Gyclopoid Copepods Associated

with Asteroid Echinoderms in

New Caledonia

ARTHUR G. HUMES

r -2 s

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 217
SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

The emphasis upon publications as a means of diffusing knowledge was expressed

by the first Secretary of the Smithsonian Institution. In his formal plan for the Institution, Joseph Henry articulated a program that included the following statement:

"It is proposed to publish a series of reports, giving an account of the new discoveries

in science, and of the changes made from year to year in all branches of knowledge."

This keynote of basic research has been adhered to over the years in the issuance

of thousands of titles in serial publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with

the following active series:

Smithsonian Annals of Flight

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoology

Smithsonian Studies in History and Technology

In these series, the Institution publishes original articles and monographs dealing

with the research and collections of its several museums and offices and of professional

colleagues at other institutions of learning. These papers report newly acquired facts,

synoptic interpretations of data, or original theory in specialized fields. These publications are distributed by mailing lists to libraries, laboratories, and other interested

institutions and specialists throughout the world. Individual copies may be obtained

from the Smithsonian Institution Press as long as stocks are available.

S. DILLON RIPLEY

Secretary

Smithsonian Institution
SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 21 7

Gyclopoid Gopepods Associated

with Asteroid Echinoderms in

New Caledonia

Arthur G. Humes

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976

Gyclopoid Copepods Associated

with Asteroid Echinoderms in

New Caledonia

Arthur G. Humes

Introduction

For the vast area of the tropical Pacific Ocean

there are only a few records of copepods living in

association with Asteroidea. Kossmann (1877) briefly

described Stellicola semperi, a species that has

remained of uncertain validity, from Ophidiaster

miliaris (actually Linckia miliaris (Linck) according to Monod and Dollfus (1932:142, footnote)) at

Isabela, Philippine Islands. Stebbing (1900) described Linckiomolgus caeruleus (now placed in

the genus Stellicola) from Linckia at Feather Island,

China Straits, eastern end of New Guinea. Humes

and Stock (1973) described Stellicola illgi from

Linckia laevigata (Linnaeus) at Urukthapel, Palau

Islands. They also recorded the presence of Stellicola pollex Humes and Ho, 1967, on Linckia diplax

(Miiller and Troschel), Linckia multiflora (Lamarck), and Linckia guildingi Gray in the Hawaiian

Islands. Humes (1970) described Stellicola acanthasteris (now placed on the genus Synstellicola)

from Acanthaster planci (Linnaeus) at Eniwetok

Atoll, Marshall Islands. Humes and Stock (1973)

reported S. acanthasteris from Acanthaster planci at

Viti Levu, Fiji, and near Noumea, New Caledonia.

Humes (1971) described Astroxynus culcitae from

Culcita novaeguineae Miiller and Troschel at Eniwetok Atoll.

This paper includes reports on the following

copepods from asteroids in New Caledonia:

Arthur G. Humes, Boston University Marine Program, Marine

Biological Laboratory, Woods Hole, Massachusetts, 02543.

STELLICOMITIDAE

1. Astroxynus culcitae Humes, 1971

from Culcita novaeguineae Miiller and Troschel

LlCHOMOLCIDAE

2. Metaxymolgus echinasteris, new species

from Echinaster luzonicus (Gray)

3. Stellicola novaecaledoniae, new species

from Linckia laevigata (Linnaeus) and Linckia guildingi Gray

4. Stellicola oreastriphilus Kossmann, 1877

from Culcita novaeguineae Miiller and Troschel (new

host)

5. Stellicola parvulipes, new species

from Culcita novaeguineae Miiller and Troschel

6. Synstellicola acanthasteris (Humes, 1970)

from Acanthaster planci (Linnaeus)

7. Synstellicola pichoni (Humes and Ho, 1966)

from Protoreaster lincki (de Blainville) (new host)

The collections in New Caledonia made during

June-August 1971, and the subsequent study of the

specimens were supported by a grant (GB—838IX)

from the National Science Foundation. Mr. Roger

C. Halverson from the University of California at

Santa Barbara assisted in the field work. I wish to

express my thanks to the staff of the Centre

ORSTOM de Noumea for their generous assistance.

I am greatly indebted to Miss Maureen E. Downey

of the National Museum of Natural History,

Smithsonian Institution, Washington D.C., for the

identifications of the host echinoderms.

All the 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 following abbreviations are used: Ax = first

antenna, A2 = second antenna, MXPD = maxilliped,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and Pi=leg 1. Specimens deposited in the National

Museum of Natural History, Smithsonian Institution, are listed under the acronym "USNM" (for

the collection numbers of the old United States

National Museum).

Astroxynus culcitae Humes, 1971

FIGURE la

SPECIMENS COLLECTED (all from the cushion star,

Culcita novaeguineae Miiller and Troschel).—101

(both sexes) from one host, Rocher a la Voile, near

Noumea, 22°18'24"S, 166°25'50"E, 13 June 1971;

63 from one host, in 2 m, Rocher a la Voile,

19 June; 24 from one host, in 1 m, east of Isle To

N'du, near Noumea, 22°10'49"S, 166°17'12"E, 12

July; 13 from 5 hosts, in 10 cm, eastern side of

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

16 July; and 5 from 7 hosts, intertidal, eastern end

of Isle Maitre, 22°20'35"S, 166°25'45"E, 31 July.

REMARKS.—These copepods agree in nearly all

features with Astroxynus culcitae Humes, 1971, as

described from Culcita novaeguineae at Eniwetok

Atoll. The only significant difference found is the

relative length of the outer seta on the coxa-basis

of legs 3 and 4. In the New Caledonian specimens

this seta (Figure la) is very long (about 90 ^im),

exceeding slightly the length of the ramus (78 \im),

and easily seen in undissected copepods, while in

the Eniwetok material the seta is short and

inconspicuous.

Culcita novaeguineae is the only known host.

Metaxymolgus echinaateris, new species

FIGURES \b-g, 2a-;, 3a-i, 4a-d

TYPE MATERIAL.—47 $ 9, 44 $ $ from 20

orange-red to brown sea stars, Echinaster luzonicus

(Gray), in a tidal pool, at the public beach west of

Paita, near Noumea, New Caledonia, 22°07'00"S,

166°12'00"E, 22 July 1971. Holotype 9, allotype,

and 82 paratypes (42 9 9, 40 $ $) deposited in

the National Museum of Natural History (under

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

OTHER SPECIMENS (all from Echinaster luzonicus).-AS 9 9, 78 $ $ , and 60 copepodids from 6

hosts, in 0.5 m, Ricaudy Reef, near Noumea,

22°19'00"S, 166°26'44"E, 5 June; 1 9 , 9 $ $ , and 2

copepodids from 5 hosts, in an intertidal pool,

Ricaudy Reef, near Noumea, 25 July; and 16 $ 9,

18 $ $ . and 12 copepodids from 7 hosts, in 2 m,

Rocher a la Voile (Pte. Pontillion), near Noumea,

22°18'24"S, 166°25'50"E, 2 June.

FEMALE.—Body (Figure lfr) with a moderately

broad prosome. Length (without the setae on the

caudal rami) 1.66 mm (1.50-1.77 mm) and the

greatest width 0.99 mm (0.96-0.02 mm), based on

10 specimens in lactic acid. Segment of leg 1 separated from the cephalosome by a dorsal transverse

suture. Epimeral areas of the segments of legs

2 and 3 rounded, those of the segment of leg 4

truncated. Ratio of the length to the width of the

prosome 1.12:1. Ratio of the length of the prosome

to that of the urosome 1.81:1.

Segment of leg 5 (Figure \c) 104 X 200 um.

Between this segment and the genital segment a

short, weak ventral sclerite. Genital segment 208 X

313 jxm, much wider than long, with rounded lateral

margins in dorsal view. Genital areas located

dorsolaterally in the posterior half of the segment.

Each area (Figure Id) bearing two naked setae

28 and 17 \im, and a small spinous process. Three

postgenital segments from anterior to posterior

83 X 117, 60 X 107, and 86 X 107 um. Posteroventral margin of the anal segment smooth.

Caudal ramus (Figure \e) 66 X 48 ^m, with the

ratio of the length to the width 1.38:1. Outer lateral

seta 99 \im and naked. Dorsal seta 50 ^m and

plumose. Outermost terminal seta 100 \im and the

innermost terminal seta 195 um, both with lateral

and ventral spinules. Two long median terminal

setae 330 \im (outer) and 515 um (inner), both with

lateral spinules along their midregions and both

inserted dorsally to a small ventral flange with a

smooth margin.

Body surface bearing small hairs (sensilla) as

indicated in Figure 1&.

Egg sac (Figure 16) elongated oval, 935 X 407

jim, reaching to the tips of the longest ramal setae.

Numerous eggs, each approximately 73 \itn in

diameter.

Rostrum (Figure \f) with an incomplete posteroventral margin.

First antenna (Figure \g) 520 \im long. Lengths

of the seven segments (measured along their posterior nonsetiferous margins): 31 (78 ^m along the

anterior margin), 205, 31, 86, 53, 40, and 23 jim
NUMBER 217

FIGURE 1.—Astroxynus culcitae Humes, 1971, female: a, leg 3 and intercoxal plate, posterior (A).

Metaxymolgus echinasteris, new species, female: b, dorsal (B); c, urosome, dorsal (C); d, genital

area, dorsal (D); e, caudal ramus, dorsal (A); /, rostrum, ventral (£); g, first antenna, dorsal (F).

(Scale A = 0.1 mm, B = 1.0 mm, C = 05 mm, D= 0.05 mm, E= 0.3 mm, F = 02 mm.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

respectively. Formula for the armature: 4, 13, 6, 3,

4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete.

Certain setae on the distal segments with lateral

spinules.

Second antenna (Figure 2a) 380 \un long. Formula for the four segments: 1, 1, 3, and II + 5.

Fourth segment 104 ^m along its outer edge, 75 urn

along its inner edge, and 34 \im wide. Two claws

very unequal, the larger claw stout and recurved,

78 urn along its axis, the smaller claw slender and

almost setiform, 26 \im. All the elements naked.

Labrum (Figure 2b) with two rather slender

divergent posteroventral lobes. Mandible (Figure

2c) having in succession on its convex margin a

scalelike area with a row of spinules, a small tooth,

and a deeply serrated hyaline fringe. Concave margin beyond the indentation with a row of long

slender spinules. Lash long and barbed distally.

Paragnath (Figure 2d) a small hairy lobe. First

maxilla (Figure 2e) bearing four elements. Second

maxilla (Figure 2/) with an unornamented first segment. Elongated second segment with a small setule

on its proximal outer (ventral) surface, a smooth

surficial posterior seta, an inner (dorsal) distal spine

with 4-5 large spines along the distal edge and a

few small barbules along the proximal edge. Segment produced to form a lash, which bears outwardly four teeth of unequal sizes followed by a

row of spinules, and inwardly a row of spinules

near its tip. Maxilliped (Figure 2g) with an

unarmed first segment. Second segment carrying

two very unequal setae, both finely barbed. Third

segment bearing a slender setule and a recurved

spine, both naked, and terminating in a small clawlike process.

Ventral area between the maxillipeds and the

first pair of legs (Figure 2h) not protuberant. A

line connecting the bases of the maxillipeds.

Legs 1-4 (Figures 2i-j, 3a-b) having the same

segmentation and armature as in other species of

Metaxymolgus, for example, M. securiger (Humes,

1964). Two of eleven females examined showing

111,1,5 instead of the usual 11,1,5 on the third segment of the exopod of leg 4. Inner coxal seta in

leg 3 distinctly shorter than in legs 1 and 2; in leg 4

this seta minute (20 \im) and naked. Inner margin

of the basis with a row of hairlike spinules in legs

1-3 but smooth in leg 4. Exopod of leg 4 187 um.

First segment of the endopod 52 X 41 jim (not

including the spiniform processes), and its inner

plumose seta 83 \im. Second segment 88 X 28.5 [Am

(greatest width, and length not including the spiniform processes). Two terminal fringed spines 33 ^im

(outer) and 66 [im (inner), with their tips bifurcated (Figure 3c). Outer margins of both segments

with a row of hairs.

Leg 5 (Figure 3d) with an elongated unornamented free segment 130 [im in length, having a

large proximal inner rounded expansion. Width at

the level of the expansion 55 um, and more distally

24 \im. Two terminal naked setae 65 and 81 \im.

Leg 6 represented by the two setae on the genital

area (Figure Id).

Living specimens in transmitted light pale orange,

the eye red, the egg sacs orange.

MALE.—Body (Figure Se) with a less broadened

prosome than in the female. Length (excluding

the setae on the caudal rami) 1.35 mm (1.31-1.38

mm) and the greatest width 0.62 mm (0.61-0.64

mm), based on 10 specimens in lactic acid. Epimera

of the segment of leg 2 subtruncated and those of

the segment of leg 4 rounded. Ratio of the length

to the width of the prosome 1.29:1. Ratio of the

length of the prosome to that of the urosome

1.19:1.

Segment of leg 5 (Figure 3/) 52 X 133 um. No

ventral intersegmental sclerite. Genital segment

270 X 270 um. Four postgenital segments from

anterior to posterior 44 X 88, 44 X 86, 31 X 81,

and 57 X 86 um.

Caudal ramus resembling that of the female, but

smaller, 47 X 35 \im.

Body surface with small hairs (Figure 3e-/) as

in the female.

Rostrum like that of the female.

First antenna similar to that of the female, but

three long aesthetes added (at points indicated by

arrows in Figure lg), the formula thus becoming:

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

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

(Figure 3g) showing sexual dimorphism. Setae on

the first and second segments, and one of the three

setae on the third segment, more prominent than

in the female and pectinate along the proximal

(inner) side. Inner surface of the second segment

bearing numerous small spinules. Fourth segment

relatively longer than in the female, its outer edge

105 um, its inner edge 66 jan, and its width 24 \im.

Larger claw 62 um and the smaller claw 24 um.

Labrum, paragnath, first maxilla, and second
NUMBER 217

FIGURE 2.—Metaxymolgus echinasteris, new species, female: a, second antenna, postero-inner (F);

b, labrum, ventral (G); c, mandible, posterior (D); d, paragnaths, posteroventral (A); e, first

maxilla, anterior (D); /, second maxilla, posterior (A); g, maxilliped, posterior (A); h, area

between maxillipeds and first pair of legs, ventral (C); i, leg 1 and intercoxal plate, anterior (F);

;, leg 2, anterior (C). (Scale G = 0.1 mm.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

maxilla like those in the female. Maxilliped (Figure

Sh) long and slender. First segment unornamented.

Second segment bearing two setae (one smooth,

the other finely pectinate along the proximal edge),

a long row of spines on the inner surface, and a

cluster of short spines at the distal anterior corner

of the segment. Small third segment unarmed. Claw

218 jim along its axis (including the terminal

lamella), incompletely divided about midway, and

bearing two proximal unequal setae, the longer

seta finely pectinate along one side.

Ventral area between the maxillipeds and the

first pair of legs like that in the female.

Legs 1-4 segmented as in the female and with

the same spine and setal formula except for the

third segment of the endopod of leg 1 (Figure 3i),

which has the formula 1,1,4 instead of 1,5 as in the

female. Outer distal spiniform process on the first

segment of this endopod smaller than in the female.

Legs 2 and 3 like those in the female. Leg 4 resembling that of the female except for slight sexual

dimorphism in the endopod (Figure 4a). Here the

two terminal spines on the second segment measuring 52 um (outer) and 57 um (inner), more nearly

equal in length than in the female. (In the specimen drawn the tip of the inner spine trifurcated,

rather than bifurcated as in all other specimens

seen.) One of the eleven males examined with the

formula 111,1,5 instead of the usual 11,1,5 on the

third segment of the exopod of leg 4.

Leg 5 (Figure 46) having a slender unornamented

free segment 55 X 11 um, with a proximal inner

expansion. Two naked terminal setae 66 and 31

um.

Leg 6 (Figure 4c) consisting of a posteroventral

flap on the genital segment bearing two naked

setae 55 and 74 um, and a minute spiniform

process.

Spermatophore (Figure 4d), attached to the

female, 260 X 125 ^m.

Living specimens paler orange than in the female,

the eye red.

ETYMOLOGY.—The specific name echinasteris is

derived from the generic name of the sea star

serving as host.

COMPARISON WITH RELATED SPECIES.—By the nature of the long free segment of leg 5 in the female,

unornamented and with a large proximal inner

expansion, Metaxymolgus echinasteris may be distinguished from all species assigned to the genus

by Humes and Stock (1973) except possibly M.

securiger (Humes, 1964). The latter differs conspicuously from the new species, however, in its

smaller size (female 1.22 mm), the nearly equal

claws on the second antenna, and the rather globular female genital segment, which is only slightly

wider than long.

The very unequal claws on the second antenna

of M. echinasteris serve to distinguish it from most

of the 23 known species in the genus, in which both

claws are distinctly clawlike and recurved though

one may be much shorter than the other. In two

species, Metaxymolgus gracilipes (A. Scott, 1909)

and M. simplex (Thompson and A. Scott, 1903),

the lesser claw seems to approach the condition in

the new species, judging from the scanty information in the original descriptions. Both of these

species differ, however, in other ways from the new

species. In M. simplex the female is only about

half as long as the New Caledonian form, the genital segment is about as long as wide, and the free

segment of leg 5 lacks a proximal expansion. In

the female of M. gracilipes the genital segment is

about as long as wide and the free segment of leg 5

lacks a proximal expansion.

Stellicola novaecaledoniae, new species

FIGURES 4e-i, ba-k, 6a-i

TYPE MATERIAL.—146 $ $, 173 $ $, and 95

copepodids from the body surface of 24 bright blue

sea stars, Linckia laevigata (Linnaeus), in 0.5 m,

on the reef at Goro, southeastern New Caledonia,

22°18'00"S, 167°02'00"E, 6 August 1971. Holotype $, allotype, and 220 para types (100 $ $,

120 $ $) deposited in the National Museum of

Natural History (USNM), Smithsonian Institution;

the remaining paratypes in the collection of the

author.

OTHER SPECIMENS.—From Linckia laevigata: 33

5 $, 42 $ $ , and 4 copepodids from 14 hosts, in

0.1-1 m, on the reef about 5 kms south of Yate,

southeastern New Caledonia, 22°HWS, 166°59'

00"E, 23 June 1971. From Linckia guildingi Gray:

75 $ $ , 77 $ $ , and 42 copepodids from 40 hosts,

in 1-2 m, west of Isle Ngou, near Noumea, 22° 13'

44"S, 166°23'01"E, 3 August; 13 $ $, 19 $ $, and

2 copepodids from 20 hosts, in 1-2 m, west of Isle

Ngou, 29 July; 19 $ $, 27 $ <J from 10 hosts, in
NUMBER 217

b

FIGURE 3.—Metaxymolgus echinasteris, new species, female: a, endopod of leg 3, with inner edge

of coxa and basis, anterior (F); b, leg 4 and intercoxal plate, anterior (F); c, outer spine on

second segment of endopod of leg 4, anterior (H); d, leg 5, dorsal (G). Metaxymolgus echinasteris,

new species, male: e, dorsal (B); /, urosome, dorsal (C); g, second antenna, postero-inner (G);

h, maxilliped, inner (G); i, endopod of leg 1, anterior (G). (Scale H = 0.02 mm).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Metaxymolgus echinasteris, new species, male: a, endopod of leg 4, anterior (G); b,

leg 5, dorsal (D); c, leg 6, ventral (F); d, spermatophore, attached to female, dorsal (D). Stellicola novaecaledoniae, new species, female: e, dorsal (B); /, urosome, dorsal (F); g, genital area,

ventral (A); h, caudal ramus, dorsal (I); i, rostrum, ventral (E). (Scale I = 0.03 mm.)
NUMBER 217

1-2 m, Ricaudy Reef, near Noumea, 19 June; and

2 9 $ , 1 $ from 2 hosts, in 2 m, west of Isle Ngou,

3 August.

FEMALE.—Body (Figure 4e) with a broad flattened prosome and a relatively short urosome.

Length (not including the setae on the caudal rami)

1.28 mm (1.25-1.35 mm) and the greatest width

0.97 mm (0.94-1.02 mm), based on 10 specimens in

lactic acid. Ratio of the length to the width of the

prosome 1:1. Segment of leg 1 incompletely fused

with the cephalosome. Epimera of the segments

bearing legs 1-3 formed as in S. illgi Humes and

Stock, 1973. Segment of leg 4 small and almost

completely covered in dorsal view by the tergum

of the preceding segment. Ratio of the length of the

prosome to that of the urosome 2.5:1.

Segment of leg 5 (Figure 4/) 125 X 247 um.

Between this segment and the genital segment no

ventral intersegmental sclerite. Genital segment

159 X 221 ^m, wider than long. Genital areas

located ventrolaterally in the posterior part of the

segment. Each area (Figure 4g) bearing a plumose

seta 55 um mounted on an elongated pedicel, a

unilaterally spinulose seta 16 um, and a small spiniform process. Three postgenital segments from

anterior to posterior 49 X 91, 44 X 82, and 53 X 68

um (the width taken at the middle). Posterior

margin of the anal segment smooth.

Caudal ramus (Figure 4h) short, 32 X 25 um.

Outer lateral seta 73 um and naked. Dorsal seta 38

urn with a few short spinules. Outermost terminal

seta 135 um with a few proximal spinules. Innermost terminal seta 122 um with spinules. Two

median terminal setae 375 um (outer) and 600 um

(inner), both with distal spinules and inserted

dorsally to a small ventral flange with minute marginal spinules. On the proximal outer edge of the

ramus a long setule (19 um).

Body surface bearing a few small hairs (sensilla)

as indicated in Figure 4e-f.

Egg sac (Figure 4e) in dorsal view elongated oval,

880 X 480 um, reaching beyond the tips of the

longest ramal setae. In side view the egg sac flattened, with only three layers of eggs at its thickest

part. General form of the egg sac leaflike. Numerous eggs each about 72 um in diameter.

Rostrum (Figure 4t) resembling that of S. illgi.

First antenna (Figure 5a) about 425 um long.

Lengths of the seven segments (measured along

their posterior nonsetiferous margins): 39 (61 um

along the anterior margin), 180, 62, 66, 32, 17.5,

and 21 um respectively. Formula for the armature

the same as in Metaxymolgus echinasteris. All the

setae naked. Third segment having an oblique

suture on its ventral surface, this suture being

absent on the dorsal surface.

Second antenna (Figure bb) 440 um long and

3-segmented. Formula for the armature: 1, 1,

3 + 1 + 5 small subterminal elements. All the

setae naked. Third segment 218 um along its outer

edge, 140 ^m along its inner edge, and 35 um wide.

Claw (Figure 5c) 91 um along its axis.

Labrum (Figure bd) with two broad divergent

posteroventral lobes. Median postoral surface immediately behind the labrum finely spinulose.

Mandible (Figure be) similar to that of S. illgi.

Paragnath a small hairy lobe. First maxilla (Figure

5/) with four naked elements. Second maxilla

(Figure 5g) resembling that of S. illgi in major

respects. Maxilliped (Figure 5h) also similar to that

species, but one of the setae on the second segment

with spinules.

Ventral surface between the maxillipeds and the

first pair of legs (Figure 5t) not protuberant.

Legs 1-4 (Figures bj-k, 6a-b) having the same

segmentation and armature as in S. illgi. Intercoxal

plate of leg 1 with a medially indented ventral margin (Figure 5;); in legs 2-4 this margin smooth.

Inner coxal seta of legs 1-3 long and plumose, but

this seta in leg 4 very small, 6 [Am, and naked.

Exopod of leg 4 78 um long. First segment of the

endopod 35 X 16.5 um, with its inner distal naked

seta 33 um. Second segment 36 X 11 um, with its

two barbed spines 26 um (outer) and 65 um

(inner) and its inner plumose seta 67 um. Hairs

present on the outer margins of both segments and

on the inner margin of the second segment.

Leg 5 (Figure 6c) having an unornamented free

segment 65 X 15.5 um (greatest length, width at

the middle) about 4.2 times longer than wide. Two

very unequal terminal elements 20 um (finely

barbed) and 115 um (naked). Seta on the body

near the insertion of the free segment about 40 um

and lightly plumose.

Leg 6 represented by the two setae on the genital

area (Figure 4g).

Most living specimens in transmitted light pale

blue, the eye red, the egg sacs brownish gray. A few

females lacking the blue color entirely and slightly

opaque. Blue color quickly changing to reddish
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

k

FIGURE 5.—Stellicola novaecaledoniae, new species, female: a, first antenna, ventral (F); b, second

antenna, posterior (F); c, claw of second antenna, posterior (A); d, labrum and spinulose postoral

area, with paragnaths indicated by broken lines, ventral (A); e, mandible, posterior (A); /, first

maxilla, posterior (D); g, second maxilla, posterior (A); h, maxilliped, posterior (A); i, area

between maxillipeds and first pair of legs, ventral (C); /, leg 1 and intercoxal plate, anterior (F);

k, leg 2 and intercoxal plate, anterior (F).
NUMBER 217 11

FIGURE 6.—Stellicola novaecaledoniae, new species, female: a, leg 3, anterior (F); b, leg 4 and

intercoxal plate, anterior (F); c, leg 5, dorsal (A). Stellicola novaecaledoniae, new species, male:

d, dorsal (B); e, urosome, dorsal (F); /, second antenna, posterior (G); g, labrum, ventral (A);

h, maxilliped, posterior (D); i, leg 6, ventral (G). Stellicola parvulipes, new species, female:

;, dorsal (E); k, lateral (E).
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

orange when the copepods were placed in 70 percent ethyl alcohol.

MALE.—Body (Figure 6d) with a broad prosome,

its outline in dorsal view more spherical than in the

female. Length (without the ramal setae) 0.83 mm

(0.75-0.89 mm) and the greatest width 0.56 mm

(0.55-0.58 mm), based on 10 specimens in lactic

acid. Ratio of the length to the width of the prosome 1:1.1, slightly wider than long. Ratio of the

length of the prosome to that of the urosome

1.78:1.

As in other species of Stellicola the segment bearing leg 5 fused with the genital segment (Figure

6e), the combined segment being 180 X 164 um.

Posterior third of the dorsal surface of this segment

covered with small spinules. Four postgenital segments from anterior to posterior 47 X 73, 39 X 65,

30 X 57, and 29 X 47 um (width at the middle).

Caudal ramus 31 X 26 um, resembling that of the

female.

Rostrum and first antenna (without added

aesthetes) like those of the female.

Second antenna (Figure 6/) 275 urn long. Two

features of sexual dimorphism evident: (1) a prominent inner spiniform process just proximal to the

seta on the second segment and (2) the distinct

curvature of the third segment. Claw 69 um.

Labrum (Figure 6g) with the configuration of

the two lobes slightly different from the female.

Mandible, paragnath, first maxilla, and second

maxilla like those of the female. Maxilliped (Figure

6h) small, its entire length only about 83 um, and

4-segmented. First segment unarmed. Second segment bearing two naked setae, with prominent

spinules near the insertion of the more proximal

seta. Third segment small and unarmed. Elongated

fourth segment (37 um long) not clawlike, but with

a broad serrated tip; bearing distally a bladelike

process bent back upon the segment and provided

with minute spinules along its outer edge. In addition, the segment carrying proximally two very

unequal naked setae, the larger one on the posterior

surface of the segment, the smaller one on the

anterior surface.

Ventral area between the maxillipeds and the

first pair of legs like that of the female.

Legs 1-4 similar to those in the female.

Leg 5 also resembling that of the female, but

the free segment smaller, 26 X 8 um.

Leg 6 (Figure 6t) a posteroventral flap on the

genital segment bearing two naked setae 30 and 39

um and a small spiniform process.

Spermatophore not observed.

Living specimens paler blue than in the female.

ETYMOLOGY.—The specific name novaecaledoniae

refers to the presence of this species in New Caledonia.

COMPARISON WITH OTHER SPECIES.—There are

presently 10 species in the genus Stellicola (Humes

and Stock, 1973). The female of the new species

may be distinguished by its elongated free segment

in leg 5 (more than 3:1) from all these species

except S. femineus Humes and Ho, 1967, and 5. illgi

Humes and Stock, 1973. It may be separated from

S. femineus, which has a large toothlike spine on

the terminal lash of the second maxilla. The male

of the new species is easily distinguished by the

form of its maxilliped from all species except

S. illgi. (Males of four species are unknown. See

Humes and Stock, 1973).

Stellicola novaecaledoniae is closer to 5. illgi than

to any other member of the genus. There are, however, several easily recognizable differences between

the two species as indicated in Table 1.

LICHOMOLGID COPEPODS ASSOCIATED WITH Linckia.—There are now four species of the lichomolgid

genus Stellicola known to live on Linckia, a genus

of sea stars widely distributed through the tropical

Indian and Pacific oceans. Three of these copepods

are definitely associated with Linckia laevigata.

They are Stellicola pollex Humes and Ho, 1967, in

Madagascar (Humes and Ho, 1967), Stellicola illgi

Humes and Stock, 1973, in the Palau Islands and

Fiji, and the new species Stellicola novaecaledoniae

in New Caledonia. A fourth copepod, Stellicola

caeruleus (Stebbing, 1900), was reported from a

"blue Linckia" in New Guinea by Stebbing (1900).

Clark (1921:636) later listed this copepod (under

the name Linckiomolgus caeruleus Stebbing) as

associated with Linckia laevigata.

Two of these four Stellicola species are not

restricted to Linckia laevigata, but occur also on

other sea stars in this genus. Humes and Stock

(1973) have reported Stellicola pollex from Linckia

diplax, Linckia multiflora, and Linckia guildingi in

Hawaii. Stellicola novaecaledoniae occurs on Linckia guildingi as reported above.

Sufficient collections are not available to establish the range of the various Stellicola on Linckia.

The occurrence of Stellicola pollex in both Mada-
NUMBER 217 13

gascar and Hawaii, and of Stellicola illgi in the

Palaus and Fiji (for records see Humes and Stock,

1973), indicates a wide distribution for these copepods.

Stellicola oreastriphilus Kossmann, 1877

SPECIMENS COLLECTED (all from the cushion sea

star Culcita novaeguineae Miiller and Troschel, a

new host).—9 9 9, 5 $ $ , and 1 copepodid from

one host, in 0.5 m, eastern end of Isle Maitre, near

Noumea, 22°20'35"S, 166°25'10"E, 8 June 1971;

1 9, 4 $ $ , and 1 copepodid, from one host, in

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

166°25'50"E, 13 June; 2 5$, 3 U from one host,

in 2 m, Rocher a la Voile, 19 June; 7 $ $ , 11 $ $ ,

and 1 copepodid from one host, in 1 m, east of

Isle To N'du, near Noumea, 22°10'49"S, 166°17'

12"E, 12 July; 11 9 9,5$$ from 5 hosts, in

10 cm, eastern end of Isle Maitre, near Noumea,

22°20'35"S, 166°25'10"E, 16 July; 23 $ 9 from 7

hosts, intertidal, eastern end of Isle Maitre, near

Noumea, 22°20'35"S, 166°25'45"E, 31 July.

REMARKS.—Stellicola oreastriphilus is a euryxenous species known to live on eight different asteroids. In the Red Sea Kossmann (1877) found it on

Asterope carinifera (Lamarck). In Madagascar it

was reported by Humes and Cressey (1961) on

Protoreaster lincki (de Blainville), Culcita schmideliana (Retzius), Pentaceraster mammillatus (Audouin), and Poraster superbus (Mobius), by Humes

and Ho (1966) on Choriaster granulatus (Liitken),

and by Humes and Ho (1967) on Protoreaster

nodosus (Linnaeus).

Stellicola parvulipes, new species

FIGURES 6j-k, 7a-k, 8a-i, 9a-k

TYPE MATERIAL.—13 9 9, 3 $ $ from one

cushion star, Culcita novae guineae Mtiller and

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

New Caledonia, 22°10/49"S, 166°17'12"E, 12 July

1971. Holotype 9, allotype, and 14 paratypes

(12 9 9, 2 $ $) deposited in the National

Museum of Natural History (USNM), Smithsonian

Institution.

OTHER SPECIMENS (all from Culcita novaeguineae).—6 9 9 from one host, in 2 m, Rocher a la

Voile, near Noumea, 22°18/24"S, 166°25'50"E, 19

June; 5 9 9 , 3 $ $ from 5 hosts, in 10 cm, eastern

end of Isle Maitre, near Noumea, 22°20'35"S,

166°25'45"E, 16 July.

FEMALE.—Body (Figure 6/-A) with an anteriorly

subtruncated prosome, which is moderately thickened dorsoventrally. Length (excluding the setae

on the caudal rami) 0.65 mm (0.64—0.66 mm) and

the greatest width 0.29 mm (0.28-0.30 mm), based

on 10 specimens in lactic acid. Ratio of the length

TABLE 1.—Selected characters useful in distinguishing Stellicola novaecaledoniae

and Stellicola illgi

Characters

FEMALE

Lateral contour of

genital segment

Ratio of length to width of

free segment of leg 5

Body length

MALE

Sexual dimorphism in

second antenna

Length of process bent back

upon fourth segment of

maxilliped

Body length

S. novaecaledoniae

slightly angular

4.2:1

1.28 mm (1.25-1.35 mm)

present

half length of segment

0.83 mm (0.75-0.89 mm)

S. illgi

rounded

3.3:1

1.00 mm (0.94-1.06 mm)

absent

short, less than one fourth

length of segment*

0.60 mm (0.55-0.64 mm)

• This small process was not mentioned in the original description of Humes and Stock

(1973). Reexamination of paratypes of S. illgi has shown that it is present, though much smaller

than in the new species and easily overlooked.
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Stellicola parvulipes, new species, female: a, urosome, dorsal (G); b, genital area,

dorsal (D); c, caudal ramus, dorsal (H); d, rostrum, ventral (F); e, first antenna, dorsal (A); /,

second antenna, posterior (A); g, labrum, ventral (A); h, mandible, posterior (I); i, first maxilla,

posterior (I); /, second maxilla, posterior (I); k, maxilliped, posterior (I).
NUMBER 217 15

to the width of the prosome 1.54:1. Segment of leg 1

separated from the cephalosome by a weak transverse dorsal furrow. Epimera of the segments of legs

1-3 rounded. Ratio of the length of the prosome to

that of the urosome 2.24:1.

Segment of leg 5 (Figure 7a) 72 X 103 um.

Between this segment and the genital segment no

ventral intersegmental sclerite. Genital segment a

little flattened dorsoventrally; in dorsal view subquadrate, 77 X 97 um, somewhat wider than long.

Genital areas situated dorsally on a pair of posteriorly projecting lobes. Each area (Figure 7b) bearing

two unequal naked setae approximately 9 and 3

um respectively. Three postgenital segments from

anterior to posterior 30 X 50, 22 X 45, and 21 X 42

um. Posterior border of the anal segment smooth.

Caudal ramus (Figure 7c) 15.5 X 13.5 um. Outer

lateral seta 25 um, the dorsal seta 33 um, the outermost terminal seta 44 um, the innermost terminal

seta 39 um, and the two long median terminal

setae 130 um (outer) and 210 um (inner), both

inserted dorsally to a small ventral flange with a

smooth margin. All the setae naked. Proximally on

the outer edge of the ramus a slender setule and

near the middle of the ramus several inner spinules.

Body surface with a few small hairs (sensilla) as

indicated in Figures 6/ and 7a.

Egg sac not observed.

Rostrum (Figure 7d) elongated and rounded

posteroventrally.

First antenna (Figure 7e) 170 um long. Lengths

of the seven segments (measured along their posterior nonsetiferous margins): 12 (30 um along the

anterior margin), 55, 22, 25, 17.5, 9, and 9.5 um

respectively. Configuration of the last two segments

suggesting their fusion, but in actuality a weak

suture existing between the segments. Formula for

the armature the same as in Stellicola novaecaledoniae. All the setae naked. Segment 3 with a

diagonal suture on its ventral surface.

Second antenna (Figure 7/) 192 um long and

3-segmented. Armature: 1, 1, 3 + I + 5 small subterminal elements. All the setae naked. Third segment 51 um along the outer edge, 35 um along the

inner edge, and 23 um wide. Claw 53 um along its

axis.

Labrum (Figure 7g) with two broad posteroventral lobes with smooth margins.

Mandible (Figure 7h) resembling in general form

that of S. novaecaledoniae, but the elements in the

fringe along the convex side of the mandible,

though slender and closely packed at first, becoming broader and separated distally. Paragnath a

small hairy lobe. First maxilla (Figure 7i) slender

with three terminal naked elements, one much

shorter than the others. At the region where a

subterminal fourth element occurs in other species,

here a slight swelling of the wall, but a distinct

element not seen. Second maxilla (Figure 7;) resembling in major respects that of S. novaecaledoniae,

but the spinules on the large inner spine here of a

more uniform size and the lash having a very large

but slender proximal tooth followed by slender

spinules. Maxilliped (Figure 7k) 3-segmented, with

the first segment unarmed, the second with two

small naked setae, and the third with one seta in

addition to the terminal spiniform process.

Ventral surface between the maxillipeds and the

first pair of legs (Figure 8a) not protuberant.

Legs 1-4 (Figure Sb-e) with the same segmentation and armature as in S. novaecaledoniae. Basis

of leg 4 not greatly elongated as it is in other species

of the genus. In this leg the inner coxal seta vestigial. Exopod 46 um long. First segment of the

endopod 16.5 X 10 urn, with its inner distal seta

45 um; the second segment 20 X 10 um, with its

three elements from outer to inner 14, 26, and

52 um.

Leg 5 (Figure Sf-g) not clearly separated from

the body and consisting of a minute lobe, corresponding to the free segment in other species, largely

hidden in dorsal view; its two terminal setae about

22 um.

Leg 6 represented by the two setae on the genital

area (Figure 76).

Living specimens in transmitted light opaque, the

eye red.

MALE.—Body (Figure Sk) with the prosome more

rounded anteriorly and somewhat less thickened

dorsoventrally than in the female. Length (without

the ramal setae) 0.59 mm (0.59-0.60 mm) and the

greatest width 0.25 mm (0.24-0.25 mm), based on 4

specimens in lactic acid. Ratio of the length to the

width of the prosome 1.27:1. Ratio of the length of

the prosome to that of the urosome 1.25:1.

Segment of leg 5 fused with the genital segment

(Figure Si), whose lateral borders are slightly irregular in dorsal view. Combined segment 125 X 130

um. Four postgenital segments from anterior to
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

h

FIGURE 8.—Stellicola parvulipes, new species, female: a, area between maxillipeds and first pair

of legs, ventral (A); b, Jeg 1 and intercoxal plate, anterior (A); c, leg 2 and intercoxal plate,

anterior (A); d, leg 3 and intercoxal plate, anterior (A); e, leg 4 and intercoxal plate, anterior

(A); /, leg 5, ventral (H); g, leg 5, lateral (H). Stellicola parvulipes, new species, male: h, dorsal

(E); i, urosome, dorsal (G).
NUMBER 217 17

FIGURE 9.—Stellicola parvulipes, new species, male: a, first antenna, dorsal (A); b, second antenna,

posterior (A); c, labrum, ventral (A); d, maxilliped, postero-inner (D); e, area between maxillipeds and first pair of legs, ventral (A); /, same, lateral (A); g, endopod of leg 1, anterior (A);

h, endopod of leg 2, anterior (A); i, endopod of leg 3, anterior (A); /, right half of fused segment

of leg 5 and genital segment, showing both leg 5 and leg 6, ventral (A); k, fused segment of leg

5 and genital segment, lateral (A). Synstellicola pichoni (Humes and Ho, 1966), female: /, endopod of leg 2, anterior (G); m, third segment of endopod of leg 3, anterior (G).
18

posterior 33 X 52, 29 X 47, 27 X 42, and 16 X 36

Caudal ramus 14 X 14 (j.m, similar to that of the

female.

Rostrum like that of the female. First antenna

(Figure 9a) 130 ^m long. Lengths of the segments

(measured as in the female): 9 (25 |xm along the

anterior edge), 37, 15.5, 17.5, 15.5, 10, and 6.5 ^m

respectively, with certain segments relatively shorter

than in the opposite sex. Armature like that in the

female. Last two segments not completely separated.

Second antenna (Figure 9fr) 155 |xm long. Third

segment relatively longer than in the female, being

57 urn along its outer edge, 42 ^m along its inner

edge, and 21 \im wide. Claw 50 ^im along its axis.

Labrum (Figure 9c) with two broad posteroventral lobes, each with a small sclerotized process

on the margin. Mandible, paragnath, first maxilla,

and second maxilla like those in the female. Maxilliped (Figure 9d) relatively short. First segment

unarmed. Second segment with two short naked

setae and a long narrowly U-shaped row of minute

spinules. Small third segment unarmed. Relatively

short claw, 58 [ira. along its axis, bearing proximally

two unequal setae.

Ventral surface between the maxillipeds and the

first pair of legs (Figure 9e) strongly protuberant.

Protuberance best seen in lateral view (Figure 9/).

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

Endopods of legs 1-3 (Figure 9g-i) differing slightly

from those in the female in the development of the

outer distal spiniform processes on segments 1 and 2.

Leg 4 like that of the female.

Leg 5 (Figure 9 j-k) a small lobe not separated

from the body, thus resembling that of the female.

Two terminal setae, however, here very unequal,

32 and 85 \im.

Leg 6 (Figure 9j-k) a posteroventral flap on the

genital segment bearing two naked setae, one 28 \im,

the other 13 \im and hyaline. Median edges of the

two flaps projecting ventrally (as in Figure 9k).

Just anterior to the sixth legs a conspicuous median

ventral process (Figure 9j-k).

Spermatophore not observed.

Living specimens colored as in the female.

ETYMOLOGY.—The specific name parvulipes, from

the Latin parviilus = very small and pes = foot,

alludes to the minute leg 5 in this species.

COMPARISON WITH OTHER SPECIES.—With the addition of the two new species of Stellicola described

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in this work there are now 12 species in the genus

Stellicola Kossmann, 1877, as defined by Humes and

Stock (1973). Stellicola parvulipes differs from all

others in three respects: (1) the female is shorter

(0.65 mm), all other species being close to 1 mm or

more in length; (2) the epimera of the metasomal

segments are rounded rather than pointed as in

other species; and (3) instead of leg 5 having a

free segment (that is, separated from the body by

an articulation), it is fused with the body.

Seven species of Stellicola have a greatly elongated

basis in leg 4 (S. pollex Humes and Ho, 1967,

S. caeruleus (Stebbing, 1900), S. illgi Humes and

Stock, 1973, S. longicaudatus (Thompson and

A. Scott, 1903), S. pleurobranchi Kossmann, 1877,

S. thorelli Kossmann, 1877, and S. novaecaledoniae

described above). The remaining four species have

a much less elongated basis, approaching the condition in the new species. These may be easily

distinguished from S. parvulipes as follows: in

S. femineus Humes and Ho, 1967, the female genital

segment is very wide with rounded lateral margins

and the male has a female-like maxilliped; in

S. oreastriphilus Kossmann, 1877, the lash of the

second maxilla has more than one large spine and

the female genital segment is much wider; in S. curticaudatus (Thompson and A. Scott, 1903) the lash

of the second maxilla has a series of graduated

spines, none markedly larger than the others; and

in S. holothuriae (Ummerkutty, 1962) the lash of

the second maxilla has several spines instead of a

single strongly developed one.

Synstellicola acanthasteris (Humes, 1970)

SPECIMENS COLLECTED (all from the crown of

thorns sea star Acanthaster planci (Linnaeus)).—

18 $$,1 1 $ «$ from one host, in 4 m, west of

Isle Ngou, northwest of Noumea, 22°13'44"S, 166°

23'01"E, 3 August 1971; 7 $2, 7 $ $ from one

host, in 2 m, Ricaudy Reef, near Noumea, 22° 19'

05"S, 166°26'28"E, 20 July. (These two records

have published by Humes and Stock, 1973, but are

repeated here for the sake of completeness.)

REMARKS.—This copepod is associated only with

Acanthaster planci. It was described from Eniwetok

Atoll, Marshall Islands (Humes, 1970) and has

since been reported from Viti Levu, Fiji (Humes

and Stock, 1973).
NUMBER 217 19

Synstellicola pichoni (Humes and Ho, 1966)

FIGURE 9l~m

SPECIMENS COLLECTED.—4 $ $,12 $ $ from one

sea star, Protoreaster lincki (de Blainville), a new

host, in 0.5 m, western side of Isle Maitre, near

Noumea, 22°20'05"S, 166°24'05"E, 8 August 1971.

REMARKS.—These copepods agree closely with

paratypic material from Choriaster granulatus (Liitken) in Madagascar. The New Caledonian specimens are a little larger, however, the female being

1.08 mm (1.05-1.11 mm) in length and 0.48 mm

(0.42-0.51 mm) in greatest width (based on 4

specimens in lactic acid), and the male 0.89 mm

(0.73-0.98 mm) in length and 0.35 mm (0.26-0.40

mm) in greatest width (based on 6 specimens in

lactic acid). The spines on the third segment of

the endopod in leg 2 (Figure 9/) and leg 3 (Figure

9m) are relatively longer than in the Madagascar

specimens. Otherwise the copepods from the two

hosts and localities are similar. (The dimensions

for the free segment of leg 5 in the female given

by Humes and Ho, 1966, page 99, are erroneus and

should read 28 X 9 \nm, instead of 65 X 21 \im).

Synstellicola pichoni is now known to live on two

species of sea stars, in localities remote from one

another (on Choriaster granulatus in Madagascar

and on Protoreaster lincki in New Caledonia).

Literature Cited

Clark, A. H.

1921. A Monograph of the Existing Crinoids, volume I:

The Comatulids, part 2. Bulletin of the United

States National Museum, 82:I-XXV, 1-795.

Humes, A. G.

1964. New Species of Lichomolgus (Copepoda, Cyclopoida)

from Sea Anemones and Nudibranchs in Madagascar. Cahiers O.R.S.T.O.M., Oceanographie, 1963, 6

(serie Nosy Be II):59-130.

1970. Stellicola acanthasteris, n. sp. (Copepoda, Cyclopoida) Associated with the Starfish Acanthaster

planci (L.) at Eniwetok Atoll. Publications of the

Seto Marine Biological Laboratory, 17(5):329-338.

1971. Cyclopoid Copepods (Stellicomitidae) Parasitic on

Sea Stars from Madagascar and Eniwetok Atoll.

Journal of Parasitology, 57(6): 1330-1343.

Humes, A. G., and R. F. Cressey

1961. Lichomolgus oreastriphilus (Kossmann) copepode

cyclopoide parasite des etoilcs de mer a Madagascar.

Memoires de I'Institut Scientifique de Madagascar,

1959, series F, 3:83-92.

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

1966. Cyclopoid Copepods Associated with the Starfish

Choriaster granulatus (Liitken) in Madagascar.

Cahiers O.R.S.T.OJM., serie Oceanographie, 4(3):

93-108.

1967. New Species of Stellicola (Copepoda, Cyclopoida)

Associated with Starfishes in Madagascar, with a

Redescription of 5. caeruleus (Stebbing, 1900). Bulletin of the British Museum (Natural History), Zoology, 15(5):201-225.

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

1973. A Revision of the Family Lichomolgidae Kossmann,

1877, Cyclopoid Copepods Mainly Associated with

Marine Invertebrates. Smithsonian Contributions to

Zoology, 127: pages i-v, 1-368.

Kossmann, R.

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

1-24 in volume 4 in Zoologische Ergebnisse einer

im Auftrage der Koniglichen Academie der Wissenschaften zu Berlin Ausgefiihrten Reise in die Kustengebiete des Rothen Meeres.

Monod, Th. and R. -Ph. Dollfus

1932. Les copepodes parasites de mollusques. Annales de

Parasitologie Humaine et Comparee, 10:129-204.

Scott, A.

1909. The Copepoda of the Siboga Expedition, Part I:

Free-swimming, Littoral, and Semi-parasitic Copepoda. Volume 29a in Siboga Expedition. Pages 1-

323.

Stebbing, T. R. R.

1900. On Crustacea Brought by Dr. Willey from the

South Seas. Pages 605-690 in part 5 in A. Willey,

Zoological results based on material from New

Britain, New Guinea, Loyalty Islands and elsewhere,

collected during the years 1895,1896 and 1897.

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.

Ummerkutty, A. N. P.

1962. Studies on Indian Copepods 5: On Eleven New

Species of Marine Cyclopoid Copepods from the

South-east Coast of India. Journal of the Marine

Biological Association of India, 1961, 3(1 & 2): 19-69.

U.S. GOVERNMENT PRINTING OFFICE: 1»7S SS9/4

Publication in Smithsonian Contributions to Zoology

Manuscripts for serial publications are accepted by the Smithsonian Institution Press, subject to substantive review, only through departments of the various Smithsonian museums. NonSmithsonian authors should address inquiries to the appropriate department. If submission is

invited, the following format requirements of the Press will govern the preparation of copy.

Copy must be typewritten, double-spaced, on one side of standard white bond paper, with

W2" top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompanied by the original artwork. Duplicate copies of all material, including illustrations, should be

retained by the author. There may be several paragraphs to a page, but each page should begin

with a new paragraph. Number consecutively all pages, including title page, abstract, text, literature cited, legends, and tables. The minimum length is 30 pages, including typescript and illustrations.

The title should be complete and clear for easy indexing by abstracting services. Taxonomic

titles will carry a final line indicating the higher categories to which the taxon is referable:

"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify

the author on the first page of text with an unnumbered footnote that includes his professional

mailing address. A table of contents is optional. An index, if required, may be supplied by the

author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further

headings may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet,

omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date:page) with a full reference at the

end of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent lines indented about three spaces. Within an entry, use a period-dash (.—) to separate

each reference. Enclose with square brackets any annotation in, or at the end of, the entry.

For references within the text, use the author-date system: "(Jones, 1910)" and "Jones

(1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

Use the metric system instead of, or in addition to, the English system.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively; however, if a group of figures is treated as a single figure, the components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 96) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on

pages separate from the text and not attached to the artwork. An instruction booklet for the

preparation of illustrations is available from the Press on request.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and

article titles, using initial caps with all words except minor terms such as "and, of, the." For

capitalization of titles in foreign languages, follow the national practice of each language.

Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume,

number, and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," "pages."

For free copies of his own paper, a Smithsonian author should indicate his requirements

on "Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will

receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.
i