ROGER F. CRESi Cope pods Parasitic on

Sharks from the

West Coast of Florida

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1970 NUMBER 38
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 38

Roger F.Cressey Copepod s Parasitic Oil

Sharks from the

West Coast of Florida

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

197O
ABSTRACT

Cressey, Roger F. Copepods Parasitic on Sharks from the West Coast of Florida.

Smithsonian Contributions to Zoology, 38:1-30. 1970.—Thirty-one species of

parasitic copepods were taken from sixteen species of sharks caught off Sarasota,

Florida. Of these, the following are described as new: Alebion lobatus, Nesippus

nana, Kroyeria longicauda, Kroyerina scottorum, and Nemesis spinulosus. It was

noted that the spermatophore attached to females of Alebion serve as one of the

best taxonomic characters for separating females of this genus.

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

in the Institution's annual report, Smithsonian Year.

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Roger F. Cressey Copepods Parasitic on

Sharks from the

West Coast of Florida

Introduction

Between 1965 and 1970 over 400 collections of parasitic copepods representing 31 species were taken

from 16 species of sharks, mostly from off Sarasota,

Florida. The work was accomplished in cooperation

with the Mote Marine Laboratory (formerly Cape

Haze Marine Laboratory) while the laboratory was

engaged in activities concerning shark biology.

Larger species of sharks were caught on trot lines

set from one to five miles off Siesta Key. Smaller

species were caught by using a modified trot line

set closer to shore off Siesta Key and also in various

locations in Tampa Bay.

This paper represents an account of those species

of parasites present on the 16 species of sharks collected. Five new species are described. No attempt

has been made, in most cases, at redescribing the

known species since it is my intention to do diis

later in revisionary works concerning several of the

genera represented here. The large collection reported here, together with collections from other

parts of the world, will enable me to more effectively do this revisionary work. Paralebion elongatus Wilson, however, has been redescribed since

it does not fit into any proposed future revisionary

work.

The material representing four species of Alebion

reported here has enabled me to demonstrate the

usefulness of the spermatophore as a taxonomic

character in this genus.

The collection also adds much to the information

Roger F. Cressey, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, D. C. 20560.

previously known regarding the host specificity of

many of the copepods.

All material is preserved in 70 percent alcohol

and deposited in the Division of Crustacea,

Smithsonian Institution.

All figures were drawn with the aid of a camera

lucida.

In the spine-and-seta formula given for new

species the Roman numerals refer to spines and the

Arabic to setae.

Acknowledgments

The author thanks Drs. Eugenie Clark, Sylvia Earle,

and Perry Gilbert, directors of the Mote (Cape

Haze) Marine Laboratory during various phases

of the work. Special thanks are given to Roberta

and Hugh Scott of the laboratory staff, who made

several collections of copepods during periods when

the author was not at the laboratory.

The holotype of Alebion carchariae Kr0yer was

loaned to me by Dr. Torben Wolff of the

Copenhagen Museum.

Miss Rosalind Weil assisted in the preparation

of the figures.

That portion of the research conducted between

July 1966 and June 1968 was supported by a grant

from the Office of Naval Research [Nonr- 1354

(12)].

Family CALIGIDAE Dana, 1852

Lepeophtheirus eurus Bere, 1936

FIGURES 1-4

MATERIAL.—Two collections from gills of Carcharinus limbatus: one containing 8 $ ? collected

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

23 May 1963 and the other containing 1 $ collected

20 August 1965.

FEMALE.—Total length of 1 specimen 3.0 mm

FIGURES 1-4.—Lepeophtheirus eurus Bere, female: 1, dorsal;

2, sternal furca; 3, endopod of leg 2; 4, tip of leg 4.

and greatest width 1.78 mm. Body form as in Figure

1. Sternal furca (Figure 2) with bifid tines; inner

branch shorter than outer, outer branch with expanded tips. Endopod of leg 2 (Figure 3) first segment with row of prominent spines along outer

edge, last segment with row of hairs internal to

outer edge. Tip of leg 4 (Figure 4) armed with 3

setae; outer 2 equal in length, inner about 2.5 times

as long as outer setae, all setae somewhat blunt at

tips and bearing heavy fringe. The features described above will serve to distinguish this species

from others in the genus.

REMARKS.—This copepod was originally described by Bere in 1936 from the "outer surface" of

the same host reported here. These three collections

from C. limbatus may indicate a possible specific

association between this copepod and its host.

Family EURYPHORIDAE Yamaguti, 1963

Paralebion elongatus Wilson, 1911

FIGURES 5-11

MATERIAL.—Fifty-five collections from body surface of four hosts as follows (number of collections

in parentheses): Carcharinus milberti (18), C.

leucas (25), C. limbatus (3), and Negaprion brevirostris (9). No seasonal variation was noted in

the abundance of this species since it was collected

during all seasons.

FEMALE.—Body form as in Figure 5. Total length

of 1 specimen 13.9 mm and greatest width 5.2 mm.

In his original description Wilson (1911, p. 629)

states that abdomen is 2-jointed and each segment

is about equal in length. Near middle of abdomen

there is a slight constriction; no suture or joint is

visible at this point. There is, however, a separation

near terminus which divides abdomen into 2 unequal segments, anterior several times longer than

posterior. Caudal rami possess 6 terminal setae

(Figure 6) rather than 4 as reported by Wilson

(innermost very small). Sternal furca (Figure 7)

double but bases of outer branches separated from

common base of inner. Second leg endopod (Figure

8) first segment with row of spines along outer edge

(similar in nature to those reported in previous

species, L. eurus); terminal segment with 2 sickleshaped sclerotized areas at base of 2 outermost setae.
NUMBER 38

9

FIGURES 5-11.—Paralebion elongatus Wilson, female: 5, dorsal; 6, caudal ramus; 7, sternal furca;

8, endopod of leg 2; 9, leg 4. Male: 10, dorsal; 11, second antenna.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Leg 3 with single well-developed spine at base of

exopod. Leg 4 (Figure 9) exopod 3-segmented, last

2 segments each with comblike fringe on outer

margin.

MALE.—Body form as in Figure 10. Total length

7.5 mm, greatest width 2.1 mm. Appendages as in

female except for second antenna (Figure 11).

Claw of second antenna short and heavily sclerotized, basal segment with corrugated areas as in

Figure 11.

This genus was placed in the family Euryphoridae by Yamaguti (1963) because of the alleged

relationship between it and Alebion. After examining material of both genera, it is apparent that they

are not closely related and the inclusion of Par alebion in the Euryphoridae is questionable. I have

continued Yamaguti's family assignment of this

genus for the present. Future work, however, may

show the Paralebion should be placed in another

family.

Alebion carchariae Krftyer, 1863

FIGURES 12, IS

MATERIAL.—Seventy-one collections from body

surface of 9 species of sharks as follows (number of

collections in parentheses): Carcharinus milberti

(37); C. leucas (11); C. obscurus (7); C. limbatus

(7); C. maculipinnis (3); Negaprion brevirostris

(3); Galeocerdo cuvieri (1); Sphyrna mokarran (1);

and S. lewini (1). All collections made between 14

October and 27 May during the cooler season.

REMARKS.—This species of Alebion was the most

common of the four reported here and as can be

seen from the list above is a common parasite of

the carcharinid sharks of the area. Other collections

from various parts of the world attest to its ubiquitous nature and will be commented on in more detail in a subsequent work revising the genus. The

females of this species can be easily separated from

other members of the genus by the form of the spermatophore attached to the ventral surface of the

genital segment. It should be pointed out, however,

that the spermatophore has two forms (Figure 12

and 13). Both types of spermatophores were often

present on different females in the same collection.

Specimens bearing the different forms are otherwise

indistinguishable. The form represented by Figure

FIGURES 12, 13.—Alebion carchariae Kroyer, female: spermatophore in place on genital segment.

12 was collected from each of the hosts recorded

above, whereas the form represented by Figure 13

was collected only from C. milberti, C. obscurus,

and C. leucas. In the 37 collections from C. milberti

there are 140 spermatophore bearing females. Of

these, 117 bear the type with the inner median sinus

(Figure 12) and 23 bear the type shown in Figure

13. Since a similar situation was found in the new

species, A. lobatus, described below, I do not feel

that these four spermatophore forms indicate four

species but that some other explanation is needed.

Both spermatophore forms were distributed

throughout the collections without regard to season,
NUMBER 38

host size, sex, or site of infestation. Consequently

no explanation can be offered at this time.

Since this species was absent from collections

made during the summer months, a clear preference

is indicated for the winter inhabitants. Several individuals of those hosts present were examined

during summer collecting periods.

Alebion gracilis Wilson, 1905

FIGURE 14

MATERIAL.—A single collection containing 1 $

from body surface of Galeocerdo cuvieri 16 August

1965.

REMARKS.—This copepod has been reported from

various parts of the world. Examination of the type

specimen of this species and a comparison with the

type of A. carchariae leads me to suspect that many

of these records are in error and that A. gracilis is

actually quite rare and A. carchariae quite common.

My own report of A. gracilis from Madagascar

(Cressey, 1967b) is in error as well as that of Lewis

(1967) in his report on copepods parasitic on

Hawaiian sharks. This confusion will be dealt with

in more detail in my forthcoming revision of the

genus Alebion.

The female is easily identified by the form of the

attached spermatophore (Figure 14).

Alebion elegans Capart, 1953

FIGURE 15

MATERIAL.—Three collections from Sphyrna

mokarran and one from S. lewini.

REMARKS.—As in other species of the genus, the

females can be easily identified by the shape and

position of the attached spermatophore (Figure

15). The genital segment lacks the prominent lateral bulges found on both A. carchariae and A.

gracilis and bears fewer spinules. The four collections that are reported here were made at various

FIGURE 14.—Alebion gracilis Wilson, female: spermatophore

in place on genital segment.

FIGURE 15.—Alebion elegans Capart, female: spermatophore

in place on genital segment.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

20 21

FIGURES 16-21.—Alebion lobatus, new species, female: 16, dorsal; 17, genital segment and abdomen; 18, spennatophore in place on genital segment; 19, spermatophore in place on genital segment, abdomen, and caudal rami; 20, first antenna; 21, oral area.
NUMBER 38

times of the year and show no seasonal preference.

This copepod, however, has been reported only

from hammerhead sharks.

Alebion lobatus, new species

FIKURES 16-28

MATERIAL.—Thirty collections from body surface

of Carcharinus milberti. Holotype $ (USNM

128492) and 7 paratype 9 9 (USNM 128493) collected from above host 25 February 1967 at Sarasota, Florida.

FEMALE.—Body form as in Figure 16. Total

length 9.45 mm (9.30-9.90 mm) and greatest width

4.37 mm (4.20-4.50 mm) based on an average of

5 type specimens. Cephalon slightly longer than

wide (4.50 x 4.95 mm, holotype), about one-half

total length. Thoracic segments bearing legs 1-3

fused, articulating with cephalon. Thoracic segment

bearing leg 4 free and bearing prominent dorsal

bilobed plate, characteristic of other species in

genus. Genital segment (Figure 17) only slightly

longer than wide, measured from anterior border

to abdomen (2.25 x 2.52 mm); genital segment 4.20

mm long when leg 5 is included (posterior processes) . Row of small spines along anterolateral edge

and another lateral row of small spines along posterior edge (segment bulges slightly at this part).

Spermatophores attached ventrally near attachment

of abdomen. Spermatophore of 2 forms. In the most

common form encountered in the collection reported here, each spermatophore bears 2 prominent

anterior lobes (Figure 18); inner surface of each

with pilose area at about midmargin; anterior

halves divergent, posterior halves parallel. Alternate

form of spermatophore (Figure 19) without lobes

and generally smaller but still exhibiting divergent

nature (see Remarks). Abdomen (Figure 19) 1-

segmented and 1.62 mm long; anterior segment

with prominent lateral winglike processes. Caudal

rami (Figure 19) bear 4 plumose terminal setae and

2 smaller subterminal setae, each ramus somewhat

longer than wide (652 M X 420 ju).

First antenna (Figure 20) 2-segmented, basal segment with several small plumose setae, terminal segment with 13 setae (2 in form of aesthetes and 2

fused at base), 4 setae plumose as indicated in figure. Oral area as in Figure 21. Second antenna in

form of claw, first 2 segments each with rugose area,

terminal claw with small seta near midpoint. Mandible of usual caligoid type with 11 teeth. First

maxilla represented by group of 3 small setae lateral

to mouth tube and anterior to prominent flangelike

process. Second maxilla as in other species of genus,

second segment with patch of hairs near distal end;

terminal processes with lateral rows of spinules,

spinules on longest process in incomplete spiral

rows. Maxilliped in form of stout claw; tip of claw

(Figure 22) thicker than in other species of genus,

bearing seta near midpoint.

Legs 1-3 biramous. Leg 1 (Figure 23) exopod 2-

segmented, basal segment with small spine on outer

distal corner, terminal segment with 3 spines and

4 setae, terminal spine in form of broad blade directed toward mid-axis of body; terminal seta much

shorter than other 3 and all setae plumose; endopod

2-segmented, terminal segment with 3 inwardly directed plumose setae. Leg 2 (Figure 24) exopod 3-

segmented, basal segment with membranous outer

sheet enveloping outer margin, last 2 segments each

bear an inwardly directed modified spine (as in leg

1) and setae as indicated in figure; endopod 3-segmented, outer margins of first 2 segments pilose

and each with inner setae as indicated in figure,

terminal segment with 6 setae. Leg 3 (Figure 25)

basipod in form of broad flange; exopod (Figure

26) 3-segmented, basal segment with few small

spinules on outer edge and spine on outer distal

corner, second segment with modified spine, terminal segment with 2 modified spines, each segment

with setae as in figure; endopod first 2 segments

with outer edges pilose, terminal segment small and

bearing 4 setae; rami closely appressed and one

overlapping the other (broad fringe on basipod lies

over rami, not shown in Figure 25). Leg 4 (Figure

27) small, 2-segmented; basal segment with outer

plumose spine, terminal segment with 3 spines

(plumosities seen on outermost only). Leg 5 represented by 2 long processes on outer distal corner of

genital segment, each tipped with several short

spines (Figure 28) and row of short spines on

dorsal surface. Leg 6 absent.

Egg strings uniseriate.

MALE.—Unknown. Because of the lack of understanding of the diagnostic features of males of

Alebion coupled with the fact that this species

commonly occurs in the same collections as A.

carchariae, I have deferred description of the male
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

24

FICUKES 22-28.—Alebion lobatus, new species, female: 22, maxilliped; 23, leg 1; 24, leg 2; 25,

leg 3; 26, endopod of leg 3; 27, leg 4; 28, tip of leg 5.

until I have completed a comprehensive revision of

the genus with a better understanding of the male

distinguishing characters.

ETYMOLOGY.—The name lobatus refers to lobes

present on spermatophore.

REMARKS.—Females of the new species can be

separated easily from other species of the genus by

the form of the attached spermatophore. They can

be further separated from all species, except A.

elegans, by the shape of the genital segment. In A.

lobatus and A. elegans the genital segment lacks the

prominent lateral bulges and is much less spinose.

The new species can be separated from A. elegans

by the following: the fifth legs of A. elegans extend
NUMBER 38

well beyond the caudal rami, whereas those of A.

lobatus extend only slightly beyond; the last segments of the second antenna and of the maxilliped

of elegans are much longer and more delicate than

those of lobatus; the spermatophores in position on

the females of elegans diverge from their posteriormost part, whereas in lobatus the divergence is only

in the anterior half.

The occurrence of two forms of the spermatophore is similar to the situation encountered in A.

carchariae. Of the 77 females bearing spermatophores, 62 were of the knobbed type and 15 of the

smooth type. This ratio (1:4.1) is close to the ratio between the types found in A. carchariae (1:5).

It seems reasonable to assume that the same mechanisms are operating in both species. It could not be

determined if this situation exists in other species

of Alebion because of the small sample size in the

other species.

All of the collections are from Carcharinus milberti, indicating a much stronger host specificity

than in most other species of Alebion. The host

is a common winter resident of the Sarasota area.

Consequently, this copepod was not collected in

the summer.

Family PANDARIDAE Edwards, 1840

Descriptions of the eight species of this family

cited below have been omitted, except for the one

new species, since they were provided earlier in the

revision of the family (Cressey, 1967a). I have,

however, included a figure of each species (female)

so that identifications of shark copepods from the

west coast of Florida can be made without reference

to the earlier paper. For other than coastal species,

the more comprehensive treatment should be

consulted.

Pandarus cranchii Leach, 1819

FIGURE 29

MATERIAL.—Two collections from pectoral fin

and tail of Car char odon carcharias, 21 January 1966

and 20 February 1967; one collection from Carcharinus leucas, 22 March 1967; and one collection

from C. obscurus, 8 November 1966.

Pandarus smithii Rathbun, 1886

FICURES 30, 31

MATERIAL.—Five collections from body surface

of Carcharinus milberti; two collections from Carcharodon carcharias; one collection from Carcharinus obscurus; and one collection from C. leucas. All

collections made during winter months except that

from C. leucas (25 July 1967).

Pandarus floridanus Cressey, 1967

FIGURE 32

MATERIAL.—Two collections from body surface

of Carcharodon carcharias, 21 January 1966 and 20

February 1967.

Pandarus sinuatus Say, 1817

FIGURE 33

MATERIAL.—Sixteen collections from Carcharinus

milberti; ten collections from C. leucas; five collections from Negaprion brevirostris; two collections

from Ginglymostoma cirratum; one collection from

Mustelus norrisi; and one collection from C.

acronotus.

REMARKS.—This parasite occurs on the body surface of its host and was collected during all seasons

of the year.

Perissopus dentatus Steenstrup and Liitken, 1861

FIGURES 34, 35

MATERIAL.—Eighty-eight collections from nine

species of hosts as follows: Carcharinus milberti

(47), C. leucas (17), Negaprion brevirostris (5),

Carcharinus acronotus (4), Mustelus norrisi (4),

Carcharinus maculipinnis (3), C. limbatus (3),

Sphyrna tiburo (2) and Scoliodon terranovae (1).

REMARKS.—In my 1967 paper (Cressey, 1967a), I

discussed the possible variation in body form of

this species and whether or not this indicated more

than one species. At that time there was not enough

available material on which to base definite conclusions. The large collection reported here, however, has confirmed my earlier conclusion that only

one species is involved and no evidence to support

establishing more than one could be found. The
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

29

32 33

FIGURES 29-33.—Pandarus cranchii Leach, female: 29, dorsal. Pandarus smithii Rathbun, female:

30, dorsal. Male: 31, dorsal. Pandarus floridanus Cressey, female: 32, dorsal. Pandarus sinuatus

Say, female: 33, dorsal.
NUMBER 38 11

37

38

FIGURES 34, 35.—Perissopus dentatus Steenstrup and Liitken,

female: dorsal.

FIGURES 36-38.—Nesippus orientalis Heller, female: 36, dorsal.

Nesippus crypturus Heller, female: 37, dorsal. Nesippus tigris

Cressey, female: 38, dorsal.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

two principal body forms are illustrated in Figures

34 and 35. These two forms, plus intermediates,

were found distributed randomly regardless of host

or site of attachment on the host. The parasite is

most often found attached near the external nares

or on the trailing edges of the fins. Those collected

from the external nares were predominantly of the

type represented in Figure 34, while both forms

were found on the fins, frequently side by side on

the same animal.

This species was collected during all seasons and

its seasonal distribution was dependent on the

seasonal occurrence of the various host species.

Nesippus orientalis Heller, 1868

FIGURE 36

MATERIAL.—Ninety-six collections from 11 species

of hosts as follows: Carcharinus milberti (39), C.

leucas (26) , C. obscurus (6), C. maculipinnis (6),

Galeocerdo cuvieri (6), Carcharinus limbatus (5),

C. acronotus (2), Negaprion brevirostris (2), Carcharodon carcharias (2), Ginglymostoma cirratum

(1), and Sphyrna mokarran (1).

REMARKS.—This species is very common, occurring in 11 of the 16 species of sharks reported here.

The copepod is always found in the mouth or on

the gill arches of the host, often in clusters of 25 or

more individuals.

Nesippus crypturus Heller, 1868

FIGURE 37

MATERIAL.—Forty-five collections from mouths of

seven species of sharks as follows: Carcharinus leucas (13), C. milberti (12) , Galeocerdo cuvieri (11),

Sphyrna mokarran (5), S. lewini (2), C. limbatus

(1), and Carcharodon carcharias (1). Collections

made during all seasons.

REMARKS.—This species is very common, frequently occurring with N. orientalis in a variety

of inshore species of sharks.

Nesippus tigris Cressey, 1967

FIGURE 38

MATERIAL.—Four collections from nasal sinuses

of Galeocerdo cuvieri. Two collections made during July 1965, one in December 1966, and the last

in January 1967, indicating no seasonal preference.

REMARKS.—This copepod was previously recorded from the same host in Malagasy Republic.

It appears to be widely distributed, occurring

wherever the host occurs.

Nesippus nana, new species

FICURES 39-51

MATERIAL.—Nine collections from gill arches of

Carcharinus milberti. Holotype $ (USNM 128494)

and 10 para type $ $ (USNM 128495) collected

20 December 1966 at Sarasota, Florida.

FEMALE.—Body form as in Figure 39. Total

length 3.77 mm (3.68-3.83 mm) and greatest width

2.12 mm (2.03-2.18 mm) based on an average of 5

specimens. Cephalon wider than long (2.10 X 1.58

mm in 1 specimen). Dorsal surface of cephalon

border with conspicuous papillae, each bearing

short hair (Figure 40). First thoracic segment fused

with cephalon. Segments 2-4 free. No dorsal plates

present on segment 4 and not expanded laterally.

Abdomen 1-segmented, wider than long (650 /* X

472 fx). Genital segment baglike and about onehalf total length, longer than wide (1.58 X .90 mm).

Caudal rami (Figure 41) extending beyond posterior edge of genital segment, longer than wide

(336 ^ X 207 p.), bearing 5 setae, all but one of

which is naked; short row of fine spinules along

posterior portion of inner edge.

Oral area as in other species of genus. Adhesion

pads associated with first antenna, second antenna,

and maxilliped; maxilliped pad lateral to mouth

tube and very conspicuous. First antenna (Figure

42) 2-segmented, 6 terminal spines of first segment

with bristles scattered over surface, all other spines

and setae naked. Second antenna (Figure 43) with

clawlike terminal segment bearing single seta near

midpoint along inner edge, weak articulation at

this point suggesting fusion of 2 segments; penultimate segment with adhesion pad. Mouth tube and

mandible as in other members of genus, mandible

with 10 teeth. First maxilla (Figure 44) with group

of 3 small setae near anterior end and short stout

spine near outer midmargin. Second maxilla (Figure 45) with terminal segment in form of weak

claw, tip bulbous, entire segment bearing longitudinal rows of fine spinules; penultimate segment
NUMBER 38 13

42

FIGURES 39-48.—Nesippus nana, new species, female: 39, dorsal; 40, edge of cephalon; 41, caudal

ramus; 42, first antenna; 43, second antenna; 44, first maxilla; 45, second maxilla; 46, distal portion of second segment of second maxilla; 47, maxilliped; 48, leg 1.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

bearing short spine and patch of hairs at distal end

(Figure 46); an internal canallike structure present

within segment and terminating at distal patch of

hairs. Maxilliped (Figure 47) with terminal segment in shape of thumblike claw; when chela is

closed, claw fits into depression on opposing segment.

Legs 1-4 biramose. Spine and seta formula as

follows:

Seg.

See.

1

2

Leg

exo

1:0

IV:3

;

end

0:0

3

Leg

exo

1:1

IV:5

2

end

0:1

7

Leg

exo

1:1

IV:4

3

end

0:1

4

Leg

exo

IV

4

end

0

Leg 1 (Figure 48) exopod 2-segmented; spine on

outer terminal corner of first segment with subdistal accessory spinule, proximal 2 spines of terminal segment same, 2 terminal spines fingerlike,

setae pinched off and shortened: endopod 2-segmented, terminal setae shortened. Leg 2 (Figure

49) rami 2-segmented; exopod with patch of spinules on outer margins of both segments, no accessory spinule on outer spines, setae shortened. Leg

3 (Figure 50) similar to leg 2 except for lack of

spinules along outer edges of exopod segments and

reduction in number of setae. Leg 4 (Figure 51)

rami 1-segmented, surface of leg wrinkled and with

many folds; exopod bearing 4 fingerlike spines distally, endopod unarmed except lor small knob at

tip. Leg 5 represented by 2 setae located lateral to

abdomen. Leg 6 absent. Most setae present on legs

1-3 shortened. Those setae still entire have conspicuous constriction near base. Apparently distal

portion breaks off at this point and seta is reduced

to short blunt process. Blunt processes would function more effectively by aiding animal in movement

over rough surface (gill bar) of host.

Egg strings long and straight.

MALE.—Unknown.

ETYMOLOGY.—The species name nana refers to

small size of this species when compared with others

of genus, except N. vespa.

REMARKS.—This species is most closely related

to N. crypturus. It can, however, be separated from

FIGURES 49-51.—Nesippus nana, new species, female: 49, leg 2; 50, leg 3; 51, leg 4.
NUMBER 38 15

it by the form of the thoracic segment bearing leg 4.

In crypturus this segment is expanded laterally near

the area of attachment of the leg, whereas in nana

it is not. It can be further separated from crypturus

by the presence in crypturus of a plumose seta on

the exopod of leg 4 and the pointed tips on the four

spines; in nana there is no plumose seta present and

the spines are fingerlike. In crypturus the caudal

rami usually do not extend beyond the genital segment, whereas in nana they extend well beyond. It

can be separated from orientalis by the presence of

dorsal plates on the fourth thoracic segment of

orientalis, from tigris by the absence of fingerlike

spines on legs 1-4 of tigris, from vespa by the

presence of setae on the fourth leg endopod of

vespa.

Family EUDACTYLINIDAE Yamaguti, 1963

Eudactylina pollex Cressey, 1967

FIGURES 52, 53

MATERIAL.—Three collections from gills of

Sphyma mokarran and one collection from S.

lewini.

REMARKS.—This species was described by Cressey

(1967b) from Malagasy Republic from the gills of

S. mokarran. Its occurrence in Florida indicates that

it may be worldwide in distribution and specific

to hammerhead sharks.

The dorsal aspect of the female is represented by

Figure 52. The species can be separated from the

others of the genus found in Florida Gulf of Mexico

waters by the nature of the exopod of leg 2 (Figure

53); in E. pollex it is relatively short, whereas in

the other species it is very long (Figure 56).

Eudactylina longispina Bere, 1936

FIGURE 54

MATERIAL.—A single collection from gill filaments of Sphyrna tiburo collected in Tampa Bay

15 January 1968.

REMARKS.—This species was reported by Bere

(1936) and Pearse (1952a) from the same host as

cited above, indicating a possible host specificity.

So far it has been collected only in the Gulf of

Mexico.

Females of E. longispina superficially resemble

E. pollex in dorsal aspect. Consequently no habit

figure of longispina is included here. It can be most

easily distinguished from E. pollex and other species

of the genus by the unusually long spine on the

endopod of leg 4 (Figure 54). Like E. pollex the

first antenna of E. longispina does not bear

the clawlike spine on the second segment found in

the other species of Eudactylina reported here.

Eudactylina pusilla Cressey, 1967

FIGURES 55, 56

MATERIAL.—Eight collections from gill filaments

of Galeocerdo cuvieri.

REMARKS.—This species is characterized by the

presence of spinules on the body surface

(Figure 55) and the prominent lobe on the

exopod of leg 2 (Figure 56).

This copepod was previously reported (Cressey

1967b.) from the same host in Malagasy Republic,

indicating a probable circumglobal distribution and

definite host specificity.

The species was collected during all seasons of

the year.

Eudactylina spinifera Wilson, 1932

FIGURE 57

MATERIAL.—Five collections from gills of Carcharinus milberti and one collection from C. aeronotus.

REMARKS.—This copepod was described by Wilson from material collected from C. milberti. Bere

(1936) reported the species collected in the Gulf

of Mexico from C. obscurus.

This species can be separated from the others of

the genus cited in this paper by the clawlike nature

of the leg 4 endopod (Figure 57).

Eudactylina aspera Heller, 1868

FIGURE 58

MATERIAL.—A single collection from gill filaments of Carcharinus limbatus during September

1966.

REMARKS.—This species looks superficially similar

to E. pusilla, but can be characterized by the nature

of the second antenna (Figure 58) with accessory
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

54

FIGURES 52-58.—Eudactylina pollex Cressey, female: 52, dorsal; 53, leg 2. Eudactylina longispina

Bere, female; 54, leg 4. Eudactylina pusilla Cressey, female: 55, dorsal; 56, leg 2. Eudactylina

spinifera Wilson, female: 57, leg 4. Eudactylina aspera Heller, female: 58, second antenna.
NUMBER 38 17

63

61

FIGURES 59-63.—Kroyeria gemursa Cressey, female: 59, end of

abdomen and caudal rami. Kroyeria spatulata Pearse, female:

60, leg 4. Kroyeria dispar Wilson, female: 61, abdomen and

caudal rami. Kroyeria papillipes Wilson, female: 62, abdomen

and caudal rami; 63, caudal rami.

spiniform processes on the first two segments and a

patch of prominent spinules on the basal segment.

Kroyeria gemursa Cressey, 1967

FIGURE 59

MATERIAL.—Four collections from between gill

filaments of Sphyrna mokarran.

REMARKS.—This species was described from the

same host cited above, collected in the Indian

Ocean. This indicates a possible host specificity and

cosmopolitan distribution.

Kroyeria gemursa can be separated from other

members of the genus by the sclerotized area at the

posterior end of the abdomen (Figure 59).

Kroyeria gracilis Wilson, 1932

MATERIAL.—A single collection from between gill

filaments of Carcharinus obscurus.

REMARKS.—This species has been reported from

a variety of shark species from the Atlantic and

Gulf of Mexico by Wilson (1932) and Bere (1936)

respectively, and apparently is not very host specific.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Additional material is needed, however, before a

definitive conclusion can be drawn.

The species is characterized by bearing only a

few lateral spines on the inner edges of the last two

endopod segments of legs 1-4 (similar to Kroyeria

longicauda, new species, Figures 70-73). It can be

easily separated from longicauda by the lengthwidth ratio of the caudal rami (Table 1) and by

the presence of a hyaline fringe on the rami of the

new species.

Kroyeria spatulata Pearse, 1948

FIGURE 60

MATERIAL.—Nine collections from between gill

filaments of Carcharinus leucas and a single collection from Negaprion brevirostris.

REMARKS.—This species reported from Scoliodon

terraenovae, Carcharias littoralis, and Carcharinus

limbatus by Pearse (1948 and 1952b) is apparently

common on many shark species of the western

Atlantic and Gulf of Mexico.

This species is characterized by the saddlelike

sclerotizations on the outer distal corners of segments 2 and 3 of the fourth leg exopod (Figure 60).

Kroyeria dispar Wilson, 1935

FIGURE 61

MATERIAL.—Fifteen collections from between

gill filaments of Galeocerdo cuvieri collected during

all seasons.

REMARKS.—This species was reported by Cressey

(1967b) from the same host as above from Malagasy Republic. In that paper (1967) I stated that

the species had been previously described by Wilson (1935) from Squalus. This record was cited in

Yamaguti (1963). Subsequent investigation has revealed that Wilson's (1935) original host designation was "shark," hence no host name should be

associated with that record. Consequently, the only

records citing a specific host name record this

species from G. cuvieri, the tiger shark.

As in the following species, there are no prominent spines on the interpodal plates of legs 1 ^ of

K. dispar. On the basis of this character and the

sclerotized distal portion of the abdomen (Figure

61), K. dispar can be separated from the other

members of the genus.

This species occurs during all seasons, frequently

in combination with K. papillipes. An analysis of

the occurrence of these two species shows K. dispar

to be predominantly a winter inhabitant and K.

papillipes to occur in heavier infestations during

the early summer.

It was also evident that for K. dispar and K.

papillipes the average size of individuals of each

species is generally greater during periods when infestations are heaviest. The average size varies by

as much as 2 mm from one season to the other. Because collections were not made with this analysis

in mind and were made by more than one collector,

the absolute data may be unreliable and is not presented here. A cursory analysis, however, revealed

the trends cited above.

Kroyeria sphyrnae Rangnekar, 1957

MATERIAL.—Two collections from between gill

filaments of Carcharinus acronotus.

REMARKS.—This species, described by Rangnekar

(1957) from Sphyrna sp. in India, is easily separated from the other species cited from Florida by

its small size (Table 1) and because the last two

segments of the fourth leg endopod bear only two

or three lateral spinules.

The two recorded host species suggest that this

copepod is not very host specific.

Kroyeria papillipes Wilson, 1932

FIGURES 62, 63

MATERIAL.—Seventeen collections from between

gill filaments of Galeocerdo cuvieri.

REMARKS.—This species described from the same

host as reported here may well be specific to that

species of shark. Wilson also records a single female

from Sphyrna zygaena. This specimen was not deposited in the Smithsonian collections, but a specimen from the same host collected in 1911 and

identified by Wilson as K. lineata is on deposit. It

seems possible that Wilson's record of K. papillipes

from the hammerhead shark is in error.

In the collections of Kroyeria from Galeocerdo

cuvieri, this species occurred with K. dispar in 9 of

23 collections. It seems apparent that both of these

species are quite host specific and, since both species

lack the prominent interpodal spines found on all
NUMBER 38 19

other species of Kroyeria, that they are closely related. Kroyeria papillipes can be easily separated

from K. dispar by the nature of the abdomen and

caudal rami (compare Figures 61, 62, and 63).

Kroyeria papillipes occurred in collections made

during all seasons of the year.

Kroyeria longicauda, new species

FIGURES 64-77

MATERIAL.—Two collections containing 10 $ $

collected from between gill filaments of Carcharinus

limbatus. Holotype (USNM 128496) and 8 paratypes (USNM 128497) from above host collected

20 August 1965 at Sarasota, Florida.

FEMALE.—Body form as in Figure 64. Total

length 3.7 mm. Greatest width 0.6 mm measured at

widest part of cephalon. Cephalon slightly wider

than long (0.6 X 0.5 mm). Thoracic segments

bearing legs 2-4 free. Segment bearing leg 1 (fused

with cephalon) with well-developed aciculum 475 fi

in length and bearing a subapical accessory process

(Figure 65). Genital segment long (2.3 mm), comprising about two-thirds body length. Abdomen

3-segmented, last 2 segments incompletely divided;

entire abdomen 0.45 mm long. Caudal rami (Figure

66) about 5 times as long as wide (224 /i X 45 ju),

each ramus with outer lateral fringe and bearing

6 setae, 3 naked subterminal and 3 plumose terminal, longest seta 51 ^ long.

Oral area as in other members of genus. First

antenna (Figure 67) 8-segmented, each armed as in

the figure; terminal segment with 12 setae and an

aesthete at its tip (Figure 68). Second antenna

(Figure 69) of usual chelate type; movable claw fits

within cup-shaped tip on immovable claw (Figure

70), short spine on inner border of movable claw.

Mouth tube without lateral lobes. Mandible (Figure 71) with few teeth, prominent apical tooth, 2

small subapical teeth, and 2 larger proximal teeth.

Second maxilla (Figure 72) with fringed apical

process, spinose subapical knob, and subapical patch

of hairs. Maxilliped (Figure 73) terminal segment

in form of prominent recurved claw.

Legs 1-4 biramose. Rami 3-segmented. Spine and

seta formula as follows:

First leg (Figure 74) exopod armed as in figure;

endopod first segment with fringe (usually row of

hairs on other species) on outer margin, 5 spinules

on outer margin of second segment, and 4 small

spinules on outer margin of terminal segment. Leg

2 (Figure 75) armed as in leg 1 except for outer

spine on second exopod segment and additional

spine on last segment. Leg 3 (Figure 76) armed as

in leg 2 except for reduced number of setae on

terminal segment of exopod. Leg 4 (Figure 77) as

in leg 3 except for inner seta present on second segment of exopod and 1 less seta on terminal segment. Leg 5 represented by group of 4 setae on

lateral midmargin of genital segment; setae in 2

groups of 2 each. Leg 6 in other species represented

by a single seta at posterior corner of genital segment. No seta at this site could be found in the

examined specimens of this species, but it could

have been broken off.

Egg strings uniseriate.

MALE.—Unknown.

ETYMOLOGY.—The name longicauda refers to

length of caudal rami relative to width.

REMARKS.—This species can be separated from

the other members of the genus by the 5:1 lengthto-width ratio of the caudal rami, the hyaline fringe

on outer lateral margin of each ramus, and the nature of the lateral spinules on the last two segments

of the endopods of legs 1-4.

TABLE 1.—Total length and caudal rami measurements of representative females of the Florida

species of Kroyeria

Species

dispar

papillipes

gemursa

spatulata

gracilis

longicauda ....

sphyrna

Total

length

(mm)

10.5

10.0

9.7

7.3

6.5

3.7

2.2

(

length

421

200

264

280

266

224

121

Caudal rami

•

X

X

X

X

X

X

X

width(ji)

131

59

94

65

74

45

30

i

- ratio

3.2:1

3.4:1

2.8:1

4.3:1

3.6:1

5:1

4.1:1

Seg.

Seg.

Seg.

1

2

3

Legl

exo

I

0:

II:

:1

1

4

end

0:1

0:0

6

Leg

exo

1:1

1:1

111:4

2

end

0:1

0:0

6

Leg

exo

1:1

1:1

111:4

3

end

0:1

0:0

4

Leg

exo

1:1

I.I

11:4

4

end

0:1

0:1

3

Kroyerina scottorum, new species

FIGURES 78-92

MATERIAL.—Holotype 9 (USNM 128498) and

7 paratypes (USNM 128499) from nasal fossae of a
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

72 .

FICURES 64-75.—Kroyeria longicauda, new species, female: 64, dorsal; 65, tip of aciculum; 66,

caudal ramus; 67, first antenna; 68, tip of last segment of first antenna; 69, second antenna; 70,

tip of second antenna chela; 71, mandible; 72, distal end of second maxilla; 73, maxilliped; 74,

leg 1; 75, leg 2.
NUMBER 38 21

single Sphyrna lewini collected at Sarasota, Florida,

10 August 1965.

FEMALE.—Body form as in Figure 78. Total

length 4.1 mm. Greatest width 0.8 mm measured at

widest part of cephalon. Cephalon slightly wider

than long. Thoracic segment bearing first pair of

legs fused with head. Thoracic segments bearing

legs 2-4 free, each about equal in length, but second and third slightly narrower than the one before. Genital segment comprising nearly half total

length (1.9 mm) and of uniform width along entire

length (0.65 mm). Abdomen 2-segmented; each segment measuring 0.37 mm and 0.51 mm in length

respectively, second segment only slightly narrower

than first. Caudal rami (Figure 79) nearly 4 times

as long as wide (312 ft X 83 ^) and bearing 3 terminal and 2 outer subterminal spines, innermost

terminal spine longest (88 p); 2 rows of fine spinules present near distal end of ventral surface, 4

terminal spines slightly bifid at tip.

First antenna (Figure 80) with 7 distinct segments, basal segment partially subdivided into 4

segments, each segment with short setae as in figure; tip of antenna (Figure 81) with 3 blunttipped setae in addition to normally shaped setae

as in figure. Area between bases of first antenna

with 2 short hornlike processes. Second antenna

(Figure 82) strongly chelate, entire appendage

relatively short and stout, strongly developed; tip

of movable claw inserts into depression on tip of

immovable claw, immovable claw with pigment

spots as indicated in figure and short seta on inner

edge. Mouth area (Figure 83) with short cone, labrum with lateral spinose knobs. Mandible of usual

caligoid type, with 8 teeth. First maxilla (Figure

83) with anteriorly directed hook and posteriorly

directed palp, bearing 2 spinulose setae. Second

maxilla (Figure 84) apparently 3-segmented, second segment with distal spinose area, patch of long

hairs, and broad spine, terminal segment with both

edges bearing row of lamellae and tip bearing 3

short spines. Maxilliped (Figure 85) in form of

claw as in the figure.

Legs 1-4 biramose. Spine and seta formula as

follows:

FIGURES 76-77.—Kroyeria longicauda, new species, female: 76,

leg 3; 77, leg 4.

Seg.

Seg.

Seg.

1

2

3

Leg

exo

1:0

1:1

11:4

1

end

0:1

0:1

6

Leg

exo

1:1

1:1

111:4

2

end

0:1

0:1

6

Lei

exo

1:1

1:1

111:4

\ 3

end

0:1

0:1

1:1:3

Leg

exo

1:1

1:1

111:4

4

end

0:1

0:1

1:1:5
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 78-86.—Kroyerina scottorum, new species, female: 78, dorsal; 79, caudal ramus; 80, first

antenna; 81, tip of first antenna; 82; second antenna; 83, first maxilla and mouth tube; 84,

second maxilla; 85, maxilliped; 86, leg 1.
NUMBER 58 23

89

FIGURES 87-92.—Kroyerina scottorum, new species, female: 87, leg 2; 88, last endopod segment of

leg 3; 89, leg 4 endopod; 90, leg 5. Male: 91, genital segment, abdomen, and caudal rami, ventral;

92, rostral area, ventral.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Leg 1 (Figure 86) coxopod with patch of spinules

on outer distal corner; basipod with outer and inner setae; both rami with striated fringe on outer

edge of each segment, no patches of spinules, setae

arranged as in figure. Leg 2 (Figure 87) coxopod

with patch of spinules as in leg 1; basipod with

outer seta only and fringe and patch of spinules

along inner margin; rami with fringes on each

segment as in leg 1, setae arranged as in figure and

each sclerotized on basal one-third, proximal twothirds areolate in appearance, path of spinules on

first endopod segment. Leg 3 similar to leg 2 except

last endopod segment (Figure 88) with terminal

fringed spine between an outer seta and 3 inner

setae. Leg 4 similar to leg 2 except endopod (Figure

89) first segment lateral hairs in 2 separate groups

and terminal segment with fringed spine between

outer seta and 2 inner setae. Leg 5 (Figure 90)

short sclerotized segment bearing 3 setae, inserted

on midlateral margin of genital segment (Figure

78) . Leg 6 absent.

No specimens were ovigerous.

MALE.—Body form generally as in female. Total

length 3.4 mm. Greatest width 0.6 mm, measured at

widest part of cephalon. Genital segment (Figure

91) relatively shorter than in female (measuring

1087 /x X 508 fi) and bearing 7 rows of fine spinules

along each edge anterior to leg 5 as shown in figure.

Abdomen (Figure 91) 3-segmented, each segment

measuring 464 fi, 363 p, and 247 /* in length respectively, each bearing patches of spinules as in figure.

Caudal rami as in female except for patches of

spinules on ventral surface as in figure.

Cephalic and thoracic appendages as in female

except leg 5 of male bears 4 long setae rather than

3. Leg 6 represented by 2 long setae at posterior

outer corners of genital segment. Area between

bases of first antennae (Figure 92) with 2 hornlike

processes as in female.

ETYMOLOGY.—This species is named for Roberta

and Hugh Scott, whose collecting efforts at times

when the author was not at the Mote Marine Laboratory significantly added to the material

incorporated in this paper.

REMARKS.—This third member of the genus

Kroyerina can be separated from the other two by

the following facts. The ratio of the length to width

of the caudal rami of the three species is as follows:

K. scottorum 4:1 (312,1 X 83,*), K. elongate 2.5:1

(295M X 112ju), K. nasuta 3:1 (192/x X 62/*). The

inner terminal spine on the caudal ramus of K.

nasuta is twice as long as the next one to it, the

outermost of the three terminal setae is spinelike.

The labrum of K. nasuta has no lateral spinose

lobes (present in both K. elongata and scottorum).

Nemesis lamna Risso, 1826

FIGURE 93

MATERIAL.—Two collections from Carcharodon

carcharias and a single collection from Cetorhinus

maximus}. (According to Stewart Springer, U.S.

Bureau of Commercial Fisheries, there is some

question regarding the specific determination of

this shark.)

This copepod is worldwide in distribution and

has been reported many times from lamnid sharks.

It has been recently redescribed by Cressey (1968).

Nemesis atlantica Wilson, 1922

MATERIAL.—Fourteen collections from gills of

Carcharinus leucas, four collections from C. limbatus, two collections from C. maculipinnis, one

collection from C. acronotus, and one collection

from Sphyrna mokarran.

REMARKS.—This copepod has been reported a

number of times from several species of sharks. It

was collected in Florida throughout the year,

showing no seasonal preference.

Nemesis pilosus Pearse, 1951

MATERIAL.—Nine collections from gills of Negaprion brevirostris, one collection from Carcharinus

maculipinnis, and one collection from C. limbatus.

REMARKS.—This copepod was described by

Pearse (1951) from Carcharias littoralis from the

Bahamas and has not been reported since.

This species can be separated from N. atlantica

by the armature of leg 3. In N. pilosus the last

endopod segment bears five spines, whereas in N.

atlantica this same segment bears only four. The

spines on legs 2 and 3 of N. pilosus are much longer

than those of N. atlantica.
NUMBER 58 25

Nemesis spinulosus, new species

FIGURES 94-110

MATERIAL.—Three hundred and six specimens

C303 9 $ , 3 tf d) collected from gills of 27 Carcharinus milberti; an additional 2 9 $ from gills

of a single Carcharinus obscurus. Holotype 9

(USNM 128500) and 31 paratype 9 9 (USNM

128501) collected 17 December 1966 from C.

milberti, Sarasota, Florida.

FEMALE.—Body form as in Figure 94. Total

length 2.98 mm, based on an average of 4 specimens

(2.93-3.00 mm). Greatest width 1.18 mm (1.12-

1.20 mm). Cephalon only slightly longer than wide

(1.2 X 1.1 mm), posterior margin overlapping first

free thoracic segment. Thoracic segments bearing

legs 2-5 free and measuring 1.5, 1.5, 1.3, and .87 mm

in width respectively. Genital segment (Figure 95)

wider than long (.54 X .35 mm). Abdomen 3-seginented, segments measuring .35, .29, and .25 mm in

width respectively; third segment about as long as

first and second combined. Caudal rami (Figure

96) each with 5 terminal setae, inner 3 much stouter than outer 2; each ramus about twice as long as

wide (188/i X 94^) and bearing row of hairs on

inner margin (not seen on all specimens examined).

First antenna (Figure 97) 13-14 segmented (terminal segment incompletely divided) and armed as

in figure. Second antenna (Figure 98); second segment with 2 patches of spinules, inner patch composed of heavier spinules than outer patch; third

segment with patch of spinules along inner border;

terminal segment in form of claw bearing 2 short

setae and median accessory process. Mandible of

usual caligoid type, bearing 9 teeth at tip. First

maxilla (Figure 99) biramose, each ramus armed

with 2 setae as in figure; each seta sclerotized about

three-fourths of its length, distal fourth weak. Second maxilla with inner patch of heavy spines, outer

patch of hairs, and short spinose claw at tip (Figure

100). Maxilliped (Figure 101) in form of stout

claw, both segments with broad-based setae as in

figure.

Legs 1-4 biramose, all rami 2-segmented. Spine

and seta formula as follows:

FIGURE 93.—Nemesis lamna Risso, female: dorsal.

Seg.

Seg.

1

2

Leg

exo

1:0

III

1

end

0:1

II

Leg

exo

1:1

VII

or

VIII

2

end

0:1

VI

Leg

exo

1:1

VII

or

VIII

B

end

0:1

V

Leg

exo

1:1

VII

or

vin

4

end

0:1

rv
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOCK

102

FIGURES 94-102.—Nemesis spinulosus, new species, female: 94, dorsal; 95, genital segment and

abdomen, ventral; 96, caudal ram us; 97, first antenna; 98, second antenna; 99, first maxilla; 100,

tip of second maxilla; 101, maxilliped; 102, leg 1.
NUMBER 38 27

108

FIGURES 103-110.—Nemesis spinulosus, new species, female: 103, leg 2; 104, leg 3; 105, leg 4. Male:

106, lateral; 107, caudal ramus; 108, leg 2; 109, leg 3; 110, leg 4.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Leg 1 (Figure 102) modified as in other species

of genus: exopod 2-segmented, each segment bearing broad spinules as in figure; endopod first segment recurved toward exopod bearing row of broad

spinules along outer border, patch of setules on

inner border, and spinose knob at tip; second segment short, bearing 3 stout spines terminally and

few spinules on each lateral margin. Leg 2 (Figure

103) with rows and patches of spinules on all segments as in figure; each ramus bearing long terminal spines. Leg 3 (Figure 104) similar to leg 2

except that terminal spines on each ramus are shorter and more heavily sclerotized. Leg 4 (Figure 105)

as in preceding 2 legs except for fewer spinules on

rami and shorter terminal spines. Leg 5 (Figure 95)

1-segmented; plumose seta arising near base of free

segment, free segment bearing 3 small, terminal

setae. Leg 6 absent.

Egg strings uniseriate, long.

MALE.—Body form as in Figure 106. Total length

2.63 mm. Greatest width 0.68 mm measured at widest part of cephalon. Cephalon slightly longer than

wide (0.82 X 0.68 mm), comprising about onefourth body length. Thoracic segments bearing legs

2-4 free, measuring 0.60, 0.60, and 0.56 mm in

width respectively, each segment somewhat overlapping anterior portion of following segment.

Genital segment as long as wide (0.52 X 0.52 mm),

narrowest anteriorly. Abdomen 4-segmented, measuring 354 IJL, 271 ix, 201 (i and 201 p in width and

295 n, 177 fi, 130 (i, and 130 p in length respectively.

Caudal ramus (Figure 107) armed as in female, but

each ramus somewhat longer in relation to width

(207 p. X 83 fi), all spines and setae plumose.

Cephalic appendages as in female except second

segment of second antenna of male has fewer spines

and no spinules on third segment.

Leg 1 as in female. Legs 2-4 with spine and seta

formula as follows:

Seg.

Seg.

1

2

Leg

exo

1:1

111:4

2

end

0:1

6

Leg

exo

1:1

111:5

end

0:1

6

Leg

exo

1:1

III:

4

end

0:1

5

Leg 2 (Figure 108) with patches of spinules as

indicated in figure, all setae unmodified and plumose. Leg 3 (Figure 109) exopod with unmodified

spines and setae as figured; endopod first segment

inner seta heavily sclerotized and thick throughout

entire length, last segment bearing terminal inwardly curved, naked, sclerotized seta in addition to 5

unmodified plumose setae of usual type. Leg 4

(Figure 110) with modified setae on endopod as in

leg 3, in addition to usual unmodified setae. Leg 5

(Figure 106) broad lobate segment, bearing 3 terminal setae as in female. Leg 6 broad lobe on outer

distal corner of genital segment, bearing 3 setae,

innermost longest.

ETYMOLOGY.—The specific name spinulosus

alludes to presence of spinules on second antenna.

REMARKS.—This species differs from all others of

the genus, except tiburo Pearse (1952a), by the

presence of spinules on the third segment of the

second antenna. The holotype of N. tiburo (USNM

93702) was compared to this new species and the

following differences were noted: only one patch of

spinules on the second segment of the second antenna of N. tiburo, as compared with two in N.

spinulosus; only four spines on the endopod of leg

3 of N. tiburo, as compared with five in N. spinulosus; caudal rami of N. tiburo are tear shaped and

spaced wide apart with proximal portion narrow,

whereas in N. spinulosus the rami are close together

and of uniform width throughout their lengths.

The variation in number of terminal spines on

the exopods of legs 2-4 of 14 females on N.

spinulosus, all from the same sample, was as

follows:

Leg 2 Leg 3 Leg 4

6 spines 7 spines 7 spines 8 spines 7 spines 8 spines

No. of

specimens 12 11

The number of terminal endopod spines was

constant.

List of Shark Species and the Copepod Parasites

(Number in parentheses after host name refers to

number of sharks from which copepods were removed. Number following copepod name indicates

number of collections.)

Carcharinus milberti (69)

Alebion lobatus (30)

Alebion carchariae (37)

Paralebion elongatus (18)

Nesippus orientalis (39)

Nesippus crypturus (12)

Nesippus nana (9)

Perissopus dentatus (47)
NUMBE

R 38

29

Pandarus smithii (5)

Pandarus sinuatus (16)

Nemesis spinulosus (27)

Eudactylina spinifera (5)

Carcharinus obscurus (9)

Alebion carchariae (7)

Nesippus orientalis (6)

Pandarus smithii (1)

Pandarus cranchii (1)

Kroyeria gracilis (1)

Nemesis spinulosus (1)

Carcharinus leucas (42)

Alebion carchariae (11)

Paralebion elongatus (25)

Nesippus orientalis (26)

Nesippus crypturus (13)

Perissopus dentatus (17)

Pandarus smithii (1)

Pandarus cranchii (1)

Pandarus sinuatus (10)

Kroyeria spatulata (9)

Nemesis atlantica (14)

Carcharinus maculipinnis (10)

Alebion carchariae (3)

Nesippus orientalis (6)

Perissopus dentatus (3)

Nemesis pilosus (1)

Nemesis atlantica (2)

Carcharinus limbatus (16)

Alebion carchariae (7)

Paralebion elongatus (3)

Lepeophtheirus eurus (2)

Nesippus orientalis (5)

Nesippus cry pturus (1)

Perissopus dentatus (3)

Kroyeria longicauda (2)

Nemesis pilosus (1)

Nemesis atlantica (4)

Eudactylina aspera (1)

Carcharinus acronotus (7)

Nesippus orientalis (2)

Perissopus dentatus (4)

Pandarus sinuatus (1)

Kroyeria sphyrnae (2)

Nemesis atlantica (1)

Eudactylina spinifera (1)

Galeocerdo cuvieri (29)

Alebion carchariae (1)

Alebion gracilis (1)

Nesippus orientalis (6)

Nesippus crypturus (11)

Nesippus tigris (4)

Kroyeria papillipes (17)

Kroyeria dispar (15)

Eudactylina pusilla (8)

Negaprion brevirostris (19)

Alebion carchariae (3)

Paralebion elongatus (9)

Nesippus orientalis (2)

Perissopus dentatus (5)

Pandarus sinuatus (5)

Kroyeria spatulata (1)

Nemesis pilosus (9)

Ginglymostoma cirratum (3)

Nesippus orientalis (1)

Pandarus sinuatus (2)

Sphyrna mokarran (9)

Alebion carchariae (1)

Alebion elegans (3)

Nesippus orientalis (1)

Nesippus crypturus (5)

Kroyeria gemursa (4)

Nemesis atlantica (1)

Eudactylina pollex (3)

Sphyrna lewini (2)

Alebion carchariae (1)

Alebion elegans (1)

Nesippus crypturus (2)

Kroyerina scottorum (1)

Eudactylina pollex (1)

Sphyrna tiburo (3)

Perissopus dentatus (2)

Eudactylina longispina (1)

Mustelus norrisi (4)

Perissopus dentatus (4)

Pandarus sinuatus (1)

Scoliodon terranovae (1)

Perissopus dentatus (1)

Carcharodon carcharias (2)

Nesippus orientalis (2)

Nesippus crypturus (1)

Pandarus smithii (2)

Pandarus cranchii (2)

Pandarus floridanus (2)

Nemesis lamna (2)

Cetorhinus maxim us? (1)

Nemesis lamna (1)
References

Bere, R.

1936. Parasitic Copepods from Gulf of Mexico Fish.

American Midland Naturalist, 17 (3):577-625.

Capart, A.

1953. Quelques cope'podes parasites des poissons marins

de la region de Dakar. Bulletin de I'Institut francais

d' Afrique noire, 15 (2):647-670.

Cressey, R.

1967a. Revision of the Family Pandaridae (Copepoda:

Caligoida). Proceedings of the United States National

Museum, 121 (3570): 1-133.

1967b. Caligoid Copepods Parasitic on Sharks of the Indian

Ocean. Proceedings of the United States National

Museum, 121 (3572): 1-21.

1968. Caligoid Copepods Parasitic on Isurus oxyrinchus

with an Example of Habitat Shift. Proceedings of

the United States National Museum, 125 (3653): 1-26.

Heller, C.

1868. Crustaceen. No. 8 in Volume 2 of Zoologischer Theil

in Reise der Oesterreichischen Fregatte Novara . . .,

280 pages, 25 plates. [The plates were printed in

1865, but the text and plates were published in

1868.]

Kreyer, H.

1863. Bidrag til Kundskab om Snyltekrebsene. Naturhistorisk Tidsskrift, 3 (2):75-426.

Leach, W.

1819. Entomostraca. In Dictionnaire des Sciences naturelles

... . 14:524-543.

Lewis, A.

1967. Copepod Crustaceans Parasitic on Elasmobranch

Fishes of the Hawaiian Islands. Proceedings of the

United States National Museum, 118 (3524):57-154.

Pearse, A.

1948. A Second Report on Parasitic Copepods Collected at

Beaufort, N.C. Journal of the Elisha Mitchell Scientific Society, 64(1):127-131.

1951. Parasitic Crustacea from Bimini, Bahamas. Proceedings of the United States National Museum,

101 (3280):341-372.

1952a. Parasitic Crustaceans from Alligator Harbor, Florida.

The Quarterly Journal of the Florida Academy of

Sciences, 15 (4): 187-243.

1952b. Parasitic Crustacea from the Texas Coast. Publications of the Institute of Marine Sciences, University of Texas, 2 (2):5^2.

Rangnekar, M.

1957. Copepod Parasites of the Families Argulidae,

Caligidae, Dichelesthidae, and Lernaeopodidae.

Journal of the University of Bombay, Section B,

26 (3):8-20.

Rathbun, R.

1886. Description of Parasitic Copepoda Belonging to the

Genera Pandarus and Chondracanthus. Proceedings

of the United States National Museum, 9:310-324.

Risso, A.

1826. Histoire naturelle des principales Productions de

I'Europe Miridionale et particulierement ce celles

des environs de Nice et des Alpes Maritimes, 5:1-

135.

Say, T.

1817. An Account of the Crustacea of the United States.

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Steenstrup, J. and C. Liitken

1861. Bidrag til Kundskab om det aabne Havs Snyltekrebs

og Lernaeer samt om nogle andre nye eller hidtil

kun ufuldstaen digt kjendte parasitiske Copepoder,

5:341-432.

Wilson, C. B.

1905. New Species of Parasitic Copepods from the Massachusetts Coast. Proceedings of the Biological Society of Washington, 18:127-132.

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New Genera and Species. Proceedings of the United

States National Museum, 39:625-634.

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the Family Dichelesthiidac. Proceedings of the

United States National Museum, 60:1-100.

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