JU-SHET Parasitic Copepods

of the Family

_ Chondracanthidae

from Fishes of

Eastern North America

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 87
SERIAL PUBLICATIONS OF TH E SMITHSONIA N 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
SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 87

Ho Parasitic Gopepods

of the Family

Chondracanthidae

from Fishes of

Eastern North America

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON
ABSTRACT

Ho, Ju-shey. Parasitic Copepods of the Family Chondracanthidae from Fishes of

Eastern North America. Smithsonian Contributions to Zoology, number 87, 39

pages. 1971.—Nineteen species representing nine genera of chondrachanthid copepods are known to occur on fishes off the Atlantic and the Gulf coast of North

America. They are: Acanthochondria cornuta*, A. cyclopsetta, A. exilipes, A.

galerita, A. phycidis, Acanthochondrites annulatus*, Berea ancoralis*, Blias prionoti*,

Chondracanthodes deflexus*, C. radiatus, Chondtacanthus cottunculi, C. janebennettae, C. merluccii, C. nodosus, C. wilsoni, Heterochondria crassicornis*, Pseudochondracanthus diceraus*, P. hexaceraus, and Rhynchochondria longa*. A redescription is given for all species except eight that are marked with an asterisk, since they

have already been treated elsewhere in a revision of the chondracanthid genera (Ho,

1970). A new form, Chondiacanthus wilsoni new species, is described and a key to

the nineteen eastern North American species is given.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1971

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washinsrton, D.C. 20402 - Price 50 cents (paper cover)
Ho Parasitic Copepods

of the Family

Chondracanthidae

from Fishes of

Eastern North America

Introduction

So far as I am aware, twenty-four species of cyclopoid

copepods belonging to the family Chondracanthidae

have been reported to occur on the fishes off the

Atlantic and the Gulf coast of North America. They

were reported by Bere (1930, 1936), Causey (1953,

1955a, 1955b, 1960), Ho (1967, 1970), Pearse (1947,

1952a, 1952b), Rathbun (1886), and Wilson (1908,

1932, 1935). Specimens of these twenty-four species,

including the type specimens studied by the above six

investigators, were deposited in the United States

National Museum (USNM).

In the summer of 1967, while I was making a revisional study of the family, all chondracanthid

copepods in the collections of USNM were reexamined. This restudy has revealed that six of the

twenty-four reported species are conspecific with other

species, that the specimens from Woods Hole, Massachusetts, reported as Or alien triglae by Wilson

(1932) are actually a species (Rebelula bouvieri) of

the family Sphyriidae, and that the specimen from

Dry Tortugas, Florida, identified as Chondiacanthus

nodosus by Wilson (1935) is actually a new species.

Consequently, in accordance with my recently revised

genera of the Chondracanthidae (Ho 1970) and with

the addition of a species newly recorded off Florida,

there are now nineteen species representing nine

genera known to occur in this region. It is worthy of

note here that they all belong to the subfamily

Chondracanthinae.

Ju-shey Ho, Department of Biology, California State College,

Long Beach, California 90801.

As pointed out by Ho (1970), the large number of

species in a family may be the result of successful

adaptive diversification, but in the case of the Chondracanthidae (about 170 species), it is, at least, partly

due to the failure of some previous investigators to

recognize variant features which are actually individual and nonspecific, and partly due to inadequate

and incomplete original descriptions which misled

subsequent investigators to describe new species from

specimens actually belonging to known species. Therefore, in the course of revising the family Chondracanthidae, it is necessary to reexamine every specimen,

regardless of type specimen or nontype specimen,

reported in the past for verifying its proposed identification. The purpose of presenting here the

redescriptions of species occurring in the eastern

North America is, accordingly, to clarify the past

confusion in specific identification and to present an

orderly basis for future study of the species of the

North American chondracanthid copepods. Since

eight of these nineteen species are type species of their

genera and have already been redescribed in the

revision of the genera (Ho 1970), the following

redescriptions deal only with the remaining eleven

species, but all nineteen species are included in the

key.

In addition to reexamination of the specimens deposited in USNM, I have also studied some chondracanthids that were collected by myself from

Apalachee Bay, Florida and on board research vessel

Oregon (during Cruise 105, off Georgia and Florida)

and research vessel Albatross IV (during Cruise 66-3,

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

off Massachusetts and Maine). The type specimens

of some species deposited in the Universitetets Zoologiske Museum (UZM), Copenhagen, Denmark,

were also studied to confirm some specific identifications.

I wish to thank Dr. Arthur G. Humes (Department

of Biology, Boston University, Boston) for arranging

financial support for this study from his grants (GB

1809, GB 5838, and GB 838IX) from the National

Science Foundation and to Dr. Roger F. Cressey

(United States National Museum, Washington, D.C.)

and Dr. Torben Wolff (Universitetets Zoologiske

Museum, Copenhagen) for making available to me

for study the chondracanthid copepods deposited in

their museums. The Bureau of Commercial Fisheries,

Fish and Wildlife Service, gave me the opportunity

to collect parasitic copepods on board R/V Oregon

and R/V Albatross IV.

Key to the Nineteen Species of Chondracanthidae Known from

Eastern North America

1. Trunk region with outgrowths in form of processes, protrusions, or knobs; posterior proccesses present 2

Trunk region without outgrowths; with or without posterior processes 9

2. Three pairs of unmodified legs present, all biramous; both sexes with an accessary

antennule Rhynchochondria longa Ho 1967

Two pairs of legs present, either lobate or rudimentary; accessary antennule either present

or absent (when present, only in male) 3

3. Two pairs of legs partially modified, with saclike protopod and rodlike rami (Chondracanthodes Wilson 1932) 4

Two pairs of legs modified, either trilobate or bilobate (Chondracanthus Delaroche

1811) 5

4. First antenna small and filiform; head without lateral processes.

Chondracanthus deflexus Wilson 1932

First antenna large and lobate; head with a pair of long posterolateral processes

C.radiatus (Muller 1776)

5. Posterodorsal portion of head protruded into a large crestlike outgrowth; abdomen greatly

elongated C. janebennettae Causey 1953

Posterodorsal portion of head smooth; abdomen not elongated 6

6. Trunk region with only one or two pairs of lateral outgrowths; legs bilobate 7

Trunk region with more than four pairs of lateral outgrowths; legs trilobate 8

7. First antenna large, with triangular basal portion; head without posterolateral processes

C. cottunculi Rathbun 1886

First antenna small, with evenly swollen basal portion; head bearing a pair of posterolateral processes C. merluccii (Holten 1802)

8. First antenna extremely small; claw of maxilliped with only one hooklet; caudal ramus

with usual long terminal process C. nodosus (Muller 1776)

First antenna large and fleshy; claw of maxilliped bearing numerous spinules; caudal ramus

lacking long terminal process C. wilsoni new species

9. Posterior processes present 10

Posterior process absent 17

10. One pair of modified legs present; male without first antennae and legs (Pseudochondracanthus Wilson 1908) 11

Two pairs of modified legs present; male with first antennae and legs 12

11. Head with only one pair of lateral processes; mandible bearing more teeth (56 on convex

side and 34 on concave side) P. diceraus Wilson 1908

Head with two pairs of lateral processes; mandible bearing less teeth (23 on convex side

and 14 on concave side) P. hexaceraus Wilson 1935

12. Legs unilobate; posterior end of body with a median lobe.

Acanthochondrites annulatus (Olsson 1886)

Legs bilobate; posterior end of body without median lobe (Acanthochondria Oakley

1927) 13

13. Rami of lobate legs extremely short and broad A. phycidis (Rathbun 1886)

Rami of lobate legs long and cylindrical 14
NUMBER 8 7

14. First antenna small and filiform; posterior process longer than the last trunk segment

A. exilipes Wilson 1932

First antenna large and lobate; posterior process distinctly shorter than the last trunk

segment 15

15. Rami of lobate legs at least four times longer than the protopod. A. cyclopaetta Pearse 1952

Rami of lobate legs at most two times longer than the protopod 16

16. Oral region flanked with a pair of small processes; labrum bearing denticles on posterior

surface; and rami of legs bluntly pointed A. galerita (Rathbun 1886)

Oral region without processes; labrum with smooth posterior surface; and rami of legs not

pointed A. cornuta (Miiller 1776)

17. Second antenna uncinate and legs unilobate. Heterochondria crassicornis (Krayer 1837)

Second antenna non-uncinate and legs biramous 18

18. Second antenna clavate and legs bilobate Berea ancoralis (Bere 1936)

Second antenna trifurcate and legs partially modified, with lobate protopod and rodlike

rami Blias prionoti Krayer 1863

Acanthochondria cyclopsetta Pearse 1952

FIGURE 1

Acanthochondria cyclopsetta Pearse 1952b: 37-38, figs. 145-

149.—Yamaguti 1963: 276, pi. 238: fig. 5.

MATERIAL STUDIED.—From USNM: 1 female

(mounted on slide) on gill of Cyclopsetta chittendeni

Norman, off Padre Island, Texas (USNM 92664,

type).

REMARKS.—Since only one specimen mounted in

balsam is known of this species, it is impossible to

make a satisfactory redescription of it. I have, however, made three illustrations (Figures \a, b, c) to

supplement the original description given by Pearse.

There is not the slightest doubt that this species is a

valid one in Acanthochondria. The following combination of characters distinguishes it from the rest of

the species in the genus: the large size (10.58 mm

long); the swollen, fleshy first antenna; the spinules

on posterior margin of the labrum; and the long,

attenuated, bilobate legs. The eggs are rather small

for such a large sized copepod, each egg being only

about 118 fx in diameter.

No pygmy male was seen on the slide, but Pearse

(1952b: 38) has noted that the male was "attached

to female, but lost."

The most outstanding feature of this species is the

bilobate, long legs, a condition which has never been

seen in any other known species of Acanthochandria.

Acanthochondria exilipes Wilson 1932

FIGURES 2—4

Acanthochondria exilipes Wilson 1932: 501-502, pi. 37:

figs, a-e.—Yamaguti 1963: 277.

MATERIAL STUDIED.—From USNM: 1 female on

gills of Lopholatilus chameleonticeps Goode and Bean,

Woods Hole, Massachusetts, 1914 (USNM 59777,

type) ; 55 females, with males attached, on gills of

L. chameleonticeps, Woods Hole, Massachusetts, 1914

(USNM 59778, paratypes) ; 11 females, with males

attached, on gills of L. chameleonticeps, south of

Nantucket, Massachusetts, 21 July 1902 (USNM

59776); 1 female, with male attached, on Lopholatilus species, off Martha's Vineyard, Massachusetts

(USNM 6084); and 7 females, with males attached,

on anterior and posterior ends of gill cavity of L.

chameleonticeps, off Martha's Vineyard, Massachusetts, 23 August 1881 (USNM 6089).

FEMALE.—Body (Figures 2a, b) rather long. Head

elliptical, distinctly longer than wide in extended

specimen (Figure 2a) but nearly as long as wide in

contracted specimen (Figure 2c). First pedigerous

segment nearly as wide as head, and second one only

slightly wider. Third pedigerous segment much larger

than fourth; a small lateral bulge located between

these two segments. Posterior processes longer than

fourth pedigerous segment. A pair of small vermiform

processes (Figure 2d) located on posterior surface of

trunk, to one of which the dwarf male is attached.

Genital segment (Figure 2d) bearing a pair of spiniform elements on midventural surface and abdomen,

a pair of setules on dorsal surface. Caudal ramus

(Figure 2e) with 3 setae, 1 knoblike process and an

abruptly narrowed spiniform terminal process; all

naked. Egg sac slightly shorter than body in extended

specimen, containing many rows of small eggs.

First antenna (Figure 2/) small, not much swollen

at basal region, and rather flattened; armature being:
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 1.—Acanthochondria cyclopsetta female: a, body of holotype, dorsal; b, head and part

of neck, dorsolateral; c, genito-abdomen, ventral. Scale: 1 mm in a; 0.1 mm in b, c.

1—1—2—8. Second antenna (Figure 2g) indistinctly

2-segmented, with usual recurved terminal hook.

Labrum with denticles on posterior surface and

shaped as in A. galerita. Mandible (Figure 3a)

2-segmented, with about 37 to 41 teeth on convex

side and 32 to 34 teeth on concave side. Paragnath

(Figure 36) bearing rather few but coarser spinules.

First maxilla (Figure 3c) bearing only 2 elements,

with spinules on both inner and outer surfaces. Second maxilla (Figure 3d) 2-segmented, of a typical

form, bearing a row of about 16 teeth on terminal

process. Maxilliped (Figure 3e) 3-segmented; first
NUMBER 8 7

FIGURE 2.—Acanthochondria exilipes female: a, body of syntype, dorsal; b, same, lateral; c,

body of a contracted specimen, dorsal; d, genito-abdomen, lateral; e, caudal ramus; /, first

antenna; g, second antenna. Scale: 1 mm in a, b, c; 0.1 mm in d, f, g; 0.01 mm in *.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Acanthochondria exilipes female: a, mandible; b, paragnath; c, first maxilla; d,

second maxilla; e, maxilliped; /, leg 1; g, slightly abnormal leg 1; h, abnormal leg 1; i, leg 2.

Scale: 0.01 mm in a, b, c, d, e; 0.1 mm in /, g, h, i.

segment unarmed, second segment with rows of denticles distributed in two patches, and terminal segment a claw with a small hooklet.

Leg 1 (Figure 3/) much smaller than leg 2 (Figure 3i). Both legs with longer endopod; exopod

shorter than protopod. A minute outer seta on both

protopods. An abnormal leg 1 (Figure 3h) seen on

left side of a specimen which has a slightly abnormal

leg (Figure 3g) on the opposite side.

MEASUREMENTS.—Body 8.74 mm in extended

specimen and 7.44 mm in contracted one; head 1.78

X 1-31 mm; genital segment 0.49 X 0-74 mm; abdomen 0. 53 X 0.38 mm; egg sacs 7.36 mm and 7.99

mm; egg 184 /*.
NUMBER 8 7

a

FIGURE 4.—Acanthochondria exilipes male: a, body, lateral; b, first and second antennae; c,

terminal blade of mandible; d, leg 1; e, leg 2. Scale: 0.1 mm in a; 0.01 mm in b, c, d, e.

MALE.—Body (Figure 4a) 0.38 X 0.41 mm, with

cephalothorax slightly longer than metasome plus

urosome. Metamerism on body rather distinct. A

small seta on lateral surface of second pedigerous

segment. Genital segment with usual ventrolateral

ridge. Caudal ramus different from female in having

rather indistinct knoblike process and evenly tapered

terminal process.

First antenna (Figure Ab) cylindrical, armature as

usual, being: 1-1-2-2-8. Second antenna (Figure

46) 2-segmented, with a seta on each segment;

terminal hook rather small. Rostral area bearing a

median protrusion.

Labrum with sexual dimorphism as observed in A.

phycidis. Mandible (Figure 4c) bearing 18 to 19

teeth on convex side and 11 to 13 teeth on concave

side, where there is an additional row of 3 teeth.

Paragnath and maxilliped resembling those of female.

First and second maxilla with same sexual dimorphism as in A. galerita.

Leg 1 (Figure 4rf) larger than leg 2 (Figure 4e),

both bearing a long outer seta on protopod. First

exopod different from second in having 2 bipartite

elements.

REMARKS.—As pointed out by Wilson (1932: 502)

this species can be easily recognized by its posterior

processes being longer than the last trunk segment

(fourth pedigerous segment). Although this character

is shared with A. soleae (Kr^yer 1838), A. exilipes

can be distinguished from it by its small first antenna

(in relation to the size of the head), the elliptical

head, and the nature of the rami on both legs.

Acanthochondria galerita (Rathbun 1886)

FIGURES 5-7

Chondracanthus galeritus Rathbun 1886: 317-319, pi. 8:

figs. 1-7; pi. 10: figs. 1-7.

Acanthochondria galerita (Rathbun).—Oakley 1930: 186.—

Wilson 1932: 505, figs. 209b.—Yamaguti 1963: 277.

A. albigutta Pearse 1952a: 225-227, figs. 103-110.—Causey

1955a: 12, pi. 1: figs. 6-7; 1960: 333.—Yamaguti 1963:

276.

MATERIAL STUDIED.—From USNM: (following

collections are all from oral cavities of Paralichthys
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 5.—Acanthochondria galerita female: a, body of syntype, dorsal; b, same, lateral; c,

body of a contracted specimen, dorsal; d, genito-abdomen, ventral; e, caudal ramus; /, first

antenna; g, second antenna. Scale: 1 mm in a. b, c; 0.1 mm in d, e, f, g.
NUMBER 8 7

dent at us (Linnaeus) in Woods Hole, Massachusetts)

14 females, with males attached, and 1 young female,

8 August 1883 (USNM 6036, co-type) ; 6 females,

with males attached, 28 August 1883 (USNM 6077,

co-type) ; 5 females, with males attached, 30 August

1883 (USNM 6082) ; 1 female, with male attached,

8 August 1883; 28 females, with males attached, 14

August 1883; 15 females, with males attached, 15 July

1904 (USNM 59782); 18 females, with males attached, 15 August 1900 (USNM 59814) ; 6 females,

with males attached, 10 July 1906 (USNM 54129);

and 2 females, with males attached, 29 July 1913

(USNM 79599).

The following two collections were labeled as

Acanthochondria albigutta Pearse and mounted on

slide: 1 females, with male attached, on P. albiguttus

Jordan and Gilbert, Alligator Harbor, Florida, 10

May 1952 (USNM 93696, type) ; and 1 young female,

with a copepodid male attached, on P. albiguttus,

Alligator Harbor, Florida, 1 June 1952 (USNM

93697).

FEMALE.—Body (Figures 5a, b, c) of varied appearance according to different states of contraction.

Head deflected backward in extended specimens

(Figure 5b) and indistinctly divided into anterodorsal

"hood" portion (with dorsal carapace) and posteroventral "cheek" portion which is the main contractile

portion of the head and bears mouth parts on its

posteroventral surface. Oral region flanked by a pair

of small processes (Figure 6a). First pedigerous segment narrower than head, but second wider. Third

pedigerous segment with rounded and slightly protruded anterior corners in contracted specimen (Figure 5c) but not in extended specimen (Figure 5a).

Posterior processes short and blunt. A pair of small

vermiform processes (Figure 5d) located on posterior

surface of trunk. Genital segment (Figure 5d) much

larger than abdomen, with a pair of hyaline, obtuse

elements on ventral surface. Abdomen (Figure 5d)

carrying a pair of setules on anterodorsal surface.

Caudal ramus (Figure 5e) bearing 3 setae and 1 blunt

knob, and terminating in a characteristic spiniform

process as shown in figure. Egg sac nearly as long as

body, containing many rows of small eggs.

First antenna (Figure 5/) large, with swollen base,

armed with 6 terminal and 2 subterminal setae. Second antenna (Figure 5g) indistinctly 2-segmented,

terminal segment a slender, recurved hook.

Labrum (Figure 6a) with denticles on posterior

surface. Mandible (Figure 6b) 2-segmented, terminal

blade bearing about 57 slender teeth on convex side

and 37 on concave side. Paragnath (Figure 6c) bilobed, with spinulose ventral lobe. First maxilla

(Figure 6d) bearing an element. Second maxilla

2-segmented, with distal process (Figure 6e) bearing

a small, simple setule, a large, spiniform seta with

hyaline tip, and a row of about 12 teeth. Maxilliped

(Figure 6/) 3-segmented, proximal 2 segments of

usual form, but terminal hook being peculiar in having 2 hooklets on concave surface and a patch of

spinules on outer surface at base.

Leg 1 (Figure 6g) much smaller than leg 2 (Figure 6h), both having bluntly pointed rami and

slightly smaller exopod. Protopod bearing a small

outer seta.

MEASUREMENTS.—Body 6.70 mm in extended

specimen and 4.65 mm in contracted one; head

(carapace) 1.36 X 1-31 mm; genital segment 0.36 X

0.46 mm; abdomen 0.30 X 0.18 mm; egg sacs 6.32

mm and 6.51 mm; egg 162 /A.

MALE.—Body (Figure 7a) 0.53 X 0.29 mm, having rather small metasome plus urosome, which

together are only about half as long as the cephalosome. Main flexure on body located between first

and second pedigerous segment. A setule on lateral

surface of third pedigerous segment. Genital segment

with usual ventro-lateral ridges. Abdomen greatly

reduced, bearing a pair of small setules on dorsal

surface. Caudal ramus resembling that of female.

First antenna (Figure 76) cylindrical, bearing usual

armature of 1-1-2-2-8. Second antenna (Figure 7b)

2-segmented, bearing one seta on each segment.

Rostral area with median protrusion located between

bases of first and second antennae.

Labrum (Figure 7c) different from female in having median protrusion and lateral conical processes.

Mandible with 33 teeth on concave side and 16 on

convex side. Paragnath as in female. First maxilla

(Figure 7d) with slight sexual dimorphism in the

three elements. Second maxilla without teeth on

terminal process. Maxilliped different from female in

terminal hook lacking basal patch of spinules and

with only one hooklet.

Leg 1 (Figure le) and leg 2 (Figure 7/) alike,

only different in size. Both with long outer seta on

protopod, bifurcate exopod, and simple endopod.

REMARKS.-—The most outstanding features of this

species are the bluntly pointed rami of the legs, the
10

a

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

d

FIGURE 6.—Acanthochondria galerita female: a, labrum and oral lobes, ventral; b, mandible;

c, paragnath, d, first maxilla; e, terminal segment of second maxilla; /, maxilliped; g, leg 1; h,

leg 2. Scale: 0.1 mm in a, g, h; 0.01 mm in b, c, d, e, f.

abruptly narrowed terminal processes on the caudal

ramus, and the hoodlike head which remains distinct

regardless of the state of contraction.

The species described by Pearse (1952a) as A.

albigutta is in all probability conspecific with the

present species. The rather pointed rami in the two

pairs of legs, the swollen first antenna, the denticulated posterior margin of the labrum, and the hoodlike

head are all reminiscent of A. galerita. The only

major difference that I can see is the possession of a

pair of relatively narrow posterior processes. Since all

available specimens in the collection of USNM

(Pearse's type specimens) are mounted in balsam on

slides, it is difficult to determine whether this difference is real or the result of distortion after mounting.

Of the two type specimens of A. albigutta, one is
NUMBER 8 7 11

a

FIGURE 7.—Acanthochondria galerita male: a, body, lateral; b, first and second antennae; c,

labrum, ventral; d, first maxilla; e, leg 1; /, leg 2. Scale: 0.1 mm in a; 0.01 mm in b, c, d, e, f.

ovigerous (No. 93693, measuring 9.60 X 3.52 mm)

and the other nonovigerous (No. 93697, measuring

4.86 X 1-39 mm). It is noteworthy that the former

carries a typical chondracanthid-form dwarf male and

the latter, which was designated by Pearse as "type"

of A. albigutta, an unmodified copepodid male. Although I did not have the opportunity to reexamine

Causey's collections of A. albigutta, his "supplementary material" (Causey 1955a: 12, pi. 1 : figs. 6, 7)

provide further evidence that A. albigutta is synonymous with A. galerita.

Acanthochondria phycidis (Rathbun 1886)

FIGURES 8-10

Chondracanthus phycidis Rathbun 1886: 320-323, pi. 9:

figs. 1-6, pi. 10: figs. 8-13.—Fowler 1912: 476.

Chondracanthus chilensis Wilson 1918: 11-15, figs. 1-6.—

Stuardo and Fagetti 1960: 200.

Acanthochondria phycidis (Rathbun).—Oakley 1930: 186.—

Wilson 1932: 504-505, figs. 299c—Yamaguti 1963: 278.

A. putputae Oakley 1930: 193-195, figs. 6A-D.—Yamaguti

1963: 278.

A. chilensis (Wilson).—Oakley 1930: 186.—Yamaguti

1963: 276.

MATERIAL STUDIED.—From USNM: 3 females and

2 young females, with males attached, off Martha's

Vineyard, Massachusetts, 1883 (USNM 6066, type);

7 females and 8 young females, with males attached,

on gills of Urophycis tenuis (Mitchill), off Martha's

Vineyard, Massachusetts, 1883 (USNM 6067, cotype) ; 1 female on "trout," Falkland Islands, 13 April

1927 (USNM 60481, labeled as A. clavata); 5

females and 2 young females, with males attached, on

"mullet" in Port Stanley, Falkland Islands, 15 March

1927 (USNM 60485, labeled as Chondracanthodes

deflexus); 3 females and 1 young female, with males

attached, on gills of Merluccius gayi (Guich), Valparaiso, Chile, March 1961 (USNM 51043). From

author's collection: 28 females, with males attached,

at bases of gill arches of U. regius (Walbaum), from
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

following ten stations of R/V Oregon Cruise 105-

5741 (1 female, 29°45'N 80°13'W -» 29°38/N

80°12'W), 5757 (3 females, 30°03'N 80°07'W -»

29°56'N 80°l(nV), 5759 (3 females, 30°02'N

80°08'W -+ 29°35'N 80°09'W), 5763 (4 females

29°59'N 80°09'W -> 29°51'N SO'ICAV), 5767 (4

females, 29°51'N 80°ll'W -• 29°44'N 80°ll'W),

5791 (1 female, 24°23'N 82°42'W -* 24°23'N

82°47/W), 5793 (3 females, 24°18/N 82°53'N ->

24°19'N 83o00'W), 5795 (3 females, 24°16/N

82°30/W -* 24O16'N 82°38'W), 5798 (3 females,

29°14'N 80°05'W -+ 29°09'N 80°03'W), and 5802

(3 females, 29°15'N 80°05"W -+ 29°09'N 80°02'W).

FEMALE.—Body (Figures 8a, b) rather short and

stout. Head nearly as long as wide, with greatest

width in its posterior two thirds, and the narrowed

anterior margin less than half of this width. Posteroventral surface of head at oral area greatly swollen,

with lateral swellings visible in dorsal view. First

pedigerous segment broadest, with posterior corners

slightly indented and forming a small lobe. Fourth

pedigerous segment forming last trunk segment, which

a

\

g

FIGURE 8.—Acanthochondria phycidis female: a, body, dorsal; b, same, lateral; c, posterior

part of trunk and genito-abdomen, dorsal; d, genito-abdomen, ventral; e, caudal ramus; /, first

antenna; g, second antenna. Scale: 1 mm in a, b, c; 0.1 mm in d, e, f, g.
NUMBER 8 7 13

FIGURE 9.—Acanthichondria phycidis female: a, mandible; b, paragnath; c, first maxilla; d,

second maxilla; e, maxilliped; /, leg 1; g, leg 2. Scale: 0.01 mm in a, b, c, d, e; 0.1 mm in /, g.

is characteristically bent ventrad in all alcoholized

specimens; protruding posterolaterally into a stout,

blunt posterior process (Figure 8c). Genital segment

(Figure 8d) with egg sac attachment area situated

dorsolaterally and bearing a pair of blunt, hyaline

setae at midventral region. Abdomen (Figure 8d)

oval, bearing a pair of setules on anterodorsal surface.

Caudal ramus (Figure 8^) of usual form, located on

anteroventral surface of abdomen. Egg sac cylindrical,

longer than body.

First antenna (Figure 8/) fleshy, only slightly

swollen at basal portion, with usual armature of

1-1-2-2-8. Second antenna (Figure 8g) 2-segmented,

terminal segment a recurved claw.

Labrum shaped as in A. galerita and fringed with

dentic'es on posterior margin. Mandible (Figure 9a)

bearing about 58 teeth on convex side and 44 teeth

on concave side. Paragnath (Figure 9b) bilobate with

a smaller naked basal lobe and a larger spinulose

terminal lobe. First maxilla (Figure 9c) an oval lobe

bearing 2 terminal elements and covered with small

spinules on dorsal (posterior) and inner surfaces.

Second maxilla (Figure 9d) of usual form, bearing a

row of 12 teeth on terminal process. Maxilliped (Figure 9e) short and stout, with usual armature.

Both leg 1 (Figure 9/) and leg 2 (Figure 9g) indistinctly bilobate, stout, and covered with very fine

spinules; outer seta on protopod very small; and

exopod indistinguishable from protopod.

MEASUREMENTS.—Body 6.32 mm; head 1.75 X

1.70 mm; genital segment 0.38 X 0-53 mm; abdomen 0.55 X 0.35 mm; egg sacs 7.06 mm and 7.95

mm; egg 232 /*.

MALE.—Body (Figure 10a) 0.80 X 0.37 mm; with

widest portion on posterior part of cephalosome which

is oval in dorsal view. Second and third pedigerous

segments rather well separated, but fourth pedigerous

segment indistinctly fused with urosome. A small seta

on dorsolateral surface of second and third pedigerous segments. Abdominal portion (Figure \0b) bearing a pair of setules on dorsal surface. Caudal ramus

(Figure 106) as in female.

First antenna (Figure 10c) elongate and cylindrical, armature as in female. Second antenna (Figure
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 10.—Acanthochondria phycidis male: a, body, lateral; b, posterior part of body, dorsal;

c, first and second antennae; d, labrum, ventral; e, second maxilla; /, leg 1; g, leg 2. Scale: 0.1

mm in a, b; 0.01 mm in c, d, e, f, g.

10c) 2-segmented, terminal segment a recurved hook

bearing a small hyaline element on outer surface. A

blunt knob in rostral area between bases of two pairs

of antennae.

Labrum (Figure \0d) shaped as in female, but

having a rounded median knob. Mandible with usual

sexual dimorphism. Paragnath and first maxilla as in

female. Second maxilla (Figure lOe) with simple

short terminal process. Maxilliped showing very slight

sexual dimorphism in having indistinct hooklet on

terminal claw.

Leg 1 (Figure 10/) more swollen than leg 2 (Figure lOg), both bearing a long outer seta on protopod,

a bluntly pointed endopod, and a smaller exopod
NUMBER 8 7 15

composed of two elements. Number of elements in

exopod of leg 1 variable, ranging from one to three,

sometimes even different between two members of

same pair.

REMARKS.—A. phycidis is distinctive in having a

small cylindrical first antenna, lateral swellings in

head, indistinctly bilobated thoracic legs, and stubby

posterior processes. The very characteristic poorly

bilobated condition of the thoracic legs is also known

in A. limandae (Kr^yer 1863), but the latter is

different from A. phycidis in having a pair of large,

swollen, fleshy first antennae.

A. pur pur ae Oakley 1930 is apparently conspecific

with this species. The only difference between the two

species is their size of the body, the former being 10.5

mm long (Oakley 1930:193), while the largest specimen of A. phycidis is only 6.32 mm in length. The

seven specimens contained in USNM 60485 are

definitely attributable to what Oakley has described

as A. purpurae. They were, however, mislabeled as

"Chondracanthodes defiexus." These specimens were

found on "mullet" from the same locality of Oakley's

A. purpurae (Falkland Islands). A closer observation

of these seven South American specimens has further

confirmed that Oakley's A. purpurae is the same

species as A. phycidis. The largest specimen measures

11.16 mm, nearly twice as long as the largest North

American specimen.

A. chilensis ( = C. chilensis) is apparently another

synonym of A. phycidis. It was first reported by Wilson

(1918) on the gills of the hake (Merluccius gayi)

from Valparaiso, Chile. When Stuardo and Fagetti

(1960) made an extensive survey of the copepod

parasites of the same species of hake in the Bay of

Valparaiso, however, they did not find again this

particular parasite. The largest type specimen of A.

chilensis in USNM 51043 measures 11.53 X 4.92

mm, which is much larger than the North Atlantic

specimens and about the size of those found in the

Falkland Islands.

Chondracanthodes radiatus (Miiller 1776)

FIGURE 11

Lernaea radiate Miiller 1776: 226; 1788: 39, pi. 33: fig.

4 .- H. Milne Edwards 1840: 504.

Anops radiatus (Miiller).—Oken 1815: 357.

Entomoda radiata (Miiller).—Lamarck 1816: 233.

Lernentoma radiata (Miiller).—Blainville 1822: 440.

Chondracanthus radiatus (Miiller).—Cuvier 1830: 258.—

Kroyer 1863: 251, pi. 14: figs, la-e.—Bassett-Smith

1899: 494.—Brian 1908: 17.—Stephensen 1913: 341.—

Wilson 1920: 6L.—Hansen 1923: 45-46.—Oakley 1930:

187.

Chondracanthoides radiatus (Miiller).—Yamaguti, 1963:

283.

MATERIAL STUDIED.—From UZM: 2 females and

1 young female on Macrurus fabricii L., Greenland.

From author's collection: 1 female on wall of operculum of Macrurus bairdii (Goode and Bean), Sta.

5753 (29°29'N 79°56'W -»29°23'N 79°57'W) of

R/V Oregon Cruise 105.

FEMALE.—Body (Figures lla, b, c) greatly swollen

and stout. Head distinctly wider than long, deflected

ventrally, and extended posterolaterally into a long

attenuated process; dorsal surface bearing a median

sclerite and a small swelling in posterior region. First

pedigerous segment incompletely separated from

cephalosome, bearing a large, rounded, dorsal swelling. Trunk voluminous, divided into three portions,

with each portion bearing a pair of processes: first

pair on anterior corner of trunk, with a small basal

swelling on anterior surface; second pair on midlateral portion of trunk and bent ventrally; and third

pair on posterior corner of trunk directed posteriorly.

Genito-abdomen and caudal ramus of usual form.

Egg sac unknown. First antenna (with right one

broken off in the specimen drawn) large, fleshy, and

tipped with 8 setae. Second antenna 2-segmented,

with uncinate terminal segment. Oral appendages as

in C. defiexus. Two pairs of modified legs present (see

Figure lie), with greatly swollen protopod protruding laterally into a blunt process and carrying a small

outer seta; both rami rod-like, covered with fine denticles; endopod longer than exopod.

MEASUREMENTS.—Body 6.51 mm; head 1.59 X

2.33 mm.

MALE.—Not seen.

REMARKS.—This species is similar to C. tuberofurcatus Kabata and Gusev 1966 in having a pair of

large, fleshy first antennae and a pair of posterolateral

processes in the head. The cephalic process in C.

radiatus, however, is much longer and attenuated and

the first pair of processes in the trunk region is not

furcated distally as in C. tuberofurcatus.

Two of Kr^yer's three type specimens of C. radiatus

deposited in UZM are ovigerous and the third one is

a young nonovigerous female. This young female
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 11.—Chondracanthodes radiatus female: a, body, dorsal; b, same, lateral; c, same,

ventral; d, leg 1 of holotype. Scale: 1 mm in a, b, c; 0.1 mm in d.
NUMBER 8 7 17

measures 5.39 mm and has all the body processes developed as in the ovigerous one, but they are shorter

and smaller. One of the two ovigerous females has

all mouth parts and the first pair of legs taken off,

and the other one has the right second antenna and

all oral appendages except the maxillipeds removed.

The dissected two first legs were kept in a separate

vial; one of them is here illustrated in Figure lid.

These two ovigerous females measure 9.38 mm and

10.21 mm in length respectively, and a detached complete egg sac is 14.98 mm long. No male was found,

although it was described by Kn£yer.

The specimen taken off Florida differs from the

ones from Greenland in having a smaller body, more

attenuated body processes, and a comparatively

swollen trunk. These differences are reg£ .*ded as

merely geographical variations.

As pointed out by Hansen (1923:46), the specimen described by Brian (1912) as C. radiatus is not

a

the same as Miiller's radiatus. From Brian's description and figures, it seems to me that he was dealing

with a specimen of C. deflexus. The possession of

short, clavate, and simple lateral processes on the

body, small first antennae, and the absence of cephalic

processes are all very convincing.

Chondracanthus cottunculi Rathbun 1886

FIGURES 12-14

Chondracanthus cottunculi Rathbun 1886: 322-324, pi. 11:

figs. 1-7.—Fowler 1912: 475.—Wilson 1932: 477-498,

fig. 297c—Yamaguti 1963: 272.

Chondracanthopsis cottunculi (Rathbun).—Oakley 1930:

196.

MATERIAL STUDIED.—From USNM: 1 female, with

male attached, on Cottunculus species, south of Block

Island, Rhode Island, 23 July 1884 (USNM 8454,

type) ; 2 females, with males attached, in branchial

FIGURE 12.—Chondracanthus cottunculi female: a, body of syntype, dorsal; b, same, lateral; c,

genito-abdomen, lateral; d, first and second antenna. Scale: 1 mm in a, b; 0.1 mm c, d.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cavities of C. thomsoni (Gunther), 1883 (USNM

6139) ; and 2 females, with males attached, in branchial cavities of C. microps Collet, off Martha's

Vineyard, Massachusetts, 21 September 1883 (USNM

6166).

FEMALE.—Body (Figures 12a, b) rather plump.

Head distinctly wider than long, with small rounded

knob at each anterior corner and swollen posterolateral surface (not in the form of a process). First

pedigerous segment distinctly narrower than head,

but second pedigerous segment protruded laterally

forming widest part of body. Third pedigerous segment larger than fourth and bearing a short, cylindrical ventral process. Posterior process blunt. Genital

segment (Figure 12c) about twice as large as abdomen, carrying a pair of midventral elements. Caudal

ram us (Figure 12c) of typical form. Egg sac only a

little shorter than body, with many rows of small eggs.

First antenna (Figure I2d) large, flattened subtriangular, and fleshy; armed with 8 terminal setae

and an isolated seta on posterior surface of distal

portion. Second antenna (Figure I2d) of usual form,

2-segmented and uncinate.

Labrum with smooth posterior margin. Mandible

(Figure 13a) 2-segmented, terminal blade bearing

about 39 teeth on convex side and 37 teeth on concave

side. Paragnath (Figure 136) a bilobate lobe with

spinules on inner surface of terminal lobe. First

maxilla (Figure 13c) bearing 2 elements and an

inner swelling. Second maxilla (Figure \3d) 2-segmented, bearing a row of about 13 teeth on terminal

process. Maxilliped (Figure 13e) 3-segmented unarmed, second segment with rows of denticles distributed in two patches, and terminal segment a short

claw with a hooklet.

Leg 1 (Figure 13/) smaller than leg 2 (Figure 13g),

both bilobate, with stout conical endopod and longer

exopod with bent tip. A small outer seta on both

protopods.

MEASUREMENTS.—Body 7.69 mm (in extended

specimen); head 1.33 X 1-73 mm; genital segment

0.48 X 0.65 mm; abdomen 0.38 X 0.31 mm; egg

sacs 6.07 mm and 6.24 mm; egg 192 /*.

MALE.—Body (Figure 14a) 0.95 X 0.49 mm, with

distinct metamerism on body; cephalosome about as

long as metasome plus urosome, but distinctly wider.

FIGURE 13.—Chondracanthus cottunculi female: a, mandible; b, paragnath, c, first maxilla; d,

second maxilla; *, maxilliped; /, leg 1; g, leg 2. Scale: 0.1 mm in a, d, e, f, g; 0.01 mm in b, c.
NUMBER 8 7 19

FIGURE 14.—Chondracanthus cottunculi male: a, body, lateral; b, urosome, ventral; c, first

and second antennae; d, mouth parts, ventral; e, leg 1; /, leg 2. Scale: 0.1 mm in a, b; 0.01 mm

in b, c, d, e, f.

A small seta on lateral surface of second and third

pedigerous segment. Genital segment and abdomen

relatively well separated (Figure 146). A pair of

spermatophores (Figure 146) seen inside body of

one male. Caudal ramus (Figure 146) as in female.

First antenna (Figure 14c) cylindrical, armature

being: 1-1-2-2-8. Second antenna (Figure 14c)

2-segmented, with terminal recurved hook bearing

a bifurcate process (accessary antennule) on its ventrolateral surface. A median protrusion located between bases of first and second antenna (see Figure

14c).

Labrum (Figure I4d) as in female, with smooth

posterior margin. Mandible (Figure I4d) and first

maxilla (Figure 14^) with usual sexual dimorphism.

Paragnath (Figure 14rf) and maxilliped as in female.

Second maxilla differing from female in having a row

of only 4 teeth on terminal process.

Leg 1 (Figure 14^) larger than leg 2 (Figure 14/),

both with rather prominent rami in the form of a

cylindrical process. Exopods larger than endopods;

tipped with 3 elements in first but only 2 in second.

A long outer seta on each protopod.

REMARKS.—The male of this species was neither

described nor figured in Rathbun's (1886) original

description; he only remarked (page 324) that "males

were attached to most of the females." Wilson (1932:

498) stated that the male was "unknown." The male

is, however, in reality rather peculiar in having an

accessary antennule-like element on the second an-
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tenna. A similar element was also found in the male

of C. pinguis Wilson 1912 by Kabata (1968).

C. cottunculi was transferred to the genus Chondracanthopsis by Oakley (1930). Heegaard (1945)

suggested that it should be included in Acanthocanthopsis. However, as stated by Ho (1970:9), these

two genera are not valid.

This species can very easily be distinguished from

other species of Chondracanthus by the following

combination of characters: without a cephalic process,

only one pair of processes on the body, short posterior

processes, and a large, subtriangular first antenna.

Chondracanthus janebennettae Causey 1953

FIGURES 15-17

Chondracanthus janebennettae Causey 1953: 10, fig. 10;

1955a: 11-12, pi. 3: figs. 13-16.—Yamaguti 1963: 273,

pi. 246: fig. 2.

MATERIAL STUDIED.—From USNM: 1 female, with

male attached, on flounder, Grand Isle, Louisiana

(USNM 79714, labeled as C. distortus). From auwith male attached, on Paralichthys lethostigmus

Jordan and Gilbert, Pascagoula, Mississippi (USNM

97622); 1 female, with male attached, on P. lethostigmus, Pascagoula, Mississippi (USNM 97623) ;

and 2 females, with males attached, on gills of P.

albiguttus Jordan and Gilbert, Grande Isle, Louisiana

(USNM 79714, labeled as C. distortus). From author's collection: 11 females, with males attached, on

wall of opercula of Ancylopsetta quadrocellata Gill,

St. George Island, Florida, 12 July 1965.

FEMALE.—Body (Figures 15a, b) rather broad and

flat. Head well-demarcated from first pedigerous

segment, with posterodorsal surface and posteroventral corners protruded into a large, blunt lobe; a

pair of small processes in oral area. An elongate

sclerite on middorsal surface anterior to dorsal lobe

(see Figure 15c). First and second pedigerous segments narrower than head, forming an indistinct neck

region. Third pedigerous segment broadest, with

posterior corners protruded backward into a long,

attenuate process. Fourth pedigerous segment (Figure

\bd) shaped as the third, but narrower; its posterior

corners protruded into a long, attenuate posterior

process. Genital segment (Figure \be) small, attached

to posterodorsal surface of trunk. Abdomen (Figure

15*) transformed into long, attenuate caudal process,

carrying caudal rami at its basal portion on lateroventral surface. Caudal ramus (Figure 15/) armed

with 3 setae (1 outer and 2 inner) on basal portion;

spinulose terminal portion with forked tip. Egg sac

cylindrical, longer than body, with only a few rows

of eggs.

First antenna (Figure 15g) fleshy, with basal portion greatly enlarged and distal half narrowed and

turned ventrally, bearing 11 small setae on terminal

and subterminal surface. Second antenna (Figure

15/J) 2-segmented, terminal segment a strongly recurved, slender hook.

Labrum (Figure 16a) bearing denticles on posterior

surface. Mandible (Figure 16&) 2-segmented, terminal segment bearing on convex side a complete row

of 106 teeth and another incomplete row of 23 teeth,

and on concave side a row of 66 teeth. Paragnath

(Figure 16c) a small lobe armed with several spinules.

First maxilla (Figure 16a1) an oval lobe bearing 2

elements. Second maxilla (Figure 16^) 2-segmented

robust and unarmed; second segment an attenuate

process, bearing 1 small simple seta, 1 large spinulose

seta, a row of about 16 teeth on posterior surface, and

a single tooth on anterior surface close to tip. Maxilliped (Figure 16/) 3-segmented; first segment unarmed; second segment bearing rows of spinules

distributed in two patches; and distal segment a claw

with a patch of denticles on base and 6 to 8 small

blunt teeth on inner edge of hook.

Leg 1 (Figure 16g) bilobate, with small protopod

bearing an outer seta, both endopod and exopod long

and attenuate. Leg 2 (Figure 16/i) as in leg 1, only

with more slender rami.

MEASUREMENTS.—Body 5.92 mm; head 1.50 X

1.68 mm; genital segment 0.38 X 0.50 mm; abdomen

1.17 X 0.23 mm; egg sacs 8.37 mm and 7.81 mm;

egg 156 ft.

MALE.—-Body (Figure 17a) 0.63 X 0.30 mm, with

widest portion in posterior part of cephalosome,

which is incompletely fused with first pedigerous segment. Metamerism on body rather distinct. Genital

segment (Figure 17b) rather wide, but abdomen

narrow and protruded into a small lobe posteriorly.

Caudal ramus (Figure 176) as in female.

First antenna (Figure 17c) cylindrical, armature

being: 1-1-1-3-8. Second antenna (Figure 17c)

2-segmented, with a recurved terminal segment.
NUMBER 8 7 21

FIGURE 15.—Chondracanthus janebennettae female: a, body, dorsal; b, same, lateral; c, head,

anterodorsal; d, posterior part of body, dorsal; e, genito-abdomen, ventral; /, caudal ramus; g,

first antenna; h, second antenna. Scale: 1 mm in a, b, c, d; 0.1 mm in e, f, g, h.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 16.—Chondracanthus janebennettae female: a, oral area and mouth parts, ventral; b,

mandible; c, paragnath; d, first maxilla; e, second maxilla; /, maxilliped; g, leg 1; h, leg 2.

Scale: 0.1 mm a, f; 0.01 mm b, c, d, e; 1 mm in f, g, h.

Labrum, paragnath, and first maxilla as in female.

Mandible and second maxilla (Figure \ld) exhibiting

usual sexual dimorphism. Maxilliped (Figure Me)

different from female in having only one hooklet on

terminal claw.

Leg 1 (Figure 17/) and leg 2 similar, biramous,

with large, robust protopod bearing a long outer seta.

Exopod a slender process with forked tip and endopod

a small lobe tipped with a conical knob.

REMARKS.—The present species is characteristic in

having, in the female, attenuated lateral processes on

trunk, a large crestlike dorsal lobe on the cephalosome,

and the abdomen prolonged into an attenuate, cylindrical caudal process. The last character is not found

in any other known species of Chondracanthus. The

presence of a large number of teeth on the mandible,

the second maxilla, and the terminal claw of the

maxilliped is also unique in the genus.
NUMBER 8 7 23

FIGURE 17.—Chondracanthus janebennettae male: a, body, lateral; b, posterior part of body,

ventral; c, first and second antenna; d, second maxilla; e, maxilliped; /, leg 1. Scale 0.1 mm

in a, b; 0.01 mm c, d, e, f.

Chondracanthus merluccii (Holten 1802)

FIGURES 18-20

Lernaea merluccii Holten 1802: 135-136, pi. 3: fig. 2.

Chondracanthus merluccii (Holten).—Kroyer 1837: 278-

280, pi. 3: figs. 9a-d.—H. Milne Edwards 1840: 503.—

Heller 1866: 756.—Olsson 1868: 34.—Richiardi 1880:

151.—Stossich 1880: 253.—Valle 1880: 73.—Cams

1885: 354.—T. Scott 1892: 262; 1900: 166-167; 1906:

372.—Bassett-Smith 1896: 161; 1899: 494.—Schimkewitsch 1896: 339, pi. 14: figs. 5-7, 11-20.—Brian 1898:

21; 1906: 97, pi. 7: fig. 4.—Norman and T. Scott 1906:

217.—T. Scott and A. Scott 1913: 180, pi. 20: fig. 10;

pi. 47: fig. 8; pi. 53: figs. 16-19.—Stock 1915: 71.—

Oakley 1930: 188, figs. 2A-D.—Wilson 1932: 498, pi. 1:

fig. a, figs. 298a-e.—El Saby 1933: 865-866, pi. 2: figs.

7-9.—Leigh-Sharpe 1934: 113.—Pesta 1934: 48, fig. 29.

—Oorde-de Lint and Schuurmans Stekhoven 1936: 109,

fig. 52.—Barnard 1948: 251; 1955: 286-287, figs. 23ab.—Carvalho 1951: 143.—Rose and Vaissiere 1952: 171.

—Nunes-Ruivo 1956: 35-36—Szidat 1955: 2; 1961:

3, fig. lb.— Markewitsch 1956: 85, fig. 39. Capart

1959: 111.—Yamaguti 1963: 273, pi. 248: fig. 5.—

O'Riordan 1966: 376.

C. xyphiae Guerin 1829-44, pi. 9: fig. 20.

C. merlangi Kroyer 1863: 331.—Yamaguti 1963: 273.

C. itramineus Wilson 1923a: 10-11, pi. 1: fig. 13> p i. 2:

figs. 14-15.—Yamaguti 1963: 274, pi. 248: fig. 2.

Chondracanthus species Wilson 1923b: 14.

MATERIAL STUDIED.—From USNM (all found in

oral cavities of Merluccius bilinearis (Mitchill) ) : 2
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

females, England, 1884 (USNM 8454); 8 females

and 3 young females, with males attached, fish market

in Washington, D. C. (USNM 37874) ; 20 females

and 8 young females, with males attached, Woods

Hole, Massachusetts, 22 July 1924 (USNM 56635) ;

19 females, with males attached, Woods Hole, Massachusetts, June 1914 (USNM 59783); 1 female,

Woods Hole, Massachusetts, 20 October 1903

(USNM 59786) ; 35 females, with males attached,

Woods Hole, Massachusetts, 13 August 1902 (USNM

59780) ; 15 females, with males attached, fish market

in Washington, D. C. (USNM 59775); 60 females,

with males attached, Woods Hole, Massachusetts, 20

October 1903 (USNM 59787); 33 females and 1

young female, with males attached, Woods Hole,

Massachusetts, 1 November 1898 (USNM 59797) ; 21

females and 4 young females, with males attached,

Woods Hole, Massachusetts, 14 August 1902 (USNM

59785) ; 34 females, with males attached, Casco Bay,

Maine, 14 August 1902 (USNM 59784) ; 19 females

with males attached, Block Island, Rhode Island, 14

June 1884 (USNM 59796) ; 2 females and 2 young

females, with males attached, Woods Hole, Massachusetts, 22 July 1924 (USNM 59815); 36 females,

with males attached, fish market in Washington, D. C,

5 January 1909; and 15 females, with males attached,

Casco Bay, Maine, 24 July 1873 (USNM 60543).

From author's collection: 6 females, with males attached, on roof of mouth cavities of M. bilinearis,

from stations 5763 (29°59'N 80°10/W -* 29°51'N

SOMtTW), 5793 (24°18'N 82°53'W -+ 24°19'N

83°OO'W), and 5795 (24°16'N 82°30'W - • 24°16'N

82°38'W) of R/V Oregon Cruise 105.

FEMALE.—Body (Figures 18a, b) with only one

pair of lateral processes. Head bearing a small process at each posterolateral corner. Anterior margin of

head (Figure 18c) truncated and much narrower

than rounded posterior margin. First pedigerous segment small, narrower than head. Second pedigerous

segment wider than head, with shoulderlike rounded

anterior corners. Third pedigerous segment broadest

carrying a large cylindrical, lateral process on each

lateroventral surface. Fourth pedigerous segment

produced posterolaterally into a blunt, cylindrical

posterior process and bearing a midventral swelling

just in front of genital segment (see Figure 186).

Genito-abdomen (Figure 18a") smaller than the ventral knob just in front of it. Abdomen (Figure 18d)

small. Caudal ramus (Figure I8e) attached ventrally

to abdomen in anterior area; basal portion bearing 1

dorsal and 2 ventral setae, and terminal portion an

attenuated process bearing spinules. Egg sac cylindrical, nearly twice as long as body.

First antenna (Figure 18/) fleshy, consisting of a

swollen basal portion and two small terminal segments, armature being: 1-1-2-2-8. Second antenna

(Figure ISg) 2-segmented, terminal segment a strong

recurved hook.

Labrum as in C. janebennettae. Mandible (Figure

19a) 2-segmented, terminal segment falcate, bearing

about 31 large teeth on convex side and 20 smaller

teeth on concave side. Paragnath (Figure 196) with

a small lobule at base; spinules only on distal surface.

First maxilla (Figure 19c) bearing two terminal elements and a small knob on inner surface. Second

maxilla (Figure 19a1) 2-segmented; first segment

robust and unarmed; second segment an attenuate

process, bearing 1 small simple seta, 1 large barbed

seta, and a row of about 9 large teeth. Maxilliped

(Figure \9e) 3-segmented; first segment unarmed;

second segment bearing rows of denticles distributed

in two patches, and distal segment a claw with a

hooklet.

Leg 1 (Figure 18c) bilobate, with bluntly pointed

rami; endopod larger than exopod; and only protopod

bearing an outer small seta. Leg 2 (Figure 19/)

nearly three times larger than leg 1, with cylindrical

rami longer than protopod, which carries a small

outer seta. Both legs covered with fine spinules.

MEASUREMENTS.—Body 10.44 mm; head 1.51 X

1.89 mm; genital segment 0.41 X 0.55 mm; abdomen

0.10 X 0.14 mm; egg sacs 11.23 mm and 10.94 mm;

192 ft

MALE.—Body (Figure 20a) 0.97 X 0.39 mm, with

widest portion on posterior part of cephalosome.

Cephalosome larger than metasome plus urosome.

Last two thoracic segments and genital completely

fused. Abdomen (Figure 206) carrying a pair of

setules on dorsal surface. Caudal ramus (Figure 206)

as in female but much smaller.

First antenna asymetrically developed in one of the

males from off Florida (Figure 20c). A normally developed first antenna with armature of 1-2-2-8.

Second antenna (Figure 20c) 2-segmented, with recurved terminal hook. Labrum and oral appendages

as in female, except mandible and second maxilla
NUMBER 8 7 25

a

FIGURE 18.—Chondracanthus merluccii female: a, body, dorsal; b, same, ventral; c, head and

leg 1, ventral; d, genito-abdomen, ventral; e, caudal ramus; /, first antenna; g, second antenna.

Scale: 1 mm in a, b, c; 0.1 mm in d, e, f, g.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Chondracanthus merluccii female: a, mandible; b, paragnath; c, first maxilla; d,

second maxilla; e, maxilliped; /, leg 2. Scale: 0.01 mm in a, b, c; 0.1 mm in d, e; 0.5 mm in /.

(Figure 20d) which exhibit usual sexual dimorphism.

A conical knob (Figure 20c) in rostral area located

between bases of two pairs of antennae.

Leg 1 (Figure 20<?) larger than leg 2 (Figure 20/),

both having an inflated protopod bearing a long outer

seta, a bluntly pointed endopod, and a rather sharply

pointed exopod carrying a seta.

REMARKS.—The considerably smaller specimens

taken from hakes caught off Florida (author's collection) are only about half of the size of specimens

taken in the other parts of the Atlantic Ocean. The

specimens from South Africa differ from those occurring in other waters in having a pair of very small

cephalic processes. This feature was emphasized by

Wilson (1932) in distinguishing them as a new species,

C. stramineous.

T. Scott and A. Scott (1913) have reported that

occasionally more than one male may be found on a

female, but, in the course of this study, I have not

encountered such a polyandrous condition.

C. palpifer Wilson 1912 is the only species of Chondracanthus that is closely related to the present

species. It was first reported by Wilson (1912) from

Nanaimo, British Columbia, and later by Fraser

(1920) from Vancouver, British Columbia, and by

Brian (1944) and Szidat (1955) from Argentina. It

is morphologically different from C. merluccii in the

absence of cephalic processes and in having unequally

developed rami in leg 2 (with the exopod distinctly

larger than the endopod). The occurrence of C.

palpifer on M. hubbsi Marini in the Western South

Atlantic and C. merluccii on M. bilinearis, M. merluccius, and M. capensis in the North Atlantic and

the Eastern South Atlantic seems to support Szidat's

(1955, 1961) hypothesis that M. hubbsi originated

from the North Pacific stock of hake (M. productus)

and not from the North Atlantic stock (M. bilinearis).

Szidat found that the parasite fauna of M. hubbsi

(occurring off the coast of Brazil and Argentina) is

nearly identical with that of M. productus in the

Eastern North Pacific but distinctly different from

that of M. bilinearis in the Western North Atlantic.

During Cruise 16 of R/V Anton Bruun, I had an

opportunity to examine more than 5000 hakes (M.

gayi gayi) caught in the trawl at twelve stations off

the Peruvian coast, but not a single one bore chondracanthid copepods. As mentioned previously in con-
NUMBER 8 7 27

FIGURE 20.— Chondracanthus merluccii male: a, body, lateral; b, posterior part of body, dorsal;

c, first and second antennae (left first antenna atrophied); d, second maxilla; e, leg 1; /,

leg 2. Scale: 0.1 mm in a, b, c; 0.01 mm in d, e, f.

nection with Acanthochondria phycidis, when Stuardo

and Fagetti (1960) made an extensive survey on the

copepod parasites of M. gayi gayi in the Bay of Valparaiso, Chile, they too did not find a single chondracanthid parasite on the hake.

Chondracanthus nodosus (Mtiller 1776)

FIGURES 21-23

Lernaea nodosa Miiller 1776: 227; 1788: 40, pi. 33: fig.

5.—Lamarck 1816: 231.

Lernentoma nodosa (Miiller).—Blainville 1822: 441.

Chondracanthus nodosus (Miiller).—Kroyer 1837: 196;

1838: 133-135, pi. 3: figs. 2a-f; 1863: 332.—H. Milne

Edwards 1840: 503.— Olsson 1868: 34.—Bassett-Smith

1899: 494.—T. Scott and A. Scott 1913: 176, pi. 48:

figs. 13-16, pi. 52: figs. 1-3.—Stephensen 1913: 341-

342.—Wilson 1920: 6L.—Hansen 1923: 45.—LeighSharpe and Perkins 1924: 291-294, figs. 4-6.—Heegaard

1947: 7, fig. 8.

C. williamsoni T. Scott 1909: 76-77, pi. 3: figs. 8-17.

Chondracanthopsis nodosus (Miiller).—Oakley 1930: 196,
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

figs. 7A-D.—Wilson 1932: 508, pi. 39: figs. 2a-g.—

Leigh-Sharpe 1933: 113.—Oorde-de Lint and Schuurmans

Stekhoven 1936: 111, figs. 60A, B.—Sinderman 1961:

HI.—Williams 1963: 510-516.—Yamaguti 1963: 284,

pi. 246: fig. 2.

MATERIAL STUDIED.—All recorded on Sebastes

marinus (Linnaeus). From UZM: 2 females, with

males attached, Greenland; 5 females, with males

attached, Greenland, December 1891; 1 female, with

FIGURE 21.—Chondracanthus nodosus female: a, body, dorsal; b, same, lateral; c, body of a

contracted specimen, lateral; d, genito-abdomen, lateral; e, caudal ramus; /, first antenna; g,

second antenna. Scale: 1 mm in a, b, c; 0.1 mm in d, f, g; 0.01 mm in e.
NUMBER 8 7 29

male attached, Norway; 10 females, with males attached, 3 July 1902, Norway; 4 females, with males

attached, outside Denmark; 6 females, with males attached, outside Denmark; 1 females, outside Denmark ; 4 females, with males attached, 11 July 1902,

Denmark; 2 females, 13 August 1868, Skagen, Denmark; and 7 females, with males attached, the

Kattegat, Denmark. From USNM: 4 females, with

males attached, Massachusetts Bay, 1878 (USNM

60541). From author's collection: 8 females, with

males attached, on wall of branchial cavities, Sta. 1-3

(41°47'N 69°34/W -> 41°46'N 69°33'W) of R/V

Albatross IV Cruise 66-3.

FEMALE.—Body (Figure 21a, b, c) with many short

and rounded protrusions. Head protruded posterolaterally into rounded swellings. Both neck and trunk

regions extensible; an extended specimen (Figure

216) with longer neck and lower dorsal knobs, but a

contracted specimen (Figure 21c) showing shorter

neck and higher dorsal knobs. Second pedigerous

segment protruded laterally into a large lobe. Trunk

region bearing five lateral knobs, two ventral protrusions (which probably indicate respectively the

third and fourth pedigerous segments), and a pair

of posterior processes; four dorsal knobs variable in

height with different state of contraction. Genital

segment (Figure 21a") carrying two midventral elements as usual in chondracanthids. Abdomen (Figglobular, carrying a pair of small setules on

dorsal surface. Caudal ramus (Figure 2\e) bearing

on basal portion 3 setae and a small lobule; terminal

portion attenuated into spinulose process. Egg sac

long, cylindrical, with many rows of eggs.

First antenna (Figure 21/) fleshy; with enlarged

basal portion carrying two elements and with small

cylindrical terminal portion, twelve elements. Second

antenna (Figure 2\g) 2-segmented, with recurved

terminal segment.

Labrum with smooth posterior margin. Mandible

(Figure 22a) bearing 47 to 54 teeth on convex side

and 40 to 43 teeth on concave side. Paragnath (Figure 226) bilobated distally, both lobe spinulose. First

FIGURE 22.—Chondracanthus nodosus female: a, mandible; b, paragnath; c, first maxillia; d,

second maxilla; e, maxilliped; /, leg 1; g, leg 2. Scale: 0.01 mm in a, b, c; 0.1 mm d, #;

0.2 mm in /, g.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

maxilla (Figure 22c) bearing 2 terminal elements and

a basal lobule. Second maxilla (Figure 22d) 2-segmented; terminal segment an attenuated process

bearing 1 small simple setule, 1 large, spiniform seta,

a row of about 9 teeth on posterior surface, and a

single tooth on anterior surface. Maxilliped (Figure

22e) 3-segmented, armature of usual form.

Both leg 1 (Figure 22/) and leg 2 (Figure 22g)

similar in structure, with coxal portion probably bilaterally protruded into a pair of large alae and basis

and both rami fused to form the so-called "boot." A

small seta on outer surface of basis. Some specimens

have exopodal portion of "boot" bluntly attenuated

and much longer than endopodal portion. Surface of

both legs sparsely scattered with fine spinules.

MEASUREMENTS.—Body 6.70 mm; head 1.62 X

2.58 mm; genital segment 0.54 X 0.62 mm; abdomen

0.38 X 0.26 mm; egg sacs 7.35 mm and 7.09 mm;

egg 298 /x.

MALE.—Body (Figure 23a) small, 0.69 X 0.35

mm. Cephalosome rather thick and globular. Metamerism on metasome rather distinct. Third pedigerous

segment bearing a setule on dorsolateral surface and

fourth pedigerous segment, a pair of setules on middorsal surface. Genito-abdomen (Figure 236) having

essentially a typical chondracanthid form. Abdomen

FIGURE 23.—Chondracanthus nodosus male: a, body, lateral; b, urosome, dorsal; c, first and

second antennae; d, mandible; e, second maxilla; /, leg 1; g, leg 2. Scale: 0.1 mm in a, b;

0.2 mm in c; 0.01 mm in d, e, f, g.
NUMBER 8 7 31

indistinguishably fused with genital segment. Caudal

ramus as in female.

First antenna (Figure 23c) elongate and cylindrical, bearing 2 stouter elements in proximal portion

and 12 more slender elements in distal portion. Some

specimens having one or both first antennae atrophied.

Second antenna (Figure 23c) 2-segmented; first segment stout, bearing a small dorsal seta; second

segment a recurved hook bearing 2 setae (1 inner and

other outer). Rostral area protruded into a blunt lobe.

Sexual dimorphism in mouth parts occurring in

mandible (Figure 23d) and second maxilla (Figure

23e), the former being different in having proportionately larger and smaller number of teeth and the

latter in absence of anterior tooth and having fewer

posterior teeth (about 2 to 4).

Both leg 1 (Figure 23/) and leg 2 (Figure 23g)

saccular, with constriction near proximal portion.

Protopod bearing a large, long, outer seta; exopod

represented by a small forked lobe and endopod, a

larger conical lobe.

REMARKS.—This species is similar to C. zei Delaroche 1811 and different from the remaining species

of Chondarcanthus in having two pairs of trilobate

legs. It can be very easily distinguished from C. zei,

however, by the nature and number of the body processes in the trunk region and in having a pair of extremely small first antennae.

The specimens from the New England coast are

somewhat smaller than those from the Greenland.

The largest Greenlandian specimen measures 9.78

mm (in UZM).

An extremely doubtful structure, the so-called "conjunctive tube," was described by Leigh-Sharpe and

Perkins (1924) in this species. According to them, the

female embeds its head in a tumor on the gill arch

and a conjunctive tube, which is bifurcate at its tip,

arises from the gill arch and is attached to the vulvae

of the female, with one bifurcation to each vulva. A

cross section of this tube showed scattered inside certain small cells consisting almost entirely of a deeply

staining nucleus with scanty cytoplasm and having

two or three projections. They suggested that these

cells were spermatozoa. They described small tumors

on the gill arches containing "the remains of what

appear to be male animals." When they traced the

conjunctive tube into the tumor, they found an encapsulated male. Although they did not definitely

draw a conclusion from their observations, it seems

doubtful to me that the epigynic male embedded in

the tumor is the one who fertilized the eggs by sending

the spermatozoa down through the "conjunctive

tube." If one accepts their findings and this implication, what, then, is the function of having another

pygmy male attached in the vicinity of the genital

segment of the female, a condition which we observe

in each species of the Chondracanthidae where both

sexes are known? I find that their discoveries are

rather difficult to accept. It is noteworthy that of the

eight females of C. nodosus studied by them, only one

had the "conjunctive tube" connected to its vulvae

and the remaining seven had no conjunctive tubes

attached.

This parasite seems to occur exclusively on red

fishes (Sebastes species) in the North Atlantic Ocean.

Sindermann (1961) and Williams (1963) have

studied the occurrence of C. nodosus on red fish of

different fishing grounds in the Western North Atlantic and the Eastern North Atlantic. Both of them

concluded that C. nodosus is a possible parasitological

indicator of different stocks of red fish, since the incidence of infestation varied on different fishing

grounds. As they have suggested, however, further detailed and extensive study on this matter is needed.

Chondracanthus wilsoni, new species

FIGURE 24

Chondracanthopsis nodosus (Muller).—Wilson 1935: 344.

MATERIAL STUDIED.—From USNM: 1 young

female, without male attached, on gill of Spariosoma

viride, Dry Tortugas, Florida (USNM 64013).

FEMALE.—Body (Figures 24a, b) without distinct

tagmatization. Cephalosome wider than long and

lacking processes. First pedigerous segment with

rounded lateral surface; nearly as wide as cephalosome. Second pedigerous segment with lateral surface distinctly protruded. Remaining pedigerous

segments indistinctly fused, bearing three pairs of

lateral protrusions, the middle one slightly bipartite.

Posterior process short and blunt. A pair of vermiform

processes (with right one broken off in specimen

drawn) (Figure 24c) located on ventral surface just

in front of genito-abdomen. Genital segment (Figure

24c) slightly larger than abdomen which is not of

usual elliptical form but squarish. Caudal ramus (Figure 24c) a fleshy process bearing three small spinules.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 24.—Chondracanthus wilsoni female: a, body of holotype, dorsal; b, same, lateral; c,

posterior end of body, ventral; d, anterior end of head, ventral; e, maxilliped; /, leg 2. Scale:

1 mm in a, b; 0.1 mm in c, d, e, f.

First antenna (Figure 24<i) large, fleshy, and

plump. Second antenna of usual form, 2-segmented,

with terminal recurved hook. Maxilliped (Figure

24e) 3-segmented, first and second segments resembling those of C. nodosus but terminal claw different

in having two rows (8 and 3) of hooklets. Two pairs

of legs similar (Figures 24b, /), trilobate and covered

with fine denticles. Protopod with large outer protrusion, exopod a rounded lobe bearing 2 spines (one

of them tripartite), and endopod a longer protrusion

bearing a spine on its basal portion.

MEASUREMENTS.—Body 4.23 mm; head 0.98 X

1.27 mm; genital segment 0.19 X 0.37 mm; abdomen 0.20 X 0.26 mm.

MALE.—Unknown (probably attached to right

vermiform process of female and lost with it).

ETYMOLOGY.—The species is named in commemoration of Dr. Charles B. Wilson who pioneered in
NUMBER 8 7 33

studying North American parasitic copepods and

discovered this parasite.

REMARKS.—Although only one young female specimen is known, it is still justifiable to recognize it as a

new species, for certain features on the body are significantly different from the other known species of

Chondracanthus. It most closely resembles C. nodosus,

but nevertheless distinctly differs from it in having a

larger, fleshy first antenna, a number of hooklets on

the claw of maxilliped, and hook-like spines on the

rami of both trilobate thoracic legs.

It is impossible to determine whether this only

known young female has attained fully developed sets

of lateral processes on the body. Although the geneial

apearance of C. wilsoni is strongly reminiscent of C.

nodosus, this young female differs from an ovigerous

female of the latter in having one less pair of lateral

processes in the trunk region.

This is the third known species of Chondr acanthus

that possesses trilobate, modified legs (the others being

C. zei and C. nodosus).

Psuedochondracanthus hexaceraus Wilson 1935

FIGURES 25-26

Pseudo chondr acanthus hexaceraus Wilson 1935: 348-350,

pi. 4: figs. 50-57.

Pseudochondracanthoides hexaceraus (Wilson).—Yamaguti

1963: 293, pi. 59: figs. 2a-g.

MATERIAL STUDIED.—From USNM: 3 females and

1 young female, with male attached to only one

female, on Spheroides splengleri (Bloch), Dry Tortugas, Florida (USNM 64014, type).

FEMALE.—Body (Figures 25a, b) rather short and

stout, with first pedigerous segment completely fused

with cephalosome and distinctly separated from rest

of body. Head possessing two pairs of lateral processes,

with posterior pair larger and bearing surficial denticles (visible only in lactic acid under high power

with oil immersion) on their distal half. Dorsal part

of cephalosome with well-developed median sclerites

(like an inverted Y) and slightly raised posteriorly.

Trunk squarish, covered with surficial scale-like

sclerotization (see Figure 25c), except ventromedical

region; posterior corners not well-protruded. Genital

segment (Figure 25c) bearing a suckerlike ventral

depression; egg sac attachment area situated dorsolaterally. Abdomen (Figure 25c) very small, completely fused with genital segment. Caudal ramus

(Figure 25c) lobular, carrying one ventral seta and

one subterminal process bearing spinules.

First antenna (Figures 25d) fleshy, with basal portion greatly swollen, bearing 6 terminal setae. Second

antenna of a typical chondracanthid form with recurved terminal segment.

Labrum (Figure 25e) with protruded posterior

corners and smooth posterior margin. Mandible (Figure 25/) indistinctly 2-segmented; terminal segment

falcate, armed with 23 teeth on convex side and 14

teeth on concave side. Paragnath (Figure 25^) a small

lobe bearing fine spinules. First maxilla (Figure 25e)

bearing 2 setae. Second maxilla (Figure 25g) 2-segmented; first segment robust and unarmed; second

segment a bilaterally denticulated process bearing on

basal portion 1 setule and 1 spiniform seta with hyaline tip. Maxilliped (Figure 25h) 3-segmented, short,

and robust. First segment unarmed, second segment

protruded at distomedial corner where it is armed

with fine denticles, and terminal segment a claw without hooklet.

Leg 1 (Figure 25i) a large, fleshy, swollen process

terminating in two small lobes (representing rami).

Protopodial region carrying an outer seta.

MEASUREMENTS.—Body 1.55 mm; head 0.63 X

0.52 mm; genital segment 0.25 X 0.22 mm; abdomen

0.05 X 0.19 mm; egg sacs 0.82 mm and 0.88 mm;

egg 120 /*.

MALE.—Body (Figure 26) 0.25 X 0.13 mm, with

metasome strongly recurved, and indistinctly segmented. First antenna not seen; second antenna and

oral appendages as in female, except usual dimorphism shown in mandible and second maxilla. Thoracic

legs not seen.

REMARKS.—In his original description of this

species, Wilson (1935) wrote that his material consisted of "four females, two of them with egg strings

and two with attached males." However, among the

four type specimens from USNM, I have found one

of the two females without "egg strings" to be an immature adult (only 0.93 mm long) and the other, a

slightly more mature one with eggs still held inside its

trunk. Only one of the two ovigerous females carried

a pygmy male; the missing male on the other female

was probably removed by Wilson. I have dissected the

mature female without egg sacs in order to study the

oral appendages. The male was removed from its
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Pseudochondracanthus hexaceraus female: a, body of syntype, dorsal; b, same,

ventral; c, posterior end of body, ventral: d, first antenna; e, mouth parts, ventral; /, mandible;

g, second maxilla; h, maxilliped; i, leg 1. Scale: 0.1 mm in a, b, c; 0.05 mm in d; 0.01 mm in

e, f, g, h, i.
NUMBER 8 7 35

FIGURE 26.—Pseudochondracanthus hexaceraus male: lateral.

Scale 0.05 mm.

female in order to make Figure 26, but no dissection

was attempted on this only known specimen. This

intact male has been returned to the original vial in

the collection of USNM.

Considerable discrepancies exist between Wilson's

original description and my observations. There are

only two pairs of lateral processes in the head region

and the Wilson's third pair is actually the modified

first pair of legs. Thus the name of the species, hexaceraus, seems inappropriate. I was not able to find

"the rudimentary legs on the free segment" as Wilson

described (on p. 349) and illustrated (in fig. 57). The

first antenna is not "four-segmented" in the terminal

portion. The second maxilla was incorrectly described

as the first maxilla and the maxilliped as the second

maxilla.

Literature Cited

Barnard, K. H.

1948. New Records and Descriptions of New Species of

Parasitic Copepoda of South Africa. The Annals

and Magazine of Natural History, series 12, 1(4):

242-254, figures 1-8.

1955. South African Parasitic Copepoda. Annals of the

South African Museum, 41(5): 223-312, figures

1-33.

Bassett-Smith, P. W.

1896. A List of the Parasitic Copepoda of Fish Obtained

at Plymouth. The Journal of the Marine Biological

Association of the United Kingdom, new series, 4:

155-163.

1899. A Systematic Description of Parasitic Copepoda

Found on Fishes, with an Enumeration of the

Known Species. Proceedings of the Zoological

Society of London, pages 438—507, plate 26.

Bere, R.

1930. The Parasitic Copepods of the Fish of the

Passamaquoddy Region. Contributions to Canadian Biology and Fisheries, Being Studies from the

Biological Stations of Canada, new series, 5(13):

423-430, plates 1-2.

1936. Parasitic Copepods from Gulf of Mexico Fish.

The American Midland Naturalist, 17(3) : 577 -

625, plates 1-12.

Blainville, H. M. D. de

1822. Me'moire sur les Lernees (Lernaea, Lin.). Journal

de Physique, de Chemie, d'Histoire naturelle et

des Arts, 95: 372-380, 437-447, figures 1-17.

Brian, A.

1898. Catalogo di Copepodi Parassiti dei Pesci della

Liguria. Atti della Societa Ligustica di Scienze

Naturali e Geografiche, 9: 1-31, plates l^t.

1906. Copepodi Parassiti dei Pesci d'ltalia. 187 pages,

21 plates, Genova.

1908. Note Preliminaire sur les Copepodes Parasites des

Poissons Provenant des Campagnes Scientifiques de

S. A. S. le Prince Albert I " de Monaco ou

Deposes dans les Collections du Mus£e Oc£anographique. Bulletin de I'Institute Ocianographique, 110: 1-19, figures 1-7.

1912. Copepodes Parasites des Poissons et des Echinides

Provenant des Campagnes Scientifiques de S. A. S.

le Prince Albert I " de Monaco (1886-1910).

Re suit at s des Campagnes Scientifiques, 38: 1—58,

plates 1-12.

1944. Copepodos Parasitos de Peces y Cetaceos del Museo

Argentino de Ciencias Naturales. Anales del Museo

Argentino de Ciencias Naturales, 41(14): 193-

220, plates 1-10.

Capart, A.

1959. Copepodes Parasites. Expedition Oetanographique

Beige dans les Eaux C&tihes Africaines de I'Atlantique Sud (1948-1949), Risultats Scientifiques,

3(5): 57-126, figures 1-37.

Cams, J. V.

1885. Prodromus Faunae Mediterranae sive Descriptio

Animalium Marts Mediterranei Incolarum. Pars I.

Coelenterate, Echinodermata, Vermes, Arthropoda.

xiii + 524 pages. Stuttgart.

Carvalho, J. de P.

1951. Notas Sobre Alquns Copepodos Parasitos de Peixes

Maritimos da Costa do Estado de Sao Paulo.

Boletino do Instituto Paulista de Oceanografia,

2(2): 135-144.

Causey, D.

1953. Parasitic Copepoda from Grand Isle, Louisiana.

Occasional Papers of the Marine Laboratory,

Louisiana State University, 7:1—18, figures 1-22.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1955a. Parasitic Copepoda from Gulf of Mexico Fish.

Occasional Papers of the Marine Laboratory,

Louisiana State University, 9: 1-19, plates 1-3.

1955b. The External Morphology of Blias prionoti Kreyer,

a Copepod Parasite of the Sea Robins (Prionotus).

Publication of the Institute of Marine Science,

4(1): 7-12, figures 1-7.

1960. Parasitic Copepoda from Mexican Coastal Fishes.

Bulletin of Marine Science of the Gulf and

Caribbean, 10(3): 323-337, figures 1-2.

Cuvier, G. L.

1830. Le Regne Animal. Nouvelle edition, volume 4,

pages 255-258.

El Saby, M. K.

1933. The Internal Anatomy of Several Parasitic

Copepoda. Proceedings of the General Meetings

for Scientific Business of the Zoological Society of

London, 4: 861-869, plates, 1-3.

Fowler, H. W.

1912. Crustacea of New Jersey. Annual Report of the

New Jersey State Museum for 1911, pages 29-650.

Fraser, CM .

1920. Copepods Parasitic on Fish from the Vancouver

Island Region. The Transaction of the Royal Society of Canada, series 3, 13(5): 45-68, plates

1-8.

GueYin-Meneville, F. E.

1829-1844. Crustaces. Iconographie du Regne Animal,

Volume 2.

Hansen, A. V.

1923. Crustacea Copepoda, 2. Copepoda Parasita and

Semiparasita. The Danish Ingolf-Expedition, 3

(7): 1-92, plates 1-5.

Heegaard, P. E.

1945. Some Parasitic Copepods from Fishes in the

Uppsala University Collections. Arkiv for Zoologi,

35A(18): 1-27, figures 1-54.

1947. Discussion of the Mouth Appendages of the

Copepods. Arkiv for Zoologi, 40A(3) : 1-8, figures

1-8.

Heller, C.

1866. Carcinologische Beitrage zur Fauna des Adriatischen Meers. Verhandlungen der KaiserlichKoniglichen Zoologisch-botanischen Gesellschaft in

Wien, 1: 723-760.

Ho, J. S.

1967. A New Cyclopoid Copepod (Chondracanthidae)

Parasitic on the Armored Sea Robin from the

Florida Straits. The Journal of Parasitology,

53(2) : 406-411, figures 1-29.

1970. Revision of the Genera of the Chondracanthidae, a

Copepod Family Parasitic on Marine Fishes. Beaufortia, Zoological Museum of the University of

Amsterdam, 17(229): 105-218.

Holten, H. S.

1802. Anmaekninger til Beskrivelsen over Zeus guttatus,

sant Beskrivelser over Tvende Nye Arter Lernaeer.

Skrivter af Naturistorie-Selskabet, Kjebenhavn,

5(2): 125-137, plate 3.

Kabata, Z.

1968. Some Chondracanthidae (Copepoda) from Fishes

of British Columbia. Journal of the Fisheries Research Board of Canada, 25(2): 321-345, figures

1-93.

Kabata, Z., and A. V. Gusev

1966. Parasitic Copepoda of Fishes from the Collection

of the Zoological Institute in Leningrad. The

Journal of the Linnean Society of London, (Zoology), 46(309): 155-207, figures 1-176.

Kroyer, H.

1837. Om Snyltekrebsene, Isaer Med Hensyn til den

Danske Fauna. Naturhistorisk Tiddskrift, Kjebenhavn, 1(2): 172-208, plate 2; 1(3): 252-304,

plate 3; 1(5): 475-506, plate 5; 1(6): 605-628,

plate 6.

1838. Om Snyltekrebsene, Isaer Med Hensyn til den

Danske Fauna. Naturhistorisk Tiddskrift, Kjebenhavn, 2(1): 8-52, plate 1; 131-157, plate 3.

1863-64. Bidrag til Kundskab om Snylterkrebsene.

Naturhistorisk Tiddskrift, Kjebenhavn, 3(2) : 75 -

426, plates 1-18.

Lamarck, J. B. P. de

1816. Histoire Naturelle des Animaux sans Vertebres.

Volume 5, 612 pages. Paris.

Leigh-Sharpe, W. H.

1933. A Second List of Parasitic Copepods of Plymouth,

with a Description of Three New Species. Parasitology, 25(1): 113-118.

1934. A Third List of Parasitic Copepods of Plymouth,

with Notes. Parasitology, 26(2): 112-113.

Leigh-Sharpe, W. H., and M. G. L. Perkins

1924. Some Parasitic Copepoda from Iceland, with an

Account of Peniculus clavatus, the Conjunctive

Tubes of Chondracanthus nodosus and the Males

of Clavella dubia. Parasitology, 16(3): 289-295,

figures 1-6.

Markewitsch, A. P.

1956. Parasitic Copepoda of Fishes of USSR. 259 pages,

figures 1-153, (in Russian). Kiev.

Milne Edwards, H.

1840. Histoire Naturelle des Crustads, Comprenant

I'Anatomie, la Physiologie et la Classification de

ces Animaux. Volume 3, 605 pages, plates 29-42.

Paris.

Muller, O. F.

1776. Zoologiae Danicae Prodromus, seu Animalium

Daniae et Norvegiae Indigenarium Characteres,

Nomina, et Synonyma Imprimis Popularium. xxxii

-+- 282 pages. Havniae.

1788. Zoologia Danica, seu Animalium Daniae et

Nowegiae Rariorum ac Minus Notorum, Descriptiones et Historia. Volume 1, pages 1—52, plate 33.

Havniae.
NUMBER 8 7 37

Norman, C. A. M., and T. Scott

1906. The Crustacea of Devon and Cornwall. 232 pages.

London: William Wesley and Son.

Nunes-Ruivo, L.

1956. Copepodes Parasites de Peixes dos Mares de

Angola. Trabalhos da Missao de Biologia Maritima.

Anais da Junta de Investigaqoes do Ultramar,

Lisboa, 9(2): 1-45, plates 1-7.

Oakley, C. L.

1930. The Chondracanthidae (Crustacea: Copepoda) ;

with a Description of Five New Genera and One

New Species. Parasitology, 22(2): 182-201, figures

1-8.

Oken, L.

1815—16. Lehrbuch Naturgescrichte, (3, Zoologie). Volume 2, pages 181 and 357.

Olsson, P.

1868-69. Prodromus Faunae Copepodorum Parasitantium

Scandinaviae. Acta Universitatis Lundensis, Lunds

Universitets Ars-Skrift for Ar 1868, III, Afdelningen for Mathematik och Naturvetenskap, 8: 1—49.

Oorde-de Lint, G. M. van, and J. H. Schuurmans Stekhoven,

Jr1936. Copepoda Parasitica. Die Tierweit der Nord und

Ostesee, 31 (10c): 73-197.

O'Riordan, C. E.

1966. Parasitic Copepods in the Collections of the National Museum of Ireland. Proceedings of the

Royal Irish Academy, 64B(20) : 371-378.

Pearse, A. S.

1947. Parasitic Copepods from Beaufort, North Carolina.

Journal of the Elisha Mitchell Scientific Society,

63(1): 1-16, figures 1-51.

1952a. Parasitic Crustaceans from Alligator Harbor,

Florida. The Quarterly Journal of the Florida

Academy of Sciences, 15(4): 187-243, figures

1-143.

1952b. Parasitic Crustaceans from the Texas Coast. Publication of the Institute of Marine Science, 2(2) :

5-42, figures 1-157.

Pesta, O.

1934. Krebstiere Oder Crustacea, I. Ruderfusser Oder

Copepoda. 4. Monstrilloida; 5. Notodelphyoida; 6.

Caligoida; 7. Lernaeoida. Die Tierweit Deutschlands und der Angrenzenden Meeresteile nach

Ihrem Merkmalen und nach Iher Lebenswise, part

29, 68 pages, figures 1-42.

Rathbun, R.

1886. Descriptions of Parasitic Copepoda Belonging to

the Genera Pandarus and Chondracanthus. Proceedings of the United States National Museum,

9: 310-324, plates 5-11.

Richiardi, S.

1880. Contribuzione alia Fauna d'ltalia. I. Catalogo

Sistematico dei Crostacei che Vivono sul Corpo

Degli Animali Acquatici. Catalog Degli Espositori

e delle Cose Esposte, Espozitione Internazionale di

Pesca in Berlino, pages 147-152. Pisa.

Rose, M., and R. Vaissiere

1952. Catalogue PrGliminaire des Cope'podes de PAfrique

du Nord. Bulletin de la Sociitt d'Histoire Naturelle

de I'Afrique du Nord, 43: 164-176.

Schimkewitsch, W.

1896. Studien uber Parasitische Copepoden. Zeitschrift

fur Wissenschaftliche Zoologie, 61(3) : 339-362 ,

plates 14-16.

Scott, T.

1892. Additions to the Fauna of the Firth of Forth.

Tenth Annual Report of the Fishery Board for

Scotland, Part III, Scientific Investigations, pages

244-272, plates 7-13.

1900. Notes on Some Crustacean Parasites of Fishes.

Eighteenth Annual Report of the Fishery Board

for Scotland, Part HI, Scientific Investigations,

pages 144-188, plates 5-8.

1906. A Catalogue of Land, Fresh-water, and Marine

Crustacea Found in the Basin of the River Forth

and Its Estuary. Proceedings of the Royal Physical

Society of Edinburgh, 16: 97-190, 267-386.

1909. Some Notes on Fish Parasites. Twenty-sixth Annual Report of the Fishery Board for Scotland,

Part III, Scientific Investigations, pages 73—92,

plate 3.

Scott, T., and A. Scott

1913. The British Parasitic Copepoda. Ray Society,

London, volume 1, pages 1—152; volume 2, plates

1-72.

Sindermann, C. J.

1961. Parasitological Tags for Redfish of the Western

North Atlantic. International Commission for the

Northwest Atlantic Fisheries, Special Publication

No. 3, pages 111-117.

Stephensen, K.

1913. Granlands Krebsdyr og Pycnogonider (Conspectus

Crustaceorum et Pynogonidorum Groenlandiae).

Meddelelser om Grenland, 22: 335-351.

Stock, V.

1915. On Some of the Parasitic Copepods of the Bay of

Fundy Fish. Contributions to Canadian Biology,

Being Studies from the Marine Biological Stations

of Canada, 1911-1914, Marine Biology, 1: 69-71.

Stossich, M.

1880. Prospetto della Fauna del Mare Adriatico. Parte

III. Class V. Crustacea. Bolletino della Societa

Adriatica di Scienze Naturali in Trieste, 6(1) :

178-271.

Stuardo, J., and E. Fagetti

1960. Copepodos Parisitos Chilenos. II. Estudio Preliminar de los Copepodes Parasitos de Merluccius

gayi gayi en la Bahia de Valparaiso. Revista de

Biologia Marina, 10(1-3): 195-200.

Szidat, L.

1955. La Fauna Parasitos de "Merluccius hubbsi" como

Caracter Auzilliar para la Solucion de los Problemas Sistematicos y Zoogeograficos del Genero
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

"Merluccius" L. Comunicaciones del Instituto

National de Investigation de la Ciencias Naturales.

Museo Argentino de Ciencias Naturales "Bernardino Rivadavia," 3(1) : 1-54.

1961. Versuch Einer Zoogeographie des Siid-Atlantik mit

Hilfe von Leitparasiten der Meeresfische. Parasitologische Schriftenreihe, 13: 1-58, figures 1—54.

Valle, A.

1880. Crostacei Parassiti dei Pesci del Mare Adriatico.

Bolletino dell a Societa Adriatic a di Scienze

Naturali in Trieste, 6(1 ) : 55-90.

Williams, I. C.

1963. The Infestation of the Redfish Sebastes marinus

(L.) and S. mentella Travin (Scleroparei:

Scorpaenidae) by the Copepods Peniculus clavatus

(Miiller), Sphyrion lumpi (Kroyer) and Chondracanthopsis nodosus (Miiller) in the Eastern

North Atlantic. Parasitology, 53(3/4): 501-525,

figures 1—3.

Wilson, C. B.

1908. North American Parasitic Copepods: A List of

Those Found Upon the Fishes of the Pacific Coast,

with Description of New Genera and Species.

Proceedings of the United States National Museum,

35: 431-481, plates 46-83.

1912. Parasitic Copepods From Nanaimo, British Columbia, Including Eight Species New to Science.

Contributions to Canadian Biology, Being Studies

from the Marine Biological Stations of Canada,

1906-1910, Marine Biology, 9: 85-101, plates 3-9.

1918. Chondtacanthus chilensis a New Species of Parasitic copepod . Anales de Zoologia Aplicada, 5(1 ) :

11-15, figures 1-6.

1920. Report on the Parasitic Copepoda Collected During the Canadian Arctic Expedition, 1913-18.

Report of the Canadian Arctic Expedition, volume

7: Crustacea, Part L: 3L-16L.

1923a. New Species of Parasitic Copepods from Southern

Africa. Meddelanden fran Goteborgs Musei Zoologiska Avdelning, 19: 1-12, plates 1-2.

1923b. Parasitic Copepods in the Collection of the Riksmuseum at Stockholm. Arkiv for Zoologi, 15A(3) :

1-15, plates 1-2.

1932. The Copepods of the Woods Hole Region, Massachusetts. United States National Museum Bulletin,

158: 1-635, figures 1-316, plates 1-41.

1935. Parasitic Copepods from the Dry Tortugas. Papers

from Tortugas Laboratory, 29: 329-347, plates 1-6.

Yamaguti, S.

1963. Parasitic Copepoda and Branchiura of Fishes, vii

4- 1104 pages, plates 1-333. New York, London,

Sydney: Interscience Publishers.
Index

[Names of new species in italics]

Acanthocanthopsis, 20

Acanthochondria, 2, 3

albigutta, 9, 10, 11

chilensis, 15

cornuta, 3

cyclopsetta, 3, 4 (fig.)

exilipes,3,5 (fig.), 6 (fig.), 7 (fig.)

galerita, 3, 4, 7, 8 (fig.), 10 (fig.), 11 (fig.)

limandae, 15

phycidis, 2, 7, 11, 12 (fig.), 13 (fig.), 14 (fig.), 15, 27

purpurae, 15

soleae, 7

Acanthochondrites annulatus, 2

albigutta, Acanthochondria, 9, 10, 11

ancoralis, Berea, 3

annulatus, Acanthochondrites, 2

Berea ancoralis, 3

Blias prionoti, 3

bouvieri, Rebelula, 1

chilensis, Acanthochondria, 15

Chondracanthodes, 2

deflexus, 2, 15

radiatus, 2, 15, 16 (fig.), 17

tuberofurcatus, 15

Chondracanthopsis, 20

Chondracanthus, 2, 22, 31, 33

cottunculi, 2, 17 (fig.), 18 (fig.), 19 (fig.)

janebennettae, 2, 20, 21 (fig.), 22 (fig.), 23 (fig.), 24

merluccii, 2, 23, 25 (fig.), 26 (fig.), 27 (fig.)

nodosus, 2, 27, 28 (fig.), 29 (fig.), 30 (fig.), 31, 32, 33

palpifer, 26

pinguis, 20

stramineaus, 26

wilsoni, 2, 31, 32 (fig.)

zei, 31,33

cornuta, Acanthochondria, 3

crassicornis, Heterochondria, 3

cyclopsetta, Acanthochondria, 3, 4 (fig.)

deflexus, Chondracanthodes, 2, 15

diceraus, Pseudochondracanthus, 2

exilipes, Acanthochondria, 3,5 (fig.), 6 (fig.), 7 (fig.)

galerita, Acanthochondria, 3, 4, 7, 8 (fig.), 10 (fig.), 11 (fig.)

Heterochondria crassicornis, 3

hexaceraus, Pseudochondracanthus, 2, 33, 34 (fig.), 35 (fig.)

janebennettae, Chondracanthus, 2, 20, 21 (fig.), 22 (fig.), 23

(fig-), 24

limandae, Acanthochondria, 15

longa, Rhynchochondria, 2

merluccii, Chondracanthus, 2, 23, 25 (fig.), 26 (fig.), 27

(fig-)

nodosus, Chondracanthus, 2, 27, 28 (fig.), 29 (fig.), 30 (fig.),

31,32,33

Oralien triglae, 1

palpifer, Chondracanthus, 26

pinguis, Chondracanthus, 20

phycidis, Acanthochondria, 2, 7, 11, 12 (fig.), 13 (fig.), 14

(fig.), 15, 27

prionoti, Blias, 3

Pseudochondracanthus, 2

diceraus, 2

hexaceraus, 2, 33, 34 (fig.), 35 (fig.)

purpurae, Acanthochondria, 15

radiatus, Chondracanthodes, 2, 15, 16 (fig.), 17

Rebelula bouvieri, 1

Rhynchochondria longa, 2

soleae, Acanthochondria, 7

stramineaus, Chondracanthus, 26

triglae, Oralien, 1

wilsoni, Chondracanthus, 2, 31, 32 (fig.)

zei, Chondracanthus, 31, 33

.S. GOVERNMENT PRINTING OFFICE: 1971 0—408-609

39



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.

Non-Smithsonian authors should address inquiries to the appropriate department. If submission

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

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

\)k" 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 all pages consecutively, including title page, abstract,

text, literature cited, legends, and tables. A manuscript should consist of at least thirty 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 a taxon, use a period-dash (.—) to separate

each entry. Enclose with square brackets any annotation in or at the end of the taxon. For

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

the colon system for page references: "(Jones 1910:122)," and abbreviate further data:

"(Jones 1910:122, fig. 3, pi. 5: 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) usually 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 individual components should be indicated by lowercase italic letters on the illustration, in the legend, and

in text references: "Figure 96." Submit all legends on pages separate from the text and not

attached to the artwork. An instruction sheet 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 fifty free copies; order forms for quantities above this amount, with instructions for

payment, will be supplied when page proof is forwarded.