Gyclopoid Copepods (Lichomolgidae)

Associated with Alcyonaceans

in New Caledoni;

ARTHUR G. HUMES

m

wtu.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 191
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 19 1

Cyclopoid Copepods (Lichomolgidae)

Associated with Alcyonaceans

in New Caledonia

Arthur G. Humes

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Humes, Arthur G. Cyclopoid Copepods (Lichomolgidae) Associated with Alcyonaceans in New Caledonia. Smithsonian Contributions to Zoology, number 191,

27 pages, 13 figures, 3 tables, 1975.—Notoxynus mundus, new genus, new species,

Metaxymolgus praelongipes, new species, and Metaxymolgus comparatus, new

species, are described from Xenia membranacea; and Metaxymolgus cincinnatus,

new species, and Metaxymolgus mimicus, new species, from Cladiella pachyclados. The two species of Metaxymolgus from Xenia and the two from Cladiella

represent pairs of species. Fifteen other species are reported also from New Caledonia, ten of these from new hosts. Eleven of these species are recorded for the

first time in the Pacific Ocean. Twenty lichomolgids are now known from soft

corals in New Caledonia.

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 5219. SERIES

COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Humes, Arthur Grovcr, 1916—

Cyclopoid copepods (Lichomolgidae) associated with alcyonaceans in New Caledonia. (Smithsonian contributions to zoology, no. 191)

1. Lichomolgidae. 2. Alcyonaria. 3. Crustacea—New Caledonia. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology no 191

QL1.S54 n. 191 [QL444.C73] 591'.08s [595'.34] 74-20787
Contents

Page

Introduction 1

Notoxynus, new genus . . . 1

N. mundus, new species . 2

Mctaxymolgus praelongipes, new species 7

M. comparatus, new species 11

M. cincinnatus, new species 16

M. mimiciis, new species . 21

New Records and Hosts 25

Literature Cited 27

i n

Cyclopoid Copepods (Lichomolgidae)

Associated with Alcyonaceans

in New Caledonia

Arthur G. Humes

Introduction

In tropical waters cyclopoid copepods are frequently associated with soft corals. Humes and

Stock (1973) listed 37 species of Lichomolgidae

from Alcyonacea in northwestern Madagascar. In

New Caledonia, however, copepods living with

these cnidarian hosts have not been investigated.

In this paper three new copepods are described

from Xenia membranacea Schenk and two from

Cladiella pachyclados (Klunzinger). In addition,

records of fifteen other species are given, ten of

them from new hosts. Eleven of these species are

reported for the first time in the Pacific Ocean.

The fieldwork in New Caledonia during JuneAugust, 1971, and the subsequent study of the copepods were supported by a grant (GB-8381X) from

the National Science Foundation. Mr. Roger C.

Halverson from the University of California at

Santa Barbara aided in making the collections. The

generous aid given by the staff of the Centre

ORSTOM de Noumea is acknowledged with

thanks.

I am indebted to Dr. J. Verseveldt, Zwolle, The

Netherlands, for the identification of the alcyonacean hosts.

All figures have been drawn with the aid of a

camera lucida. The letter after the explanation of

each figure refers to the scale at which it was

drawn. The abbreviations used are: A, = first anArthur C. Humes, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts 02543.

tenna, A2 = second antenna, MXPD = maxilliped,

and Px = 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.

Notoxynus, new genus

DIAGNOSIS.—Lichomolgidae. Urosome in the female 5-segmented, in the male 6-segmented. Caudal

ramus with six setae. Rostrum broadly rounded

posteroventrally. First antenna 7-segmented, in the

female with the formula 4, 13, 6, 3, 4 + 1 aesthete,

2 + 1 aesthete, and 7 + 1 aesthete; in the male 4,

13 + 2 aesthetes, 6, 3 + 1 aesthete, 4 + 1 aesthete,

2 + 1 aesthete, and 7 + 1 aesthete. Second antenna

4-segmeVited, with the armature 1,1,3 and 1 + 6,

there being a single terminal claw.

Labrum with two lobes separated by a median

cleft. Mandible with the base divided by an indentation into proximal and distal parts; convex edge

with a scalelike area bearing a row of small

spinules and leading to a pointed apex. Lash moderately long. Paragnath a small hairy lobe. First

maxilla with three elements and a small thornlike

process. Second maxilla of the usual lichomolgid

type, but the second segment set at an angle to the

first. Maxilliped 3-segmented in the female, 4-

segmented in the male (assuming that the proximal

part of the claw represents a fourth segment).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Comparison of Notoxynus with other lichomolgid genera having

more than two elements on the 2-segmented endopod of leg 4

Genera

Notoxynus, new genus

Ascidioxynus Humes and Stock, 1972

Astericola Rosoll, 1889

Indomolgus Humes and Ho, 1966

Paclabius Kossmann, 1877

Stellicola Kossmann, 1877

Synstellicola Humes and Stock, 1972

Number of

segments in

second antenna

4

4

3

4

3

3

3

Number of claws

on last segment

of second antenna

1

2

2 or 3

2

1

1

1

Formula for

second segment

of leg 4 endopod

11,1

11,1

2,1

11,2; 1,3; 11,3

11,1

11,1 (II in S.

pollex Humes

and Ho, 1967c)

11,1

Legs 1-4 with 3-segmented rami, except for the

endopod of leg 4 which is 2-segmented. Armature

of the usual lichomolgidiform type, but leg 4 endopod 0-1; 11,1. Third segment of leg 4 exopod

11,1,5- No sexual dimorphism in leg 1. Leg 5 with

a small free segment bearing two setae.

Other features as in the species described below.

Associated with alcyonaceans.

TYPE-SPECIES.—Notoxynus mundus, new species.

ETYMOLOGY.—The name is a combination of the

Greek votog ( = the south), alluding to its occurrence in the southern hemisphere, and |wo? (= a

companion).

COMPARISON WITH RELATED GENERA.—Six genera

of the Lichomolgidae have, as in Notoxynus, a

2-segmented endopod in leg 4 with the second segment bearing more than two elements. Notoxynus

may be distinguished from these genera by the

features compared in Table 1.

Notoxynus mundus, new species

FIGURES 1, 2, 3a-h

TYPE MATERIAL.—19 $ $, 21 $ $ from 6 colonies of the alcyonacean Xenia membranacea

Schenk, in 15 cm, on the reef about 5 km south of

Yate, southeastern New Caledonia, 22oH'00"S,

166°59'00"E, 23 June 1971. Holotype $, allotype,

and 28 para types (13 9 $, 15 i $) deposited in

the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (under the

catalog numbers of the old United States National

Museum: USNM); the remaining paratypes in the

collection of the author.

FEMALE.—Body (Figure la) moderately large,

with the prosome neither greatly expanded nor

flattened. Length 2.00 mm (1.76-2.18 mm) and the

greatest width 0.82 mm (0.76-0.86 mm), based on

10 specimens. Segment of leg 1 separated from the

head by a dorsal transverse furrow. Epimera of the

segments of legs 1-3 elongated posteriorly. Segment

of leg 4 with a rectangular dorsum without pronounced epimera. Ratio of the length to the width

of the prosome 1.32:1. Ratio of the length of the

prosome to that of the urosome 1.31:1.

Segment of leg 5 (Figure \b) 130 X 352. Between

this segment and the genital segment no ventral

sclerite. Genital segment elongated, 297 long and

266 in greatest width (in its anterior third). Genital

areas situated laterally near the junction of the

anterior two-thirds of the segment. Each area (Figure \c) with two small naked setae 12 and 6. Three

postgenital segments from anterior to posterior

101 X 156, 57 X 136, and 104 X 150. Posteroventral border of the anal segment smooth.

Caudal ramus (Figure Id) elongated, 200 X 50,

or 4 times longer than wide. Six setae relatively

short, none longer than the ramus, and naked.

Outer lateral seta 78 and placed distally. Dorsal

seta 50 and placed marginally. Outermost terminal

seta 112, innermost terminal seta 130, outer median

terminal seta 170, and inner median terminal seta

180.

Body surface with a few very small hairs (sensilla).

Egg sac (Figure la) oval 440 X 330, containing

about 22 eggs, each approximately 110 in diameter

but of rather irregular shape.

Rostrum (Figure \e) weak and broadly rounded

in ventral view.
FIGURE 1.—Notoxynus mundus, new genus, new species, female: a, dorsal (A); b, urosome,

dorsal (B); c, genital area, dorsal (C); d, caudal ramus, dorsal (D); e, rostrum, ventral (B);

/, first antenna, with arrows indicating positions of three aesthetes added in male, anterodorsal

(D); g, second antenna, anterior (D); /«, labrum, with paragnaths indicated by broken lines,

ventral (D); i, mandible, posterior (C).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Notoxynus mundus, new genus, new species, female: a, first maxilla, anterior (C);

b, second maxilla, posterior (E); c, maxilliped, antero-outer (E); d, area between maxillipeds

and first pair of legs, ventral (B); e, leg 1 and intercoxal plate, anterior (D); f, leg 2, anterior

(D); g, endopod of leg 3, anterior (D); /i, leg 4 and intercoxal plate, anterior (D).
NUMBER 191

First antenna (Figure 1/) 450 long. Lengths of

the seven segments: 5 (115 along the anterior margin), 117, 35, 58, 65, 43, and 18 respectively. Formula for the armature: 4, 13, 6, 3, 4 + 1 aesthete,

2 + 1 aesthete, and 7 + 1 aesthete. All setae naked.

Second antenna (Figure lg) 4-segmented and 375

long, with the formula 1, 1, 3, and 1 + 6. Fourth

segment 88 along the outer side, 41 along the inner

side, and 39 wide. Terminal claw short, 44 along its

axis. Other terminal elements consisting of two

large obtuse setae and four minute setules. All

setae naked.

Labrum (Figure 1/J) with two rounded posteroventral lobes. Mandible (Figure It) with its concave margin having an indentation followed by a

row of slender spinules; with its convex margin

bearing a scalelike area with a row of minute

spinules leading to a pointed apex followed by a

striated serrate fringe. Lash moderately long and

barbed. Paragnath (Figure Ih) a small hairy lobe.

First maxilla (Figure 2a) relatively large, with

three elements and a small thornlike process. Second maxilla (Figure 2b) 2-segmented. First segment

with a group of outer distal setules. Second segment set at an angle to the first and bearing a

conspicuously barbed inner (dorsal) seta and a

surficial posterior barbed seta; the segment terminating in a swollen lash dentate along its outer

(ventral) margin and bilaterally barbed distally.

Maxilliped (Figure 2c) 3-segmented. First segment

unarmed. Second segment with two unequal setae,

the longer one finely barbed. Third segment bearing two unequal setae and terminating in a bilaterally spinulose pointed apex.

Ventral area between the maxillipeds and the

first pair of legs (Figure 2d) not protuberant. Sclerotized line connecting the bases of the maxillipeds.

Legs 1-4 (Figure 2e-h) with 3-segmented rami

except for the endopod of leg 4 which is 2-

segmented. Armature as follows:

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

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

1-0 exp 1-0; 1-1; 111,1,5

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

1-0 exp 1-0; 1-1; 111,1,5

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

P2 coxa 0-1 basis ._...,.._ , ,

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

P3 coxa 0-1 basis ' ' ' " * '-

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

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

enp 0-1; 11,1

Inner element on the coxa of legs 1-3 a large

feathered seta, but in leg 4 reduced to a small

naked seta 20. Inner margin of the basis in leg 1

smooth, but with a row of hairlike spinules in legs

2-4. Exopod of leg 4 with a length of 234. First

segment of the endopod 52 X 54 (not including

the spinitorm processes) and the inner distal plumose seta 117. Second segment 117 long (without

the spiniform processes); greatest width 47 and

least width 26. Two terminal barbed spines 46

(outer) and 99 (inner). Both segments with an

outer row of hairs and the second segment with a

row of inner hairs proximal to the seta.

Leg 5 (Figure 3a,b) with a small unornamented

free segment 33 X 22. Two terminal setae 39 and

25. All setae naked.

Leg 6 represented by the two setae on the genital

area (Figure \c).

Color of living specimens in transmitted light

slightly brownish, the eye red, the egg sacs gray.

MALE.—Body (Figure 3c) resembling in general

form that of the female. Length 1.59 mm (1.50-

1.70 mm) and the greatest width 0.61 mm (0.56-

0.65 mm), based on 10 specimens. Ratio of the

length to the width of the prosome 1.36:1. Ratio

of the length of the prosome to that of the urosome

1.21:1.

Segment of leg 5 (Figure 3d) 65 X 247. No ventral intersegmental sclerite. Four postgenital segments from anterior to posterior 44 X 94, 44 X 91,

29 X 86, and 65 X 96.

Caudal ramus similar to that of the female but

smaller, 161 X 39, ratio 4.13:1.

Body surface ornamented with hairs as in the

female.

First antenna as in the female but three long

aesthetes added, two on the second segment and

one on the fourth segment, so that the formula is

4, 13 + 2 aesthetes, 6, 3 + 1 aesthete, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete. Second

antenna as in the female.

Labrum, mandible, paragnath, first maxilla, and

second maxilla similar to those in the female.

Maxilliped (Figure 3e) 4-segmented (assuming that

the proximal part of the claw represents a fourth

segment). First segment lacking elements but with

an inner distal recurved thornlike process. Second

segment with two naked setae and a group of short

spines. Small third segment unarmed. Claw 250

along its axis (including the small terminal lamella) and without trace of subdivision; proximally
FIGURE 3.—S'otoxymis mutidus, new genus, new species. Female: a, leg 5, dorsal (C): b, free

segment of leg :">, flat outer view (F). Male: c, dorsal (A); d, urosome, dorsal (B); e, maxilliped,

inner (D); /, leg 4, anterior (D); g, free segment of leg 5, flat outer view (F); h, leg 6, ventral

(G). Metaxymolgus praelongipes, new species, female: i, dorsal (H); /, urosome, ventral (B);

k, genital segment, lateral (B); /, genital area, dorsal (E).
NUMBER 191

bearing two unequal setae, the longer one with

minute barbules.

Ventral area between the maxillipeds and the

first pair of legs like that in the female.

Legs 1-4 segmented and armed as in the female.

No sexual dimorphism in leg 1. Inner coxal element of leg 4 a sparsely feathered seta 60 (Figure

3/). Two terminal spines on the endopod of leg 4,

36 (outer) and 84 (inner).

Leg 5 resembling that of the female. Free segment (Figure 3g) 28 X 21, with the two terminal

setae 14 and 33.

Leg 6 (Figure 3/J) a posteroventral flap on the

genital segment bearing two naked setae 39 and 26.

Spermatophore not observed.

Color in living specimens as in the female.

ETYMOLOGY.—The specific name mundus from

Latin (= neat, elegant) recalls the well-proportioned

form of this species.

Metaxymolgus praelongipes, new species

FIGURES 3/-/, 4, 5, Ga-d

TYPE MATERIAL.—15 $ $ , 9 $ $ from 6 colonies

of Xenia membranacea Schenk, in 15 cm, on the

reef about 5 km south of Yate, southeastern New

Caledonia, 22°ll'00"S, 166°59'00"E, 23 June 1971.

(These are the same colonies from which Notoxynus mundus was recovered.) Holotype $ , allotype,

and 17 paratypes (11 $ $ , 6 & $) deposited in the

National Museum of Natural History, Smithsonian

Institution, Washington, D.C. (under the catalog

numbers of the former United States National

Museum: USNM); the remaining paratypes (dissected) in the collection of the author.

FEMALE.—Body (Figure Si) moderately elongated

with the prosome not unusually thickened dorsoventrally. Length 2.14 mm (1.76-2.30 mm) and the

greatest width 0.79 mm (0.67-0.86 mm), based on

10 specimens. First pedigerous segment separated

dorsally from the cephalosome by a transverse suture. Ratio of the length to the width of the prosome 1.56:1. Ratio of the length of the prosome to

that of the urosome 1.35:1.

Segment of leg 5 (Figure 3;) 122 X 363. Between

this segment and the genital segment no ventral

sclerite. Genital segment in dorsal view elongated,

374 long, 286 in greatest width anteriorly and 197

wide posteriorly; in lateral view (Figure 3/c) not

swollen. Ratio of the length to the average width

about 1.55:1. Genital areas located dorsolaterally

in the anterior half of the segment. Each area

(Figure 3/) with two small naked setae 22 and 20

and a spiniform process. Three postgenital segments from anterior to posterior 125 X 159, 70 X

140, and 117 X 148. Anal segment bearing on each

side a posteroventral row of minute spinules.

Caudal ramus (Figure 4a) elongated, 177 in

greatest dorsal length, 187 in greatest ventral

length, and 65 wide. Ratio of dorsal length to

width 2.72:1. Outer lateral seta 160 and naked.

Dorsal seta 44 and naked. Outermost terminal seta

230, innermost terminal seta 340, and the two

median terminal setae 510 (outer) and 780 (inner),

both inserted between dorsal (smooth) and ventral

(with marginal denticles) flanges. All four terminal

setae plumose.

Body surface with a few hairs (sensilla) and refractile points (Figure Si,j).

Egg sac (Figure 3*) elongated oval, 780 X 415

in the specimen drawn, reaching to the middle of

the caudal ramus and containing approximately 25

eggs about 125-140 in diameter though of irregular

shape.

Rostrum (Figure 4b) subtriangular in ventral

view.

First antenna (Figure 4r) 682 long. Lengths of

the seven segments: 60 (130 along the anterior

margin), 169, 42, 143, 112, 81, and 31 respectively.

Formula for the armature: 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete. All the

setae naked.

Second antenna (Figure -id) 429 long and 4-

segmented, with the armature 1, 1, 3, and II + 5.

Fourth segment 82 along the outer side, 46 along

the inner side, and 36 wide. Two terminal claws

very unequal, the larger 46 along its axis, the

smaller 17. Five small subterminal setules.

Labrum (Figure 4e) with two broadly rounded

posteroventral lobes. Mandible (Figure 4/) having

a strongly constricted base. Distal to the constriction on the convex margin a scalelike area with a

row of blunt spinules followed by two small teeth

or none (Figure 4g) and then a serrated fringe: on

the convex margin a row of long slender spinules.

Lash long and barbed. Paragnath (Figure 4e) a

small hairy lobe. First maxilla (Figure 4h) relatively

large; terminally with two slender setae, a stout

hyaline element, and subterminally having a small
1

FIGURE 4.—Metaxymolgus praelongipes, new species, female: a, caudal ramus, dorsal (D); b,

rostrum, ventral (G); c, first antenna, with arrows indicating positions of three aesthetes added

in male, dorsal (G); d, second antenna, postero-inner (D); e, labrum, with paragnaths indicated

by broken lines, ventral (D); /, mandible, posterior (E); g, distal edge of mandible, posterior

(E); h, first maxilla, posterior (E); i, second maxilla, posterior (E); /, maxilliped, postero-inner

(E); k, area between maxillipeds and first pair of legs, ventral (B); /, leg 1 and intercoxal plate,

anterior (D).
NUMBER 191

fingerlike process. Second maxilla (4?) with a large

unarmed first segment. Second segment armed with

a small setule on its proximal outer (ventral) surface, a surficial posterior seta barbed along one

edge, and an inner (dorsal) distal spine bilaterally

with spinules. This segment produced to form a

lash whose proximal half is expanded on the outer

(ventral) side, where it bears strong marginal teeth

grading into small barbules distally. Maxilliped

(Figure 4;) 3-segmented. First segment with an

inner cluster of slender spinules. Second segment

with two unequal finely barbed setae and a similar

cluster of spinules. Third segment with a smooth

spine, a small naked seta, and a stout terminal

bilaterally barbed spiniform process (lacking a distinct articulation with the segment); in addition to

the three elements a small but distinct adjacent

marginal knob.

Area between the maxillipeds and the first pair

of legs (Figure 4k) not protuberant.

Legs 1-4 (Figures 41, ba-c) segmented and armed

as in other species in the genus. Coxa of leg 1 with

an outer prominence on the posterior surface (Figure 4/). Leg 4 (Figure 5c) with the inner coxal seta

50 and naked. Inner margin of the basis smooth.

Exopod 290 long. First segment of the endopod 65

long without the spiniform processes and 68 wide,

its distal inner seta plumose and 100. Second endopod segment 161 long without the processes (172

including the processes), 58 in greatest width, 43

in least width; outer terminal spine 57, inner spine

73 with a bifurcated tip; both spines barbed.

Leg 5 (Figure bd) elongated, reaching to the first

postgenital segment. Free segment in dorsal view

424 long, 95 in greatest width proximally at the

slight expansion. In flat ventral view the free segment as in Figure be. Two naked terminal setae 40

and 117.

Leg 6 represented by the two setae on the genital

area (Figure SI).

Living specimens in transmitted light opaque,

the eye red, the egg sacs grayish to opaque.

MALE.—Body (Figure bf) more slender than in

the female. Length 1.77 mm (1.66-1.86 mm) and

the greatest width 0.54 mm (0.51-0.54 mm), based

on 8 specimens. Ratio of the length to the width

of the prosome 1.73:1. Ratio of the length of the

prosome to that of the virosome 1.07:1.

Segment of leg 5 (Figure bg) 73 X 187. No ventral intersegmental sclerite. Genital segment elongated, 363 X 297. Four postgenital segments from

anterior to posterior 73 X 125, 75 X 110, 44 X 94,

and 84 X 104.

Caudal ramus resembling that of the female, but

smaller, 112 in greatest dorsal length, 122 in greatest ventral length, and 44 wide.

Body surface sparsely ornamented as in the

female.

Rostrum as in the female. First antenna like that

of the female but three long aesthetes added, two

on segment 2 and one on segment 4. Second antenna similar to that of the female except for

numerous short spines on the inner surface of the

first and second segments (Figure bh).

Labrum, mandible, and paragnath like those of

the female. First maxilla (Figure 5i) with the stout

terminal element less enlarged than in the female

and relatively longer. Second maxilla similar to

that in the female. Maxilliped (Figure 5;) with the

first and third segments unarmed. Second segment

ornamented with two rows of spines and bearing

two setae, one smoothly tapered, the other with a

hyaline inset distal portion. Claw 360 along its axis

including the terminal lamella, showing a weak

subdivision about midway, and bearing proximally

two very unequal setae, the larger one with an

obliquely striated tip.

Area between the maxillipeds and the first pair

of legs like that of the female.

Legs 1-4 segmented as in the female and with

the same spine and setal formula as in that sex

except for the third segment of the endopod of leg

1 (Figure 6a) where the formula is 1,1,4 instead of

1,5 as in the female. Rest of leg 1, legs 2 and 3, and

the exopod of leg 4 resembling the female. Second

segment of the endopod of leg 4 (Figure 6b) a

little shorter than in the female, 117 long including

the processes, 43 in greatest width, 32 in least width.

Two terminal spines more unequal than in the

opposite sex, outer 34, inner 52.

Leg 5 (Figure 6c) with the free segment 70 X 18,

the two terminal setae 28 and 17.

Leg 6 (Figure 6d) the usual posteroventral flap

on the genital segment, bearing two naked setae

33 and 24.

Spermatophore not observed.

Color in living specimens like that of the female.

ETYMOLOGY.—The specific name praclongipcs,

from Latin praelongns (= very long) and pea

(= foot), alludes to the elongated fifth leg.
FIGURE b.—Metaxymolgus praelongipes, new species. Female: a, leg 2, anterior (D); b, third

segment of endopod of leg 3, anterior (D); c, leg 4 and intercoxal plate, anterior (D); d, leg 5,

dorsal (B); e, free segment of leg ">. Hat ventral view (B). Male: /, dorsal (H); g, urosome,

dorsal (B); It, second antenna, antero-outer (E); i, first maxilla, posterior (C); /, maxilliped,

inner (D).
NUMBER 191 11

COMPARISON WITH OTHER SPECIES IN THE GENUS.—

The genus Metaxyrnolgns contains 25 species.

Twenty-three of these were listed by Humes and

Stock (1973). Two new species have since been

described: M. echinasteris Humes, in press, from

the asteroid Echinaster luzonicus (Gray) in New

Caledonia, and M. micropus Humes, 1973a, from

the ophiuroid Astroboa nuda (Lyman) in Madagascar.

Metaxymolgus praelongipes, a species whose

large size is noteworthy, may be distinguished from

all other members of the genus by a combination

of three characters: (1) the elongated female genital segment with small anteriorly located expansions in dorsal view, (2) the long free segment of

the female leg 5 (more than twice as long as any

other previously described species) reaching to the

first postgenital segment (in all other species it is

clearly shorter than the genital segment), and (3)

the very unequal claws on the second antenna in

both sexes.

Sexual dimorphism in the first maxilla, evident

in the new species, does not occur is other species

of Metaxymolgus as far as known; however, the

first maxilla in both sexes has not been described

in several species.

Metaxymolgus comparatus, new species

FIGURES 6e-l, 7, 8a-g

TYPE MATERIAL.—14 $ $ , 18 $ $ from 6 colonies of Xenia membranacea Schenk, in 15 cm, on

the reef 5 km south of Yate, southeastern New

Caledonia, 22°ll/00"S, 166°59'00"E, 23 June 1971.

(These are the same colonies from which Notoxynus mundus and Metaxymolgus praelongipes were

recovered.) Holotype 9 , allotype, and 25 paratypes

(10 99 , 15 $ $) deposited in the National Museum of Natural History (under USNM numbers),

Washington; the remaining paratypes (dissected)

in the collection of the author.

Features not mentioned in the description below

may be assumed to be similar to those in the preceding species.

FEMALE.—Body (Figure 6e) resembling that of

M. praelongipes. Length 2.06 mm (1.95-2.24 mm)

and the greatest width 0.74 mm (0.70-0.78 mm),

based on 10 specimens. Ratio of the length to the

width of the prosome 1.60:1. Ratio of the length

of the prosome to that of the urosome 1.32:1.

Segment of leg 5 (Figure 6/) 130 X 330. Between

this segment and the genital segment no ventral

sclerite. Genital segment in dorsal view elongated,

286 long, 242 in greatest width far anteriorly and

180 wide posteriorly; in lateral view shaped as in

Figure 6g. Ratio of the length to the average width

1.35:1. Each genital area (Figure 6h) with two

small naked setae 16 and 10 and a spiniform process. Three postgenital segments from anterior to

posterior 156 X 161, 83 X 146, and 143 X 154.

Caudal ramus (Figure 6i) moderately elongated,

148 in greatest dorsal length, 156 in greatest ventral length, and 70 wide. Ratio of the length (based

on the dorsal length) to the width 2.11:1. Outer

lateral seta 143 and naked. Dorsal seta 33 and

naked. Outermost terminal seta 179, innermost terminal seta 268, and the two median terminal setae

415 (outer) and 670 (inner), both inserted between

dorsal (smooth) and ventral (with prominent marginal denticles) flanges. All four terminal setae

plumose.

Egg sac (Figure 6e) elongated, 179 X 81, containing approximately 22 eggs about 138 in diameter.

Rostrum (Figure 6;) broadly rounded in ventral

view with clusters of minute retractile points.

First antenna (Figure 6k) 670 long. Lengths of

the seven segments: 55 (117 along the anterior

margin), 177, 47, 130, 101, 68, and 39 respectively.

Armature as in M. praelongipes.

Second antenna (Figure 6/) 418 long. Fourth

segment 97 along the outer side, 55 along the inner

side, and 39 wide. Two terminal claws subequal,

43 and 51 along their axes. One of the five subterminal setules large.

Labrum and paragnaths as in Figure la. Mandible and second maxilla identical to those in M.

praelongipes. First maxilla (Figure 7b) with the

finely barbed element more slender than in that

species. Maxilliped (Figure 7c) resembling that of

M. praelongipes but without spinules on the first

segment, the two setae on the second segment much

more unequal in length, and the distal marginal

knob on the third segment larger.

Area between the maxillipeds and the first pair

of legs as in M. praelongipes.

Legs 1-4 resembling those in M. praelongipes

but the third segment of the endopods of legs 1-3

a little shorter (Figure Id-f). Leg 4 (Figure Ig)

with the inner coxal seta 35 and naked. Exopod

253 long. First segment of the endopod 73 without
1

FIGURE 6.—Metaxymolgns praelongipes, new species, male: a, endopod of leg 1, anterior (D);

b, endopod of leg 4, anterior (D); r, leg "J, dorsal (E); d, leg 6, ventral (G). Metaxymolgus

comparatus, new species, female: e, dorsal (A); /, urosome, dorsal (B); g, genital segment, lateral (B); h, genital area, lateral (C); i, caudal ramus, dorsal (D); /, rostrum, ventral (G);

k, first antenna, dorsal (G); /, second antenna, posterior (D).
FIGURE 7.—Metaxymolgus comparatus, new species. Female: a, labrum, with paragnaths indicated by broken lines, ventral (D); b, first maxilla, posterior (E); c, second maxilla, posteroinner (E), d, endopod of leg 1, anterior (D); e, endopod of leg 2, anterior (D); /, third segment

of endopod of leg 3, anterior (D); g, leg 4 and intercoxal plate, anterior (D); h, leg 5, dorsal

(G); i, free segment of leg 5, flat ventral view (G). Male: ;, dorsal (A); k, urosome, dorsal (B);

/, second antenna, posterior (D).
FIGURE 8.—Metaxymolgus comparatus, new species, male: a, maxilliped, inner (D); b, seta on

outer surface of second segment of maxilliped, outer (I); c, endopod of leg 1, anterior (D);

d, endopod of leg 2, anterior (D); e, leg 5, dorsal (E); /, leg 6, ventral (G); g, pair of spermatophores, attached to female, dorsal (B). Metaxymolgus cincinnatus, new species, female:

h, dorsal (A); i, urosome, dorsal (G); ;', genital segment, lateral (G); k, genital area, dorsal

(E); /, caudal ramus, dorsal (E); m, egg sac, dorsal (B); n, rostrum, ventral (G).
NUMBER 191 15

the spiniform processes and 55 wide, its distal inner

plumose seta 115. Second endopod segment 164

including the processes, 44 in greatest width, 31 in

least width. Outer terminal spine 44, inner spine

73.

Leg 5 (Figure Ih) slightly arcuate in dorsal view,

the free segment 297 long and 78 wide at the small

proximal inner expansion. In a flat ventral view

the free segment 352 X 110 and shaped as in Figure li. Two naked terminal setae 153 and 75.

Leg 6 represented by the two setae on the genital

area (Figure 6h).

Living specimens with the same color as in M.

praelongipes.

MALE.—Body (Figure If) shaped as in M. praelongipes. Length 1.75 mm (1.66-1.82 mm) and the

greatest width 0.55 mm (0.51-0.58 mm), based on

10 specimens. Ratio of the length to the width of

the prosome 1.63:1. Ratio of the length of the

prosome to that of the urosome 1.13:1.

Segment of leg 5 (Figure Ik) 75 X 200. No ventral intersegmental sclerite. Genital segment 374 X

374. Four postgenital segments from anterior to

posterior 72 X 127, 83 X 109, 49 X 101, and 104 X

120.

Caudal ramus similar to that of the female, but

smaller, 109 in dorsal length, 117 in ventral length,

and 58 wide.

Rostrum as in the female. First antenna like that

of the female but three long aesthetes added (Figure If), two on segment 2 and one on segment 4.

Second antenna (Figure 7/) like that of the female

except for small inner spines on the first two

segments.

Labrum, mandible, paragnath, first maxilla, and

second maxilla like those of the female. Maxilliped

(Figure Sa) with the claw 200 along its axis. Larger

of the two elements on the base of the claw shorter

than in M. praelongipes but similarly striated. Seta

on the outer surface of the second segment barbed

as in Figure Sb.

Legs 1-4 as in the female except for sexual

dimorphism in the third endopod segment of legs

1 and 2. Formula for the third endopod segment

of leg 1 (Figure 8c) 1,1,4. Three spines on the third

endopod segment of leg 2 (Figure Sd) shorter than

in the female (female 39, 44, and 55; male 26, 34,

and 44 from outer to inner).

Leg 5 (Figure Se) with a slender free segment

75 X 18. Two naked terminal setae 36 and 39.

Leg 6 (Figure 8/) with the two naked setae about

30.

Spermatophore (Figure 8g) attached to the female in pairs, 340 X 160, not including the neck.

Color as in the female.

ETYMOLOGY.—The specific name comparatus

from Latin (= matched or paired) refers to the

many similarities between this species and M.

praelongipes.

COMPARISON WITH RELATED SPECIES.—Metaxymolgus comparatus, along with M. praelongipes, is

larger than any other species in the genus. M. comparatus may be distinguished from all other species

except M. praelongipes by the following features:

(1) the elongated female genital segment with small

anteriorly located lateral expansions in dorsal view,

and (2) the long free segment of the female leg 5

(more than half again as long as in other species).

Several points of distinction between M. comparatus and M. praelongipes are shown in Table 2.

Aside from the differences which establish the

separate identities of the two new species of Metaxymolgus from Xenia, there are several striking similarities. The two species are, in fact, more like each

other in many ways than they are like other species

in the genus and may be regarded as paired species.

TABLE 2.—Chief points of distinction between Metaxymolgus praelongipes and

Metaxymolgus comparatus

Characters

Caudal ramus of $

Claws on second antenna

First maxilla

Spines on third segment of endopod of leg 2

Length of free segment of leg 5 in $

M. praelongipes M. comparatus

177 x 65, ratio 2.72:1 148 x 70, ratio 2.11:1

extremely unequal subequal

sexually dimorphic same in both sexes

similar in both sexes sexually dimorphic

424 (reaching to first 297 (not reaching to

postgenital segment) first postgenital

segment)
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

In the female, M. praelongipes and M. comparatus

have in common: (1) an elongated genital segment

with small anteriorly located lateral expansions in

dorsal view; (2) a similar mandible; (3) the first

maxilla with three terminal elements, one broader

than the other two, and a subterminal fingerlike

process; (4) a similar second maxilla; (5) a small

marginal knob on the third segment of the maxilliped; (6) the inner coxal seta of leg 4 relatively

long and naked; and (7) the free segment of leg 5

relatively long, approaching or reaching the first

postgenital segment and at least one half again as

long as other species in the genus. In the male the

two species have: (1) similar sexual dimorphism in

the second antenna, and (2) an obliquely striated

seta on the proximal part of the claw of the

maxilliped.

Metaxymolgus cincinnatus, new species

FIGURES 8h-n, 9, 10, \la-i

TYPE MATERIAL.—94 $ $, 139 $ $ from one

colony of the alcyonacean Cladiella pachyclados

(Klunzinger), in 0.5 m, Isle aux Serpents, near

Noumea, New Caledonia, 26°16'52"S, 166°25'12"E,

19 July 1971. Holotype $ , allotype, and 225 paratypes (90 9 9, 135 $ <?) deposited in the National

Museum of Natural History (under USNM numbers), Washington; the remaining paratypes (dissected) in the collection of the author.

OTHER SPECIMENS (all from Cladiella pachyclados).—3 $$, 2 $ $ , from one colony, in 2 m,

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

166°25'50"E, 28 June 1971; 19 $ 9, 17 $ $, from

one colony and 4 $ $ from another colony, in 1 m,

west of Isle To N'du, near Noumea, 22°10'42"S,

166°16'30"E, 29 June; 10 $ $ , 37 $ $ from one

colony, in 2 m, west of Isle Ngou, near Noumea,

22°13'44"S, 166°23'01"E, 29 July.

FEMALE.—Body (Figure 8/i) rather slender, with

the prosome not unusually thickened dorsoventrally.

Length 1.46 mm (1.34-1.63 mm) and the greatest

width 0.49 mm (0.47-0.54 mm), based on 10 specimens. Segment of leg 1 separated from the head by

a dorsal transverse furrow. Epimera of the segments

of legs 1-4 variously rounded posteriorly. Ratio of

the length to the width of the prosome 1.70:1. Ratio

of the length of the prosome to that of the urosome

1.23:1.

Segment of leg 5 (Figure Si) 100 X 180, with a

rounded lobe overhanging the proximal inner part

of the free segment of leg 5. Between this segment

and the genital segment no ventral sclerite. Genital

segment elongated, in dorsal view 234 long, 166

wide in the expanded anterior two-thirds and 105

wide posterior to the transverse sclerotization. This

dorsal subdivision of the segment not evident laterally or ventrally (Figure 8;). Genital areas located

dorsolaterally in the posterior half of the expanded

anterior part of the segment. Each area (Figure 8k)

bearing two small naked setae, 8 and 11, with an

adjacent minute spiniform process. Three postgenital segments from anterior to posterior 91 X 91,

65 X 83, and 65 X 83. Posteroventral border of the

anal segment with a row of minute spinules on

both sides.

Caudal ramus (Figure 8/) elongated, 169 X 35,

with the ratio of length to width 4.83:1. Outer lateral seta 86 and naked. Dorsal seta 55 and naked.

Outermost terminal seta 120, innermost terminal

seta 164, and the two median terminal setae 265

(outer) and 375 (inner), both inserted between

dorsal (smooth) and ventral (with minute marginal

spinules) flanges. All four terminal setae plumose.

Body surface with a few small hairs (sensilla).

Egg sac with approximately 15 eggs arranged

loosely as in Figure Sh (size of right sac 506 X

220) or more compactly as in Figure 8m (size 385

X 253). Egg diameter about 117.

Rostrum (Figure Sn) broadly rounded in ventral

view; in lateral view (Figure 9a) slightly projected.

First antenna (Figure 96) 427 long. Lengths of

the seven segments: 35 (67 along the anterior

margin), 118, 31, 68, 65, 47, 31 respectively. Armature as in M. praelongipes. All setae naked.

Second antenna (Figure 9c) 4-segmented and 265

long, with the formula 1,1,3, and II + 5. Fourth

segment 88 along the outer side, 57 along the inner

side and 31 wide. Two terminal claws very unequal,

larger claw 55 along its axis, smaller claw 13. All

setae naked.

Labrum (Figure 9d) with two rounded posteroventral lobes with hyaline margins. Mandible (Figure 9e) with its convex margin having a scalelike

area with 5-6 dentiform spines followed by a serrated fringe; with its concave margin beyond the

indentation bearing a row of slender spinules. Lash

long and barbed. Paragnath (Figure 9d) a small

lobe with slender spinules. First maxilla (Figure
FIGURE 9.—Metaxymolgus cincinnatus, new species, female: a, rostrum, lateral (G); b, first

antenna, dorsal (D); c, second antenna, posterior (E); d, labrum, with paragnaths indicated by

broken lines, ventral (E); e, mandible (F); /, first maxilla, anterior (F); g, second maxilla,

posterior (F); h, maxilliped, posterior (C); i, area between maxillipeds and first pair of legs,

ventral (D).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Metaxymolgus cincinnatus, new species, female: a, leg 1 and intercoxal plate, anterior (E); b, leg 2, anterior (E); c, third segment of endopod of leg 3, anterior (E); d, leg 4

and intercoxal plate, anterior (E); e, leg 5, dorsal (E). Metaxymolgus cincinnatus, new species,

male: /, dorsal (A).
NUMBER 191 19

9/) with four elements. Second maxilla (Figure

9g) 2-segmented. First segment unornamented. Second segment armed with a small setule on its proximal outer (ventral) surface, a surficial posterior

seta finely barbed along one edge, and an inner

(dorsal) distal spine with a row of conspicuous

spinules unilaterally. This segment produced to

form a lash with graded marginal teeth along one

side. Maxilliped (Figure 9h) 3-segmented. First segment unornamented. Second segment with a group

of small slender spinules and two very unequal

setae, the larger seta with two short rows of

spinules. Third segment with a smooth seta, a finely

barbed spine, and a terminal barbed spiniform

process.

Area between the maxillipeds and the first pair

of legs (Figure 9i) only slightly protuberant.

Legs 1-4 (Figure \0a-d) segmented and armed

as in other species in the genus. Coxa of leg 1 with

an outer prominence on its posterior surface (Figure 10a). Inner coxal seta in legs 1-3 with its tip

strongly curled posteriorly. Leg 4 (Figure lOd) with

the inner coxal seta 30, lightly plumose, and not

curled posteriorly as in the preceding legs. Inner

margin of the basis smooth. Exopod 180 long. First

segment of the endopod 44 long, not including the

spiniform processes and 31 wide, its distal inner

plumose seta 75. Second endopod segment 85 long

without the processes, 22 in greatest width, 19 in

least width; outer terminal spine 29, inner spine

60, both barbed. In another female these spines

31 and 69. Outer margins of both endopod segments with a row of short spinules instead of long

hairlike spinules as in legs 1-3.

Leg 5 (Figure lOe) scarcely reaching to the genital areas. Free segment 117 X 44, the width taken

at the rounded proximal inner expansion. Two

smooth terminal elements 30 and 101. Dorsal seta

on the body near the insertion of the free segment

52 and lightly feathered. Free segment ornamented

dorsally and outwardly with small spines.

Leg 6 represented by the two setae on the genital

area (Figure 8k).

Living specimens in transmitted light opaque,

the eye red, the egg sacs gray.

MALE.—Body (Figure 10/) slender. Length 1.25

mm (1.18-1.34 mm) and the greatest width 0.35

mm (0.33-0.37 mm), based on 10 specimens. Ratio

of the length to the width of the prosome 1.88:1.

Ratio of the length of the prosome to that of the

urosome 1.20:1.

Segment of leg 5 (Figure lla) 52 X 107. No ventral intersegmental sclerite. Genital segment elongated 265 X 216. Four postgenital segments from

anterior to posterior 44 X 75, 49 X 70, 36 X 65,

and 40 X 65.

Caudal ramus resembling that of the female, but

smaller, 114 X 30.

Body surface sparsely ornamented as in the female.

Rostrum as in the female. First antenna resembling that of the female but three long aesthetes

added (Figure 10/), two on segment 2 and one on

segment 4. Second antenna (Figure 116) with the

first and second segments bearing inwardly two

rows of short stout spines and a pectinate seta;

otherwise this appendage similar to that in the

female.

Labrum, mandible, paragnath, first maxilla, and

second maxilla like those in the female. Maxilliped

(Figure lie) with the first and third segments unarmed. Second segment bearing two setae, one of

them smooth, the other pectinate along one side,

and two rows of spines. Claw 228 along its axis

including the terminal lamella, weakly divided

about midway, and bearing proximally two very

unequal setae. Larger of these two setae 62 with a

slightly expanded and bilaterally spinulose distal

portion, the ratio of its length to the length of the

claw 1:3.66.

Area between the maxillipeds and the first pair

of legs like that of the female.

Legs 1-4 segmented as in the female and similarly armed except for the third segment of the

endopod of leg 1 (Figure 11 d-e) where the formula

is 1,1,4 instead of 1,5 as in the female. Sexual dimorphism also evident in the third segment of the

endopod of leg 2 (Figure 11/) where, although the

formula is the same as in the female, the terminal

spiniform processes and the fine ornamentation are

distinctive. Leg 3 without sexual dimorphism. Dimensions of the endopod of leg 4 (Figure llg): first

segment 34 long, not including the spiniform processes, and 25 wide, with the distal inner seta 82;

second segment 60 without the processes, 19 in

greatest width, 14 in least width, with the outer

terminal spine 33, the inner spine 69. In other

respects legs 1-4 resembling those of the female.

Leg 5 (Figure \\h) with the free segment 44 X
FIGURE 11.—Metaxymolgus cincinnatus, new species, male: a, urosome, dorsal (G); b, second

antenna, posterior (E); c, maxilliped, outer (E); d, endopod of leg 1, anterior (E); e, third

segment of endopod of leg 1, posterior (C); /, third segment of endopod of leg 2, anterior (C);

g, endopod of leg 5, anterior (E); h, leg 5, dorsal (C); i, leg 6, ventral (D). Metaxymolgus

mimicus, new species, female: ;', dorsal (A); k, urosome, dorsal (G).
NUMBER 191 21

12, without fine ornamentation; the two terminal

setae 35 and 41.

Leg 6 (Figure Hi) the usual posteroventral flap

on the genital segment, bearing two naked setae 34

and 29.

Spermatophore not observed.

Color in living specimens like that of the female.

ETYMOLOGY.—The specific name cincinnatus from

Latin ( = having curled hair) alludes to the curled

inner coxal setae on legs 1-3.

COMPARISON WITH RELATED SPECIES.—Metaxymolgus cincinnatus may be separated from other species

in the genus by two easily recognizable features.

The caudal ramus in the female is relatively longer

(ratio 4.83:1) than in any other species except M.

micropus Humes, 1973a (ratio 4.84:1), an associate

of the basket star Astroboa nuda (Lyman) in Madagascar. The minute leg 5 of M. micropus (11 X 11)

distinguishes it immediately, however, from the

new species.

The extremely unequal claws on the second antenna in both sexes distinguish the new species

from all others in the genus except M. echinasteris

Humes, in press, associated with the sea star Echinaster luzonicus (Gray) in New Caledonia, and

M. praelongipes Humes, described above. In contrast to M. cincinnatus, however, these two species

have in the female an undivided genital segment of

a different configuration.

Although the exact nature of the inner coxal seta

in legs 1-3 is not known in several species, the

curled tip of this seta seen in M. cincinnatus seems

distinctive.

Metaxymolgus mimicus, new species

FICURES 11/-*, 12, 13

TYPE MATERIAL.—15 9 9, 12 $ $ from one colony of Cladiella pachyclados (Klunzinger), in 0.5

m, Isle aux Serpents, near Noumea, New Caledonia,

22°16'52"S, 166°25'12"E, 19 July 1971. (This is the

same colony from which Metaxymolgus cincinnatus

was recovered.) Holotype 9 , allotype, and 19 paratypes (11 $9, 8 $ s) deposited in the National

Museum of Natural History (under USNM numbers), Washington; the remaining para types (dissected) in the collection of the author.

OTHER SPECIMENS (all from Cladiella pachyclados).—2 $ S , 1 copepodid from one colony, in 2 m,

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

166°25'50"E, 28 June 1971; 2 9 9,2 $ $ from one

colony, in 1 m, Isle aux Serpents, west of Pte.

Denouel, near Noumea, 22°16'52//S, 166°25'12"E,

19 July; 2 9 9 , 8 <j ,5 from one colony and 5 9 9,

14 $ $ from another colony, in 1 m, west of Isle

To N'du, near Noumea, 22°10'42"S, 166°16'30"E,

29 June; 4 9 9, 52 $ $ from one colony, in 2 m,

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

166°23'01"E, 29 July.

FEMALE.—Body (Figure 11;) with a slightly

broader prosome than in Af. cincinnatus. Length

1.59 mm (1.50-1.73 mm) and the greatest width

0.61 mm (0.58-0.64 mm), based on 10 specimens.

Ratio of the length to the width of the prosome

1.45:1. Ratio of the length of the prosome to that

of the urosome 1.56:1.

Segment of leg 5 (Figure Ilk) 104 X 238, with a

broad lobe overhanging the base of the free segment of leg 5. Between this segment and the genital

segment no ventral sclerite. Genital segment elongated, in dorsal view 252 long, 200 wide in the

expanded anterior third and 125 wide posterior to

the transverse sclerotization. This dorsal subdivision of the segment absent laterally and ventrally.

Genital areas located dorsolaterally just posterior

to the widest part of the segment. Each area (Figure 12a) with two small naked seta about 11 and a

minute spiniform process. Three postgenital segments from anterior to posterior 73 X 114, 44 X

105, and 44 X 117. Posteroventral border of the

anal segment with a row of very small spinules on

both sides.

Caudal ramus (Figure 126) 130 X 55, with the

ratio of length to width 2.36:1. Outer lateral seta

122 and naked. Dorsal seta 55 and sparsely feathered. Outermost terminal seta 170, innermost terminal seta 225, and the two median terminal setae

390 (outer) and 550 (inner), both inserted between

dorsal (smooth) and ventral (with small marginal

spinules) flanges. All four terminal setae plumose.

Body surface with small hairs (sensilla) as in

Figure 11;.

Egg sac (Figure 11;) elongated oval, 572 X 341,

reaching nearly to the ends of the caudal rami and

containing about 47 eggs approximately 105 in

diameter.

Rostrum (Figure 12c) rounded posteroventrally.

First antenna 507 long and having the same armature as in Af. cincinnatus. Lengths of the seven
FIGURE 12.—Metaxymolgus mimicus, new species, female: a, genital area, dorsal (E); b, caudal

ramus, dorsal (E); c, rostrum, ventral (G); d, second antenna, posterior (E); e, endopod of leg

1, anterior (E); /, third segment of endopod of leg 2, anterior (E); g, leg 4 and intercoxal plate,

anterior (D); It, leg 5, dorsal (D). Metaxymolgus mimicus, new species, male: i, dorsal (A);

/", urosome, dorsal (G).
NUMBER 191

segments: 36 (83 along the anterior margin, 135,

35, 75, 81, 66, and 32 respectively.

Second antenna (Figure I2d) generally similar

to that of M. cincinnatus, but the two terminal

claws unequal, both 54 along their axes. Fourth

segment 99 along the outer side, 63 along the inner

side, and 31 wide.

Labrum, mandible, paragnath, first maxilla, second maxilla, maxilliped, and the ventral area between the maxillipeds and the first pair of legs

resembling those in M. cincinnatus.

Legs 1-4 similar to those of M. cincinnatus except that the inner coxal seta of legs 1-3 has a

straight tip, not curled posteriorly and the exopod

spines of legs 1-3 have less strongly developed

spinous fringes. Endopod of leg 1 (Figure \2.e) with

the outer terminal seta on the third segment

slightly stronger than the other four setae (as in

M. cincinnatus). Third segment of the endopod of

leg 2 as in Figure 12/. Leg 4 (Figure 12g) with a

naked inner coxal seta 35. Exopod 208 long. First

endopod segment 44, not including the spiniform

processes, and 39 wide, with its inner distal plumose

seta 73. Second endopod segment 99 long without

the processes, 34 in greatest width, and 25 in least

width; outer terminal spine 34, inner spine 82,

both barbed. Outer margins of both endopod segments with minute spinules as in M. cincinnatus.

Second endopod segment having a few inner hairs

and, in some specimens, a small outer marginal

notch.

Leg 5 (Figure 12/i) with the free segment 161

long and 70 wide at the basal inner expansion.

Two naked terminal setae 60 and 94. Dorsal seta

on the body near the insertion of the free segment

about 50 and lightly feathered. Free segment ornamented dorsally and outwardly with small spines.

Leg 6 represented by the two setae on the genital

area (Figure 12a).

Living specimens in transmitted light opaque,

the eye red, the egg sacs gray.

MALE.—Body (Figure 12i) slender. Length 1.14

mm (1.08-1.22 mm) and the greatest width 0.37

mm (0.34-0.39 mm), based on 10 specimens. Ratio

of the length to the width of the prosome 1.73:1.

Ratio of the length of the prosome to that of the

urosome 1.30:1.

Segment of leg 5 (Figure 12;) 55 X 130. No ventral intersegmental sclerite. Genital segment longer

than wide, 286 X 242. Four postgenital segments

from anterior to posterior 36 X 78, 34 X 78, 21 X

75, and 26 X 83.

Caudal ramus resembling that of the female, but

smaller, 91 X 43.

Body surface ornamented as in the female.

Rostrum as in the female. First antenna like that

of the female but three long aesthetes added (Figure 12^) as in the male of M. cincinnatus. Second

antenna (Figure 13a) showing sexual dimorphism

similar to that in M. cincinnatus.

Labrum, mandible, paragnath, first maxilla, and

second maxilla like those of the female. Maxilliped

(Figure 136) similar to that of M. cincinnatus, but

the larger of the two setae on the proximal part of

the claw relatively longer, 91, the ratio of its length

to the length of the claw 1:2.62. Length of the claw

234.

Area between the maxillipeds and the first pair

of legs like that of the female.

Legs 1^1 segmented and armed as in the female

except for the third segment of the endopod of

leg 1 (Figure 13c) where the formula is 1,1,4 instead of 1,5 as in the female. A spiniform process

between the two spines. Sexual dimorphism in the

third endopod segment of leg 2 (Figure 13d), the

process between the two terminal spines with a

peculiar digitiform extension. Leg 3 lacking sexual

dimorphism. Dimensions of the endopod of leg 4

(Figure \Se): first segment 32 long, not including

the spiniform processes, with the distal inner seta

55; second segment 64 without the processes, 22 in

greatest width, 17 in least width, with the outer

terminal spine 27, the inner spine 66. Second segment lacking an outer marginal notch.

Leg 5 (Figure 13/) with the free segment 57 X

15, with small outer spinules; the two terminal

naked setae 57 and 20 and the sparsely plumose

dorsal seta near the insertion of the segment about

40.

Leg 6 (Figure 18g) the usual posteroventral flap

on the genital segment, bearing two naked setae

36 and 52.

Spermatophore not observed.

Color in living specimens like that of the female.

ETYMOLOGY.—The specific name mimicus from

Latin (= imitative) refers to the many similarities

of this species to M. cincinnatus.

COMPARISON WITH OTHER SPECIES.—The elongated dorsally divided female genital segment distinguishes Metaxymolgus mimicus from all other
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Metaxymolgus mimicus, new species, male: a, second antenna, posterior; (E); b,

maxilliped, inner (E); c, endopod of leg 1, anterior (E); d, third segment of endopod of leg 2,

anterior (C); e, endopod of leg 4, anterior (E); /, leg 5, dorsal (C); g, leg 6, ventral (D).

species in the genus except M. hirsutipes (T. Scott,

1893) and M. cincinnatus. The new species may be

separated from M. hirsutipes chiefly by details of

the mandible, maxilliped, and leg 5, and from

M. cincinnatus by the features indicated in Table 3.

Although the two species are clearly distinct,

there are many similarities between them. Conspicuous among these are the elongated female

genital segment with a dorsal subdivision, the form

of the mouthparts, the outer terminal seta on the

third segment of the endopod of leg 1 being slightly

stronger than the other, the outer margins of the

two segments of the endopod of leg 4 having short

spinules instead of long hairlike spinules, the free

segment of leg 5 in the female having a prominent

inner proximal expansion, the sexual dimorphism

in the second antenna of the male, the nature of

the large proximal seta on the claw of the maxilliped, and the sexual dimorphism in the endopod

of leg 1.

Metaxymolgus mimicus and M. cincinnatus resemble each other more closely than they do other

species in the genus, and represent a pair of species

associated with a single host species.

PAIRED SPECIES ASSOCIATED WITH A SINGLE HOST

SPECIES.—Several instances are known where pairs
NUMBER 191 25

TABLE 3.—Comparison of certain features of Metaxymolgus mimicus and

M. cincinnatus

Characters

Caudal ramus

Claws on second antenna

Inner coxal seta on legs 1-3

Ratio of length of larger proximal seta on $

maxilliped claw to length of claw

Process between two terminal spines on third

segment of $ leg 2 endopod

M. mimicus M. cincinnatus

$ = 2.36:1

$ =2.12:1

equal size

tip straight

1:2.62

with a digitiform

process

$ =4.83:1

$ = 3.80:1

very unequal size

tip curled

posteriorly

1:3.66

smoothly attenuated

of species of lichomolgid copepods are associated

with one host genus. Paramolgus constrictus (Humes,

1969) and Paramolgus insectus (Humes, 1969) occur

on the antipatharian genus Antipathes. Paramolgus

politus (Humes and Ho, 1967b) and Paramolgus

simulans (Humes and Ho, 1967b) live with the

actiniarian Rhodactis rhodostoma (Ehrenberg).

Plesiomolgus organicus (Humes and Ho, 1967a)

and Plesiomolgus conjunctus (Humes and Ho,

1967a) are associated with the stoloniferan Tubipora musica (Linnaeus). Metaxymolgus praelongipes and Metaxymolgus comparatus, both described

above, occur with the alcyonacean Xenia membranacea Schenk.

The explanation for such pairing of species

would seem to lie in the evolutionary history of the

association between the copepods and the hosts.

Presumably in each of these pairs of copepods an

ancestral copepod associate, confronted by some

kind of barrier inhibiting free genetic exchange

throughout the population, evolved into two forms

now sufficiently distinct to be recognized as separate species but retaining a number of characteristics in common. Such an evolution is suggested

by the interpretation by Bocquet, Stock, and Louise

(1963) of speciation in Asterocheres on regular echinoids in the Mediterranean Sea and on the Atlantic

coast of Europe.

New Records and Hosts

(New hosts are preceded by an asterisk)

Acanthomolgus varirostratus (Humes and Ho, 1968b)

Acanthomolgus exilipes (Humes and Ho, 1968b)

Acanthomolgus gentilis (Humes and Ho, 1968b)

From Dendronephthya mucronata (Putter): many

specimens from one colony, in 1.5 m, Rocher a la

Voile, Noumea, 22°18'24"S, 166°25'50"E, 13 June

1971; many specimens from 5 colonies, in 4 m,

Ricaudy Reef, 22°19'05"S, 166°26'28"E, 28 July.

These three species have been previously known

only from Madagascar (Humes and Stock, 1973).

Anisomolgus insolens (Humes and Ho, 1968a)

From Lobophytum crassum Von Marenzeller:

1 9 , 3 $ s from one colony, intertidal, on reef 5

km south of Yate, southeastern New Caledonia,

22oll'00"S, 166°59'00"E, 23 June 1971.

This species has been recorded from both Madagascar (Humes and Stock, 1973) and Eniwetok

Atoll (Humes, 1973b).

Anisomolgus protentus (Humes and Frost, 1964)

From *Sarcophyton elegans Moser: many specimens from one colony, in 2 m, west of Isle Mando,

near Noumea, 22°18'59"S, 166°09'30"E, 1 July 1971.

This species has until now been reported only

from Madagascar (Humes and Stock, 1973).

Colobomolgus dentipes (Thompson and A. Scott,

1903)

From *Sinularia polydactyla (Ehrenberg): many

specimens including copepodids from one colony,

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

22°18'59"S, 166°09'30"E, 1 July 1971.

This species has previously been reported from
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ceylon (Thompson and A. Scott, 1903) and Madagascar (Humes and Stock, 1973).

Metaxymolgus aculeatus (Humes and Ho, 1968b)

From *Stereonephthya inordinata (TixierDurivault): 36 9 9 , 47 $ $ , from one colony in

30 m, outside Recif Mtere, west of Noumea,

22°20'40"S, 166°13'55"E, 23 July 1971.

This species is known from both Madagascar

Humes and Stock, 1973) and Eniwetok Atoll

(Humes, 1973b).

Metaxymolgus spinulifer (Humes and Frost, 1964)

From Paralemnalia thyrsoides (Ehrenberg): 4

9 9 , 1 $ from one colony, in 3 m, eastern side of

Isle Maitre, near Noumea, 22°20'35"S, 166°25'10"E,

8 June 1971.

From Lemnalia elegans (May): 17 9 $ , 10 $ $

from one colony, in 3 m, north of Isle Maitre,

22°19y30"S, 166°24'35"E, 13 July 1971.

This species has been previously known only

from Madagascar (Humes and Stock, 1973).

Paradoridicola sinulariae Humes and Stock, 1973

From *Sinidaria flexibilis (Quoy and Gaimard):

198 99 , 304 s <$ , 134 copepodids from one colony, in 3 m, west of Isle Mando, near Noumea,

22°18'59"S, 166°09'30"E, 15 July 1971.

This species has previously been known only

from Madagascar (Humes and Stock, 1973).

Paradoridicola squamiger (Humes and Frost, 1964)

Paradoridicola adelphns (Humes and Ho, 1968a)

From Sinularia polydactyla (Ehrenberg): many

specimens from one colony, in 0.5 m, Ricaudy Reef,

22°19'05"S, 166°26'28"E, 5 June 1971; many specimens from one colony, in 2 m, western side of Isle

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

June; few specimens from one colony, in 2 m, eastern side of Ricaudy Reef, Noumea, 22°19'00"S,

166°27'18"E, 18 June; many specimens from one

colony, in 0.5 m, Isle To N'du near Noumea,

22°10'42"S, 166°16'30"E, 29 June; many specimens

from one colony, in 1 m, west of Isle Mando, near

Noumea, 22°18'59"S, lee^^CE , 1 July; few

adults, more than 400 copepodids from one colony,

in 1 m, east of Isle To N'du, near Noumea,

22°10'49"S, 166°17'12"E, 12 July; many specimens

from one colony, in 1 m, west of Isle Mando,

22°18/59"S, 166°09'30"E, 12 July; many specimens

from one colony, in 0.5 m, west of Paita, northwest

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

specimens from one colony, in 20 cm, intertidal,

eastern end of Isle Maitre, 22°2O'35"S, 166°25'10"E,

31 July.

Paradoridicola squamiger has been known only

from Madagascar, but P. adelphus has been reported from both Madagascar (Humes and Stock,

1973) and Eniwetok Atoll (Humes, 1973b).

Paramolgus clavatus (Humes and Ho, 1968c)

From *Stereonephthya inordinata (TixierDurivault): 1 9 from the above-mentioned colony

with which Metaxymolgus aculeatus was associated.

This species has previously been known only

from Madagascar (Humes and Stock, 1973).

Paramolgus eniwetokensis Humes, 1973b

From *Lobophytum crassum Von Marenzeller:

1 9 from the above-mentioned colony with which

Anisomolgus insolens was associated.

This species has been reported only from Eniwetok Atoll (Humes, 1973b).

Paramolgus spathophorus (Humes and Ho, 1968a)

Paramolgus eniwetokensis Humes, 1973b

Anisomolgus insolens (Humes and Ho, 1968a)

From *Lobophytum crebriplicatum Von Marenzeller: many specimens from one colony, in 2 m,

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

166°25'50"E, 28 June 1971; many specimens from

one colony in 2 m, Rocher a la Voile, 19 June;

many specimens (not including A. insolens) from

one colony in 3 m, eastern side of Isle Maitre, near

Noumea, 22°20'35"S, 166°25'45"E, 8 June.

Paramolgus eniwetokensis has been reported from

Eniwetok Atoll (Humes, 1973b), P. spathophorus
NUMBER 191 27

from Madagascar (Humes and Stock, 1973) and A.

insolens from both Madagascar (Humes and Stock,

1973) and Eniwetok Atoll (Humes, 1973b).

Paredromolgus decorus (Humes and Frost, 1964)

From *Cladiella pachyclados (Klunzinger): 22

$ $ , 15 $ $ from one colony, in 0.5 m, Isle aux

Serpents, west of Pointe Denouel, near Noumea,

22°16'52"S, 166°25'12"E, 19 July 1971; 11 9 9,

9 $ $ from one colony, in 2 m, north of Rocher

a la Voile, Noumea, 22°18'24"S, 166°25'50"E, 28

June; 32 9 9 , 26 $ $ from one colony, in 1 m,

Isle aux Serpents, west of Pointe Denouel, near

Noumea, 22°16'52"S, 166O25'12"E, 19 July; 49 9

from one colony and 50 9 9 , 30 $ $ from a second colony in 1 m, west of Isle To N'du, near

Noumea, 22°10'42"S, 166°16'30"E, 29 June.

This species until now has been reported only

from Madagascar (Humes and Stock, 1973).

Literature Cited

Bocquet, C, J. H. Stock, and F. Louise

1963. Copepodes parasites d'invertebres des cotes de

France, XV: Le probleme syst&natique d'Asterocheres violaceus (Claus) et d'Asterocheres minutus

(Claus). Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen-Amsterdam,

series C, 66(l):37-53.

Humes, A. G.

1969. Cyclopoid Copepods Associated with Antipatharian

Coelenterates in Madagascar. Zoologische Mededelingen, 44(1): 1-30.

1973a. Cyclopoid Copepods Associated with the Ophiuroid

Astroboa nuda in Madagascar. Beaufortia, 21(275):

25-35.

1973b. Cyclopoid Copepods (Lichomolgidae) from Octocorals at Eniwetok Atoll. Beaufortia, 21 (282): 135-

151.

MS. Cyclopoid Copepods Associated with Sea Stars in

New Caledonia.

Humes, A. G., and B. W. Frost

1964. New Lichomolgid Copepods (Cyclopoida) Associated with Alcyonarians and Madreporarians in Madagascar. Cahiers ORSTOM, se'rie Oce'anographie, 1963

(serie Nosy Be II): 131-212.

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

1966. New Lichmolgid Copepods (Cyclopoida) from Zoanthid Coelenterates in Madagascar. Cahiers ORSTOM,

se'rie Oceanographie, 4(2):3-47.

1967a. New Lichomolgid Copepods Associated with the

Alcyonarian Coral Tubipora musica (Linnaeus) in

Madagascar. Proceedings of the United States National Museum, 121 (3573):I-24.

1967b. Two New Species of Lichomolgus (Copepoda, Cyclopoida) from an Actiniarian in Madagascar. Cahiers ORSTOM, serie Oceanographie, 5(1):3-21.

1967c. New Species of Stellicola (Copepoda, Cyclopoida)

Associated with Starfishes in Madagascar, with a

Redescription of S. caeruleus (Stebbing, 1900).

Bulletin of the British Museum (Natural History),

Zoology, 15(5):201-255.

1968a. Cyclopoid Copepods of the Genus Lichomolgus

Associated with Octocorals of the Family Alcyoniidae in Madagascar. Proceedings of the Biological

Society of Washington, 81:635-692.

1968b. Cyclopoid Copepods of the Genus Lichomolgus

Associated with Octocorals of the Family Nephtheidae in Madagascar. Proceedings of the United

States National Museum, 125 (3661): 1-41.

1968c. Cyclopoid Copepods of the Genus Lichomolgus

Associated with Octocorals of the Families Xeniidae,

Nidaliidae, and Telestidae in Madagascar. Proceedings of the Biological Society of Washington, 81:

693-750.

Humes, A. G., and J. H. Stock

1972. Preliminary Notes on a Revision of the Lichomolgidae: Cyclopoid Copepods Mainly Associated with

Marine Invertebrates. Bulletin Zoologisch Museum,

Universiteit van Amsterdam, 2 (12): 121—133.

1973. A Revision of the Family Lichomolgidae Kossmann, 1877; Cyclopoid Copepods Mainly Associated

with Marine Invertebrates. Smithsonian Contributions to Zoology, number 127, pages i-v, 1-368.

Kossman, R.

1877. Entomostraca (1. Theil: Lichomolgidae). Pages 1-

24 in Halfte IV of Zoologische Ergebnisse einer im

Auftrage der Koniglichen Academic der Wissenschaften zu Berlin Ausgefuhrten Reise in die Kiistengebiete des Rothen Meeres. 6 plates.

Rosoll, A.

1889. Uber zwei an Echinodermen lebende parasitische

Copepoden: Ascomyzon comatulae und Astericola

clausii. Sitzungsberichte der Mathematischnaturwissenschaftlichen Class der Kaiserlichen Akademie

der Wissenschaften, 97 (1): 188-202.

Scott, T.

1893. Additions to the Fauna of the Firth of Forth, Part

V. Eleventh Annual Report of the Fishery Board

for Scotland, 3:197-219.

Thompson, I. C, and A. Scott

1903. Report on the Copepoda Collected by Professor

Herdman, at Ceylon, in 1902. 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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