fc

Cyclopoid Copepods

(Nanaspididae and Sabelliphilidae)

Associated with Holothurians

in New Caledonia

RTHUR G. HUMES

'l\

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 202
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 20 2

Gyclopoid Copepods

(Nanaspididae and Sabelliphilidae)

Associated with Holothurians

in New Caledonia

Arthur G. Humes

ISSUED

G 22 1975

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Humes, Arthur G. Cyclopoid Copepods (Nanaspididae and Sabelliphilidae)

Associated with Holothurians in New Caledonia. Smithsonian Contributions to

Zoology, number 202, 41 pages, 24 figures, 1975.—Nanaspis mixta, new species,

from Microthele nobilis; Chauliolobion bulbosum, new genus, new species, from

Actinopyga echinites and Actinopyga obesa palauensis; Chauliolobion halodeimatis, new genus, new species, from Halodeima atra; Scambicornus tylotus, new

species, from Bohadschia argus; Scambicornus sewelli, new species, from Halodeima edulis, Halodeima coluber, Halodeima atra, Holothuria curiosa, Actinopyga echinites, and Microthele nobilis; Scambicornus calcaratus, new species,

from Actinopyga miliaris, Actinopyga obesa palauensis, Actinopyga plebeja,

Actinopyga serratidens, Actinopyga echinites, and Actinopyga lecanora. Six

species of Scambicornus are reported from New Caledonia for the first time, their

ranges being extended from the western Indian Ocean (Madagascar and Mauritius): Scambicornus tuberatus from Bohadschia xntiensis and from a new host

Bohadschia argus; Scambicornus subtilis from Halodeima edulis and Holothuria

curiosa, and from the new hosts Halodeima coluber, Halodeima atra, Actinopyga

echinites, and Microthele nobilis; Scambicornus campanulipes from Actinopyga

lecanora and a new host Actinopyga plebeja; Scambicornus modestus from

Stichopus chloronotus and Stichopus variegatus; Scambicornus poculiferus from

Synapta maculata; Scambicornus idoneus from Holothuria leucospilota, Halodeima atra, Actinopyga echinites, and Microthele nobilis. The Indian species

Scambicornus brachysetosus Reddiah, 1968, is briefly redescribed and figured

from paratypes.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5337. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus.)

Library of Congress Cataloging in Publication Data

Humes, Arthur Grover, 1916-

Cyclopoid copepods (Nanaspididae and Sabelliphilidae) associated with holothurians in New

Caledonia.

(Smithsonian contributions to zoology, no. 202)

Includes bibliographical references.

Supt. of Docs, no.: SI 1.27:202

1. Sabelliphilidae. 2. Nanaspididae. 3. Crustacea. I. Title. II. Series: Smithsonian contributions to zoology, no. 202.

QL1.S54 no. 202 [QL444.C73] 591'.08s [595'.34] 75-619000
Contents

Page

Introduction 1

Nanaspididae 2

Nanaspis 2

Nanaspis mixta 2

Sabelliphilidae 6

Chauliolobion 6

Chauliolobion bulbosum 7

Chauliolobion halodeimatis 14

Scambicornus 19

Scambicornus tylotus 19

Scambicornus sewelli 25

Scambicornus calcaratus 32

New Records and Hosts 37

Notes on Scambicornus brachysetosus Reddiah, 1968 38

List of New Caledonian Holothurians and Their Associated Copepods . . 40

Literature Cited 40

Cyclopoid Copepods

(Nanaspididae and Sabelliphilidae)

Associated with Holothurians

in New Caledonia

Arthur G. Humes

Introduction

Cyclopoid copepods are frequently associated

with Holothuroidea, especially in tropical waters.

The genus Nanaspis Humes and Cressey, 1959, containing ten species, is known from holothurians

from the West Indies, the western coast of Sweden,

the Mediterranean coast of France, Madagascar,

and Eniwetok Atoll. Several sabelliphilid genera

are known from holothurians, chiefly in the West

Indies, Madagascar, and the Red Sea. These records have been summarized by Humes and Stock

(1973). Associations of copepods and holothurians

have not previously been reported in New Caledonia.

Twelve copepods are here recorded from various

shallow-water holothurians in New Caledonia.

These comprise six new species, and six already

known species for which there are six new hosts

(preceded by an asterisk in the list below). The

twelve species are:

Nanaspis mixta, new species

from Microthele nobilis (Selenka)

Chauliolobion bulbosum, new genus, new species

from Actinopyga echinites (Jaeger) and Actinopyga obesa

palauensis Panning

Chauliolobion halodeimatis, new species

from Halodeima atra (Jaeger)

Scambicornus tylotus, new species

from Bohadschia argus (Jaeger)

Arthur G. Humes, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts, 02543.

Scambicornus sewelli, new species

from Halodeima edulis (Lesson), Halodeima coluber (Semper), Halodeima atra (Jaeger), Holothuria curiosa Ludwig, Actinopyga echinites (Jaeger), and Microthele

nobilis (Selenka)

Scambicornus calcaratus, new species

from Actinopyga miliaris (Quoy and Gaimard), Actinopyga

obesa palauensis Panning, Actinopyga plebeja (Selenka),

Actinopyga serratidens Pearson, Actinopyga echinites

(Jaeger), and Actinopyga lecanora (Jaeger)

Scambicornus tuberatus (Humes and Cressey, 1961)

from Bohadschia vitiensis (Semper) and 'Bohadschia argus

(Jaeger)

Scambicornus subtilis (Humes and Cressey, 1961)

from Halodeima edulis (Lesson), *Halodeima coluber

(Semper), *Halodeima atra (Jaeger), Holothuria curiosa

Ludwig, 'Actinopyga echinites (Jaeger), and 'Microthele

nobilis (Selenka)

Scambicornus campanulipes (Humes and Cressey, 1961)

from *Actinopyga plebeja (Selenka) and Actinopyga lecanora (Jaeger)

Scambicornus modestus (Humes and Cressey, 1961)

from Stichopus chloronotus (Brandt) and Stichopus variegatus Semper

Scambicornus poculiferus (Humes and Cressey, 1961)

from Synapta maculata (Chamisso and Eysenhardt)

Scambicornus idoneus (Humes and Cressey, 1961)

from Holothuria leucospilota (Brandt), Halodeima atra

(Jaeger), Actinopyga echinites (Jaeger), and Microthele

nobilis (Selenka)

The field work in New Caledonia during JuneAugust, 1971, and the ensuing study of the copepods were made possible by a grant (GB-8381X)

from the National Science Foundation. Mr. Roger

C. Halverson from the University of California at
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Santa Barbara assisted in the field work. I wish to

acknowledge with thanks and appreciation the aid

provided by the staff of the Centre ORSTOM de

Noumea.

For the identification of the holothurian hosts

I am greatly indebted to Monsieur G. Cherbonnier,

Museum National d'Histoire Naturelle, Paris. I

wish also to thank Shri G. Ramakrishna, Superintending Zoologist, Zoological Survey of India, for

sending me paratypes of Scambicornus brachysetosus Reddiah, 1968.

All figures have been drawn with the aid of a

camera lucida. The letter after the explanation of

each figure refers to the scale at which it was

drawn. The abbreviations used are: At = first antenna, A2 = second antenna, MXPD = maxilliped,

and Pj = leg 1.

The measurements were made on specimens in

lactic acid and are expressed in microns unless

otherwise stated. The body length does not include

the setae on the caudal rami. The lengths of the

first antennal segments were measured along their

posterior nonsetiferous margins.

Specimens that are deposited in the National

Museum of Natural History, Smithsonian Institution, Washington, D.C., are listed under the catalog numbers of the former United States National

Museum (USNM).

NANASPIDIDAE Humes and Cressey, 1959

Nanaspis Humes and Cressey, 1959

Nanaspis mixta, new species

FIGURES 1-3

TYPE MATERIAL.—327 9 9 , 223 $ $, and 41

copepodids from two holothurians, Microthele

nobilis (Selenka), in 2 m, west of Isle Mando, near

Noumea, New Caledonia, 22°18'59"S, 166°09'30"E,

3 July 1971. Holotype 9, allotype, and 380 paratypes (250 9 9, 130 $ s ) deposited in the National Museum of Natural History, Smithsonian

Institution, Washington, D.C.; 80 paratypes (40

9 9. 40 $ <J ) in the Zoologisch Museum, Amsterdam; the remaining paratypes in the collection of

the author.

OTHER SPECIMENS.—17 9 9 from two Microthele

nobilis, in 0.5 m, off Goro, southeastern New Caledonia, 22°18'00"S, 167°02'00"E, 6 August 1971.

FEMALE.—Body (Figure la-c) ovoid and moderately flattened dorsoventrally. Length of the shieldlike prosome (not including the urosome, which is

folded ventrally in specimens preserved in ethyl

alcohol) 450 (440-462) and the greatest width 327

(319-330), based on 10 specimens. Prosome with a

dorsal transverse suture separating the anterior

cephalosome (including the segment of leg 1) from

the posterior metasome (the segments of legs 2-4).

Posterior margin of the metasome truncated.

Urosome (Figure \d,e) with three distinct segments (that of leg 5, the genital segment, and the

anal segment). Segment of leg 5 (Figure \e) with

four transverse ventral rows of spinules. Genital

area (Figure If) with a small naked seta 5.5 long.

Anal segment rather deeply incised.

Egg sac (Figure la) 200 X 120 not including the

neck, and containing two eggs.

Caudal ramus (Figure lg) minute, 14 X 11, bearing six naked setae, the two longest 65 and 140.

Ramus with a few distal inner spinules.

Arrangement of the appendages as in Figure \c.

Rostral area not well developed (Figure lc).

First antenna (Figure \h) 82 long and probably

5-segmented, though the second and third segments

are incompletely separated. Lengths of the segments:

12 (19 along the anterior margin), 14, 5, 4.5, 4.5,

6.5, 6.5, and 20 respectively. Armature: 1, 1, 1, 2, 2,

3, and 7 + 2 aesthetes. All setae naked. Second antenna (Figure 2a) about 130 long, 4-segmented, the

first segment unornamented, the second segment

with a few distal outer spinules, the elongated

third segment with spinules as illustrated, the small

fourth segment with a minute seta and a minutely

setose pad with a recurved tip.

Oral cone (Figure 2b) projecting ventrally (Figure \b). Mandible (Figure 2c) a simple blade 23

long. First maxilla (Figure 2d) with the inner

branch having one spinulose seta and the outer

branch having three naked setae. Seta external to

the base of the first maxilla and near the base of

the second antenna large and spinulose. Second

maxilla (Figure 2e) with the large first segment

bearing a few slender outer spinules; claw 73 along

its axis and reflexed distally. Maxilliped (Figure

2/) 164 long. Ventral area between and immediately

behind the maxillipeds as in Figure 2g.

Legs 1 and 2 biramous. Both rami of leg 1 a single segment (Figure 2h). I>eg 2 (Figure 2;) with

both rami 3-segmented. Legs 3 and 4 uniramous
NUMBER 202

FIGURE 1.—Nanaspis mixta, new species, female: a, dorsal (A); b, lateral (A); C, ventral (A);

d, urosome, dorsal (B); e, urosome, ventral (B); /, genital area, dorsal (c); g, caudal ramus,

ventral (c); h, first antenna, ventral (D).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 2.—Xanaspis mixta, new species, female: a, second antenna, posteroventral (D); b, oral

cone, ventral (B); C, mandible, ventral (E); d, first maxilla and adjacent seta, posterior (B);

e, second maxilla, posterior (B); /, maxilliped, anterior (B); g, area between and behind bases of

maxillipeds, ventral (B); h, leg 1, anterior (D); i, leg 2 and intercoxal plate, posterior (B);

;, leg 3, anterior (B); k, leg 4, anterior (B); /, leg 5, ventral (c).
NUMBER 202

(the endopods absent), the ramus of leg 3 3-

segmented (Figure 2;), that of leg 4 2-segmented

(Figure 2k).

Spine and setal formula for legs 1-4 as follows

(the Roman numerals indicating spines, the Arabic

numerals setae):

P.

p2

ps

p«

protopod

coxa

coxa

coxa

1-0

0-0

0-0

0-0

basis

basis

basis

0-0

0-0

0-0

exp

enp

exp

enp

exp

exp

4

1

1-0;

0-1;

1-0;

1-0;

I-l

0-1;

I-l;

I.H.2

111,3

1,3

111,3

In leg 1 (Figure 2h) the protopod bearing an

outer seta 20 long. Exopod with four terminal

setae from outer to inner 17, 38, 70, and 8. Endopod

bearing only one seta 39 long and ornamented with

a few small spinules. All setae on leg 1 smooth.

Leg 2 (Figure 2i) with delicately fringed spines

and plumose setae. Leg 3 (Figure 2;) with the two

terminal spines on the third segment inserted very

close together and partly overlapping. Leg 4 (Figure 2k) with the spines having only a few small

distal barbules and the setae weakly plumose. Both

segments of the ramus ornamented with spinules.

Leg 5 (Figure 21) oval and flattened, 34 X 26 in

ventral view, and held in preserved specimens ventral to the edge of the genital segment (Figure

\d,e); bearing two naked setae 19 and 12 and

ornamented marginally with many long slender

spinules.

Leg 6 probably represented by the single small

seta on the genital area (Figure If).

Living specimens in transmitted light slightly

opaque, the eye red, the egg sacs gray.

MALE.—Body (Figure 3a) ovoid and flattened

dorsoventrally, with the posterior margin of the

metasome rounded. Length of the prosome (not

including the urosome, which is folded ventrally)

FIGURE 3.—Nanaspis mixta, new species, male: a, dorsal (A); b, urosome, dorsal (B); C, urosome,

ventral (B); d, urosome and leg 5, lateral (B); e, first antenna, antero-outer (B); /, last two

segments of first antenna, antero-outer (D).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

367 (360-385) and the greatest width 270 (264-

275), based on 10 specimens.

Urosome (Figure 5b-d) 3-segmented. Segment of

leg 5 smooth dorsally but with five transverse rows

of spinules ventrally. Genital segment longer dorsally than ventrally and unornamented. Anal segment deeply incised and ornamented with slender

spinules.

Caudal ramus similar to that of the female, but

having a few small spinules along the outer margin.

Rostral area undeveloped as in the female. First

antenna (Figure $e,f) 5-segmented, the first two

segments nearly equal in size, the third segment

small, the fourth segment swollen and bearing two

small inner dentate ridges, and the fifth segment

clawlike. Formula for the armature: 1, 14, 6, 5, and

10 + 2 aesthetes. Two of the elements on the claw

short and obtuse.

Second antenna, oral cone, mandible, first maxilla, second maxilla, maxilliped, and legs 1-4 as in

the female.

Leg 5 (Figure 3d) consisting of a slender segment

about 26 long. Two naked terminal setae and a

large hyaline lamellate element arising from the

middle of the ventral edge of the segment.

Leg 6 not identifiable. Genital openings probably between the fifth pedigerous and genital segments (as suggested by Stock, Humes, and Gooding,

1962, in their original description of Nanaspis pollens).

Spermatophore not seen.

Living specimens colored as in the female.

ETYMOLOGY.—The specific name mixta from

Latin (= mixed or combined) alludes to the combination in the new species of certain characters

found in West Indian and European species of

Nanaspis.

COMPARISON WITH OTHER SPECIES IN THE GENUS.

—In the four West Indian species of Nanaspis described by Stock, Humes, and Gooding (1962),

namely N. pollens, N. exigua, N. media, and N.

truncata, the endopod of leg 2 is 3-segmented as

in the new species. In all other species (N. tonsa

Humes and Cressey, 1959, N. mediterranea Stock

and Kleeton, 1963, Ar. ninae Bresciani and Liitzen,

1962. N. manca Humes, 1973, Ar. spinifera Humes,

1973, and N. pusilla Humes, 1973) this endopod is

2-segmcnted. A similar distinction is seen in the

number of setae on the exopod of leg 1, where

there are four in the new species and in the West

Indian species, but three in all others. In respect

to the formula for leg 4, N. mixta differs from the

West Indian species, where this formula is 1-0;

IV, 1.

The well-formed leg 5 in both sexes of the new

species serves as a useful recognition character. In

no other species is the fifth leg of the female as

large and as clearly set off from the body segment;

in no other species does the fifth leg of the male

bear a large hyaline lamellate element.

SABELLIPHILIDAE Gurney, 1927

Chauliolobion, new genus

DIAGNOSIS.—Sabelliphilidae. Body modified, elongated. Urosome 5-segmented in the female, 6-

segmented in the male. Caudal ramus with the two

median terminal setae relatively short. Rostrum

not well defined. First antenna 7-segmented, with

the formula in both sexes 4, 13, 6, 3, 4 + 1 aesthete,

2 + 1 aesthete, and 7 + 1 aesthete and with a few

small spines on all but the last segment. Second

antenna 4-segmented, with the formula 1, 1, 3 +

one claw, and 7.

Labrum with two posteroventral lobes. Mandible

with two long setiform processes arising near the

base of the lash. Paragnath a small hairy lobe.

First maxilla with two elements. Second maxilla

similar to that in other sabelliphilid genera. Maxilliped in the female 3-segmented with a pointed

tip; in the male 4-segmented (if the proximal part

of the claw is considered as a fourth segment).

Legs 1-4 in the female with both rami 3-

segmented, in the male with the exopods 3-

segmented but the endopods 2-segmented. In both

sexes a long digitiform process on the distal outer

corner of the coxa of legs 2 and 3. In the female

the endopod of leg 3 with the formula 0-1; 0-1;

1,1,2; that of leg 4 with 0-1; 0-1; 11,1 (or 1,2). In

the male these endopods 0-1; 1,1,2,2 and 0-1; 1,2,1

respectively. Exopod of leg 4 in both sexes 1-0;

1-1; 11,1,5. Leg 5 with a free segment bearing two

terminal elements. Leg 6 in the female represented

by the two setae near the genital openings, in the

male by a posteroventral flap on the genital segment bearing two setae.

Other features as in the species described below.

Endoparasitic in holothurians.
NUMBER 202

TYPE-SPECIES.—Chauliolobion bulbosum, new

species.

ETYMOLOGY.—The generic name is a combination

of the Greek j(auAioc; (projecting) and Xo|3iov (a

small lobe), alluding to the digitiform processes on

the coxa of legs 2 and 3.

COMPARISON WITH THE MOST CLOSELY RELATED

SABELLIPHILID GENUS.—Chauliolobion is close to

Lichothuria Stock, 1968, a genus which contains

the single species L. mandibularis Stock, 1968 (partially redescribed by Humes and Ho, 1969), endoparasitic in holothurians in the Gulf of Aqaba

and in Madagascar. As in Lichothuria the body of

Chauliolobion is elongated and slender, the terminal setae on the caudal ramus are relatively short,

and leg 5 is placed ventrally rather than dorsally.

In the female the two genera have the formula

1,1,2 on the third endopod segment of leg 3 and

three elements on the third endopod segment of

leg 4. In the male the endopods of legs 1-4 of both

genera are 2-segmented and the second segment

of the endopod of leg 4 has the formula 11,1,1.

In spite of these similarities, however, there are

several major features that occur in the two species

of Chauliolobion but are not found in Lichothuria.

Chief among these are the long digitiform process

on the distal outer corner of the coxa in legs 2 and

3, the two long setiform processes arising near the

base of the attenuated lash of the mandible, the

presence of small spines on most of the first antennal segments, and the claw of the male maxilliped being much longer than the second segment.

Chauliolobion bulbosum, new species

FIGURES 4-8

TYPE MATERIAL [all from Actinopyga echinites

(Jaeger)].—Holotype $ , allotype $ , from 22 hosts,

in 2 m, eastern end of Ricaudy Reef, near Noumea,

New Caledonia, 22°19'00"S, 166O27'18"E, 18 June

1971. Paratypes: 1 $, 2 S S from 1 host, in 1-2

m, Rocher a la Voile, Noumea, 22°18'24"S,

166°25'50"E, 2 June 1971; 2 $ $>, 1 $ from 12

hosts, in 2 m, Rocher a la Voile, 10 June 1971; 1 8

from 35 hosts, intertidal, Ricaudy Reef, 22°19'00"S,

166°26'44"E, 21 July 1971; 2 $ $ from 2 hosts, in

2 m, north of Isle Maitre, near Noumea, 22°19'00"S,

166°24'35"E, 13 July 1971. All types deposited in

the National Museum of Natural History, Smithsonian Institution, Washington, D.C., except 1 $

and 1 $ (dissected) in the collection of the author.

OTHER SPECIMENS.—4 $ $, 17 $ $ from one

Actinopyga obesa palauensis Panning in 3 m, north

of Isle Maitre, 22°19'30"S, 166°24'35"E, 13 July

1971.

FEMALE.—Body (Figure 4a,b) elongated and

slender, with the urosome not much thickened

dorsoventrally. Length 1.27 mm (1.09-1.40 mm)

and the greatest width 0.39 mm (0.39-0.41 mm),

based on 4 specimens. Epimeral areas of the segments of legs 1-4 broadly rounded. Ratio of the

length to the width of the prosome 2.14:1. Ratio

of the length of the prosome to that of the urosome

1.46:1.

Segment of leg 5 (Figure 4c,d) 99 X 221. Between this segment and the genital segment no ventral sclerite. Genital segment 234 X 244, slightly

wider than long, tapered posteriorly in dorsal view,

and ornamented ventrally with a posterior transverse row of spines. Genital areas located dorsolaterally near the midregion of the segment. Each

area (Figure 4e) bearing two naked setae 31 and

17. Three postgenital segments from anterior to

posterior 83 X 120, 73 X 94, and 49 X 78. First

and second postgenital segments with a ventral

transverse row of spines, that on the second segment continued dorsally. Anal segment with posterolateral spines and posteroventrally with two

short transverse rows of spines.

Caudal ramus (Figure 5a) 55 X 30, nearly two

times longer than wide. Outer lateral seta 50, dorsal

seta 22, outermost terminal seta 44, and innermost

terminal seta 44. Two median terminal setae relatively short and stout, the outer 138, the inner 216.

All setae spinulose as illustrated except the dorsal

seta which is smooth. Ramus ornamented with

small spines near the insertions of the outer lateral

seta and the outermost terminal seta, and with

small spines along the edge of the small dorsal

flange covering the insertions of the two median

setae.

Body surface with very few hairs (sensilla) as in

Figure 4a.

Egg sac unknown.

Rostral area (Figure 5?;) not protuberant.

First antenna (Figure 5c) 264 long. Lengths of

its seven segments: 24 (53 along the anterior edge).

46, 28, 39, 42, 32, and 24 respectively. Formula for

the armature: 4, 13, 6, 3, 4 + 1 aesthete, 2+ 1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Chauliolobion bulbosum, new genus, new species, female: a, dorsal (F); b, lateral

(F); C, urosome, dorsal (A); d, urosome, ventral (A); e, genital area, dorsal (B).
NUMBER 202

FIGURE 5.—Chauliolobion bulbosum, new genus, new species, female: a, caudal ramus, dorsal

(D); b, rostral area, ventral (c); c, first antenna, dorsal (H); d, second antenna, anterior (B);

e, second antenna, posterior (B); /, mandible, posterior (c); g, first maxilla, posterior (c);

h, second maxilla, posterior (D); i, tip of second maxilla, anterior (D).
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

aesthete, and 7+ 1 aesthete. Many of the setae

unilaterally with spinules. First segment with a

distal dorsal row of small spines. A few spines on

segments 2-5 and groups of spinules on the posterior surface of segments 5 and 6.

Second antenna (Figure 5d,e) 4-segmented, 180

long including the claw. Armature: 1, 1, 3 + claw,

and 7. Setae on the first two segments and two setae

on the fourth segment spinulose. Fourth segment

much shorter than the claw which is about 45 along

its axis. Outer surface of the second segment covered with small spines. Transverse rows of slender

spinules on the postero-outer surface of the third

and fourth segments.

Labrum with two small rounded posteroventral

lobes as in Figure bb.

Mandible (Figure 5/) having on the concave

margin near the base of the bispinulose lash a

pointed process and on the convex margin two

long setiform processes. Paragnath a small hairy

lobe. First maxilla (Figure 5g) with two unequal

terminal elements, the longer element sparsely

FIGURE 6.—Chauliolobion bulbosum, new genus, new species, female: a, maxilliped, antero-inner

(D); b, area between maxillipeds and leg 1, ventral (c); c, leg 1 and intercoxal plate, anterior

(B); d, leg 2 and intercoxal plate, anterior (B); e, outer process on coxa of leg 2, posterior (B).
NUMBER 202 11

a

FICURE 7.—Chauliolobion bulbosum, new genus, new species. Female: a, leg 3 and intercoxal

plate, anterior (B); b, leg 4 and intercoxal plate, anterior (B); C, leg 5, ventrolateral (B). Male:

d, dorsal (F).
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Chauliolobion bulbosum, new genus, new species, male: a, urosome, dorsal (A);

b, urosome, ventral (A); C, maxilliped, postero-inner (B); d, endopod of leg 1, anterior (D);

e, endopod of leg 2, anterior (D); /, endopod of leg 3, anterior (D); g, endopod of leg 4, anterior

(D); h, leg 5, ventral (B); i, leg 6, ventral (H).
NUMBER 202 13

barbed. Second maxilla (Figure 5h) 2-segmented,

the large first segment unarmed. Second segment

having a small proximal setule on its outer (ventral) margin, a finely barbed seta on its posterior

surface, and inner (dorsal) spiniform element with

prominent lateral spinules; segment terminating

in a moderately short lash with a row of long spines

along the distal edge and a row of spines on the

anterior surface (Figure 5i). Maxilliped (Figure

6a) 3-segmented, the first segment unarmed, the

second with two short naked spines, and the third

with two small setules and having a clawlike tip.

Area between the maxillipeds and the first pair

of legs (Figure 66) not protuberant (see Figure

4b).

Legs 1-4 (Figures 6c,d; 7a,b) with 3-segmented

rami throughout. Armature as follows (the Roman

numerals indicating spines, the Arabic numerals

setae):

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,4

enp 0-1; 0-1; 1,5

P2 coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,5

enp 0-1; 0-2; 1,11,3

Ps coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,5

enp 0-1; 0-2; 1,1

P4 coxa 0-1 basis 1-0 exp 1-0; 1-1; II,

_ » « f\ 1 . A 1 . TT

exp 1-0; 1-1; 11,1,5

enp 0-1; 0-1; 11,1

Intercoxal plate of legs 1 and 2 with a pair of

prominent pointed ventral processes. These processes on leg 3 small and rounded; absent on leg 4.

Inner coxal seta in legs 1—3 proximally swollen and

setulose; in leg 4 this seta not swollen. Distal outer

corner of the coxa of legs 2 and 3 bearing a digitiform process (Figure 6e) about 80 long. Endopod

of leg 2 with the inner seta on the first segment

setulose on its posterior surface; endopod of leg 3

with the inner setae on the setae on the first and

second segment similarly setulose. Leg 4 with the

exopod 125 long; the endopod 75, with the three

spines from outer to inner 11, 47, and 39. Two

inner spines of this trio plumose proximally but

barbed distally.

Leg 5 (Figure 7c) placed somewhat ventrally.

Free segment 68 X 28, about 2.43 times longer

than wide. Two terminal elements unequal, one a

slender barbed seta 46 and the other a stout spiniform finely barbed element 50 long. Rows of small

spines on the segment near the insertions of these

elements. Seta on the body near the insertion of

the free segment situated ventrally and only 15

long.

Leg 6 probably represented by the two setae on

the genital area (Figure 4e).

Color in life in transmitted light opaque, the

eye red.

MALE.—Body (Figure 7d) slender. Length 1.13

mm (1.02-1.27 mm) and the greatest width 0.28

mm (0.23-0.31 mm), based on 6 specimens. Ratio

of the length to the width of the prosome 2.38:1.

Ratio of the length of the prosome to that of the

urosome 1.29:1.

Segment of leg 5 (Figure 8a,b) 68 X 120. Between this segment and the genital segment no

ventral intersegmental sclerite. Genital segment

subquadrate, 174 X 156. Four postgenital segments

from anterior to posterior 57 X 86, 62 X 70, 57 X

60, and 29 X 60, ornamented with rows of spinules

as in the female.

Caudal ramus similar to that of the female but

shorter, 40 X 21.

Body surface ornamented with very few hairs

(sensilla).

Rostral area, first antenna, second antenna,

labrum, mandible, paragnath, first maxilla, and

second maxilla like those in the female. Maxilliped

(Figure 8c) 4-segmented, assuming that the proximal part of the claw represents a fourth segment.

First segment unarmed. Second segment with two

short setae and two rows of spines. Small third

segment unarmed. Claw 110 along its axis, bearing

proximally two unequal naked setae and a row of

small hyaline teeth on the concave margin; terminal lamella small.

Area between the maxillipeds and the first pair

of legs like that of the female.

Legs 1-4 resembling those of the female except

for the segmentation of the endopods, which are

2-segmented rather than 3-segmented (Figure

8d-g), the two distal segments fused, in legs 1 and 4

the original separation very indistinct, in legs 2 and

3 the point of separation indicated by a transverse

row of spinules. Endopod of leg 4 (Figure &g) 57

long, its three spines from outer to inner 17.5, 36,

and 35.

Leg 5 (Figure 8/J) with the free segment 34 X

15.5, its two subequal terminal elements 52 and 57.

Leg 6 (Figure 8?) a posteroventral flap on the

genital segment bearing two unequal naked setae

39 and 22.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Spermatophore (Figure 4a) oval, 135 X 70.

Color as in the female.

ETYMOLOGY.—The specific name bulbosum, Latin

(= containing bulbs), alludes to the bulblike swelling of the inner coxal setae on legs 1-3.

Chauliolobion halodeimatis, new species

FIGURES 9-12

TYPE MATERIAL.—5 $9, 3 $ $ from 8 Halodeima atra (Jaeger), in 1.5 m, Rocher a la Voile,

Noumea, New Caledonia, 22°18'24"S, 166°25'50"E,

2 June 1971. Holotype $ , allotype, and 6 paratypes

(4 $ $ , 2 $ $ ) deposited in the National Museum

of Natural History, Smithsonian Institution, Washington, D.C.

OTHER SPECIMENS.—4 $ $, 3 $ $ from 31

Halodeima atra, in 4-5 m, on the reef between

Isle Ndie and Mt. Kumuru, northwest of Noumea,

22°13'24"S, 166°24'11"E, 29 July 1971.

FEMALE.—Body (Figure 9a) elongated and slender. Length 1.01 mm (0.91-1.13 mm) and the greatest width 0.24 mm (0.23-0.26 mm), based on 8

specimens. Epimeral areas of the segments of legs

1-4 variously formed as in the figure. Ratio of the

length to the width of the prosome 2.30:1. Ratio of

the length of the prosome to that of the urosome

1.33:1.

Segment of leg 5 (Figure 9b,c) 57 X 140. Between this segment and the genital segment no ventral sclerite. Genital segment 150 X 160, slightly

wider than long, in dorsal view widest in its anterior third and tapered posteriorly, and bearing

ventrally a posterior transverse row of spines. Genital areas situated laterally near the midregion of

the. segment. Each area (Figure 9d) having two

naked setae 22 and 33. Three postgenital segments

from anterior to posterior 52 X 88, 60 X 73, and

35 X 65. First and second postgenital segments

with a ventral transverse row of spines. Anal segment with a short posteroventral row of spines on

each side.

Caudal ramus (Figure 9e) 53 X 25, 2.12 times

longer than wide. Outer lateral seta 62, dorsal seta

20, outermost terminal seta 77, innermost terminal

seta 60. Two median terminal setae relatively short,

the outer 122, the inner 156. All setae spinulose

except the smooth dorsal seta. Ramus with small

spines near the outer lateral seta and the outermost

terminal seta; small terminal dorsal flange smooth.

Body surface with very few hairs (sensilla) as in

Figure 9a.

Egg sac unknown.

Rostral area similar to that in C. bulbosum.

First antenna (Figure 9/) 174 long. Lengths of

its seven segments: 12 (29 along the anterior edge),

37, 15, 30, 29, 17.5, and 16.5 respectively. Armature

as in C. bulbosum; only one group of small spines

on the fifth segment.

Second antenna (Figure 9g) 4-segmented, 138

long including the claw. Armature as in C. bulbosum. Fourth segment much shorter than the claw

which is 34 along its axis. Spinules on the inner

surface of the first segment, on both inner and

outer surfaces of the second segment, and on the

outer surface of the fourth segment.

Labrum (Figure 10a) with the two lobes widely

divergent.

Mandible (Figure 10&) resembling that in C.

bulbosum; the outer setiform process hyaline.

Paragnath a small hairy lobe (Figure 10a). First

maxilla (Figure 10a) with two very unequal naked

elements. Patch of small spines on the ventral surface of the head at either side of the labrum. Second maxilla (Figure 10c) similar in basic form to

that of C. bulbosum, but the seta on the posterior

surface of the second segment smooth and the

inner (dorsal) spiniform element on that segment

stout and spinulose with a blunt tip. Maxilliped

(Figure 10rf) resembling that in C. bulbosum, but

the third segment more attenuated than in that

species.

Area between the maxillipeds and the first pair

of legs (Figure lOe) not protuberant.

Legs 1-4 (Figures 10/,g; \\a,b) segmented as in

C. bulbosum and possessing the same spine and

setal formula as in that species with the exception

of the third endopod of leg 4, which is 1,2. Distal

outer corner of the coxa of legs 2 and 3 bearing

a prominent digitiform process or lobe, which in

leg 2 is about as long as the exopod (Figure 10g).

Leg 4 with the exopod 96 long. Endopod of this

leg 62, with the last two segments not completely

separated; terminal armature consisting of an outer

spine 18 and two setae 36 (outer) and 52 (inner).

Leg 5 (Figure lie) placed ventrally as in C.

bulbosum. Free segment 42 X 17, about 2.47 times

longer than wide. Two unequal terminal elements

39 and 68. A few small spines near the insertion of
NUMBER 202 15

FIGURE 9.—Chauliolobion halodeimatis, new genus, new species, female: a, dorsal (F); b, urosome,

dorsal (c); c, urosome, ventral (c); d, genital area, lateral (B); e, caudal ramus, dorsal (D);

/, first antenna, ventral (B); g, second antenna, anterior (D).
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Chauliolobion halodeimatis, new genus, new species, female: a, labrum, paragnaths,

and first maxillae, ventral (D); b, mandible, posterior (c); c, second maxilla, anterior (c);

d, maxilliped, antero-inner (c); e, area between maxillipcds and leg 1, ventral (B); /, leg 1 and

intercoxal plate, anterior (D); g, leg 2, anterior (D).
NUMBER 202 17

a

FIGURE 11.—Chauliolobion halodeimatis, new genus, new species. Female: a, leg 3 and intercoxal

plate, anterior (D); b, leg 4 and intercoxal plate, anterior (D); C, leg 5, ventral (D). Male: d,

dorsal (F).
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

h

FIGURE 12.—Chauliolobion halodeimatis, new genus, new species, male: a, urosome, dorsal (c);

b, maxillipcd, postero-inner (D); C, endopod of leg 1, anterior (D); d, endopod of leg 2,

anterior (D); e, endopod of leg 3, anterior (D); /, endopod of leg 4, anterior (D); g, leg 5,

ventral (D); h, leg 6, ventral (H).
NUMBER 202 19

the smaller seta. Seta on the body near the insertion of the free segment situated ventrally and 16

long.

Leg 6 probably represented by the two setae on

the genital area (Figure 9d).

Color in life in transmitted light opaque, the

eye red.

MALE.—Body (Figure lid) slender. Length 0.92

mm (0.86-0.98 mm) and the greatest width 0.20

mm (0.19-0.21 mm), based on 7 specimens. Ratio

of the length to the width of the prosome 2.41:1.

Ratio of the length of the prosome to that of the

urosome 1.09:1.

Segment of leg 5 (Figure 12a) 52 X 117. Between

this segment and the genital segment no ventral

sclerite. Genital segment quadrate, 148 X 146.

Four postgenital segments from anterior to posterior 55 X 82, 65 X 73, 62 X 65, and 34 X 63,

ornamented ventrally with rows of spinules as in

the female.

Caudal ramus resembling that of the female and

of about the same size.

Body surface, rostal area, first antenna, second

antenna, labrum, mandible, paragnath, first maxilla, and second maxilla like those in the female.

Maxilliped (Figure 126) 4-segmented, considering

the proximal part of the claw to represent a fourth

segment. First segment unarmed. Second segment

with two short naked setae, a large inner patch of

spines, and a short row of spines. Small third segment unarmed. Claw 105 along its axis including

the terminal lamella, bearing proximally two very

unequal naked setae.

Area between the maxillipeds and the first pair

of legs like that of the female.

Legs 1-4 similar to those of the female except

for the segmentation of the endopods, which are

2-segmented rather than 3-segmented (Figure

12c-/). In these endopods the two distal segments

fused, in legs 2-4 the original line of separation

indicated by a row of small spines. Endopod of

leg 4 (Figure 12/) 62 long, the three terminal elements from outer to inner 20, 31, and 44.

Leg 5 (Figure \2g) with the free segment 39 X

13, its two unequal terminal elements 39 and 62.

Leg 6 (Figure 12/J) a posteroventral flap on the

genital segment bearing two setae 33 and 44.

Spermatophore unknown.

Color as in the female.

ETYMOLOGY.—The specific name halodeimatis is

the genitive form of the generic name of the host.

COMPARISON WITH C. bulbosum.—In Chauliolobion halodeimatis the inner coxal setae of legs 1-3

are not swollen proximally as they are in C. bulbosum, and the terminal armature of the endopod

of leg 4 is 1,2 rather than 11,1. In the female the

inner element on the free segment of leg 5 is long

and setiform, while in C. bulbosum it is stout and

spiniform. In the male the concave margin of the

claw of the maxilliped is smooth rather than provided with hyaline teeth as in the species from

Actinopyga.

Scambicornus Heegaard, 1944

Scambicornus tylotus, new species

FIGURES 13-16

TYPE MATERIAL.—62 $ $, 72 $ $ from one

holothurian, Bohadschia argus (Jaeger), in 2 m,

western side of Isle Maitre, near Noumea, New

Caledonia, 22°20'05" S, 166°24'05"E, 11 June 1971.

Holotype $, allotype, and 110 paratypes (50 2 $,

60 $ $) deposited in the National Museum of

Natural History, Smithsonian Institution, Washington, D.C.; the remaining paratypes in the collection of the author.

OTHER SPECIMENS.—7 $ $, 5 $ $ from two

Bohadschia argus, in 4 m, west of Isle Ngou, near

Noumea, 22°13'44"S, 166°23'01"E, 3 August 1971.

Both lots of specimens occurred in company with

Scambicornus tuberatus (Humes and Cressey, 1961).

FEMALE.—Body (Figure 13a) with the prosome

moderately broad. Length 1.19 mm (1.12-1.24 mm)

and the greatest width 0.69 mm (0.65-0.73 mm),

based on 10 specimens. Segment of leg 1 separated

from the head by a dorsal furrow. Epimeral areas

of the segment of leg 1 angular, those of the segments of legs 2-4 rounded. Tergum of the segment

of leg 4 hidden in dorsal view under that of leg

3. Ratio of the length to the width of the prosome

1.18:1. Ratio of the length of the prosome to that

of the urosome 2.18:1.

Segment of leg 5 (Figure 136) 78 X 211. Between

this segment and the genital segment no ventral

sclerite. Genital segment 122 X 172, wider than

long, with its anterior three-fourths laterally expanded and the posterior fourth abruptly narrowed

(91 wide). Genital areas located dorsolaterally just
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Scambicornus tylotus, new species, female: a, dorsal (i); b, urosome, dorsal (c); c,

genital area, dorsal (D); d, caudal ramus, dorsal (D); e, rostral area, ventral (F); /, first antenna,

dorsal (c); g, second antenna, anterior (H); h, third and fourth segments of second antenna,

posterior (H); i, labrum with paragnaths indicated by broken lines, ventral (H).
NUMBER 202 21

behind the middle of the segment. Each area (Figure 13c) bearing two naked setae approximately

19 long and a small spiniform process. Three postgenital segments from anterior to posterior 42 X

76, 34 X 65, and 52 X 62. Anal segment with a

row of minute spinules on its posteroventral margin on each side.

Caudal ramus (Figure 13d) elongated, 73 X 29

in greatest dimensions, about 2.5 times longer than

wide. Outer lateral seta (situated dorsally on the

ramus) 83 and naked. Dorsal seta 90 and weakly

haired distally. Outermost terminal seta 96 and

innermost terminal seta 112, both naked. Two

long median terminal setae 308 (outer) and 610

(inner), both with lateral spinules in their midregions and both inserted between dorsal (smooth)

and ventral (with a marginal row of minute spinules) flanges.

Body surface with a few small hairs (sensilla)

as in Figure 13a.

Egg sac (Figure 13a) elongated oval, 594 X 198,

with many eggs, each about 52 in diameter.

Rostral area (Figure \Se) not well defined posteroventrally, but bearing numerous retractile points.

First antenna (Figure 13/) 443 long. Lengths of

its seven segments: 34 (70 along the anterior edge),

159, 27, 78, 47, 36, and 26 respectively. Small sclerite on the ventral surface of the third segment suggesting an intercalary segment. Formula for the

armature: 4, 13 (5 + 8), 6, 3, 4 + 1 aesthete, 2 +

1 aesthete, and 7 + 1 aesthete. Setae naked except

for four with lateral spinules on the last segment.

Second antenna (Figure 13g) 4-segmented, 270

long including the claw. Armature: 1,1,3 + claw,

and 7. All setae naked. Small fourth segment much

shorter than the claw, which is 78 along its axis.

Posterior surface of the third and fourth segments

ornamented with a row of small spines (Figure

Labrum (Figure 13/) with two rather slender

posteroventral lobes.

Mandible (Figure 14a) similar to that in other

species, for example, Scambicornus lobulatus

Humes, 1967. Paragnath (Figure 13t) a small lobe

with a few inconspicuous spinules. First maxilla

(Figure 146) with three terminal elements and a

subterminal hyaline process, also similar to that in

S. lobulatus and other species in the genus. Second

maxilla (Figure 14c) 2-segmented, with the large

first segment unarmed. Second segment having a

proximal setule on its outer (ventral) margin, a

naked seta on its posterior surface and an inner

(dorsal) seta unilaterally barbed; segment terminating in a short lash with two large proximal teeth

followed by smaller spines, the third and fourth

spines in the series being larger than the rest (Figure 14d). Maxilliped (Figure 14e) 3-segmented, the

first segment unarmed, the second segment with

two short naked setae and a patch of small spines,

and the third segment with two small naked setae

and having a pointed tip.

Area between the maxillipeds and the first pair

of legs (Figure 14/) not protuberant.

Legs 1-4 (Figures I4g,h; 15a,&) with 3-segmented

rami. Armature as follows (the Roman numerals

indicating spines, the Arabic numerals setae):

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,4

enp 0-1; 0-1; 1,5

P2 coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,5

enp 0-1; 0-2; 1,11,3

Ps coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,13

enp 0-1; 0-2; 1,1115

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 11,1,5

enp 0-1; 0-1; 1,11,11

Inner seta on the coxa of legs 1 and 2 long and

plumose, but in leg 3 shorter, and in leg 4 reduced

to a small naked seta 16 long. Outer seta on the

basis of legs 1 and 2 shorter than the first exopod

segment. This seta in legs 3 and 4 much longer

(130), exceeding slightly the length of the entire

exopod. Inner margin of the basis with a row of

long slender spinules in legs 1-3, but smooth in

leg 4. In the exopod of leg 1 the most proximal

outer spine on the third segment distinctly more

slender than the other exopod spines (Figure 14g).

Endopod of leg 4 (Figure Ibb) 143 long, with both

first and second segments having a short spinel ike

inner element instead of a true seta. Five spines on

the third segment from outer to inner 26, 30, 95,

49, and 29.

Leg 5 (Figure 15c) with a relatively large free

segment 65 X 31, ratio 2.1:1. Outer terminal seta

96 with short lateral spinules. Inner terminal seta

47, swollen proximally, and with short lateral

barbules. Distal rim of the segment ornamented

with a few small spinules. Seta on the body near

the insertion of the free segment 68 with short

lateral spinules.

Leg 6 probably represented by the two setae on

the genital area (Figure 13c).
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Scambicornus tylotus, new species, female: a, mandible, anterior (D); b, first maxilla,

anterior (D); C, second maxilla, posterior (B); d, tip of second maxilla, anterior (c); e, maxilliped, antero-inner (B); /, area between maxillipeds and leg 1, ventral (c); g, leg 1 and intercoxal

plate, anterior (H); h, leg 2, anterior (H).
NUMBER 202 23

a

FIGURE 15.—Scambicornus tylolus, new species. Female: a, leg 3, anterior (H); b, leg 4 and

intercoxal plate, anterior (H); C, leg 5, dorsal (B). Male: d, dorsal (i); e, urosome, dorsal (G);

/, second antenna, anterior (H).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Color in life in transmitted light translucent to

yellowish, the eye red, the egg sacs light gray.

MALE.—Body (Figure 15d) resembling in general form that of the female, though the prosome

is narrower. Length 1.03 mm (0.96-1.11 mm) and

the greatest width 0.51 mm (0.50-0.55 mm), based

on 10 specimens. Ratio of the length to the width

of the prosome 1.30:1. Ratio of the length of the

prosome to that of the urosome 1.98:1.

Segment of leg 5 (Figure 15<?) 55 X 133. Between

this segment and the genital segment no ventral

sclerite. Genital segment quadrate, 160 X 160.

Four postgenital segments from anterior to posterior 36 X 70, 36 X 65, 31 X 62, and 43 X 64.

Caudal ramus (Figure 15e) resembling that of

the female, but shorter, 53 X 25, or 2.12 times

longer than wide.

Body surface ornamented with hairs and retractile points as in the female.

Rostral area and first antenna like those in the

female.

Second antenna (Figure 15/) with two groups of

small bosses on the second segment and a row of

several similar bosses on the third segment; otherwise the second antenna resembling that of the

female.

Labrum, mandible, paragnath, first maxilla, and

second maxilla like those in the female. Maxilliped

(Figure 16a) 4-segmented, assuming that the proximal part of the claw represents a fourth segment.

First segment unarmed. Second segment with two

short naked inner setae surrounded by a patch of

large stout blunt spines. Small third segment unarmed. Claw 222 along its axis, bearing two unequal naked setae proximally and a small terminal

lamella. Concave margin of the claw undulating,

with three knoblike processes, the claw distinctly

narrowed between the distal two of these processes

(Figure 16&).

FICURE 16.—Scambicornus tylotus, new species, male: a, maxilliped, posterior (H); b, claw of

maxilliped, anterior (H); C, endopod of leg 1, anterior (H); d, endopod of leg 2, anterior (H);

e, leg 5, dorsal (D); /, leg 6, ventral (H).
NUMBER 202 25

Area between the maxillipeds and the first pair

of legs resembling that of the female.

Legs 1-4 like those of the female except for the

endopods of legs 1 and 2. Endopod of leg 1 (Figure 16c) 2-segmented and relatively longer (130)

than the exopod (99). (In the female these two

rami are: endopod 135, exopod 117.) Endopod of

leg 2 (Figure 16d) also 2-segmented and relatively

longer (146) than the exopod (122). (In the female

these two rami are: endopod 151, exopod 133.)

Second segment of the endopod showing a slight

indication of subdivision.

Leg 5 (Figure \6e) with the unornamented free

segment 39 X 13, three times longer than wide.

Outer terminal seta 81, inner terminal spiniform

seta 70. Seta on the body near the free segment 44.

Leg 6 (Figure 16/) a posteroventral flap on the

genital segment bearing two naked setae 50 and 75.

Spermatophore not observed.

Color as in the female.

ETYMOLOGY.—The specific name tylotus, from

the Greek TUXOOTOI; (= knobbed), refers to the knoblike processes on the claw of the male maxilliped.

COMPARISON WITH RELATED SPECIES.—Humes

and Stock (1973) listed 21 species in the genus

Scambicornus. These fall in three groups based on

the armature of segments 3 and 4 of the second

antenna (Stock, 1964). Scambicornus tylotus may

be distinguished from Stock's group b, containing

S. propinquus (Nicholls, 1944) and S. armoricanus

(Uocquet, Stock, and Kleeton, 1963), where segment 3 has a moderately prehensile element and

where segment 4 is normally developed. The new

species differs from Stock's group c, comprising S.

finmarchicus (T. Scott, 1903), S. tenuicaudis (Sars,

1918), and S. brevicauda (Sewell, 1949), where prehensile elements on segment 3 are absent or setiform and where segment 4 is normally developed.

The remaining 16 species, representing Stock's

group a, have as in the new species a strongly prehensile and usually 2-segmented claw on segment 3

of the second antenna. Two of these, S. poculiferus

(Humes and Cressey, 1961) and S. nicobaricus

(Sewell, 1949), have an elongated fourth second

antennal segment (longer than the third segment).

In 4 of the remaining 14 species, 5. idoneus (Humes

and Cressey, 1961), S. petiti (Stock and Kleeton,

1963), S. prehensilis (Sars, 1918), and S. serendibicus

(Thompson and A. Scott, 1903), the female caudal

ramus is more than four times longer than wide;

and in 3 species, S. campanulipes (Humes and

Cressey, 1961), S. subtilis (Humes and Cressey,

1961), and S. robustus (Thompson and A. Scott,

1903), the ratio is only 2:1 or less.

The 7 species that remain show certain important differences from S. tylotus. In S. modestus

(Humes and Cressey, 1961), S. subgrandis (Humes

and Cressey, 1961), S. adduensis (Sewell, 1949), and

S. lobulatus Humes, 1967, the female genital segment has laterally expanded wings. In S. hamatus

(Heegaard, 1944) the second antenna is stouter and

more robust than in the new species, and the female caudal ramus is about 3:1. In S. tuberatus

(Humes and Cressey, 1961) the female leg 5 is much

smaller both in absolute size and in relation to the

genital segment than in S. tylotus. The final species,

S. brachysetosus Reddiah, 1968, may be distinguished from the new species on the basis of its

small size alone ( $ 0.6 mm, $ 0.4 mm).

The 3-knobbed claw of the male maxilliped in

S. tylotus serves as a useful distinguishing feature.

Among those 15 species where males are known, 13

have maxillipedal claws with a smooth concave

surface. 5. tuberatus has a single knoblike protuberance on the proximal part of the claw. S.

brachysetosus has a spiniform projection near the

middle of the concave side of the claw (Figure 24k).

Undissected specimens of S. tylotus and S. tuberatus in lactic acid may be quickly separated, the

females by the relative size of leg 5 and the males

by the condition of the claw of the maxilliped.

Scambicornus sewelli, new species

FIGURES 17-20

TYPE MATERIAL.—161 9 9 , 204 $ $ from 50

holothurians, Halodeima edulis (Lesson), in 3 m,

near reef between Isle Ndie" and Mt. Kumuru,

northwest of Noumea, New Caledonia, 22°13'40"S,

166°24'18"E, 9 July 1971. Holotype 9, allotype.

and 240 paratypes (100 9 9 , 140 $ $ ) deposited in

the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; 117 paratypes (57 9 9 , 60 $ $ ) in the Zoologisch Museum,

Amsterdam; the remaining paratypes (dissected)

in the collection of the author.

OTHER SPECIMENS.—From Halodeima edulis: 4

9 9 , 1 $ from 1 host, in intertidal pool, Ricaudy

Reef, Noumea, 22°19'00"S, 166°26'44"E, 9 June
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

d

FIGURE 17.—Scambicornus sewelli, new species, female: a, dorsal (i); b, urosome, dorsal (c);

c, genital area, dorsal (B); d, genital area, lateral (B); e, caudal ramus, dorsal (B); /, rostral area,

ventral (F); g, first antenna, dorsal (c); h, second antenna, posterior (H); i, labrum with

paragnaths indicated by broken lines, ventral (H).
NUMBER 202 27

1971; 3 $2, 5 $ $ from 1 host, in 2 m, western

side of Isle Maitre, near Noumea, 22°20'05"S,

166°24'05"E, 11 June; 7 $2, 1 $ from 6 hosts,

in 1 m, western side of Isle Maitre, 20 June; 5 $ $

from 1 host, in intertidal pool, southwestern side

of Port Ngea, 2 kms north of Ricaudy Reef,

22°18'18"S, 166°26'47"E, 8 July.

From Halodeima coluber (Semper): 3 2 2, 4

$ $ from 1 host, intertidal, southwestern side of

Port Ngea, 2 kms north of Ricaudy Reef, 8 July

1971; 22 2 2, 57 $ $ from 6 hosts, intertidal,

eastern side of Isle Maitre, near Noumea,

22°20'35"S, 166°25'10"E, 16 July; 14 2 2 from 6

hosts, intertidal, eastern side of Isle Maitre, 31

July.

From Halodeima atra (Jaeger): 11 2 2, 5 $ $

from 8 hosts, in 1.5 m, Rocher a la Voile, 22° 18'24"S,

166°25'50"E, 2 June 1971; 56 2 2, 18 $ $ from

10 hosts, intertidal, Ricaudy Reef, 9 June; 137

2 2 , 65 $ $ from 31 hosts, in 4-5 m, reef between

Isle Ndi£ and Mt. Kumuru, northwest of Noumea,

22°13'40"S, 166°24'18"E, 29 July.

From Holothuria curiosa Ludwig: 16 2 2 , 10

$ $ from 1 host, in 1 m, Ricaudy Reef, 9 June

1971; 19 2 2 , 20 $ $ from 1 host, intertidal, eastern side of Isle Maitre, 16 July; 14 2 2, 9 $ $

from 1 host, intertidal pool, eastern side of Isle

Maitre, 16 July.

From Actinopyga echinites (Jaeger): 4 2 2 from

2 hosts, intertidal, Ricaudy Reef, near Noumea,

9 June 1971.

From Microthele nobilis (Selenka): 1 2 from 2

hosts, in 2 m, west of Isle Mando, 22°18'59"S,

166°09'30"E, 3 July 1971.

In all but one of these collections the new species

was accompanied by S. subtilis (Humes and Cressey,

1961).

FEMALE.—Body (Figure 17a) with the prosome

a little less widened than in S. tylotus. Length 1.14

mm (1.07-1.20 mm) and the greatest width 0.60

mm (0.57-0.63 mm), based on 10 specimens. Epimera of the segments of legs 1 and 2 angular, those

of the segments of legs 3 and 4 rounded. Ratio of

the length to the width of the prosome 1.33:1.

Ratio of the length of the prosome to that of the

urosome 2.21:1.

Segment of leg 5 (Figure \7b) 91 X 156. Between

this segment and the genital segment no ventral

sclerite. Genital segment 94 in midlength, 140 in

greatest length, and 180 wide, posterolaterally expanded to the level of the second postgenital segment. Genital areas situated laterally in the posterior half of the segment. Each area (Figure 17c,d)

with two unequal naked setae 50 and 21, and a

small spiniform process. Margin of the segment

medial to the genital area with a small inwardly

turned flange. Three postgenital segments from

anterior to posterior 39 X 75, 34 X 70, and 55 X

69. Anal segment with a row of small spinules on

its posteroventral margin on each side.

Caudal ramus (Figure 17e) elongated, 68 X 31

in greatest dimensions, about 2.19 times longer

than wide. Outer lateral seta (situated dorsally on

the ramus) 300 and naked. Dorsal seta 210 and

naked. Outermost terminal seta 220 and naked,

innermost terminal seta 150 with a few proximal

inner spinules. Two long median terminal setae

540 (outer) and 870 (inner), both with a few delicate lateral spinules (easily broken off) in their

midregions and both inserted between small dorsal

(smooth) and ventral (with a marginal row of

small spinules) flanges.

Body surface with a few small hairs (sensilla) as

in Figure 17a.

Egg sac (Figure 17a) elongated oval, 570 X 250,

with numerous eggs, each about 60 in diameter.

Rostral area (Figure 17/) not well defined.

First antenna (Figure I7g) 380 long. Lengths of

its seven segments: 24 (57 along the anterior edge),

130, 34, 66, 41, 29, and 23 respectively. Formula for

the armature as in S. tylotus. All setae naked.

Second antenna (Figure \7h) 286 long. Segmentation and armature as in S. tylotus. All setae

naked. Claw 80 along its axis. Third and fourth

segments without fine ornamentation.

Labrum (Figure 17/) similar to that in S. tylotus.

Mandible (Figure 18a) with a small conical process preceding the serrations on the convex margin

and with unusually long proximal spinules on the

concave margin; bifurcation at the tip unequal.

Paragnath (Figure 17/) a small hairy lobe. First

maxilla (Figure 18b) resembling that of S. tylotus

but with a spiniform process adjacent to the longest terminal seta. Second maxilla (Figure ISr)

armed as in S. tylotus. Second segment with the

posterior surficial seta naked, the inner (dorsal)

seta with prominent unilateral barbs (Figure \$d).

and the second and third teeth on the lash longer

than the rest. Maxilliped (Figure 18c) similar to
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 18.—Scambicornus sewelli, new species, female: a, mandible, posterior (D); b, first maxilla,

anterior (D); C, second maxilla, anterior (B); d, spine on second segment of second maxilla, posterior (E); e, maxilliped, antero-inner (B); /, area between maxillipeds and leg 1, ventral (G);

g, area between maxillipeds and leg 1, lateral (c); h, leg 1 and intercoxal plate, anterior (H);

i, leg 2, anterior (H); ;', third segment of endopod of leg 3, anterior (H).
NUMBER 202 29

FIGURE 19.—Scambicornus sewelli, new species. Female: a, leg 4 and intercoxal plate, anterior

(H); b, leg 5, dorsal (B). Male: c, dorsal (F); d, urosome, dorsal (H); e, segments 1-3 of first

antenna, dorsal (H); /, second antenna, anterior (B); g, maxilliped, posterior (B); h, claw of

maxilliped, posterior (B); i, tip of claw of maxilliped, anterior (D).
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

that in S. tylotus. Tip of the third segment recurved.

Area between the maxillipeds and the first pair

of legs (Figure 18/) slightly protuberant in lateral

view (Figure 18g).

Legs 1—4 (Figures ISh-j; 19a) segmented and

armed as in S. tylotus. Leg 4 with the inner coxal

seta 11 and naked. Outer seta on the basis 130.

Exopod 118. Endopod 109, with the inner element

on segments 1 and 2 a plumose seta about 40. Five

spines on the third segment from outer to inner

21, 28, 59, 61, and 31.

Leg 5 (Figure 196) with a small free segment

22 X 16, ratio 1.37:1. Outer terminal seta 42 and

smooth. Inner terminal seta 61, slightly swollen

proximally and angular. Segment ornamented distally with a few small spinules. Seta on the body

near the insertion of the free segment 65 and

naked.

Leg 6 probably represented by the two setae on

the genital area (Figure 17d).

Color in life in transmitted light as in S. tylotus.

MALE.—Body (Figure 19c) with the prosome

more slender than in the female. Length 0.71 mm

(0.66-0.75 mm) and the greatest width 0.31 mm

(0.30-0.32 mm), based on 10 specimens. Ratio of

the length to the width of the prosome 1.59:1.

Ratio of the length of the prosome to that of the

urosome 1.89:1.

Segment of leg 5 (Figure \9d) 31 X 76. No ventral intersegmental sclerite. Genital segment 109

X 117, slightly wider than long. In dorsal view

the greatest width of the segment occurring in its

posterior third. Four postgenital segments from

anterior to posterior 26 X 49, 29 X 48, 26 X 47,

and 23 X 45.

Caudal ramus (Figure 19d) similar to that of

the female, but shorter, 29 X 21, or 1.38 times

longer than wide.

Fir.iRF. 20.—Scambicornus sewelli, new species, male: a, leg 1, anterior (B); b, terminal spine on

endopod of leg 1, anterior (c); c, endopod of leg 2, anterior (B); d, endopod of leg 3, anterior

(B); e, leg 5, dorsal (D); /, leg 6, ventral (B); g, spermatophore, ventral (B).
NUMBER 202 31

Body surface ornamented with hairs as in the

female.

Rostral area like that of the female.

First antenna 250 long, resembling that of the

female but the second segment relatively a little

shorter (Figure \9e). Lengths of the segments: 20

(39 along the anterior edge), 81, 21, 46, 26, 19, and

18 respectively.

Second antenna (Figure 19/) with two groups of

small spinules on the second segment and a few

spinules on the third segment; otherwise as in the

female.

Labrum, mandible, paragnath, first maxilla, and

second maxilla like those in the female. Maxilliped

(Figure 19g) with a somewhat enlarged first segment. Second segment more slender with two inner

naked setae and ornamented with a patch of

slender spines and a distal group of minute spinules. Small third segment unarmed. Claw (Figure

\9h) 145 along its axis (including the hyaline

terminal lamella), bearing proximally two unequal

setae, the longer with minute barbules along one

side, the other naked. Convex margin of the claw

with the sclerotization interrupted at five points

where, in a favorable view, the margin is seen to

protrude on a slight knob. Near the tip of the

claw on the anterior surface a row of minute

spinules (Figure 19i).

Legs 1-4 segmented as in S. tylotus, with the

spine and setal formula as in that species. Leg

1 (Figure 20a) with the exopod 65 and the 2-

segmented endopod 82 long. Terminal spiniform

element on the endopod with lateral hairs proximally but with a barbed fringe distally (Figure

20&). Endopod of leg 2 (Figure 20c) also 2-

segmented and relatively longer (99) than the

exopod (73). Legs 3 and 4 with the outer seta on

the basis much shorter than in the female, only

about half as long as the exopod. Endopod of leg

3 (Figure 20d) longer (104) than the exopod (87).

Outer terminal spine on the third segment with

an attenuated rather than a blunt tip and deflected

slightly toward the more proximal spine on the

outer side of the segment. Leg 4 as in the female.

Leg 5 (Figure 20e) with a minute free segment

10 X 7.5 armed as in the female.

Leg 6 (Figure 20/) a posteroventral flap on the

genital segment bearing two naked setae about 25,

a small spiniform process, and a ventral row of

slender spinules.

Spermatophore (Figure 20g) oval, 78 X 44, not

including the neck.

Color as in S. tylotus.

ETYMOLOGY.—This species is named for the late

Colonel R. B. Seymour Sewell, who described several species of Scambicornus (under the name Preherrmannella) from the Indian Ocean.

COMPARISON WITH RELATED SPECIES.—On the

grounds mentioned above in connection with S.

tylotus, the new species from Halodeima may be

separated from S. propinquus, S. armoricanus, S.

finmarchicus, S. tenuicauda, S. brevicauda, S. poculiferus, S. nicobaricus, S. idoneus, S. petiti, S. prehensilis, and S. serendibicus.

Eight of the eleven other species in the genus

may be distinguished from S. sewelli by selected

characters as follows: In S. campanulipes the female has a bell-shaped fifth leg. In S. subtilis, S.

modestus, S. subgrandis, and S. lobulatus the female genital segment has laterally expanded wings.

In S. hamatus the second antenna is stouter and

more robust and the female caudal ramus is about

3:1. In S. tuberatus leg 5 in the female is much

smaller (28 X 18) and the claw of the male maxilliped has a proximal inner digitiform process. In

S. tylotus the female leg 5 is much larger, both

alsolutely and in relation to the genital segment,

and the claw of the male maxilliped has three

inner knoblike processes.

The three species that remain are less well

known and therefore more difficult to compare

with the new species. In S. robustus, of which only

the female is known, the genital segment has evenly

rounded sides instead of being posterolaterally expanded. In S. brachysetosus both sexes are much

smaller than in S. sewelli, the female genital segment has evenly rounded sides, and the concave

side of the claw of the male maxilliped has a

spiniform projection.

The two females and two males described as

Preherrmannella adduensis by Sewell (1949) may

comprise two species. His females represent a

species close to 5. subtilis, but distinguished from

that species by the longer fourth segment on the

second antenna and by the longer caudal ramus

(nearly 3:1, instead of 1.67:1 as in S. subtilis).

Sewell's males show an irregular convex margin on

the claw of the maxilliped (his fig. 21 n1) suggesting

the unusual nature of the maxillipedal claw of

S. sexoelli. Although it is impossible to be certain
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

from Sewell's meager description and figures, the

possibility exists that Sewell's males of Preherrmannella adduensis may actually represent Scambicornus sewelli.

Examination of type material of Preherrmannella adduensis has not been possible. The specimens on which the original description was based,

collected by Sewell while on board the Investigator

in the Indian Ocean, no longer exist. Colonel

Sewell informed me by letter of 26 April 1963 that

the specimens from the Investigator were deposited

in the collections of the Zoological Survey of India.

During World War II the collections of the Indian

Museum (containing the specimens from the Investigator) were transferred to Benares. There they

were exposed to a severe flood, with the result that

the labels were washed off.

Scambicornus calcaratus, new species

FICURES 21-23

TYPE MATERIAL.—179 9 $, 120 $ $, and 43

copepodids from five holothurians, Actinopyga

miliaris (Quoy and Gaimard), in 3 m, reef between

Isle Ndie" and Mt. Kumuru, northwest of Noumea,

New Caledonia, 22°13'40"S, 166°24'18"E, 9 July

1971. Holotype $ , allotype, and 291 para types (175

9 9, 116 $ $) deposited in the National Museum

of Natural History, Smithsonian Institution, Washington, D.C.; the remaining para types (dissected)

and the copepodids in the collection of the author.

OTHER SPECIMENS.—From Actinopyga miliaris:

72 9 9 , 23 $ $ , and 3 copepodids from one host,

in 2 m, west of Isle Mando, near Noumea,

22°18'59"S, 166°09'30"E, 17 July 1971.

From Actinopyga obesa palauensis Panning: 11

9 9, 7 $ <J from one host, in 3 m, north of Isle

Maitre, near Noumea, 22°19'30"S, 166°24'35"E, 13

July 1971.

From Actinopyga plebeja (Selenka): 6 9 9, 4

$ $ from one host, in 2 m, northwestern side of

Isle Maitre, 22°20'05"S, 166°24'05"E, 11 June 1971;

9 9 9, 1 S from one host, in 2 m, north of Isle

Maitre, 13 July; 4 9 9, 3 $ $ from one host, in 2

m, west of Isle Mando, near Noumea, 15 July.

From Actinopyga serratidens Pearson: 10 9 9 ,

5 $ $ from 3 hosts, in intertidal pool, Ricaudy

Reef, 22°19'00"S, 166°26'44"E, 10 July 1971.

From Actinopyga echinites (Jaeger): 23 9 9 , 14

$ $ , and 43 copepodids from one host, in 2 m,

Rocher a la Voile, Noumea, 2 June 1971; 2 9 9

from two hosts, intertidal, Ricaudy Reef, near

Noumea, 9 June; 28 9 9, 7 $ $ , and 1 copepodid

from 12 hosts, in 2 m, Rocher a la Voile, Noumea,

10 June; 2 9 9, 5 $ $ from 22 hosts, in 2 m,

eastern end of Ricaudy Reef, near Noumea,

22°19'00"S, 166°27'18"E, 18 June; 22 9 9, 15 $ $

from two hosts, in 2 m, north of Isle Maitre, near

Noumea, 22°19'30"S, 166°24'35"E, 13 July.

From Actinopyga lecanora (Jaeger): 66 9 9 , 36

$ $ from one host, in 0.5 m, eastern side of Isle

Maitre, 22°20'05"S, 166°24'05"E, 8 June 1971; 52

9 9, 24 s $ from one host, in 2 m, north of Isle

Maitre, near Noumea, 13 July; 2 9 9 from one

host, in tide pool, beach west of Paita, northwest

of Noumea, 22°07'10"S, 166°12'00"E, 22 July; 41

9 9, 18 $ $ from one host, in tide pool, eastern

side of Isle Maitre, 31 July.

FEMALE.—Body (Figure 21a) with a moderately

broad prosome. Length 1.30 mm (1.21-1.39 mm)

and the greatest width 0.62 mm (0.58-0.64 mm),

based on 10 specimens. Tergum of leg 3 longer

than that of leg 2. Epimeral areas varied as illustrated in the figure. Ratio of the length to the

width of the prosome 1.37:1. Ratio of the length

of the prosome to that of the urosome 1.84:1.

Segment of leg 5 (Figure 21 b) 130 X 185. Between this segment and the genital segment no

ventral sclerite. Genital segment 122 X 143, slightly

wider than long and in dorsal view only slightly

expanded laterally. Genital areas situated dorsolaterally in the posterior half of the segment. Each

area (Figure 21c) bearing two naked setae about

12 long and a small spiniform process. Three postgenital segments from anterior to posterior 57 X

76, 52 X 74, and 62 X 70. Anal segment with a row

of extremely small spinules on the posteroventral

margin on each side.

Caudal ramus (Figure 21d) elongated, 81 X 29

in greatest dimensions, 2.79 times longer than wide.

Outer lateral seta (situated dorsolaterally on the

ramus) 104 and smooth. Dorsal seta 140 and lightly

feathered distally. Outermost terminal seta 73 and

the innermost terminal seta 83, both smooth. Two

long median terminal setae 275 (outer) and 638

(inner), both with sparse lateral spinules and both

inserted between small smooth dorsal and ventral

flanges.
NUMBER 202 33

FIGURE 21.—Scambicornus calcaratus, new species, female: a, dorsal (i); b, urosome, dorsal (A);

c, genital area, dorsal (D); d, caudal ramus, dorsal (B); e, first antenna, ventral (o); /, second

antenna, anterior (H); g, labrum with paragnaths indicated by broken lines, ventral (H); h,

mandible, anterior (D); i, first maxilla, posterior (D); /, second maxilla, posterior (B).
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 22.—Scambicornus calcaratus, new species, Female: a, maxilliped, antero-inner (B); b,

area between maxillipcds and leg 1, ventral (c); c, leg 1 and intercoxal plate, anterior (H); d,

leg 2, anterior (H); e, third segment of endopod of leg 3, anterior (H); /, leg 4 and intercoxal

plate, anterior (H); g, leg 5, dorsal (B). Male: h, dorsal (i).
NUMBER 202 35

Body surface with a few minute hairs (sensilla)

as in Figure 21a.

Egg sac (Figure 21a) elongated, 800 X 280, with

numerous eggs, each about 83 in diameter.

Rostral area as in S. sewelli.

First antenna (Figure 21e) 385 long. Lengths

of its seven segments: 39 (60 along the anterior

edge), 143, 23, 68, 39, 26, and 26 respectively.

Formula for the armature as in 5. tylotus and S.

sewelli. Setae naked except for two on the last

segment which are feathered.

Second antenna (Figure 21/) 4-segmented, 300

long including the claw. Armature as in the two

preceding species. Claw 83 along its axis. Third

segment with a few small inner spinules, but the

fourth segment without fine ornamentation.

Labrum (Figure 21g) with two rather elongated

posteroventral lobes.

Mandible (Figure 21 h) differing from other

species only in minor details. Paragnath (Figure

21g) a small lobe with a few hairlike spinules. First

maxilla (Figure 21t) in general form resembling

the two preceding species. Second maxilla (Figure

21;") armed basically as in S. tylotus and S. sewelli.

Lash with two large proximal teeth and only 3 or

4 smaller distal spines. Maxilliped (Figure 22a)

similar to that of S. tylotus, but the second segment

without fine ornamentation.

Area between the maxillipeds and the first pair

of legs (Figure 22&) very slightly protuberant.

Legs 1-4 (Figure 22c-/) segmented and armed

as in the two previous species. Leg 1 with the second and third outer spines on the third exopod

segment setiform and unusually elongated, the second spine 50 long and the third 105; exopod 105.

Outer seta on the basis of legs 3 and 4 not greatly

elongated, in the third leg 96 long and the exopod

148, in the fourth leg 99 and the exopod 138. Leg

4 with the inner coxal seta 16 and naked. Endopod

112, with the inner element on segments 1 and 2

a slender spine 27 long. Five spines on the third

segment from outer to inner 31, 38, 68, 47, and 30.

Leg 5 (Figure 22g) with a small unornamented

free segment 29 X 18, ratio 1.61:1. Two terminal

setae 83 and 78, both finely barbed. Seta on the

body near the insertion of the free segment 73 and

lightly feathered.

Leg 6 probably represented by the two setae on

the genital area (Figure 21c).

Color in life in transmitted light similar to that

of die two preceding species.

MALE.—Body (Figure 22h) with a slender prosome. Length 1.13 mm (1.21-1.39 mm) and the

greatest width 0.62 mm (0.58-0.64 mm), based on 10

specimens. Ratio of the length to the width of the

prosome 1.57:1. Ratio of the length of the prosome

to that of the urosome 1.60:1.

Segment of leg 5 (Figure 23a) 39 X 117. No ventral intersegmental sclerite. Genital segment 174

X 178. Four postgenital segments from anterior to

posterior 36 X 70, 44 X 70, 39 X 69, and 55 X 65.

Caudal ramus (Figure 23a) resembling that of

the female, but shorter, 62 X 29, or 2.14 times

longer than wide.

Body surface ornamented with small hairs as in

the female.

Rostral area and first antenna like those of the

female.

Second antenna (Figure 236) similar to that of

the female, but two groups of small bosses added

on the inner surface of the second segment.

Labrum, mandible, paragnath, first maxilla, and

second maxilla like those of the female. Maxilliped

(Figure 23c) long and slender. First segment unarmed. Second segment with two short naked inner

setae, a large postero-inner patch of spines, and

many small bosses along its outer surface. Small

third segment obscure and unarmed. Claw 213

along its axis (including the minute terminal

lamella) with a weak line of subdivision about midway and bearing proximally two very unequal

setae.

Legs 1-3 segmented as in S. tylotus and S. sewelli,

with the spine and setal formula as in those species.

Leg 1 (Figure 23d) with the second and third outer

spines on the third exopod segment elongated, 47

and 73 respectively; exopod 112. Endopod 156

long. Third segment of the exopod of leg 2 (Figure

23e) with the third outer spine and the terminal

spine 45 and 90 respectively (in the female these

spines 34 and 52). Endopod of leg 2 (Figure 23/)

143 long. Seta on the basis of legs 3 and 4 relatively

short, 65 long in leg 4 where the endopod is 130.

Leg 4 exopod as in the female. Leg 4 endopod

(Figure 23g) unusual in being 2-segmented, the

second and third segments fused, with their original junction indicated only by breaks in the sclerotization.

Leg 5 (Figure 23A) with a small slender unorna-
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23.—Scambicornus calcaratus, new species. Male: a, urosome, dorsal (A); b, second antenna,

anterior (H); C, maxilliped, posterior (H); d, leg 1, anterior (H); e, third segment of endopod of

leg 2, anterior (H); /, endopod of leg 2, anterior (H); g, endopod of leg 4, anterior (H); h, leg

5, dorsal (B); i, leg 6, ventral (c); /', spermatophore, ventral (c).
NUMBER 202 37

mented free segment 44 X 13. Two terminal setae

unequal, 68 and 127. Seta on the body near the

insertion of the free segment 47 long.

Leg 6 (Figure 23i) a posteroventral flap on the

genital segment bearing two naked setae 50 and

65, and a small spiniform process.

Spermatophore (Figure 23;) oval, 155 X 107,

not including the short neck.

Color as in the previous species.

ETYMOLOGY.—The specific name calcaratus, derived from the Latin calcar (a spur), alludes to the

two long spurlike elements on the third segment

of the exopod of leg 1.

New Records and Hosts

(New hosts are preceded by an asterisk)

Scambicornus campanulipes

(Humes and Cressey, 1961)

SPECIMENS COLLECTED.—From *Actinopyga piebe ja (Selenka): 1 9 from one host, in 2 m,

northwestern side of Isle Maitre, near Noumea,

22°20'O5"S, 166°24'05"E, 11 June 1971.

From Actinopyga lecanora (Jaeger): 1 9 from

one host, in 2 m, north of Isle Maitre, 22°19/30"S,

166°24'35"E, 13 July 1971.

This species has been recorded from Madagascar

(Humes and Stock, 1973) and Mauritius (Humes,

1975).

New Caledonia, 22°18'00"S, 167°02'00"E, 6 August

1971; 4 $ $ from two hosts, in 2 m, west of Isle

Mando, near Noumea, 22°18'59"S, 166°09/30"E,

3 July.

This species is known from Madagascar (Humes

and Stock, 1973) and Mauritius (Humes, 1975).

Scambicornus modestus

(Humes and Cressey, 1961)

SPECIMENS COLLECTED.—From Stichopus chloronotus (Brandt): 8 9 $, 11 $ $, and 9 copepodids plus 3 pairs in amplexus from 8 hosts, in

2 m, Rocher a la Voile, Noumea, 22°18'24"S,

166°25'50"E, 2 June 1971; 7 $9, 9 $ $ , and 4

copepodids plus 6 pairs in amplexus from 8 hosts,

in 1-2 m, Rocher a la Voile, 3 June; 54 9 9, 28

$ $ , and 11 copepodids plus 18 pairs in amplexus

from 15 hosts, in 2 m, Rocher a la Voile, 10 June.

From Stichopus variegatus Semper: 3 $ $ from

1 host, in 4 m, reef between Isle Ndie and

Mt. Kumuru, northwest of Noumea, 22°13'24"S,

166°24'11"E, 29 July 1971; 1 9, 1 & , and 3 copepodids from 1 host, in 1 m, eastern side of Isle

Maitre, near Noumea, 22°20'35"S, 166O25'45"E( 31

July; 1 9 plus 1 pair in amplexus from 2 hosts, in

2 m, west of Isle Ngou, near Noumea, 22°13'44"S,

166°23'0l"E, 3 August.

Scambicornus modestus has thus far been known

only from Madagascar (Humes and Stock, 1973).

Scambicornus idoneus

(Humes and Cressey, 1961)

SPECIMENS COLLECTED.—From Holothuria leucospilota (Brandt). 115 9 9 , 107 $ $, and 32 copepodids from 5 hosts, in 1.5-2 m, Rocher a la Voile,

Noumea, 22°18'24"S, 166°25'50"E, 2 June 1971.

From Halodeima atra (Jaeger): 18 9 9 , 13 $ S

plus 9 pairs in amplexus from 8 hosts, in 1.5 m,

Rocher a la Voile, 2 June 1971; 16 9 9 , 16 $ $

plus 1 pair in amplexus from 10 hosts, intertidal,

Ricaudy Reef, 9 June; 3 9 9, 4 $ $ from 31 hosts,

in 4-5 m, reef between Isle Ndie and Mt. Kumuru,

northwest of Noumea, 29 July.

From Actinopyga echinites (Jaeger): 1 9 , 1 <?

from 12 hosts, in 2 m, Rocher a la Voile, Noumea,

10 June 1971.

From Microthele nobilis (Selenka): 23 9 9, 13

$ $ from two hosts, in 0.5 m, Goro, southeastern

Scambicornus poculiferus

(Humes and Cressey, 1961)

SPECIMENS COLLECTED.—From Synapta maculata

(Chamisso and Eysenhardt): 3 9 9, 2 $ $ from 1

host, in 10 cm, reef 5 kms south of Yate, southeastern New Caledonia, 22°irOO"S, 166°59'00"E,

23 June 1971; 29 9 9 , 25 $ $ , and 23 copepodids

from 2 hosts, in 0.5 m, 1 km offshore from Goro

village, southeastern New Caledonia, 22°18'00"S,

167°02'00"E, 6 August.

This species is known only from Madagascar

(Humes and Stock, 1973).

Scambicornus sub til is

(Humes and Cressey, 1961)

SPECIMENS COLLECTED.—From Halodeima edulis

(Lesson): 4 9 9 , 9 <$• £ from 50 hosts, in 3 m, near
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

reef between Isle Ndie" and Mt. Kumuru, northwest

of Noumea, 22°13'40"S, 166°24'18"E, 9 July 1971;

3 $5, 6 $ $ from 1 host, intertidal, Ricaudy

Reef, Noumea, 22°19'00"S, 166°26'44"E, 9 June;

6 $ $ from 2 hosts, in 2 m, Rocher a la Voile,

Noumea, 22°18'24"S, 166°25'50"E, 10 June; 2 9 $,

13 $ $ from 1 host, in 2 m, western side of Isle

Maitre, near Noumea, 22°20'05"S, 166°24'05"E, 11

June; 5 $ $ from 6 hosts, in 1 m, western side of

Isle Maitre, near Noumea, 20 June; 1 $, 2 $ $

from 1 host, intertidal, southwestern side of Port

Ngea, 2 kms north of Ricaudy Reef, near Noumea,

22°18'18"S, 166°26'47"E, 8 July.

From *Halodeima coluber (Semper): 1 9, 2

$ $ from 6 hosts, intertidal, eastern side of Isle

Maitre, 22°20'35"S, 166°25'45"E, 16 July 1971; 2

2 9 from 6 hosts, intertidal, eastern side of Isle

Maitre, 31 July.

From *Halodeima atra (Jaeger): 12 9 9, 3 $ $

from 8 hosts, in 1.5 m, Rocher a la Voile, Noumea,

2 July 1971; 8 9 $, 6 $ $ from 10 hosts, intertidal, Ricaudy Reef, 9 June; 1 9,2$$ from 31

hosts, in 4-5 m, reef between Isle Ndie" and Mt.

Kumuru, northwest of Noumea, 29 July.

From Holothuria curiosa Ludwig: 2 2 $ from

1 host, in 0.5 m, Ricaudy Reef, 5 June 1971; 5 9 $ ,

14 $ $ plus 1 pair in amplexus from one host, in

1 m, Ricaudy Reef, 9 June; 4 9 9,3 $ $ from 1

host, intertidal, eastern end of Isle Maitre, near

Noumea, 16 July; 4 9 9 from 1 host, intertidal

pool, eastern side of Isle Maitre, 16 July.

From *Actinopyga echinites (Jaeger): 2 9 9

from 2 hosts, intertidal, Ricaudy Reef, 9 June 1971.

From *Microthele nobilis (Selenka): 2 9 9, 5

$ $ from two hosts, in 0.5 m, Goro, southeastern

New Caledonia, 22°18'00"S, 167°02'00"E, 6 August

1971; 2 9 9 from 2 hosts, in 2 m, west of Isle

Mando, near Noumea, 22°18'59"S, 166°09'30"E, 3

JulyUntil now S. subtilis has been known only from

Madagascar (Humes and Stock, 1973).

Scambicornus tuberatus

(Humes and Cressey, 1961)

SPECIMENS COLLECTED.—From Bohadschia vitiensis (Semper): 12 9 9,4 $ $ from 1 host, in 2-3

m, Anse Vata, Noumea, 22°18'27"S, 166°26'30"E,

4 June 1971; 44 9 9 , 35 $ $, and 2 copepodids

from 1 host, Anse Vata, 7 June.

From *Bohadschia argus (Jaeger); 23 9 2 , 85

$ $ from 1 host, in 2 m, western side of Isle

Maitre, near Noumea, 22°20'05"S, 166°24'05"E, 11

June 1971; 52 9 9, 76 $ $ from 2 hosts, in 4 m,

west of Isle Ngou, near Noumea, 22°13'44"S,

166°23'01"E, 3 August.

This species has previously been known only

from Madagascar (Humes and Stock, 1973).

Notes on Scambicornus brachysetosus

Reddiah, 196S

FIGURE 24

The description and figures of this small copepod

found on Holothuria (= Halodeima) atra Jaeger

at Kilakarai (Ramnad District, Madras State, India)

lack certain details essential for the correct recognition of the species. The following notes are based

on two paratypes, one female and one male, in the

collection of the Zoological Survey of India, Calcutta, generously sent to me by Shri G. Ramakrishna.

FEMALE.—Length 0.63 mm, greatest width 0.30

mm. Caudal ramus (Figure 24a) 38 X 18, 2.1 times

longer than wide. First antenna 159 long. Second

antenna (Figure 24b) 101 long, the claw about 22

along its axis. Labrum, paragnath, and first maxilla

as in Figure 24c. Mandible, second maxilla, and

maxilliped as in Figure 24d,e,f respectively.

Endopod of leg 4 (Figure 24g) 64 long, with both

first and second segments having a spiniform element instead of a true seta. Five spines on the third

segment from outer to inner 16.5, 28.5, 59, 36, and

17.5.

Leg 5 (Figure 24a) with the free segment 27.5

X 14, ornamented distally with a few minute spinules. Outer terminal seta slender and 42 long, inner

terminal element stout and spiniform, 45 long;

both elements smooth. Dorsal seta 30 long.

MALE.—Length 0.43 mm, greatest width 0.20

mm. Caudal ramus (Figure 24/j) smaller than in

the female, 19 X 13, 1.46 times longer than wide.

Second antenna (Figure 24i) showing sexual dimorphism in the presence of rows of small spines

on the second and third segments. Maxilliped (Figure 24;) with the concave surface of the claw finely

denticulated and having a distinct proximally directed spiniform process near the midregion (Figure 24A). Claw 70 along its axis.
NUMBER 202 39

FIGURE 24.—Scambicornus brachysetosus Reddiah, 1968, paratypes. Female: a, urosome, dorsal

(H); b, second antenna, posterior (D); C, labrum, paragnaths, and first maxillae, ventral (c); d,

mandible, posterior (c); e, second maxilla, posterior (D); /, maxilliped, postero-outer (D); g,

endopod of leg 4, anterior (D). Male: h, urosome, ventral (H); i, second antenna, anterior (D);

/, maxilliped, posterior (c); k, claw of maxilliped, anterior (c); /, endopod of leg 1, anterior

(D); m, endopod of leg 2, anterior (D).
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Endopod of leg 1 (Figure 24/) 2-segmented and

only a little shorter (45) than the exopod (50).

Endopod of leg 2 (Figure 24m) also 2-segmented

and similarly a little shorter (46) than the exopod

(50).

Leg 5 (Figure 24h) with the free segment 22 X

7.5.

List of New Caledonian Holothurians

and Their Associated Copepods

Actinopyga echinites (Jaeger)

Chauliolobion bulbosum, new genus, new species

Scambicornus sewelli, new species

Scambicornus calcaratus, new species

Scambicornus subtilis (Humes and Cressey, 1961)

Scambicornus idoneus (Humes and Cressey, 1961)

Actinopyga lecanora (Jaeger)

Scambicornus calcaratus, new species

Scambicornus campanulipes (Humes and Cressey, 1961)

Actinopyga miliaris (Quoy and Gaimard)

Scambicornus calcaratus, new species

Actinopyga obesa palauensis Panning

Chauliolobion bulbosum, new genus, new species

Scambicornus calcaratus, new species

Actinopyga plebeja (Selenka)

Scambicornus calcaratus, new species

Scambicornus campanulipes (Humes and Cressey, 1961)

Actinopyga serratidens Pearson

Scambicornus calcaratus, new species

Bohadschia argus (Jaeger)

Scambicornus tylotus, new species

Scambicornus tuberatus (Humes and Cressey, 1961)

Bohadschia vitiensis (Semper)

Scambicornus tuberatus (Humes and Cressey, 1961)

Halodeima atra (Jaeger)

Chauliolobion halodeimatis, new genus, new species

Scambicornus sewelli, new species

Scambicornus subtilis (Humes and Cressey, 1961)

Scambicornus idoneus (Humes and Cressey, 1961)

Halodeima coluber (Semper)

Scambicornus sewelli, new species

Scambicornus subtilis (Humes and Cressey, 1961)

Halodeima edulis (Lesson)

Scambicornus sewelli, new species

Scambicornus subtilis (Humes and Cressey, 1961)

Holothuria curiosa Ludwig

Scambicornus sewelli, new species

Scambicornus subtilis (Humes and Cressey, 1961)

Holothuria leucospilota (Brandt)

Scambicornus idoneus (Humes and Cressey, 1961)

Microthele nobilis (Selenka)

Nanaspis mixta, new species

Scambicornus sewelli, new species

Scambicornus subtilis (Humes and Cressey, 1961)

Scambicornus idoneus (Humes and Cressey, 1961)

Stichopus chloronotus (Brandt)

Scambicornus modestus (Humes and Cressey, 1961)

Stichopus variegatus Semper

Scambicornus modestus (Humes and Cressey, 1961)

Synapta maculata (Chamisso and Eysenhardt)

Scambicornus poculiferus (Humes and Cressey, 1961)

Literature Cited

Bocquet, C, J. H. Stock, and G. Kleeton

1963. Copepodes parasites d'invertebres des cotes de

France, X: Cyclopoides poecilostomes associes aux

annelides polychetes, dans la region de Roscoff.

Archives de Zoologie Experimental et Generate,

102 (notes et revue, number l):20-40.

Bresciani, J., and J. Liitzen

1962. Parasitic Copepods from the West Coast of Sweden

Including Some New or Little Known Species.

Videnskabelige Meddelelser fra Dansk Naturhistorisk

Forening i K0benhavn, 124:367-408.

Gurney, R.

1927. Zoological Results of the Cambridge Expedition to

the Suez Canal, 1924, XXXIII: Report on the

Crustacea: Copepoda (Littoral and Semi-Parasitic).

Transactions of the Zoological Society of London,

22 (4):451-477.

Heegaard, P.

1944. A New Copepod (Scambicornus hamatus) Parasitic

on a Japanese Holothurian. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i K0benhavn, 107:359-366.

Humes,

1967.

1973.

1975.

Humes,

1959.

1961.

Humes,

1969.

A. G.

A New Species of Scambicornus (Copepoda, Cyclopoida, Lichomolgidae) Associated with a Holothurian in Madagascar, with Notes on Several

Previously Described Species. Beaufortia, 14(173):

135-155.

Nanaspis (Copepoda: Cyclopoida) Parasitic on the

Holothurian Thelenota ananas (Jaeger) at Eniwetok

Atoll. Journal of Parasitology, 59 (2):384-395.

Cyclopoid Copepods Associated with Marine Invertebrates in Mauritius. Zoological Journal of the

Linnean Society of London, 56(3).

A. G., and R. F. Cressey

A New Family and Genus of Cyclopoid Copepods

Parasitic on a Holothurian. Journal of Parasitology,

45:209-216.

Copepodes cyclopoides du Genre Preherrmannella

Parasites d'Holothuries et d'un Oursin a Madagascar. Me'moires de I'Institut Scientifique de Madagascar, 1959, series F, 3:25-65.

A. G., and J.-S. Ho

Cyclopoid Copepods Parasitic in Holothurians in

Madagascar. Journal of Parasitology, 55:877-894.
NUMBER 202 41

Humes,

1973.

Nicholls

1944.

Reddiah

1968.

Sars, G.

1918.

Scott, T.

1903.

A. G., and J. H. Stock

A Revision of the Family Lichomolgidae Kossmann,

1877, Cyclopoid Copepods Mainly Associated with

Marine Invertebrates. Smithsonian Contributions

to Zoology, 127: v + 368.

, A. G.

Littoral Copepods from South Australia (II):

Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida and Caligoida. Records of the South Australian Museum, 8(1): 1-62.

, K.

Scambicornus brachysetosus n. sp. (Copepoda:

Lichomolgidae) from a Holothurian in the Gulf of

Mannar. Journal of the Marine Biological Association of India, 1967, 9 (1):126-131.

O.

An Account of the Crustacea of Norway with Short

Descriptions and Figures of All the Species, volume

VI: Copepoda, Cyclopoida, parts XIII, XIV,

Lichomolgidae (concluded), Oncaeidae, Corycaeidae,

Ergasilidae, Clausiidae, Eunicicolidae, Supplement,

pages 173-225. Bergen.

Notes on Some Copepoda from the Arctic Seas Collected in 1890 by the Rev. Canon A. M. Norman,

F.R.S. In A. M. Norman, Notes on the Natural

History of East Finmark. Annals and Magazine of

Natural History, series 7, 11 (61):4-32.

Sewell, R. B. S.

1949. The Littoral and Semi-Parasitic Cyclopoida, the

Monstrilloida, and Notodelphyoida. In John Murray Expedition, 1933-34, Scientific Reports, 9(2):

17-199.

Stock, J. H.

1964. On Scambicornus Heegaard, 1944, a Senior Synonym of Preherrmannella Seymour Sewell, 1949

(Copepoda, Cyclopoida). Beaufortia, 10(123): 183-

192.

1968. Copepoda Endoparasitic of Tropical Holothurians.

Bulletin Zoologisch Museum, Universiteit van Amsterdam, 1:89-105.

Stock, J. H., and G. Kleeton

1963. Copepodes ectoassocies aux invertebres des c6tes

du Roussillon. Vie et Milieu, 13 (4):681-702.

Stock, J. H., A. G. Humes, and R. U. Gooding

1962. Copepoda Associated with West Indian Invertebrates, I: The Genus Nanaspis (Siphonostoma,

Nanaspidae). In Studies on the Fauna of Curagao

and Other Caribbean Islands, 13:1-20.

Thompson, I. C, and A. Scott

1903. Report on the Copepoda Collected by Professor

Herdman, at Ceylon, in 1902. In Report to the

Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, supplementary reports,

7:227-307.

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