARIDAE
§Arctides guineensis (Spengler) Scyllarides aequinoctialis (Lund)
S. nodifer (Stimpson)

INFRAORDER ANOMURA
SUPERFAMILY THALASSINOIDEA
FAMILY AXIIDAE

§*Axiopsis n. sp.

FAMILY CALLIANASSIDAE

Callianassa branneri (Rathbun) C. longiventris A. Milne Edwards

SUPERFAMILY PAGUROIDEA
FAMILY PAGURIDAE

Pagurus brevidactylus (Stimpson) §Iridopagurus N. sp.

§P. impressus (Benedict) §New genus, new species

§Iridopagurus violaceus De Saint Laurent-| §New genus (?), new species
Dechancé § Unidentified pagurid genus, species

FAMILY PARAPAGURIDAE

Parapagurus pilosimanus Smith

SUPERFAMILY COENOBITOIDEA
FAMILY COENOBITIDAE

Coenobita clypeatus (Herbst)

FAMILY DIOGENIDAE

Allodardanus bredini Haig & Provenzano C. antillensis Stimpson

Calcinus tibicen (Herbst) C. tricolor (Gibbes)

*C. verrillii (Rathbun) §Dardanus imperator (Miers)

§Cancellus sp. 8D. insignis (de Saussure)

§Clibanarius anomalus A. Milne Edwards & — D. venosus (H. Milne Edwards)
Bouvier §Dardanus sp.

§Petrochirus diogenes (Linnaeus)

SUPERFAMILY GALATHEOIDEA
FAMILY GALATHEIDAE
SUBFAMILY GALATHEINAE

*Munida beanii Verrill §M. simplex Benedict

SUBFAMILY MUNIDOPSINAE
Munidopsis serratifrons (A. Milne Edwards)

FAMILY PORCELLANIDAE
Petrolisthes armatus (Gibbes)
SUPERFAMILY HIPPOIDEA
FAMILY ALBUNEIDAE

Albunea paretii Guérin

1272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

FAMILY HIPPIDAE
Hippa testudinaria (Herbst)
INFRAORDER BRACHYURA
SECTION DROMIACEA

SUPERFAMILY DROMIOIDEA
FAMILY DROMIIDAE

Dromia erythropus (Edwards) Dromidia antillensis Stimpson
SUPERFAMILY HOMOLOIDEA
FAMILY HOMOLIDAE
§Homola barbata (Fabricius)
SUPERFAMILY ?
FAMILY HAPALOCARCINIDAE

Troglocarcinus corallicola Verrill

SUPERFAMILY CALAPPOIDEA
FAMILY CALAPPIDAE
SUBFAMILY CALAPPINAE

§Calappa angusta A. Milne Edwards C. gallus (Herbst)
C. flammea (Herbst) C. ocellata Holthuis

Cycloes bairdii Stimpson
SUBFAMILY MATUTINAE
§Osachila antillensis Rathbun
FAMILY LEUCOSIIDAE
SUBFAMILY LEUCOSIINAE
§Iliacantha subglobosa Stimpson Callidactylus asper Stimpson
SUPERFAMILY RANINOIDEA
FAMILY RANINIDAE
8Symethis variolosa (Fabricius)
SECTION OXYRHYNCHA

FAMILY MAJIDAE
SUBFAMILY INACHINAE

Stenorhynchus seticornis (Herbst) Podochela riisei Stimpson

SUBFAMILY TYCHINAE
§Pitho sp.
SUBFAMILY ACANTHONYCHINAE
Acanthonyx petiverii H. Milne Edwards Epialtus bituberculatus H. Milne Edwards
§E. longirostris Stimpson
SUBFAMILY PISINAE

§Nibilia antilocapra (Stimpson) Chorinus heros (Herbst)
?Libinia emarginata Leach

VOLUME 93, NUMBER 4

1273

SUBFAMILY MAJINAE

Mithrax (Mithrax) acuticornis Stimpson

M. (Mithrax) cornutus de Saussure

M. (Mithrax) hispidus (Herbst)

§M. (Mithrax) pleuracanthus Stimpson

8M. (Mithrax) spinosissimus (Lamarck)

M. (Mithraculus) forceps (A. Milne
Edwards)

Stenocionops furcata coelata (A. Milne
Edwards)

§S. spinosissima (de Saussure)

Macrocoeloma subparallelum (Stimpson)
M. trispinosum nodipes (Desbonne)
§Macrocoeloma sp.

Microphrys bicornutus (Latreille)

FAMILY PARTHENOPIDAE
SUBFAMILY PARTHENOPINAE

§Parthenope fraterculus (Stimpson)
P. granulata (Kingsley)

§P. pourtalesii (Stimpson)
P. serrata (H. Milne Edwards)

SECTION CANCRIDEA
FAMILY CANCRIDAE
SUBFAMILY CANCRINAE

Cancer borealis Stimpson

SECTION BRACHYRHYNCHA
SUPERFAMILY PORTUNOIDEA
FAMILY PORTUNIDAE
SUBFAMILY PORTUNINAE

Portunus (Portunus) anceps (de Saussure)
P. (Portunus) sayi (Gibbes)

P. (Achelous) depressifrons (Stimpson)

P. (Achelous) ordwayi (Stimpson)

P. (Achelous) sebae (H. Milne Edwards)
§P. (Achelous) spinicarpus (Stimpson)

P. (Achelous) spinimanus Latreille

Callinectes danae Smith

C. exasperatus (Gerstaecker)

C. marginatus (A. Milne Edwards)
C. ornatus Ordway

C. sapidus Rathbun

Arenaeus cribrarius (Lamarck)
§Cronius ruber (Lamarck)

C. tumidulus (Stimpson)

SUPERFAMILY XANTHOIDEA
FAMILY XANTHIDAE

Carpilius corallinus (Herbst)
Paraliomera dispar (Stimpson)
Platypodiella spectabilis (Herbst)
§Actaea acantha (H. Milne Edwards)
§A. palmeri Rathbun

§Actaea ? sp.

§Allactaea lithostrota Williams
§Paractaea nodosa (Stimpson)
Platyactaea setigera (H. Milne Edwards)
§Platyxanthus sp.

§Cycloxanthops vittatus (Stimpson)
Cataleptodius floridanus (Gibbes)
§Pseudomedaeus agassizi (A. Milne

Edwards)

§Leptodius sp.

Xanthodius americanus (de Saussure)
Xantho denticulatus (White)

Panopeus bermudensis Benedict & Rathbun

P. herbstii H. Milne Edwards

P. occidentalis de Saussure

Eurypanopeus depressus (Smith)

?Eurytium limosum (Say)

§Micropanope sculptipes Stimpson

M. spinipes A. Milne Edwards

§M. urinator (A. Milne Edwards)

§Pilumnus dasypodus Kingsley

§P. floridanus Stimpson

§P. longleyi Rathbun

§P. miersi A. Milne Edwards

§P. sayi Rathbun

Lobopilumnus agassizii (Stimpson)

Heteractaea ceratopus (Stimpson)

Eriphia gonagra (Fabricius)

Domecia acanthophora acanthophora
(Desbonne & Schramm)

1274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

FAMILY GERYONIDAE
Geryon quinquedens Smith
FAMILY GONEPLACIDAE
SUBFAMILY CARCINOPLACINAE

§Pilumnoplax? sp.

SUBFAMILY EURYPLACINAE
Euryplax nitida Stimpson

FAMILY GRAPSIDAE
SUBFAMILY GRAPSINAE

Grapsus grapsus (Linnaeus) Pachygrapsus gracilis (de Saussure)
Geograpsus lividus (H. Milne Edwards) P. transversus (Gibbes)
Goniopsis cruentata (Latreille) Planes minutus (Linnaeus)

SUBFAMILY VARUNINAE

§Euchirograpsus americanus A. Milne Edwards

SUBFAMILY SESARMINAE

Sesarma (Holometopus) miersii Rathbun §Aratus pisonii (H. Milne Edwards)
S. (Holometopus) ricordi H. Milne Edwards Cyclograpsus integer (H. Milne Edwards)

SUBFAMILY PLAGUSIINAE
Plagusia depressa (Fabricius) Percnon gibbesi (H. Milne Edwards)

FAMILY GECARCINIDAE

Cardisoma guanhumi Latrerlle Gecarcinus (Gecarcinus) lateralis lateralis
(Freminville)

SUPERFAMILY OCYPODOIDEA
FAMILY OCYPODIDAE
SUBFAMILY OCYPODINAE

Ocypode quadrata (Fabricius)

SUPERFAMILY ?
FAMILY PALICIDAE

§Palicus affinis (A. Milne Edwards & §P. alternatus Rathbun
Bouvier)

Discussion

The accompanying list includes 276 species (when the ‘‘Synalpheus spp.”’
are handled as one species) belonging to 48 families. Of these, 69 species
(25.0%), including 5 known to be undescribed species, are reported from
Bermuda for the first time. Only 6 species (2.2%) are endemic to Bermuda,
though if some of the unknown species prove to be new, they may raise that
number slightly. With the removal of the 10 dubious species records, the
total number becomes 266 species, which are distributed among 7 infraor-
ders as indicated in Table 1.

VOLUME 93, NUMBER 4 1275

Table 1.—Crustacea Decapoda of Bermuda—summaries by infraorder after removal of 11
questionable records.

Infraorder Number of species % of total Number of endemics New records
Penaeidea 29 10.9 0 1
Stenopodidea 3 il 0 1
Caridea 89 3355 3 15
Astacidea 1 0.4 0 0
Palinura 8 3.0 0 1
Anomura 30 eS 3 14
Brachyura ; 106 39.8 0 37

TOTALS 266 100.0 6 69

Acknowledgments

Many of the collections were made by the authors, but we are also deeply
indebted to J. R. H. Lightbourn, A. T. Guest, W. E. Sterrer and numerous
staff members and visitors to the Bermuda Biological Station for supplying
much valuable material. People who supplemented, confirmed or corrected
our identifications, updated names and made other helpful comments on our
work include L. G. Abele, F. A. Chace, Jr., J. Garcia-Gomez, R. H. Gore,
J. Haig, L. B. Holthuis, W. G. Lyons, R. B. Manning, P. A. McLaughlin,
A. J. Provenzano, Jr., W. E. Sterrer and A. B. Williams. J. J. McDermott
appreciates financial support provided by a Sidney L. Wright Fellowship
from the B.B.S. and an Institutional Grant from F and M College.

Literature Cited

Bate, C. S. 1888. Report on the Crustacea Macrura collected by H. M. S. Challenger during
the years 1873—76.—Rept. Sci. Results Voy. H. M. S. Challenger, Zool. 24(Part 90):i-
xc, 1-942.

Cole, G. W. 1901. Bermuda and the Challenger Expedition: A bibliography giving a summary
of the scientific results obtained by that expedition at and near Bermuda in 1873.—
Printed for private distribution, Boston, pp. 1-16.

Glaessner, M. F. 1969. Decapoda. Jn R. C. Moore, ed., Treatise on invertebrate paleontology,
Part R Arthropoda 4. Vol. 2.—Geological Society of America and University of Kansas,
Lawrence, Kansas, pp. R399-R533.

Godet, T. L 1860. Bermuda, its history, geology, climate, products, agriculture, commerce
and government.—Smith, Elder & Co., London, xv + 271 pp.

Heilprin, A. 1888. Contributions to the natural history of the Bermuda Islands.—Proc. Acad.

Nat. Sci. Philadelphia 1888:302-328.

. 1889. The Bermuda Islands: A contribution to the physical history and zoology of the

Somers Archipelago, with an examination of the structure of coral reefs. Researches

undertaken under the auspices of the Academy of Natural Sciences of Philadelphia.

Published by the author, Philadelphia, 231 pp.

1276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Henderson, J. R. 1888. Report on the Anomura collected by H. M. S. Challenger during the
years 1873-76.—Rept. Sci. Results Voy. H. M. S. Challenger, Zool. 27(Part 69):i-xi,
1-221.

Holthuis, L. B. 1955. The recent genera of the caridean and stenopodidean shrimps (Class
Crustacea, Order Decapoda, Supersection Natantia) with keys for their determination.—

_ Zool. Verhand. 26:1—-157.

Hurdis, J. L. 1897. Rough notes and memoranda relating to the natural history of the Ber-
mudas.—R. H. Porter, London, vi + 408 pp.

Jones, J. M. 1859. The naturalist in Bermuda; a sketch of the geology, zoology, and botany,
of that remarkable group of islands; together with meteorological observations.—Reeves
& Turner, London, vi + 200 pp.

Miers, E. J. 1886. Report on the Brachyura collected by H. M. S. Challenger during the years
1873-76.—Rept. Sci. Results Voy. H. M. S. Challenger, Zool. 17(Part 49):i-l, 1-362.

Ortmann, A. 1893. Decapoden und Schizopoden.—Ergebnisse Plankton-Exped. Humboldt-
Stiftung 2(G)(b): 1-120.

Rankin, W. M. 1900. The Crustacea of the Bermuda Islands, with notes on the collections
made by the New York University Expeditions in 1897 and 1898.—Ann. New York
Acad. Sci. 12(12):521-548.

Rathbun, M. J. 1918. The grapsoid crabs of America.—Bull. U.S. Natl. Mus. 97:i-xxii, 1-

461.

. 1925. The spider crabs of America.—Bull. U.S. Natl. Mus. 129:i-xx, 1-613.

. 1930. The cancroid crabs of America of the families Euryalidae, Portunidae, Atele-

cyclidae, Cancridae and Xanthidae.—Bull. U.S. Natl. Mus. 152:i-xvi, 1-609.

. 1937. The oxystomatous and allied crabs of America.—Bull. U.S. Natl. Mus. 166:i-
vi, 1-278.

Vermrill, A. E. 1908. Decapod Crustacea of Bermuda; I.—Brachyura and Anomura. Their

distribution, variations, and habits.—Trans. Connecticut Acad. Arts Sci. 8:299-474.

. 1922. Decapod Crustacea of Bermuda. Part II. Macrura.—Trans. Connecticut Acad.

Arts Sci. 26:1-179.

Young, C. G. 1900. The stalk-eyed Crustacea of British Guiana, West Indies and Bermuda.—
John M. Watkins, London, xix + 514 pp.

(JCM) Arch Cape Marine Laboratory, Arch Cape, Oregon 97102; (JJM)
Department of Biology, Franklin and Marshall College, Lancaster, Penn-
sylvania 17604.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1277-1294

CORALAXIUS ABELEI, NEW GENUS AND NEW SPECIES
(CRUSTACEA: DECAPODA: THALASSINIDEA:
AXIIDAE): A CORAL-INHABITING SHRIMP
FROM THE FLORIDA KEYS AND THE
WESTERN CARIBBEAN SEA

Brian Kensley and Robert H. Gore

Abstract.—Coralaxius abelei, gen. et sp. nov., is described from speci-
mens collected from 11 to 76 m on coral reefs or coralline rock habitat, off
Carrie Bow Cay, Belize, and Key Largo in the Florida Keys. The new genus
closely resembles Axiopsis (Paraxiopsis) but is separated chiefly by the
reduced gill formula, a broad bilobed appendix masculina in the male, biun-
guiculate dactyls on the last 3 pereopods, and eyestalks longer than the
rostrum. The species also exhibits striking cheliped armature, differing from
all other Axiidae in the western Atlantic.

In May and early June 1979 the Office of Coastal Zone Management
undertook a geophysical and biological survey for NOAA of the deep water
coral reefs within the Key Largo Florida Marine Sanctuary. The Sanctuary
extends from the State of Florida Territorial Sea, off Key Largo in the upper
Florida Keys, seaward to the 100 m isobath, and encloses an area 20 miles
long by 5 miles wide (approximately 33 x 8 km). The 53 specimens of deca-
pod crustaceans collected during reconnaissance dives using a manned sub-
mersible in this survey were placed at the first author’s disposal. The col-
lection contained several important range extensions and at least 4 new
species. One of the specimens, a female, was a thalassinidean shrimp with
affinities to the genus Axiopsis (Paraxiopsis), which differed from the 19
other axiids recorded from the western North Atlantic (de Man 1925; Boesch
& Smalley 1972; Williams 1974) in the genera Axius, Axiopsis, and Calocaris
oll:

The description of the new species had already been accepted for publi-
cation when the second author, in conversation with the first, discovered
that the latter had found and prepared a description of the same species
using material collected by the Smithsonian Institution’s Investigations of
Marine Shallow-Water Ecosystems team at Carrie Bow Cay, Belize. The
first author had placed his material in a new genus based primarily on
branchial formula and the form of the male pleopod 2. Accordingly, we
combined our respective descriptions into this report, thereby preventing
certain synonymy.

1278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Abbreviations are as follows: Rel (rostral spine-carapace length), cw (wid-
est part of carapace), ch (highest part of carapace), TI (rostral spine tip to
telson posterior margin). Paratypes have been deposited in the Rijksmuseum
van Natuurlijke Historie, Leiden (RNHL) and British Museum (Natural
History) (BMNH).

Coralaxius, new genus

Diagnosis.—Uropodal exopod with distinct, spined suture; posterior dor-
sal carapace region rounded, smooth, without median carina; rostrum sharp-
ly produced, triangular, margins unarmed, shorter than eyestalks, situated
at lower level from and discontinuous with gastric region; antennal acicle
small, flattened, distally trispinose; eyes pigmented; pereopods 3-5 biun-
guiculate; chelipeds unequal, fingers of larger chela longer or shorter than
palm; maxilliped 2 with simple epipodite; maxilliped 3 with simple podo-
branch and epipodite; 2 arthrobranchs on maxilliped 3, and pereopods 1-4,
gills hardly branched; pleopod 1 of male bilobed, with reduced stylamblys,
pleopod 2 with rod-shaped appendix interna and broad, bilobed appendix
masculina.

Remarks.—The new genus is close to Calocaris Bell, 1853, Calastacus
Faxon, 1893, and the genus Axiopsis s.1. Borradaile, 1903. For Coralaxius,
the greatest weight is given to the very different gill arrangement (Table 1),
to the unusual form of the appendix masculina of pleopod 2 in the male, and
to the presence of biunguiculate dactyls on the last 3 pairs of pereopods.
These and other differences are summarized in Table 2 using data from
original descriptions or examination of type-specimens in the taxa under
consideration. For several of the characters information was not available
in the description, and the figures provided could be misconstrued, hence
the data are somewhat provisional.

Etymology.—The generic name is derived from “*Coral’’ where the shrimp
were found, the *‘axius’’ from the type-genus in the Axiidae. Gender: mas-
culine.

Coralaxius abelei, new species
Figs. 1-6

Material.—Holotype: female, Rcl 4.5, cw 2.6, ch 2.7, Tl 12.8 mm; Atlantic
Ocean, French Reef, off Key Largo, Monroe Co., Florida; 25°02'N,
80°19’'W; 76 m; by manipulator from Research Submersible Johnson-Sea-
Link I; 3 June 1979; G. Shinn collector. USNM 173629.

Allotype: Male, Rcl 2.7, cw 1.3, ch 1.4, Tl 7.6 mm; Caribbean Sea,
Carrie Bow Cay, Belize; 11 m; by hand, from live specimen of Porites coral;
2 February 1978; R. Larson collector. USNM 170857.

Paratypes: (All localities Carrie Bow Cay, Belize.) 1 female, Rcl 3.5, cw

VOLUME 93, NUMBER 4 1279

Fig. 1. Coralaxius abelei: Holotype female in dorsal view.

1280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

er \
eS = Sha

Pres Pty

=

|
> a
: PQ

Fig. 2. Coralaxius abelei: Adult in lateral view, pereopods 3—5 removed.

1.7, ch 1.7, Tl 9.6 mm; 15 m; 7 February 1978; R. Larson & B. Kensley
collectors. USNM 170856.—1 male (dissected & illustrated), Rcl 2.8, cw
1.3, ch not available, Tl 9.5 mm; 2 females, Rcl 3.3, 2.6, cw 1.5, 1.1, ch 1.6,
1.4, Tl 9.6, 7.5 mm, respectively; 15 m; 7 February 1978; R. Larson & B.
Kensley collectors. USNM 170858.—1 male, Rcl 2.9, cw 2.1, ch 1.5, Tl 9.0;
2 females, Rcl 4.5, 4.3, cw 1.8, 2.4, ch 2.5, 2.2, Tl] 13.4, 10.5 mm, respec-
tively; 26 m; 23 March 1978; R. Larson & B. Kensley collectors. USNM
171765.—1 male, Rcl 2.7, cw 1.4, ch 1.5, Tl 7.7 mm; 26-30 m; 26 March
1978; R. Larson & B. Kensley collectors. USNM 171766.—1 male, Rel 2.3,
cw 0.9, ch 1.3, Tl 6.4 mm; | female, Rcl 3.8, cw 1.5, ch 2.0, Tl 9.9 mm; 26
m; 27 March 1978; R. Highsmith & B. Kensley collectors. USNM 171764.—
1 male, Rel 3.5, cw 1.5, ch 2.0, Tl 9.0 mm; 13 m; 28 March 1978; R. High-
smith & B. Kensley collectors. USNM 171763.—2 males, Rcl 4.5 (gonopod
illustrated), 4.4, cw 1.4, 2.2, ch 2.0, 2.2, Tl ca. 10.5 (telson broken), 12.2

VOLUME 93, NUMBER 4

Fig. 3. Coralaxius abelei: Live adult from Carrie Bow Cay, Belize (Photograph by R.
Larson).

1282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Coralaxius abelei: a, Sternal anterior and posterior processes; b, Larger cheliped,
inner view; c, Smaller cheliped, outer view; d, Telson and uropoda.

mm, respectively; 3 females (1 ovigerous), Rcl (5.0), 4.3, 2.5, cw (2.2), 2.2,
1.3, ch (2.2), 2.1, 1.7, Tl (12.5), 11.0, 6.3 mm, respectively; 21 m; 22 March
1979; B. Kensley collector. RMHNL.—2 males, Rel 4.1, 3.8, cw 2.1,
1.9, ch 1.7, 2.1, Tl 11.8, 11.3 mm, respectively; 28 m; 12 March 1980. B.
Kensley collector. BMNH.

Diagnosis.—Characters as for genus with the following additions, sus-
pected to be diagnostic at the species level: gastric region with low, thin,

VOLUME 93, NUMBER 4 1283

median carina joining raised, posteriorly bifurcate tooth from which extends
posteriorly a rounded ridge; 2 semicircular rows of 5 spines each join with
rostral spine and pyramidal tooth to form distinct gastric corona; minor
chela with 3 elongate fanglike teeth on cutting edge of fixed finger; major
and minor chelipeds heavily armed with short sharp spines and tubercles.

Description.—Carapace smooth, shining; cervical groove distinct, no pos-
terior median carina dorsally; rostrum sharply produced, acutely triangular,
margins entire, curving anterodorsally, reaching about half length of eye-
stalk; carapace in lateral view with dorsum depressed posterior to rostrum,
but rising to meet gastric tooth; gastric region sloping anteroventrally, a low
median carina joining raised tooth, latter bifurcating gently posteriad, merg-
ing along dorsal midline of carapace into low, rounded, carinate ridge be-
coming obsolete towards cephalic groove; shallow depression either side of
this ridge; on either side of rostral spine, a semicircular row of 5 spines,
extending posteriorly and forming with the rostral spine and gastric tooth,
a sparsely setose gastric corona; outer orbital angle produced into blunt
tooth; anterior carapace margin smooth, oblique ventrally, unarmed; pos-
terior margin rounded dorsally, becoming sinuously concave below, with
row of small, plumose setae extending dorsally along this margin to just
inside large median notch of posterodorsal margin.

Abdominal somites smooth, shining, with scattered setae dorsally and
laterally, either singly or in small tufts; somites 1-5 of about equal length
middorsally, somite 6 slightly longer; fifth and sixth, medially carinate, ca-
rina of sixth stronger than fifth. Ventrolateral margins of all somites except
first and sixth truncately rounded, fringed with hairs; pleuron of first pro-
duced posteroventrally into bluntly rounded tooth, that of sixth with pos-
terolateral margin angled, emarginate to accommodate uropodal base.

Eyestalk subglobose proximally, becoming more or less cylindrical dis-
tally; cornea well developed, pigmented, wider than stalk, just reaching
distal end of basal antennular segment.

Antennular peduncle 3-segmented, extending well beyond tips of eyes;
basal article equal to, or slightly overreaching eyestalk, with small spine
distoventrally, 3 short spines on medial margin, single spine on convex outer
margin; stylocerite about half eyestalk length, with single spinule on distal
margin; second article subequal to, third article about 4 of, eyestalk length,
both unarmed. Flagella longer than carapace length, heavily setose through-
out, inner flagellum thicker than outer.

Antennal peduncle compressed, 3-segmented, basal article with single
ventral spine medial to antennal gland opening; second article about half
length of first, dorsally bearing small, flattened, bi- or trispinose scalelike
acicle, second small articulating scale ventrolaterally; third subequal to first,
unarmed; terminal article *4 length of first; flagella heavily setose throughout
length, about twice carapace length.

WS . NX |
SSCA SSSR SEES ‘ y
SSS WY,

— 7

EE
I

\\
LK

Sys

e, Max-

Maxilliped 1;

9

d

1284

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\ ‘
\ LAN

>

c, Maxilla;

Fig. 5. Coralaxius abelei: a, Mandible; b, Maxillule;

illiped 2; f, Antennular peduncle; g, Antennal peduncle.

VOLUME 93, NUMBER 4 1285

Mandibular palp 3-segmented, 2 basal segments subequal in length, ter-
minal segment somewhat longer, armed with simple marginal setae; cutting
edge with straight hyaline border, short triangular tooth at outer and inner
distal angles.

Maxillule bilobed, endopod lacking, basal endite armed with simple spines
and setae, coxal endite tipped with fringed spines. Maxilla well developed,
endopod present, basal and coxal endites bilobed, heavily armed with nu-
merous setae and spines; scaphognathite as illustrated, posterior lobe bear-
ing elongate modified seta armed with tiny spinules proximally, becoming
setulose distally. Maxilliped 1 bearing well developed epipod, posterior lobe
carrying several hooked spines; exopodal lobe elongate, endopod much re-
duced; basal and coxal endites armed with numerous processes. Maxilliped
2 with unbranched epipod; exopod distally setose, endopod heavily setose
and spinose, terminal segment bearing fringed spines. Maxilliped 3 with
unbranched podobranch plus epipod and 2 arthrobranchs; endopod pedi-
form, outer 5 segments densely setose ventrally and mesially, with long
translucent setae; coxa with tuft of setae plus single large spine mediodis-
tally, basis similarly armed, ischium and merus with row of several spines
along inner margin, merus with additional spine at outer distal angle, carpus
with large spine at inner distal angle, dactyl with same at tip; exopod
2-segmented, unarmed, extending nearly to tip of antennal peduncle.

First pereopods large, inflated, chelate, asymmetrical; larger robust,
smaller more scissorslike, both heavily covered with numerous large teeth,
spines, and sharp or rounded tubercles; merus thin, flattened, armed on
medial margin with about 10 large, forward-directed spines, increasing in
size distally, with interspersed translucent setae; posterior margin notice-
ably dentate, more spinulose; carpus about half meral length, inflated, armed
dorsomedially and posteriorly with irregular rows of acute spines, largest
at dorsodistal margin, and outer distal angle. Smaller chela slightly shorter,
much narrower than larger chela; elongate, outer surface of hand with nu-
merous large and small, forward-directed, blunt or sharp tubercles, placed
generally in 3 longitudinal rows; dorsal margin a thin, crenulate crest formed
by connected blunt, inwardly-curving teeth; inner palm with scattered
curved spinelike teeth and larger tubercles, 5 long, translucent setae ex-
tending from oblique posterior row of spines; similar row of setae along
inner ventral margin, arising from bases of acute curved teeth delineating
ventral margin of palm and fixed finger; fingers up to twice as long as palm,
tips curved inward, meeting along entire length except for small gape near
articulation; outer surface of fixed finger with 2 rows of elongate, thin,
knifelike tubercles, cutting edge irregularly dentate, with 3 long, thin, fang-
like spines, curving outwardly and over upper outer margin of dactyl; inner
surface with longitudinal row of small, blunt teeth extending to finger tip;
dactyl with high, thin longitudinal crest formed by series of about 9 sharp

1286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inwardly-curving, connected teeth; thin rounded ledge with series of blunt
teeth paralleling those on both outer and inner surfaces; second series of
subacute tubercles along lower inner margin of dactyl; immediately above
latter, 9 elongate, translucent setae curving down and inwardly over gape,
meshing with lower setae on propodus with fingers closed. Larger chela
robust, inflated proximally, thinning distally; outer surface of hand armed
with numerous, thick, rounded or subacute, raised tubercles, those on lower
margin and near gape more acute, directed obliquely forward; dorsal margin
a crenulate crest of inwardly-curving tubercles; inner palm covered with
small, pustular tubercles and spinules; row of thin, translucent setae extends
from lower inner margin of palm upward toward dactylar-propodal junction,
this area distinguished by large, blunt white tooth in some specimens; fingers
short, about equal to or shorter than length of palm, curved inward, upper
tip crossing over lower, meeting along entire length, without gape; outer
surface of fixed finger with irregular row of elongate tubercles; cutting edge
bluntly dentate in addition to 2 large rounded teeth; inner surface with sev-
eral small, elongate blunt teeth; dactyl with superior dorsal crest formed by
overlapping platelike teeth; inner surface flattened, smooth, with scattered
tubercles and long, translucent setae as in minor chela; outer surface with
rough, longitudinal ridge formed by irregularly fused, blunt tubercles, be-
coming thin rounded ridge distally toward finger tip; cutting edge irregularly
dentate, with 3 large teeth proximally at propodal junction.

Second pereopod chelate, more or less flattened, almost reaching distal
margin of carpus of pereopod 1. Ischium with distoventral spine; merus
thin, anteriorly with 3 or 4 unequal ventral spines, plus another at postero-
ventral angle; carpus somewhat expanded, dentate dorsal crest supporting
long, translucent setae, single strong ventral spine; hand inflated, with den-
tate, setose dorsal crest, plus crenulate ridge on outer surface at dactylar
junction; fingers slightly longer than palm, serrate on cutting edges, upper
tip crossing over lower, without gape; fixed finger with several large teeth
along ventral margin; dactyl with dentate crest bearing translucent setae.

Pereopods 3 and 4 generally flattened, subequal in length; ischia with
rectangular lobe on ventrodistal margin; pereopod 3 merus similar to that
of pereopod 2, but 3 ventral spines on posterior margin instead of anteriorly,
lacking spine on posteroventral angle; carpus unarmed, about half length of
propodus; propodus with 7-9 ventral spinules plus 2 larger spines at disto-
ventral angle; dactyl 7/s length of propodus, with 5 movable spinules, biun-
guiculate.

Pereopod 4 merus and carpus unarmed; propodus with 6 or 7 ventral
spinules, plus 3 elongate fringed spines and thick tuft of setae posteriorly
at distoventral angle; dactylus half length of propodus, otherwise similar in
armature to pereopod 3; biunguiculate.

Pereopod 5 shortest, more cylindrical than preceding, reaching to about

VOLUME 93, NUMBER 4 1287

Fig. 6. Coralaxius abelei: a, Maxilliped 3; b, Pereopod 2; c, Pereopod 5; d, Pereopod 3
distal propodus (some spines removed) and dactylus; e, Pereopod 4; f, Pereopod 3; g, Pleopod
1 ¢;h, Pleopod 2 ¢.

end of carpus of pereopod 4, all segments unarmed except propodus, latter
bearing ventrodistal cluster of simple spines, and distoventral tuft of setae
more elongate than preceding leg; dactylus slightly less than half propodal
length, biunguiculate, with 2 ventral and 2 lateral spinules. Pereopods 1-4
each with 2 arthrobranchs, gills absent from pereopod 5.

1288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Large sternal plate situated between coxae of pereopods 3 and 4; anterior
process smaller, raised, appearing inversely cordate, strongly bifurcate,
with 2 pronounced median ridges coalescing anteriorly into point; posterior
process much larger, similarly shaped, with bifurcation less pronounced,
developed posteriorly into 2 inflated lobes meeting with deep medial cleft.

Pleopods on somites | through 5; those on somite 1 uniramous in female,
bilobed in male, remaining pleopods biramous in both sexes, longer endo-
pods of each with appendix interna. Male pleopod 1 2-segmented, distally
spatulate, distal inner marginal lobe with about 12 tiny hooked spines (re-
duced stylamblys); pleopod 2 endopod with short cylindrical appendix in-
terna bearing several tiny hooked spines, appendix masculina broad, bi-
lobed, armed with numerous simple distal spines.

Tail fan heavily spinose/setose, number and position of spines somewhat
variable. Uropodal exopod with 2—4 spines on outer margin, larger movable
spine at transverse suture; latter with 5-7 heavy spines of varying size;
dorsal surface with 2 distinct, rounded longitudinal ridges, outer bearing 2-4
spines, inner unarmed; uropodal endopod with single dorsal ridge medially,
bearing 4 spines plus spine on endopodal margin; no transverse suture.
Telson armed with 2 proximal spines terminating submedial carinae, 2 distal
spines terminating intermedial carinae, 2 marginal spines on each side; pos-
terior margin with 5 movable spines, longest submarginally at posterolateral
angle; series of stiff setae on posterolateral margin from level of second
marginal spine distally to posterolateral angle; 4 or 5 short setae on each
side of posterior median spine, superior to elongate marginal setae; several
long setae arising from dorsal surface of telson at or between bases of dorsal
spines.

Color.—The specimens from Carrie Bow Cay when alive were blood red,
with the legs and antennal and antennular flagella having alternating red and
white bands. The Florida specimen when received was overall light
ochreous yellow, overlain with a marbled pattern of cadmium yellow on the
cephalothorax. The antennular and antennal flagella were irregularly banded
with dark reddish-purple. Tubercles, sharp spines, and the superior crests
on either hand of the first chelipeds were variously red, reddish-brown,
reddish-tan, or shining white. The elongate translucent setae on the meri of
pereopods 1 and 2, as well as on the telson, appeared clear golden under
the microscope. Eye corneas were black. These colors, with the exception
of the banding on the sensory flagella, have almost completely faded in
alcohol.

Remarks.—Coralaxius abelei is immediately distinguishable from all oth-
er western Atlantic axiids by a combination of morphological characters.
These include the short, sharp rostrum, the reduced, trispinose antennal
acicle, the corona of gastric spinules, the lack of submedian and lateral
carinae on the gastric region, by the armature of fanglike spines on the

VOLUME 93, NUMBER 4 1289

minor cheliped, the heavy spinous and tuberculate armature on both hands
of pereopod 1, by the reduced gill formula (Table 1) and by the shape of the
male second pleopod. From a perusal of the literature, it seems possible
that the minor cheliped of the first pereopod may eventually assume greater
taxonomic importance when genera of the group are subsequently redefined.
Certainly, the species described here is distinct from all other axiids in this
respect.

Sexuality.—Some axiids exhibit hermaphroditism, but the specimens of
C. abelei gave no indication of such. The males all possessed an appendix
masculina, the females ovipores on the coxae of the third pereopods. The
presence in females of a small pair of uniramous pleopods on the first ab-
dominal somite is also easily discernible. The latter feature, however, is not
always restricted to females in other axiid genera. For example, according
to de Man (1925), Axiopsis (Paraxiopsis) brocki males may have rudimen-
tary first pleopods, but are distinguished by the appendix masculina on
pleopod 2; females have uniramous first pleopods and ovipores on pereopod
3 coxae. Axiopsis (Paraxiopsis) bisquamosa males lack first pleopods, pos-
sess an appendix masculina, while the females bear uniramous first pleopods
and the usual ovipores. The female gonopores were not described for either
species. In Coralaxius, the male first pleopod is simple, but is certainly not
rudimentary.

The situation is more variable in Calocaris and Calastacus. Boesch &
Smalley (1972) noted the presence of uniramous pleopods on the first ab-
dominal somite of Calocaris (Calastacus) hirsutimanus, but stated that all
five of their specimens were males, based on the presence of an appendix
masculina on pleopod 2. Williams (1974) found uniramous pleopods only in
(apparently) functional females of Calocaris (Calastacus) jenneri; the males
of that species, and those of C. (Calastacus) oxypleura, lacked pleopods
on the first abdominal somite. St. Laurent (1972) described as an hermaph-
roditic male the new species CalJastacus laevis, which had a modified uni-
ramous first pleopod, no appendix masculina on pleopod 2, easily observed
gonopores on the coxae of both pereopods 3 and 5, and (based on remnants
of ovulary membranes on the pleopods) had apparently been ovigerous
shortly before its capture. Clearly, there is substantial variation in regard
to this feature within the genera as presently defined in the Axiidae.

Ecology.—The Belize specimens inhabit living coral, especially species
of Porites, living on the outer slope of the barrier reef off Carrie Bow Cay.
The Florida specimen inhabited a crevice in a piece of coralline limestone
rock collected from a bottom of coralline rubble and shell hash, with local-
ized dense algal cover. The depth range for the species extends from 11
—76 m.

The long, forward-directed setae along the lower margins of the meri of
the first and second chelipeds, and the basketlike effect produced by similar

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 93, NUMBER 4 1291

setae on the fingers of the major chelipeds, suggest that C. abelei uses such
setae either as a mechanical or sensory aid, perhaps by shovelling and sifting
through benthic detritus as it moves along the bottom. The large, fanglike
spines on the minor cheliped also suggest a carnivorous habit. The rocky,
limestone habitat seaward of a large, deep, coral bioherm is a biotope that
has been little sampled by conventional trawling and dredging gear from the
surface. Further collections by SCUBA or submersible may produce more
specimens and concomitant observations on their ecology.

Discussion.—Coralaxius shares with the subgenus Paraxiopsis several
characters which allow both taxa to be distinguished from other genera and
subgenera. This combination includes 1) the transverse suture on the urop-
odal exopods; 2) the reduced antennal acicle; 3) the pigmented, well-devel-
oped eyes; and 4) the depressed gastric region of the cephalothorax (fide de
Man 1925). The absence of a median dorsal keel on the posterior carapace
surface is also a character shared by Coralaxius and Paraxiopsis, and may
remain a valid character within both these taxa, although Boesch & Smalley
(1972), and Williams (1974) deprecate its taxonomic value in the closely
related Calocaris and Calastacus. There remains much uncertainty, how-
ever, regarding the limits of these genera and subgenera, with Calocaris
(Calocaris), Calocaris (Calastacus), Axiopsis (Axiopsis), and Axiopsis
(Paraxiopsis) being defined on characters which for the most part are ex-
clusive, but which also seem to be shared by 1 or 2 species within each
taxon (see Tables 1 and 2). Boesch & Smalley, for example, point out that
the similarities between some members of Calocaris (Calastacus) and Ax-
iopsis (Axiopsis) are greater than those between the former taxon.and the
nominate subgenus of Calocaris. Recently St. Laurent (1972), in describing
a new species of Calastacus, provided an emended diagnosis and re-estab-
lished the genus Calastacus, sensu Faxon (1893). However, in applying her
emended definitions and thereby separating the genera Calocaris and Ca-
lastacus, only 2 of the 9 species known at that time could be relegated to
Calastacus. The remaining 7 species of Calocaris (Calastdcus) s.l. plus the
2 subsequently described by Williams (1974), await the establishment of one
or more new genera, if St. Laurent’s revision is to be accepted.

The subgenus Axiopsis (except A. (A.) habereri Balss) is at present sep-
arated from Calocaris/Calastacus chiefly by the absence of a postcephalic-
groove carina on the cephalothorax. Unfortunately, Calocaris (Calocaris)
aberrans Bouvier also lacks this carina, thus differing from other members
of its genus. Axiopsis (Axiopsis) differs from A. (Paraxiopsis) in possessing
an elongate, well-developed antennal acicle, a character which also distin-
guishes species of C. (Calastacus) from C. (Calocaris). It is clear that these
genera and subgenera have been established using ambivalent morphological
characters, a fact long noted by students of the family, including de Man
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1292

VOLUME 93, NUMBER 4 1293

Although creation of subgenera within the family Axiidae serves to indi-
cate close relationships, the characters presently used (as noted by St. Lau-
rent, 1972) may eventually prove to be superficial, or of such variability
within populations of shrimp as to be practically without value. In her com-
parison of Calocaris and Calastacus, for example, St. Laurent (1972:353)
defined Calocaris macandreae as possessing symmetrical chelipeds. How-
ever, a specimen of this species from the Firth of Clyde (USNM 19614)
shows unequal chelae. Squires (1965:7) in describing the closely related
Calocaris templemani, noted that in this species, as well as in C. macan-
dreae *‘both chelae of the first pereopods in the same species are essentially
similar but may be slightly unequal in some.’’ Thus, a prerequisite to taxo-
nomic stability in the family is evaluation of several morphological features
presently shared at both the generic and subgeneric levels.

Etymology.—The species is named for Dr. Lawrence G. Abele, noted
carcinologist and esteemed colleague.

Acknowledgments

We thank Mr. Ron Larson, formerly of the Smithsonian Institution, the
Office of Coastal Zone Management, NOAA, Washington, D.C., and Dr.
Robert S. Jones, Science Director, Harbor Branch Foundation Laboratory,
Ft. Pierce, Florida, for providing collections containing the new species.
We also extend our thanks to Drs. Fenner A. Chace, Jr., and Raymond B.
Manning, National Museum of Natural History, and Austin B. Williams,
National Marine Fisheries Service, Systematics Laboratory, NMNH,
Washington, D.C., for providing literature concerning the new species, and
critically reading an earlier draft of the manuscript.

This is contribution number 65 of the Smithsonian Institution’s Investi-
gations of Marine Shallow-Water Ecosystems Project.

Literature Cited R

Bell, T. 1853. A history of the British stalk-eyed Crustacea.—London.

Boesch, D. F., and A. E. Smalley. 1972. A new axiid (Decapoda, Thalassinidea) from the
northern Gulf of Mexico and tropical Atlantic.—Bulletin of Marine Science 22:45-—52.

Borradaile, L. A. 1903. On the classification of the Thalassinidea.—Annals and Magazine of
Natural History (7) 12:534—-551.

de Man, J. G. 1888. Bericht tiber die im indischen Archipel von Dr. J. Brock gesammelten

Decapoden und Stomatopoden.—Archiv ftir Naturgeschichte 53:215-600.

. 1925. The Decapoda of the Siboga-Expedition. Part IV. The Axiidae collected by the

Siboga-Expedition.—Siboga-Expedition Monograph 39a5: 1-127.

Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America
to the Galapagos, to the west coast of Mexico, and in the Gulf of California, by the U.S.
Fish Commission Steamer ‘‘Albatross,’’ during 1891 VI. Preliminary descriptions of
new species of Crustacea.—Bulletin of the Museum of Comparative Zoology, Harvard
University 24:149-220.

1294 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Milne Edwards, A. 1873. Description de quelques crustacés nouveaux ou peu connus pro-
venant du Musée de M. C. Godeffroy.—Journal des Muséum Godeffroy 4:77-88.

de Saint Laurent, M. 1872. Un Thalassinide nouveau de Golfe de Gascogne, Calastacus laevis
sp. nov. Remarques sur le genre Calastacus Faxon (Crustacea Decapoda Axiidae).—
Bulletin de Muséum d’ Histoire Naturelle, Paris (3) Zoologie 29:347-356.

Squires, H. J. 1965. A new species of Calocaris (Crustacea: Decapoda, Thalassinidea) from
the northwest Atlantic.—Journal of the Fisheries Research Board of Canada 22:1-11.

Williams, A. B. 1974. Two new axiids (Crustacea: Decapoda: Thalassinidea: Calocaris) from
North Carolina and the Straits of Florida.—Proceedings of the Biological Society of
Washington 87:451-464.

(BK) Department of Invertebrate Zoology, Smithsonian Institution,
Washington, D.C. 20560; (RHG) Smithsonian Institution, Fort Pierce Bu-
reau, Ft. Pierce, Florida 33450.

PROC. BIOL. SOC. WASH.
93(4), 1980, pp. 1295-1313

RIFTIA PACHYPTILA, NEW GENUS, NEW SPECIES,
THE VESTIMENTIFERAN WORM FROM THE
GALAPAGOS RIFT GEOTHERMAL
VENTS (POGONOPHORA)!

Meredith L. Jones

Abstract.—Riftia pachyptila, new genus, new species, is described from
geothermal vents of the Galapagos Rift and from a geothermal area on the
East Pacific Rise at 21°N. These vestimentiferan worms represent a new
family, Riftiidae, in a new subphylum, Obturata, in the phylum Pogonoph-
ora. Differentiation from the two other vestimentiferan species is at the
familial level and concerns the orientation of tentacular lamellae on the
obturaculum, the number of openings of excretory ducts, and the relative
lengths of certain body regions. A most curious structure, the trophosome,
occupies much of the trunk region and is essentially a vascularized, bacteria-
filled sac; it is also the site of deposition of crystals of elemental sulfur.

In 1966 specimens of a worm, Lamellibrachia barhami Webb, were col-
lected off the coast of California at a depth of 1,125 m. These animals,
designated as a new class of Pogonophora by Webb (1969a), have subse-
quently been the object of continuing detailed research by him (Webb,
1969b, c, 1971, 1975, 1977). Inclusion in the Pogonophora was based “‘. . .
primarily on the shape, structure, and protein content of the tube,’’ as well
as **. . . distinctive pogonophoran characters: the very long trunk region,
the metasoma, and the absence of a gut’’ (Webb, 1969a, p. 41). Webb’s new
pogonophoran class, Afrenulata, was characterized merely by the lack of
a bridle (frenulum), while his new order, Vestimentifera, was erected on the
basis of: 1. Simple paired excretory ducts, without connections with the
tentacular coelom, which open exteriorly by a single median excretory pore
at the base of the ‘‘tentacular crown’’; 2. An undifferentiated trunk, pro-
vided with papillae; 3. A tentacular crown formed of numerous fused ten-
tacles surrounding a paired, centrally placed ‘‘lophophoral organ,’’ which
is in turn surrounded by a number of tentacular sheaths; 4. A so-called
vestimental region between the tentacular crown and the trunk, provided
with lateral folds which meet to form a vestimental chamber between the
vestimentum and the tube; and 5. Genital apertures which open into the

1 Contribution No. 8 of the Galapagos Rift Biology Expedition, supported by the National
Science Foundation.

1296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

vestimental chamber, with male apertures provided with ciliated grooves.
A second collection of L. barhami was reported by Webb (1977) from off
the coast of Oregon.

In 1970 a second species of vestimentiferan was collected from a depth
of 500 m on the continental slope off Guyana. Lamellibrachia luymesi van
der Land and Ngrrevang was described from a single male specimen, and
has been reported upon in two papers (van der Land and Ngrrevang, 1975,
1977). Specific differences between L. barhami and L. luymesi cited by van
der Land and N@grrevang (1975) were: 1. The relative length of the vesti-
mental region; 2. The number of tentacular sheaths; 3. The relative prox-
imity of male genital apertures; 4. The structure of the tube; and 5. The
presence of various grooves and lines of pigment on certain parts of the
trunk and vestimentum.

The descriptions of the two species of Lamellibrachia point up a con-
tinuing controversy concerning the dorso-ventral orientation of the Pogo-
nophora. On the one hand Webb considers the nerve cord to be dorsal,
while van der Land and N@rrevang assume that it is ventral; the former
view implies a relationship to the hemichordates et al., and the latter, to
the annelids. Three sets of neutral terms have been suggested: ‘‘neural’’
and ‘“‘tentacular,’’ “‘neural’’ and ‘“‘antineural,’’ ‘“‘adneural’’ and ‘‘antineur-
al,’ (Southward, 1975a, b; Anon., 1975, p. 143, respectively). In this
paper I decline to use the neutral terms of orientation, for there is no un-
equivocal distribution of tentacles in the vestimentiferans and, while ‘‘neur-
al,’ ‘‘adneural,’’ and ‘‘antineural’’ are unequivocal, I find them awkward
and I suspect that they may be susceptible to confusion. As will be noted
below, it is my opinion that the vestimentiferans, as well as the Pogono-
phora, stand in close relationship to the Annelida and I follow van der Land
and Ngrrevang in considering the nerve cord to be ventral and the major
blood vessel, provided with a thick muscular layer, to be dorsal.

In 1977 and 1979 during expeditions to the Galapagos Rift and East Pacific
Rise areas the submersible Alvin obtained a number of vestimentiferans
which are the subject of this paper. At the Galapagos sites, water coming
from geothermal vents may be as warm as 23°C and may contain hydrogen
sulfide to the extent of 160 umole per liter, and, at temperatures above 10°C,
the vent waters are anoxic; ambient temperature of bottom water near the
vents is about 2°C, it is oxygenated, and there is no hydrogen sulfide present
(Corliss et al., 1979). A total of 63 worms, collected at three geothermal
vents in the Galapagos Rift area and one vent on the East Pacific Rise, have
been deposited in the collections of the Division of Worms, National Mu-
seum of Natural History (USNM), Smithsonian Institution, and these form
the basis for the following description. Paraffin sections were cut at 5 wm
and stained with routine staining procedures indicated below. The identity
of sulfur crystals was confirmed by an ARL-SEMQ electron microprobe

VOLUME 93, NUMBER 4 1297

and by X-ray diffraction utilizing CuK X-radiation (nickel-filtered) with a
Gandolfi 114.6 mm camera and a polycrystalline sample.

Riftiidae, new family

Type-genus.—Riftia, new genus.

Diagnosis.—Vestimentiferans with tentacular lamellae at right angles to
obturaculum; without tentacular sheaths; obturaculum at feast 11% of total
length; with separated, paired, external openings of excretory ducts; with
essentially cylindrical tube.

Riftia, new genus

Type-species.—Riftia pachyptila, new species.

Gender.—Feminine.

Diagnosis.—Characters of the family (above).

Etymology.—From Danish and Norwegian, rift (rent, fissure) + -ia, in
reference to the Galapagos Rift.

Riftia pachyptila, new species
Figs. 1-4

Worm(s).—Ballard, 1977:41, 43 (with fig.)—Anon., 1979:11 (with fig.).—
The Editor, 1979:680, 681 (fig.).—Ballard and Grassle, 1979:689 (with
fig.).—Schlee, 1979:41.

Tube worm(s).—Corliss and Ballard, 1977:441 (with fig.), 449.—Corliss et
al., 1979:1079.—Schlee, 1979:42 (with fig.).—The Editor, 1979:688.—Bal-
lard and Grassle, 1979:699 (with fig.), 705 (fig.).—RISE Project Group,
1980:1421, 1432.

Pogonophora(ns).—Corliss and Ballard, 1977:441 (with fig.).—Corliss et al.,
1979:1075, 1079-1080, figs. 4, S.

Vestimentifera(ns).—Corliss et al., 1979:1079.—Ballard and Grassle,
1979:689 (with fig.).—-Galapagos Biology Expedition Participants, 1979:6
(with fig.), 7, 8.—Jannasch and Wirsen, 1979:592, 594-595.—RISE Proj-
ect Group, 1980:1424, fig. 5—Hekinian et al., 1980:1435.—Terwilliger
et al., 1980:531 et seq.

Diagnosis.—Characters of the family (above).

Etymology.—From Greek, pachys (thick) + Greek, ptilon (feather),
in reference to the aspect of the anterior plume of the worm (combina-
tion treated as an adjective).

Type-locality.—‘‘Rose Garden’’ geothermal vent (RG), Galapagos Rift
(00°48'15"N; 86°13'29"W), 2,450 m depth.

Other localities of paratypes.—Galapagos Rift—‘‘Dandelions’’ geother-
mal vent (D) (00°47'42’N; 86°08’00"W), 2,496 m depth; ‘‘Garden of Eden’”’

1298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

geothermal vent (GE) (00°47'42"N; 86°07'44"W), 2,485 m depth. In addition,
worms have been noted, but not collected, at ‘‘Mussel Bed’’ geothermal
vent (00°48'06"”N; 86°07'00’”W), 2,490 m depth. East Pacific Rise geothermal
vent (EPR) (20°51’00’N; 109°04'54”W), 2,595 m depth.

Material examined.—Holotype (USNM 59951), RG, Alvin Dive (AD) 889,
14 Feb 1979, adult female. Paratypes: 1 adult, D, AD 723, 27 Feb 1977; 1
sectioned adult (USNM 59953), 1 sectioned juvenile (USNM 59954), 8 adults
and juveniles (USNM 59955), GE, AD 733, 16 Mar 1977; 3 adults (USNM
59956), GE, AD 884, 25 Jan 1979; 1 adult (USNM 59957), 1 sectioned young
adult (USNM 59958), 1 young adult (USNM 59959), 3 sectioned juveniles
(USNM 59960-62), 7 juveniles (USNM 59963), RG, AD 889, 14 Feb 1979;
1 adult (USNM 59964), EPR, AD 915, 22 Apr 1979; 2 adults (USNM 59965),
RG, AD 983, 30 Nov 1979; 2 adults (USNM 59966-67, RG, AD 984, 1 Dec
1979; 1 adult (USNM 59968), RG, AD 988, 5 Dec 1979; 1 adult, 1 young
adult (USNM 59969), 1 adult, 1 young adult, 1 juvenile (USNM 59970), 1
adult, 2 young adults (USNM 59971), 1 adult, 1 juvenile (USNM 59972), 7
adults (USNM 59973-79), RG, AD 990, 7 Dec 1979; 1 adult, 2 young adults
(USNM 59980), 2 adults (USNM 59981-82), GE, AD 993, 10 Dec 1979.
Non-Types: 2 empty tubes (USNM 59996-97), GE, AD 983, 30 Nov 1979;
1 empty tube (USNM 59998), RG, AD 990, 7 Dec 1979; 1 empty tube with
adult fragment and 7 juveniles (USNM 59999), GE, AD 993, 10 Dec 1979.

Description.—Dimensions, relative lengths and certain meristic charac-
ters of the holotype and a representative series of paratypes are listed in
Table 1.

With four body regions: (1) Anterior tentacular plume on obturaculum;
(2) Winged vestimentum; (3) Trunk; (4) Segmented posterior opisthosome
(terminology: Webb, 1969a; van der Land and N@grrevang, 1975, 1977) (Fig.
1B).

Central axial obturaculum of plumed region paired and fused, splayed
apically (Fig. 1C); with non-ciliated dorsal groove and ventral ridge along
length (Fig. 2B); with tentacular lamellae perpendicular to axis, extending
nearly to anterior tip, giving impression of thick feather (Fig. 1D); paired

=>

Fig. 1. Riftia pachyptila (A: USNM 59999; B: USNM 59951; C, E: USNM 59957; D:
USNM 59956). A, Tube with four young individuals attached; apical aperture, upper left; closed
basal end, right center; tube cut to obtain animal; folds in tube, during initial storage/preser-
vation, give misleading appearance to relatively uniform tube diameter; B, Overall view of
holotype; C, Obturacular plume, lateral view; splayed distal end, above; D, Same, ventral
view; E, Vestimentum, ventral view; BN: bifurcated ventral nerve; JU: juvenile; LT: large
tentacle; OB: obturaculum; OP: opisthosome; TR: trunk; VC: ventral ciliated field; VE: ves-
timentum; VR: ventral ridge; VW: vestimental wing; YA: young adult. Scale bars (lower
right)—A, B: 10 cm; C—E: 5 cm.

VOLUME 93, NUMBER 4 1299

1300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Riftia pachyptila: Comparison of specimens of various lengths showing relative
lengths of obturaculum (OB), vestimentum (VE), trunk (TR), and opisthosome (OP), with
numbers of paired tentacular lamellae and of opisthosomal segments.

Percentage body length

Total length No. tent. No. op.

Cat. number Sex OB VER TR OP (mm) lamellae segs.
USNM 59951 g 11 7 80 2 1,502.00 334 96
USNM 59957 fe) 16 13 69 3 1,056.00 302 103
USNM 59964 2 34 13 49 3 513.00 257 86
USNM 59952 2 17 17 59 7 320.00 170 - 85
USNM 59955 2 192 18 53 7 273.00 193 74
USNM 59955 2 26 jap) 45 if 215.00 166 66
USNM 59959 2 26 12 56 6 199.00 116 72
USNM 59953 3 7022 16 >» 6 171.00 113 95
USNM 59959 2 27 11 7 5 169.00 111 78
USNM 59958 2 24 14 56 6 111.00 113 73
USNM 59959 3 29 13 49 9 77.00 95 82
USNM 59959 3 21/ 13 53 7 55.00 63 71
USNM 59954 B 34 24 30 13 15.50 39 36
USNM 59963 i 30 26 33 21 6.10 12 19
USNM 59963 & 33 22 28 17 6.00 23 25
USNM 59963 uy 28 19 35 19 4.30 8 18
USNM 59960 ? 28 18 34 20 1.60 1? 11
USNM 59963 v 35 17 29 19 0.75 — 2+

halves of apical split end forming operculum upon withdrawal into tube. In
life vestimentum with dorsolateral flaps/wings overlapping one another with-
in tube, with free anterior margin of flaps meeting and slightly overlapping
ventrally (Fig. 1E); free posterior margin of flaps continuous and entire,
ventrally (Fig. 1B, E); paired genital apertures at posterior third of dorsal

=>

Fig. 2. Riftia pachyptila (A: USNM 59958; B, C, E-G: USNM 59953; D: USNM 59951).
A, Vestimental wing, transverse section; Masson’s Trichrome stain; outer surface when over-
lapped, above; inner surface, below; B, Tentacular lamella, complete, from mid-region of
obturaculum; obturaculum, upper right; Mallory’s Triple stain (ML); C, Obturaculum with
associated tentacular lamellae, transverse section; Chlorazol Black; D, Opisthosome, ventral
view; E, Opisthosomal septum, complete, from one side (ML); F, Trunk body wall, transverse
section, showing plaqued papilla; Toluidine Blue; G, Trunk, transverse section (ML). CC:
circular muscle/connective tissue; CO: coelom; CT: compartmented tissue; CV: connecting
vessel; DB: anterior branch of dorsal vessel; DG: dorsal groove; DM: dorsal mesentery; DV:
dorsal vessel; FM: “‘feather’’ longitudinal muscle; GP: gland papilla; IE: inner epithelium; LG:
‘‘long’’ gland; LT: large tentacle; MM: medial mesentery; MO: muscle strands of obturacular
matrix; NL: ‘“‘normal’’ longitudinal muscle; OF: outer epithelium; OP: opisthosome; OV: ob-
turacular vessel; PG: pyriform gland; PM: parasagittal muscle; PP: plaqued papilla; PS: partial
septum; SG: “‘short’’ gland; TP: trophosome; TR: trunk; TS: testes; VB: anterior branch of
ventral vessel; VM: ventral mesentery; VN: ventral nerve; VR: ventral ridge; VV: ventral
vessel. Scale bars (lower right) A, F: 100 um; B, C, E, G: 5 mm; D: 10 mm.

VOLUME 93, NUMBER 4 1301

i ;

é
om KS
agg we

= i. ORE sem :
Mag wile: PRG PUM
LEI

1302 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

surface in both sexes; male aperture associated with ciliated ridges, con-
verging but not meeting anteriorly; female apertures lacking ciliated ridges;
ventral ciliated field somewhat pear-shaped, delimited by paired ventral
nerves and neurular tubes (Fig. 1E); papillar openings of internal pyriform
glands on surface, apart from ventral ciliated field, and on outer surface of
overlapped flaps (Fig. 2A); dorsal cavity formed by overlapping flaps nearly
lacking such papillar openings, with cuticle, lacking specialized epithelium.
Trunk with similar papillae, with united single ventral nerve and neurular
tube extending throughout length (Figs. 2G, 3A). Opisthosome with variable
number of segments, ending in rounded posterior tip (Fig. 2D); anterior
segments (about '4 length), each completely encircled with paired single
rows of setae, becoming incomplete posteriorly.

No setae on vestimentum or trunk; setae with long shafts with two toothed
areas extending free of body surface; many teeth in posterior area, pointing
anteriorly; fewer in anterior area, pointing posteriorly; identical with girdle
setae of Pogonophora (Fig. 4B); no peg-like setae.

Papillae most numerous on outer surface of vestimental wings and on
posterior trunk, progressively fewer papillae anteriorly along trunk, very
few papillae on dorsal surface of vestimentum and inner surface of wings;
papillae with plaques about three to five times more numerous than those
without plaques, latter probably exit for secreted tube material.

Tube white, flexible, extremely sturdy, essentially cylindrical (Fig. 1A);
basally blind-ending, approximate shape of opisthosome; apical opening
thinning to apertural margin; main part of tube commonly 2-3 mm thick;
tube material laid down incrementally forming rudimentary ‘‘collars’’ or

=

Fig. 3. Riftia pachyptila (A-—E, G-H: USNM 59953; F: USNM 59958). A, Trunk, ventral
body wall, transverse section; Toluidine Blue; B, Compartmented tissue adjacent to obturacular
matrix and basal origins of five consecutive tentacular lamellae, transverse section; Mallory’s
Triple stain (ML); C, Tentacles, distal free ends; Grenacher’s Borax Carmine/Fast Green; D,
Tentacles, fused as a lamella, cross-section (ML); E, Tentacle, free, showing connection of
transverse intraepithelial loop with longitudinal tentacular blood vessel (arrow), trending to-
ward adjacent pinnule, cross-section (ML); F, Dorsal vessel in posterior vestimentum, im-
mature specimen, transverse section; Paraldehyde Fuchsin; G, Tentacles, fused as a lamella,
longitudinal section (ML); H, Obturacular vessel in obturacular matrix, transverse section
(ML). AE: anterior extension of trunk coelom; CG: coagulated coelomic fluid, CL: cilia; CO:
coelom; CT: compartmented tissue; CU: cuticle; DB: anterior branch of dorsal vessel; DV:
dorsal vessel; FM: ‘‘feather’’ longitudinal muscle; LT: large tentacle; MO: muscle strands of
ebturacular matrix; NT: neurular tube; OV: obturacular vessel; PM: parasagittal muscle; PN:
pinnule; PV: perivascular cavity; RM: ring muscle; TC: tentacular coelom; TE: tentacular
epithelium; TL: vessel from tentacular lamella ‘‘2’’; TP: trophosome; TS: testes; TV: tentacular
vessel; VB: anterior branch of ventral vessel; VL: intraepidermal transverse vascular loop;
VM: ventral mesentery; VN: ventral nerve; VV: ventral vessel; X: site of formation of newest
‘‘compartment’’; 1-5: serially more proximal tentacular lamellae. Scale bars (lower right)—A,
B, F: 0.5 mm; C: 250 um; D, H: 100 wm; E: 25 wm; G: 50 um.

VOLUME 93, NUMBER 4 1303

1304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

flanges; tube secretion adhering to solid materials (rocks, limpet shells) and
to other tubes (Fig. 1A); in longer tubes, basal partitions separating worm’s
living space from unoccupied, basal, older portions (tube, presumed that of
holotype, 2,654 mm long with basal end ragged, as if torn; at 1,245 mm from
apical end of tube, secreted partition restricting occupant to distal portion
of tube; between this partition and basal end, 11 partitions previously se-
creted).

Internally, obturaculum with gelatin-like, semi-solid matrix invested with
thick cuticle (Fig. 2B, C); sagittal fusion of obturacular halves with double
layer of cuticle; internal from cuticle as many as 500 bundles of muscles,
appearing to be longitudinal, actually as many as 250 muscle rings, in para-
sagittal planes (Figs. 2C, 3B, 4C, D:PM); matrix penetrated by paired ob-
turacular blood vessels, blind-ending near split anterior region of obturac-
ulum, originating from two anterior branches of dorsal vessel in vestimental
region (Figs. 2B, C, 3H, 4C—F); obturacular vessels with thin inner lining
of muscle cells, then thick layer of connective tissue, surrounded externally
by closely applied ring muscles (Fig. 3H); vessels supported by two mes-
entery-like structures in perivascular cavity, lacking connection to any other
body cavity. Vestimental region quite solid with mass of muscle strands and
connective tissue elements intermixed (Fig. 3F); with close-packed pyriform
glands in discrete layer, opening to external surface of overlapped flaps
(dorsolateral) and vestimentum, proper (lateral and ventral) (Fig. 2A); brain
anteroventral, internal to ventral overlapping of flaps; ventral extension of
brain tissue extending to cuticular covering of body, containing paired neu-
rular tubes arising de novo in brain tissue (Fig. 4F); ventral ciliated field
formed of transversely fused cilia, emerging from epithelial cells not con-
tinuously in contiguous contact, forming considerable lacunae or sinuses;
dorsal surface of posterior %4 of vestimentum with anterior extension of
trunk and its coelomic cavities overlapping, with genital ducts moving for-

—>

Fig. 4. Riftia pachyptila (A-F: USNM 59953). A, Tentacles, fused as a lamella, showing
contracted ring muscle (center) and two relaxed ring muscles (above and below), cross-section;
Mallory’s Triple stain (ML); B, Seta in body wall of opisthosome, frontal section; Mayer’s
Hematoxylin-Eosin; C, Compartmented tissue near base of obturaculum, transverse section
(ML); D, Same, more posterior, showing beginning of transformation of compartmented tissue;
bulge to left is initiation of connection of tissue to other side; transverse section (ML); E,
Same, yet more posterior, with more extensive transformation of compartmented tissue to
brain tissue; connection of both halves complete; transverse section (ML); F, Brain at level of
descent of nerve tissue to form ventral nerve, transverse section (ML). AF: anterior ‘‘flap’’ of
vestimental wing; BR: brain; CT: compartmented tissue; CU: cuticle; DB: anterior branch of
dorsal vessel; NT: neurular tube; OV: obturacuiar vessel; PM: parasagittal muscle; PV: peri-
vascular cavity; RM: ring muscle; TC: tentacular coelom; TE: tentacular epithelium. Scale
bars (lower right)—A, B: 25 4m; C-F: 0.5 mm.

VOLUME 93, NUMBER 4 1305

1306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ward then dorsally to open through genital apertures; excretory organ (pos-
sibly paired) posterior and ventral to brain. Trunk with internal surface of
body wall with longitudinal ‘‘feather’’ muscles (van der Land and Ngrre-
vang, 1975, 1977) internal to thin layer of more conventional longitudinal
muscles, in turn internal to thin layer of mixed circular muscle and connec-
tive tissue, separated by basement membrane from layer of epithelial cells
covered by external cuticular layer (Figs. 2F, G, 3A); paired coelomic cav-
ities extending throughout entire length of trunk, separated by medial dor-
soventral mesenteries investing dorsal and ventral blood vessels, attached
to but not investing centrally located gonad and ‘‘trophosome’”’ (van der
Land and Nogrrevang, 1975, 1977) (Figs. 2G, 3A); internal broad bases of
pyriform glands project centripetally between rows of feather muscles (Fig.
2G:PG). Opisthosomal segments separated internally by two-layered septa;
with only “‘normal’’ longitudinal muscles, ‘‘feather’’ longitudinal muscles
lacking; segmental cavities separated medially by dorsoventral mesentery;
anterior faces of septa with two kinds of glands, short broad and long slen-
der, histologically different from pyriform glands of vestimentum and trunk;
long slender glands invested by blood vessels which interconnect with other
similar vessels and move centripetally to center of septum; segments also
with low, thin, multilayered circular bands of muscles forming “‘partial sep-
ta’’ (Fig. 2E); ventral nerve here, well-formed, rounded, lacking neurular
tube.

Tentacles covered with cuticle of varying thickness throughout length;
fused basally for from 50 to 75% of their length to form tentacular lamellae,
with tips of tentacles free (Fig. 2B); with two rows of multiciliated epithelial
cells, one on posteroventral face, other on anterodorsal face, extending
about 90% of basal length of each tentacle (Fig. 3D); with two rows of
multicellular pinnules on posterior surface of each tentacle, along distal 45%
of length, not extending to tips (Fig. 3C); central tentacular coelomic (?)
cavity with two blood vessels, enclosed by basement membrane of single
layer of epithelial cells (Fig. 3D, G); blood vessels connected along tentac-
ular length by transverse intraepithelial vascular loops communicating with
central cavities of pinnules (Fig. 3E); no nervous tissue seen; as many as
340 tentacles per lamella and 335 lamellae on each side of obturaculum (in
holotype), about 2.28 x 10° tentacles in plume; along length of each tentac-
ular blood vessel *‘O’’- or ‘‘U’’-shaped ring muscles appear to act as series
of sphincters (Figs. 3G, 4A); in basal region of lamellae, cuticle of fused
tentacular bases thickened, successive lamellae fused to one another for
short distance, serially (Fig. 3B:1—5); internally, cuticular matrix disappears
and blood vessels of all tentacles of a given lamella join single transverse
blood vessels which join anterior axial extensions of dorsal and ventral
vessels of main body (Fig. 3B: DB, VB); immediate fate of coelomic ten-
tacular space and of presumed tentacular nerves obscured and unknown;
basal contents of all tentacles of given lamella unite, and their residuum

VOLUME 93, NUMBER 4 1307

moving centrally, forming thin layer directed posteriorly toward base of
obturaculum, in company with residua of more apical tentacular lamellae
(Fig. 3B: X, CT); up to three tentacles of some lamellae lacking both rows
of cilia and rows of pinnules, with 50% larger diameter and up to twice
length of ‘‘normal’’ tentacles (Figs. 1C, 2B, 3C, D).

Dorsal vessel in trunk thin-walled with thin connective tissue layer cov-
ered with thin layer of circular (?) muscles (Fig. 2G); connecting branches
to trophosome, mesenterial and other vessels traversing medial mesentery;
heart body (corpus cardiacum) in dorsal vessel in midtrunk region; ante-
riorly, at level of anterior extension of trunk into vestimental region, dorsal
vessel with thin muscle lining, with succeeding thick layer of connective
tissue, surrounded by layer of striated, circular, ring muscles, supported in
perivascular cavity by thickened dorsal and ventral mesentery-like struc-
tures, whole complex supported by second dorsoventral mesentery in an-
terior extension of trunk coelom (Fig. 3F); more anteriorly, in vestimental
region, connective tissue layer of dorsal vessel lost, wall of apparently un-
oriented muscle fibers, free in perivascular cavity (mesenteries lost), trunk
coelom not present; perivascular cavity surrounding dorsal vessel not con-
nected to trunk coelom or any other coelomic cavity; just posterior to level
of brain, dorsal vessel branching, each branch moving anteriorly to run
length of obturaculum, ventromedially, receiving single vessels from trans-
verse tentacular lameilar vessels (Figs. 3B, 4C, E: DB); at level of branch-
ing, dorsal vessel giving rise to paired obturacular vessels, ending blindly
near anterior margin of splayed obturaculum, undergoing up to 25 dorso-
ventral loops through obturacular matrix (Figs. 2B, C, 3H, 4C-—F). Anterior
branches of ventral vessel run length of obturaculum, ventrolaterally, in
company with branches of dorsal vessel, receiving branches from each
transverse tentacular lamellar vessel (Fig. 3B); at base of obturaculum, pos-
terior to level of brain, branches unite, possibly forming valve of some kind,
proceeding posteriorly as single vessel embedded in loose connective tissue,
with no muscular layer. Fate of anterior branches of dorsal and ventral
vessels unknown—perhaps blind-ending or connecting near anterior margin
of obturaculum.

Blood directly from dorsal vessel of living animals, two hours after col-
lection, in laboratory of surface ship, of dark red color and viscosity of port
wine; coelomic fluid from trunk and opisthosome of similar color and vis-
cosity.” |

Residua of tentacular lamellae moving to bases of obturacula, augmented

2 The following papers concerned with the blood of Riftia pachyptila have been submitted
to Science, as of this writing: A. Arp and J. Childress, observations on the functional aspects
of oxygenation characteristics of extracellular hemoglobin; and J. Wittenberg, R. Morris, Q.
Gibson, and M. Jones, data on the kinetic constants for combination and dissociation rates of
the hemoglobin.

1308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by residua of successively more basal tentacular lamellae, forming paired
ventromedial masses of narrow compartments (Figs. 3B, 4C: CT); in ante-
rior part of vestimentum, rigid linearity of compartments breaking down and
lamellar residua becoming ‘‘transformed’’ into brain tissue (Fig. 4D, E); just
anterior to branching of dorsal and ventral vessels, “‘pillar’’ of nervous
tissue with paired neurular tubes, each in own cavity, moving ventrally to
just internal to cuticle, becoming flattened to form band of nervous tissue
with separated neurular tubes (Fig. 4F); posteriorly, band of nervous tissue
branching laterally at about 10% length of vestimentum, and area between
branches of ventral nerve, ventral ciliated field (Fig. 1D, E); at about 60-
70% of length of vestimentum, divergence of nerve bands (and width of
ventral ciliated field) at maximum, followed by rapid convergence of nerve
tissue and neurular tubes, meeting at about 85% vestimental length, at pos-
terior margin of ventral ciliated field; ventral nerve band continuing as such,
with single neurular tube, to the posterior end of trunk (Fig. 3A); in opis-
thosome ventral nerve changing from band-like to compact, rounded cord,
lacking neurular tube, proceeding to basal end of opisthosome (Fig. 2D).

Presumed excretory organ just posterior to brain at level of branching of
dorsal and ventral vessels; main mass of ciliated tubules with no obvious
separation into halves at mid-line; tubules with no apparent orientation, no
apparent internal openings, uniting with other tubules and opening into non-
ciliated cavities joining with others forming paired, non-ciliated canals, mov-
ing anterolaterally, then dorsally, at level of anterior margin of vestimentum,
then opening into basal area of dorsal groove of obturaculum by paired
excretory pores.

Sexes separate; single external morphological difference—ciliated grooves
anterior from paired male genital apertures, lacking in females; paired gen-
ital apertures of both sexes on dorsal surface at approximately same loca-
tion, about 70% vestimental length; eggs just inside female aperture at ger-
minal vesicle stage, about 78 ~m diameter, spherical except for slight
distortion due to packing; sperm just inside genital aperture with elongate
bodies, corkscrew-shaped, about 9 um long, 0.6 wm diameter, tails about
9 wm long; no spermatophores seen; sperm ducts with ridge of epithelium
along length; no eggs or developing embryos or larvae found in tubes of
either females or males; none found in dorsal cavity formed by overlapping
vestimental wings.

Trophosome of many lobules with central lumina, well-vascularized, with
vascular plexi and lacunae over surfaces;? blood vessels forming lumina

3 The following papers concerned with the trophosome of Riftia pachyptila have been sub-
mitted to Science, as of this writing: H. Fehlbeck, an account of enzymes possibly attributable
to the trophosomal bacteria; G. Rau, data on '°C to !*C ratios pertaining to these bacteria; and
C. Cavanaugh, S. Gardiner, M. Jones, H. Jannasch, and J. Waterbury, a description of and
observations on these organisms.

VOLUME 93, NUMBER 4 1309

from which capillaries ramify throughout lobular mass; ‘‘tissue’’ of tropho-
some actually close-packed bacteria; surface of trophosomal lobules with
numerous fine pigment granules/spots; trophosome in intimate association
with gonads and gonoducts (Fig. 2G); trophosome of most specimens with
crystals of free elemental sulfur in ‘‘tissue’’; trophosome present in juvenile
as small as 1.44 mm total length (USNM 59961).

In smallest juvenile (USNM 59963), total length 0.75 mm, no mouth, no
gut, no anus; setae in two rows on opisthosome identical to setae of adults
but of smaller size.

Color in life. —Obturacular region: tentacles, red; obturaculum, whitish,
occasionally with greenish tinge; dorsal groove with white margins, interior,
pink; ventral ridge, white; splayed apical surfaces pink with dark red ob-
turacular vessels near surface. Vestimental region: anterior dorsal margin
with green suffusion; inner and outer surfaces of overlapped flaps, red;
ciliated tracts of male, brown/green; ventrally united anterior nerve, green;
separated nerves at level of ciliated field, white with lateral red lines; united
posterior nerve, green; ventral ciliated field, dark red; papillae, beige against
pinkish beige background. Trunk: beige to pink beige, with green line on
anterior dorsal surface at junction with vestimentum. Opisthosome: pink/
dark red.*

Faunal associates.—Other animals present with Riftia in the immediate
vicinity of the geothermal vents include Calyptogena magnifica Boss and
Turner (vesicomyid clam), Bythograea thermydron Williams (crab), a my-
tilid mussel, several shrimp, several limpet-like gastropods, a number of
calanoid, cyclopoid, and harpacticoid copepods, at least six families of poly-
chaetes, and a brotulid fish. Of these, the crab and the several shrimp appear
to be predators on Riftia, based on direct observations (fide J. Childress).

Feeding.—The obturacular plume of Riftia, with its elaborate vascular-
ization, would seem to be the most likely site for the uptake of at least small
organic molecules. The extreme number of tentacles, each with up to 100
pairs of transverse intraepidermal vascular loops and the large cumulative
surface of the vascularized pinnules (up to 100 pairs on each tentacle),
comprise an admirable organ for molecular uptake. There is a possibility
that similar uptake takes place across the body wall of the trunk, but nu-
trients would have to be carried in the water in the tube; due to the thickness
of the tube wall, ruling out a diffusion across this wall, and the fact that
such water could be renewed only during the occasional withdrawal of Riftia
into its tube, this alternative seems much less likely. The trophosomal bac-
teria may also play a role in the nutrition of Riftia. The similarity of ratios
of !8C to !2C in the trophosome (bacteria) and in the vestimental musculature

4 Color observations were made on specimens returned to the surface vessel laboratory,
about two hours after collection; the worms probably were moribund at the time.

1310 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

found by Rau (personal communication) suggests that Riftia may utilize the
bacteria and/or their metabolites as a source of organic carbon.

Comparisons .—Riftia pachyptila is closely related to the recognized ves-
timentiferan species Lamellibrachia barhami Webb (1969a) and L. luymesi
van der Land and N@grrevang (1975). The presence of an obturaculum with
its associated tentacles basally fused to form tentacular lamellae (Figs. 1B,
2B), with the presumed excretory ducts on the anterodorsal margin of the
vestimentum near the base of the obturaculum, the presence of the vesti-
mentum with its dorsolateral flaps (Fig. 1B), with a ventral ciliated field set
off by a bifurcation of the ventral nerve (Fig. 1E), with both female and
male genital apertures opening on the posterodorsal surface of the vesti-
mentum from an anterior extension of the trunk, with a trunk containing an
extensive trophosome and primary longitudinal muscle of the ‘‘feather”’
type (Fig. 2F, G), and with neurular tubes in association with the ventral
nerve (Figs. 3A, 4F), all leave no doubt as to this relationship. Unfortu-
nately, there was no opportunity to compare the opisthosome of Riftia with
those of either species of Lamellibrachia, for in the one case (van der Land
and N@grrevang, 1975) the opisthosome was lost at the time of collection; in
the other case (Webb, 1969a) a detailed description has yet to be recorded
and this account leaves me with some doubt as to whether the opisthosome
has actually been observed. I have had the opportunity to examine speci-
mens of L. barhami deposited here by E. Barham (USNM 55162), com-
prising part of his original collection. In all specimens examined, the pos-
terior ends, usually 1 mm or less in diameter, were either poorly preserved
or had dried and were flattened against the inner surface of the tube; I was,
thus, unable to identify an opisthosome in these specimens. I have also
examined specimens of L. barhami collected by trawl by A. Carey, Oregon
State University, 30 Oct 1967, at 1,400—1,600 m depth, at about 44°35.7’N,
125°11.0’W (in part, USNM 61103). The posterior ends of these specimens,
also, appear to have been poorly preserved or to have dried so that confir-
mation of an opisthosome was not possible.

Of the differences so far observed in comparing R. pachyptila with the
descriptions of both species of Lamellibrachia and with specimens of L.
barhami, the following characters of Riftia are deemed to be of most im-
portance. Obturacular region: Tentacular lamellae are free from one another
for most of their length (Fig. 1D); tentacular lamellae arise perpendicularly
from the obturaculum (Fig. 1D); there are no “‘tentacular sheaths’? (Webb,
1969a; van der Land and N@grrevang, 1975, 1977); obturaculum comprises
no less than 11% of the total length (Table 1). Vestimental region: Bears
two separate and distinct external openings of the ducts from the presumed
excretory organ(s); vestimentum may comprise from 7 to 22% of the total
length of young or older adults (of greater than 50 mm total length) (Table
1). Tube: With one opening, nearly cylindrical, with very little tapering

VOLUME 93, NUMBER 4 1311

Table 2.—Comparison of diagnostic characters of Lamellibrachiidae Webb and Riftiidae new
family.

Lamellibrachiidae Riftiidae
Orientation of tentacular Axial, parallel to At right angles to
lamellae obturaculum obturaculum
Tentacular sheaths Present Absent
Relative length of 4% total length 10% total length
obturacular region
External opening(s) of One Two
excretory ducts
Shape of tube Tapered Essentially cylindrical

toward the closed basal end (Fig. 1A). Taken as a whole these differences
suggest that Riftia should be separated from Lamellibrachia at the familial
level (Table 2).

Although the arguments of van der Land and N@grrevang (1975, 1977) are
persuasive and their points are well-taken, at the present time, I must con-
sider the basic regionation of Lamellibrachia and Riftia as indicating a close
relationship with the frenulate Pogonophora. The presence of demonstrable
coelomic cavities in the first and second regions of the latter and the absence
of such in Lamellibrachia and Riftia might well be a consequence of the
development of the solid obturaculum and the heavy muscularization of the
vestimentum. Other differences, e.g., the vestimental wings, the presence
of the trophosome, neurular tubes, and multicellular pinnules, the absence
of spermatophores, the medial dorsoventral mesentery of the opisthosome,
the tube morphology, in my present view, might be accommodated by con-
sidering them to discriminate between subphyla. While this decision might
appear to be somewhat frivolous or gratuitous, I feel that it best summarizes
my view of the relationship of Riftia and Lamellibrachia, with respect to
the other pogonophorans, i.e., it admits the unifying character of the basic
body plan of four regions, but emphasizes the exceptional and unique dif-
ferentiating characters listed above. Further studies of Riftia and Lamelli-
brachia may temper my judgement in this respect, but for now, I here
propose two subphyla for the phylum Pogonophora: Obturata new subphy-
lum (from Latin, obturo (close up, stop up), in reference to the operculum-like
function of the obturaculum of Lamellibrachia and Riftia); and Perviata
new subphylum (from Latin, pervius (affording a passage, open, penetrable),
in reference to the lack of an operculum-like structure in the remaining
pogonophorans).

Concerning the relationships of the Pogonophora I concur with van der
Land and N@rrevang (1975, 1977), Southward (197Sa, b), and George (1973)

1312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in considering the phylum to be most closely related to the phylum Annelida.
The opisthosome, with its repeated similar segments provided with chitinous
setae, as well as the morphology of the latter, is the single most compelling
feature contributing to my conclusion.

There is no question that all vestimentiferan specimens from the various
geothermal vents of the Galapagos Rift are Riftia pachyptila. I am less
certain of the identity of the single specimen available from the East Pacific
Rise; superficially there is little difference between this specimen and those
from the Galapagos Rift, some 3,400 km distant. So far I have not chosen
to dissect this single specimen; more material should be available from the
Rise area in the coming year, when a detailed examination should shed light
on the taxonomic status of the northern specimens. The relatively large
obturacular region (Table 1, USNM 59964) suggests that a different species
may be present there, but for now, I must conclude that Riftia is a mono-
specific genus.

Acknowledgments

I thank J. B. Corliss, Oregon State University, for placing the first spec-
imens of Riftia at my disposal, J. F. Grassle, Woods Hole Oceanographic
Institution, for further specimens, R. Rieger and S. Gardiner, University of
North Carolina, and W. D. Hope, M. H. Pettibone and K. Fauchald,
National Museum of Natural History, for stimulating discussions, and the
latter two for reviewing this manuscript, L. Cullen, of J. Harshbarger’s
Registry of Tumors of Lower Animals, National Museum of Natural
History, for assistance in histological procedures, and E. Jarosewich
and P. Dunn, National Museum of Natural History, for determination
of sulfur crystals. Travel support, which allowed me the opportunity of
making personal observations of living Riftia, was from the Research Fund
of the Secretary, Smithsonian Institution; Alvin dive time was made available
through the kindness of D. Cohen, National Marine Fisheries Service NOAA.

Literature Cited

Anon. 1975. Discussion. Pp. 139-143 in A. Ngrrevang (ed.), ‘‘The Phylogeny and Systematic
Position of Pogonophora.’’—Zeitschrift fiir zoologische Systematik und Evolutionsfor-
schung, Sonderheft 1, 143 pp.

. 1979. Alvin grabs giant sea worm.—Sea Secrets 23(3):11.

Ballard, R. D. 1977. Notes on a major oceanographic find.—Oceanus 20(3):35—44.

, and J. F. Grassle. 1979. Return to oases of the deep.—National Geographic

156(5):689-705.

Corliss, J. B., and R. D. Ballard. 1977. Oases of life in the cold abyss.—National Geographic
152(4):440-453.

———.,, J. Dymond, L. I. Gordon, J. M. Edmond, R. P. von Herzen, R. D. Ballard, K. Green,
D. Williams, A. Bainbridge, K. Crane, and T. H. van Andel. 1979. Submarine thermal
springs on the Galapagos Rift.—Science 203: 1073-1083.

VOLUME 93, NUMBER 4 1313

The Editor. 1979. Strange world without sun.—National Geographic 156(5):680-688.

Galapagos Biology Expedition Participants. 1979. Galapagos ’79: Initial findings of a deep-sea
biological quest.—Oceanus 22(2):2—10.

George, J. D. 1973. The Pogonophora and their affinities. —Microscopy 32:242-252.

Hekinian, R., M. Fevrier, J. L. Bischoff, P. Picot, and W. C. Shanks. 1980. Sulfide deposits
from the East Pacific Rise near 21°-N.—Science 207: 1433-1444.

Jannasch, H. W., and C. O. Wirsen. 1979. Chemosynthetic primary production at east Pacific
sea floor spreading centers.—BioScience 29:592-598.

Land, J. van der, and A. Ngrrevang. 1975. The systematic position of Lamellibrachia [sic]

(Annelida, Vestimentifera). Pp. 86-101 in A. Ngrrevang (ed.), ‘‘The Phylogeny and

Systematic Position of Pogonophora.”’ Zeitschrift fiir zoologische Systematik und Evolu-

tionsforschung, Sonderheft 1, 143 pp.

1977. Structure and relationships of Lamellibrachia (Annelida, Vestimentifera).—

Kongelige Danske Videnskabernes Selskab, Biologiske Skrifter 21(3):1-—102.

RISE Project Group. 1980. East Pacific Rise: Hot springs and geophysical experiments.—
Science 207:1421-1433.

Schlee, S. 1979. 21°N.—Science 80, No. 1:36-46.

Southward, E. C. 1975a. Fine structure and phylogeny of the Pogonophora. Pp. 235-251 in

E. J. W. Barrington and R. P. S. Jefferies (eds.), ‘“Symposia of the Zoological Society

of London,’’ No. 36.—Academic Press, New York, xii + 361 pp.

. 1975b. Pogonophora. Pp. 129-156 in A. C. Giese and J. S. Pearse (eds.), “‘Repro-

duction of marine invertebrates. Vol. II, Entoprocts and lesser coelomates.’’—Academ-

ic Press, New York, xii + 344 pp.

Terwilliger, R. C., N. B. Terwilliger, and E. Schabtach. 1980. The structure of hemoglobin
from an unusual deep sea worm (Vestimentifera). Comparative Biochemistry and Phys-
iology 65B:531-—535. ,

Webb, M. 1969a. Lamellibrachia barhami, gen. nov., sp. nov. (Pogonophora), from the north-

east Pacific.—Bulletin of Marine Science 19:18—47.

. 1969b. An evolutionary concept of some sessile and tubiculous animals.—Sarsia 38:

1-8.

. 1969c. Regionation and terminology of the pogonophoran body.—Sarsia 38:9-24.

1971. The morphology and formation of the pogonophoran tube and its value in

systematics.—Zeitschnift fiir zoologische Systematik und Evolutionsforschung 9:169-

181.

. 1975. Studies on Lamellibrachia barhami [sic] (Pogonophora). I. The excretory organs.

Pp. 102-111 in A. Ngrrevang (ed.), ‘““The Phylogeny and Systematic Position of Pogon-

ophora.”’ Zeitschrift fiir zoologische Systematik und Evolutionsforschung, Sonderheft

1, 143 pp.

1977. Studies on Lamellibrachia barhami (Pogonophora). II. The reproductive or-
gans.—Zoologische Jahrbticher, Abteilung fiir Anatomie und Ontogenie der Tiere

97:455—-481.

Division of Worms, Department of Invertebrate Zoology, National Mu-
seum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

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(Contents continued from back cover)

Notes on Axiopsis (Axiopsis) serratifrons (A. Milne Edwards) (Crustacea: Decapoda:
Thalassinidea) Brian Kensley 1253
Obandyella, a new name for Hirsutella Bandy, 1972 (Foraminiferida), non Cooper and
Muir-Wood, 1951 (Brachiopoda) D. Haman, R. W. Huddleston, and J. P. Donahue 1264
A tabulation of the Crustacea Decapoda of Bermuda
John C. Markham and John J. McDermott 1266
Coralaxius abelei, new genus and new species (Crustacea: Decapoda: Thalassinidea:
Axiidae): a coral-inhabiting shrimp from the Florida keys and the western Caribbean

Sea Brian Kensley and Robert H. Gore 1277
Riftia pachyptila, new genus, new species, the vestimentiferan worm from the Galapagos
Rift geothermal vents (Pogonophora) Meredith Jones 1295

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CONTENTS

A new species of pikeblenny (Pisces: Chaenopsidae: Chaenopsis) from the western

Atlantic Philip A. Hastings and Robert L. Shipp

The relationships of the South American pygmy characoid fishes of the genus Elacho-
charax, with a redescription of Elachocharax junki (Teleostei: Characidae)

Stanley H. Weitzman and J. Gery

New records of Stereomastis sculpta pacifica (Faxon) (Decapoda: Polychelidae) in

the eastern Pacific Ocean Mary K. Wicksten
The identity of the frog Eupsophus vanzolinii from Ramadillas, Nahuelbuta Range,
southern Chile J. R. Formas

The status of the Central American leptodactylid frogs Eleutherodactylus melano-
stictus (Cope) and Eleutherodactylus platyrhynchus (Gunther)

Jay M. Savage and James E. DeWeese

Philander and four-eyed opossums once again Philip Hershkovitz
Polydora and Boccardia species (Polychaeta: Spionidae) from western Mexico, chiefly
from calcareous habitats James A. Blake
A new fish species of the genus Vanderhorstia (Teleostei: Gobiidae) from the Amir-
ante Islands, Indian Ocean Ernest A. Lachner and James F. McKinney

Symethinae, new subfamily, and Symethis garthi, new species, and the transfer of
Raninoides ecuadorensis to Notosceles (Raninidae: Brachyura: Gymnopleura)
Gary D. Goeke
Paramunna quadratifrons, new species, the first record of the genus in the north
Pacific Ocean (Crustacea: Isopoda: Pleurogoniidae)
Ernest W. Iverson and George D. Wilson
Parapercis diplospilus (Pisces: Mugiloididae), a new species from the Philippine Is-
lands Janet R. Gomon
The South American fish genus Rachoviscus, with a description of a new species
(Teleostei: Characidae) Stanley H. Weitzman and Carlos Alberto Goncalves da Cruz
Studies on decapod crustacea from the Indian River region of Florida. XIX. Larval
development in the laboratory of Lepidopa richmondi Benedict, 1903, with notes on
larvae of American species in the genus (Anomura: Albuneidae)
Robert H. Gore and Cindy Lee Van Dover
Review of Hysterothylacium and Iheringascaris (both previously = Thynnascaris)
(Nematoda: Anisakidae) from the northern Gulf of Mexico
Thomas L. Deardorff and Robin M. Overstreet
Syllidae (Polychaeta), principally from Florida, with descriptions of a new genus and

twenty-one new species Thomas H. Perkins
Barbidrilus paucisetus, new genus, new species (Oligochaeta: Enchytraeidae), from
eastern North America Michael S. Loden and Steven M. Locy

The tadpole of the Costa Rican fringe-limbed tree-frog, Hyla fimbrimembra
Jay M. Savage

Common inshore barnacle cyprids of the Oregonian Faunal Province (Crustacea:

Cirripedia) Jon D. Standing
A new species and a new genus of Cerviniidae (Copepoda: Harpacticoida) from the
Beaufort Sea, with a revision of the family Paul A. Montagna
Two new records of the Caribbean marine tubificid Kaketio ineri Right and Kanner
(Oligochaeta) Christer Erséus
A new species of the marine amphipod genus Gammaropsis from the southeastern
United States (Photidae) Walter G. Nelson

Cestode parasites of some Venezuelan stingrays
Monte A. Mayes and Daniel R. Brooks
Cestode parasites in Myliobatis goodei Garman (Myliobatiformes: Myliobatidae) from
Rio de la Plata, Uruguay, with a summary of cestodes collected from South American
elasmobranchs during 1975—1979
Daniel R. Brooks, Monte A. Mayes, and Thomas B. Thorson

(Contents continued inside back cover)

875

887

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943

947

963

971

982
989

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1035
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1173
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1220
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