Caligus solea n. sp. (Copepoda: Caligidae) parasiticon the common sole Solea solea (Linnaeus)from the north-eastern Mediterranean off the Turkish coast

Ibrahim Demirkale • Argun A. Ozak •

Alper Yanar • Geoffrey Allan Boxshall

Received: 8 May 2014 / Accepted: 12 June 2014

� Springer Science+Business Media Dordrecht 2014

Abstract A new species of caligid copepod, Caligus

solea n. sp., is described from the common sole Solea

solea (Linnaeus) caught off the north-eastern Medi-

terranean coast of Turkey. Both sexes of the parasite

were collected from all over the upper body surface of

its host. The new species belongs to the macarovi-

group of species as established by Boxshall & Gurney

(Bull Br Mus (Nat Hist) (Zool), 39:161–178, 1980),

with which it shares the following four characters:

(i) leg 4 with two-segmented exopod, distal segment

carrying three apical spines but no lateral spine; (ii)

distal exopodal segment of leg 1 with three plumose

setae posteriorly plus four distal margin elements,

spine 1 naked, spines 2 and 3 with accessory process

and spine 4 about twice length of the others; (iii)

females with one-segmented abdomen while males

with two-segmented abdomen; (iv) male maxilliped

with myxal process opposing the tip of the subchela.

However, the new species differs from its congeners

within the macarovi-group in the number of sensillae

on each papilla on and around the postantennal process,

and also in the absence of serrations along the distal

margin of the maxilla. This is the twenty-eighth species

of Caligus to be reported from the Mediterranean Sea.

Introduction

The family Caligidae Burmeister, 1835 currently consists

of 31 valid genera according to the recent revision by

Dojiri & Ho (2013). Among these, the genus Caligus

Muller, 1785 is by far the largest, currently comprising

approximately 250 valid species (Hayes et al., 2012).

Members of the genus Caligus, commonly known as sea

lice, infest both wild and cultured fish species living in

brackish and marine habitats (Johnson et al., 2004).

Infection with sea lice has a direct pathological effect on

the host and may result in secondary microbial infections

(Nylund et al., 1994; Cruz-Lacierda et al., 2011). Correct

identification of these parasites is therefore important in

the development of effective fish health management

strategies, especially in marine finfish aquaculture.

In the Mediterranean the genus Caligus is repre-

sented by 27 species reported from 16 different

families of teleost fishes (Raibaut et al., 1998; Ozak

et al., 2012, 2013). To the best of our knowledge, only

four of these 27 species, Caligus apodus (Brian, 1924),

C. brevicaudatus A. Scott, 1901, C. diaphanus von

Nordmann, 1832 and C. elongatus von Nordmann,

I. Demirkale � A. A. Ozak (&)

Department of Fish Diseases & Aquaculture, Faculty

of Fisheries, University of Cukurova, 01330 Balcali,

Adana, Turkey

e-mail: ozargun@cu.edu.tr

A. Yanar

Department of Basic Sciences,Faculty of Marine Sciences

and Technology, Mustafa Kemal University, Meydan Mh.

512 Sk., 31200 Iskenderun, Turkey

G. A. Boxshall

Department of Life Sciences, The Natural History

Museum, Cromwell Road, London, SW7 5BD, UK

123

Syst Parasitol (2014) 89:23–32

DOI 10.1007/s11230-014-9505-4

1832 have thus far been reported from the common

sole, Solea solea (Linnaeus) (see Kabata, 1979; Mar-

ques et al., 2009; Ozak et al., 2013). Among these four,

only C. apodus and C. brevicaudatus have been

reported from this host in the eastern Mediterranean.

However, the reported prevalence rates of these species

(3% and 27% respectively; Ozak et al., 2013) on S.

solea were lower than the prevalence of the new species

described below. The new species reported here may be

the most common caligid on S. solea in northeastern

Mediterranean waters off the Turkish coast.

Materials and methods

Parasites were collected from all over the upper body

surface of S. solea caught off Karatas (36�30001.8800N,

35�23014.6000E) and Konacık (36�21051.2300N, 35�450

46.7400E) in Iskenderun Bay, Turkey. Fish (n = 2,025;

total body length range 10–23 cm) were caught by sole

trammel nets. Parasitic copepods removed from the

infested fish were immediately preserved in 70%

ethanol. Specimens were cleared in lactic acid for 2 h

prior to examination using an Olympus SZX16 dissect-

ing microscope and Olympus BX51 compound micro-

scope. Subsequently, the specimens were dissected on

glass-slides and mounted as temporary preparations in

lactophenol. Measurements were made using an ocular

micrometer and drawings were made with the aid of a

drawing tube. All measurements are in millimetres and

are presented as the range followed by the mean in

parentheses. The scientific and common names of fishes

follow Froese & Pauly (2013), the morphological

terminology for the copepods follows Huys & Boxshall

(1991), and parasitological terms follow Bush et al.

(1997). The type-material is stored in the collection of

the Natural History Museum, London and in the second

author’s personal collection.

Family Caligidae Burmeister, 1835

Genus Caligus Muller, 1785

Caligus solea n. sp.

Type-host: Solea solea (Linnaeus) (Soleidae).

Type-locality: Northeastern Mediterranean waters off

Karatas and Konacık in Iskenderun Bay, Turkey;

collected by A. A. Ozak (4.xi.2011, 15.xii.2012; depth

range 18–55 m; mean surface water temperature

13.5�C; salinity 35 ppt).

Site on host: Upper body surface.

Prevalence: 68% (1,377 fish infected out of a total of

2,025 examined).

Type-material: Holotype female [BMNH 2014.619]; 5

paratype females [BMNH 2014.610-614] and 3 para-

type males [BMNH 2014.615-617]; the remaining

paratype material (54 females and 13 males) is in the

collection of AAO.

Etymology: The species name refers to the host genus.

Description (Figs. 1–3)

Adult female. Body (Fig. 1A) comprising caligiform

cephalothorax, incorporating first to third pedigerous

somites, free fourth pedigerous somite, genital com-

plex and suboval 1-segmented abdomen. Body length

3.9–4.22 (4.13) (n = 10) excluding caudal setae.

Cephalothoracic shield slightly wider than long,

1.89–2.01 9 1.90–2.07 (1.95 9 1.97) excluding mar-

ginal hyaline membranes. Frontal plates bearing

paired lunules. Free thoracic zone of shield comprising

almost half length of cephalothorax, wider than long,

0.92–1.03 9 1.38–1.5 (0.99 9 1.44). Posterior margin

of free thoracic zone straight, extending beyond

posterior ends of lateral zones. Fourth pedigerous

somite 0.20–0.27 9 0.51–0.60 (0.22 9 0.56), dis-

tinctly separated from genital complex. Genital com-

plex 1.10–1.21 9 0.97–1.16 (1.15 9 1.08), with

slightly rounded anterior angles, parallel sides and

straight posterior margin. Abdomen suboval, 1-seg-

mented, longer than wide, 0.7–0.78 9 0.48–0.55

(0.74 9 0.51), c.60% of length of genital complex.

Caudal rami (Fig. 1B) c.1.45 times longer than wide,

armed with 6 pinnate setae, ornamented with fine

setules on inner margin.

Antennule (Fig. 1C) 2-segmented, proximal seg-

ment distinctly wider than distal, armed with 25

plumose setae on anterior and antero-ventral surfaces

plus 2 unarmed setae located dorsally; distal segment

armed with 1 subterminal seta on posterior margin and

11 setae plus 2 aesthetascs on distal margin.

Antenna (Fig. 1D) uniramous, 3-segmented; prox-

imal segment produced into blunt spinous process;

middle segment unarmed; distal segment forming

sharply curved claw with spine-like seta proximally

and distal seta. Postantennal process (Fig. 1E) weakly

24 Syst Parasitol (2014) 89:23–32

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Fig. 1 Caligus solea n. sp. female (holotype). A, Habitus, dorsal view; B, Caudal rami; C, Antennule; D, Antenna; E, Postantennal

process and sensillae (black arrows); F, Mandible; G, Maxillule; H, Maxilla; I, Maxilliped; J, Sternal furca. Scale-bars: A, 0.5 mm; B–J,

100 lm

Syst Parasitol (2014) 89:23–32 25

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Fig. 2 Caligus solea n. sp. female. A, Leg 1; B, Leg1 endopod; C, Spines at outer distal corner of first exopodal segment; D, Tip of

distal segment of leg 1; E, Leg 2; F, Leg 3; G, Leg 4; H, Leg 5. Scale-bars: A–C, E–H, 100 lm; D, 50 lm

26 Syst Parasitol (2014) 89:23–32

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curved, carrying 2 papillae each with 2 sensillae;

similar papilla with 2 sensillae located on body surface

adjacent to postantennal process.

Mandible (Fig. 1F) with 12 teeth on one side near

apex. Maxillule (Fig. 1G) comprising dentiform pro-

cess tapering towards tip and anterior papilla bearing 3

unequal setae. Maxilla (Fig. 1H) 2-segmented, brach-

iform; proximal segment (lacertus) large, unarmed;

slender distal segment (brachium) bearing small

subterminal hyaline membrane on outer margin plus

short canna and long calamus distally. Maxilliped

(Fig. 1I) comprising robust proximal segment (cor-

pus) and distal subchela representing fused endopodal

segments plus claw; subchela armed with small seta at

base of claw. Sternal furca (Fig. 1J) with square box

and bluntly pointed parallel tines.

Swimming leg 1 (Fig. 2A) biramous, with 2-seg-

mented exopod and unsegmented vestigial endopod.

Sympod armed with lateral plumose seta and inner

seta. Endopod (Fig. 2B) vestigial, with minute spin-

iform process at apex. First exopodal segment orna-

mented with row of setules along free posterior margin

and bearing small spine plus adjacent strong spinule

(Fig. 2C) at outer distal corner. Distal exopodal

segment (Fig. 2D) with 3 plumose setae posteriorly

plus 4 terminal elements; outermost element (spine 1)

finely serrated, middle 2 elements (spines 2 and 3)

each bearing single accessory process, ornamented

with fine serrations along inner margins, innermost

element (seta 4) with setules along inner margin, about

twice as long as spines. Pectens present at bases of

spines 1, 2 and 3 (Fig. 2D).

Leg 2 (Fig. 2E) biramous, with 3-segmented rami.

Coxa small, with large pinnate seta on posterior

margin. Basis with 1 small spine at outer distal angle

plus membrane along free posterior margin and

dorsally-reflexed membrane along anterior margin.

First exopodal segment c.1.6 times longer than

second; both segments with pinnate seta on inner

margin and long oblique spine at outer distal corner

reflexed over surface of segment. Dorsally reflexed

membrane present along free outer margin of first

exopodal segment and extending distally to cover

second segment. Third exopodal segment with 3 outer

spines, each fringed with hyaline membrane, and 5

pinnate setae. First endopodal segment with inner

pinnate seta; second endopodal segment with 2 inner

pinnate setae, ornamented with rows of long setules on

outer margin; third segment with 6 pinnate setae.

Leg 3 (Fig. 2F) with coxa and basis fused into

flattened apron-like sympod. Sympod and intercoxal

sclerite with extended strips of hyaline membrane

along lateral and free posterior margins. Crescent-

shaped patch of spinules present anteriorly, adjacent to

lateral margin. Coxal seta and outer basal seta both

pinnate. Exopod 3-segmented, with outer spine on first

segment just longer than segment, extending more-or-

less parallel with long axis of ramus. Second exopodal

segment with outer spine and inner plumose seta.

Third exopodal segment with 3 outer spines and 4

short pinnate setae. Endopod 2-segmented; first seg-

ment with long inner pinnate seta, second with 6

pinnate setae, ornamented with rows of long setules

along outer margin.

Leg 4 (Fig. 2G) uniramous. Protopodal segment

with outer seta derived from basis. Exopod 2-seg-

mented; first segment with 1 distal spine extending just

beyond middle of margin of second exopodal seg-

ment; second segment with 3 apical spines along

oblique distal margin, longest spine about twice as

long as shortest spine, each spine with pecten at base.

Spine (Roman numerals) and seta (Arabic numer-

als) formula of legs 1–4 as follows:

Exopod Endopod

Leg 1 I-0; III,1,3 vestigial

Leg 2 I-1; I-1; II,I,5 0-1; 0-2; 6

Leg 3 I-0; I-1;III,4 0-1; 6

Leg 4 I-0; III absent

Leg 5 (Fig. 2H) located at posterolateral corner of

genital complex, represented by 2 papillae; outer

papilla bearing single plumose seta; inner (exopodal)

papilla bearing 2 plumose setae and located close to

egg-sac attachment area.

Adult male. Body (Fig. 3A) 3.74 mm (3.61–3.94)

long, excluding caudal setae. Cephalothoracic shield

longer than wide, 1.99–2.07 9 1.78–1.89 (2.02 9

1.85), excluding marginal hyaline membranes. Free

thoracic zone of shield wider than long, 0.91–1.00 9

1.20–1.29 (0.96 9 1.25). Fourth pedigerous somite

0.18–0.23 9 0.39–0.56 (2.21 9 0.50), distinctly

divided from genital complex. Genital complex

(Fig. 3B) 0.69–0.77 9 0.6–0.66 (0.72 9 0.63), with

rounded corners and slightly convex to parallel sides.

Syst Parasitol (2014) 89:23–32 27

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Fig. 3 Caligus solea n. sp. male. A, Habitus, dorsal view; B, Genital complex, ventral view; C, Antenna; D, Postantennal process and

sensillae (black arrows); E, Maxillule; F, Maxilliped; G, Maxilliped claw; H, Sternal furca; I, Legs 5 and 6. Scale-bars: A, 0.5 mm;

B–H, 100 lm; I, 50 lm

28 Syst Parasitol (2014) 89:23–32

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Abdomen comprising 2 somites; first free abdominal

somite 0.18–0.24 9 0.35–0.39 (0.22 9 0.37), shorter

than anal somite, 0.47–0.52 9 0.35–0.39 (0.50 9

0.37); combined length of first abdominal somite and

anal somite about equal to length of genital complex.

Caudal rami c.1.7 times longer than wide, bearing 6

pinnate setae.

Antennule as in female. Antenna (Fig. 3C) 3-seg-

mented; proximal segment long, narrow, with corru-

gated adhesion pad on mid-outer surface; middle

segment largest, with corrugated pads on medial and

distal surfaces; distal segment of antenna forming

recurved, tapering claw, armed with 2 slender basal

setae. Postantennal process (Fig. 3D) larger, more

strongly curved than that of female, carrying 2 papillae

each with 3 sensillae; similar papilla with 2 sensillae

located on body surface near postantennal process.

Maxillule (Fig. 3E) with basal corrugated pad and

small dentiform knob located medially on posterior

spinous process. Mandible and maxilla as in female.

Maxilliped (Fig. 3F) with massive corpus carrying

conspicuous conical process distally on myxal margin,

opposing tip of claw; myxal process with minute pore

at apex; subchela armed with small seta at base of claw

(Fig. 3G). Sternal furca (Fig. 3H) with diverging tines

and square box. Legs 1–4 as in female. Leg 5 (Fig. 3I)

represented by 2 papillae located on posterolateral

margins of genital complex, outer papilla with 1 and

inner papilla with 2 plumose setae. Leg 6 (Fig. 3I)

represented by a single papilla bearing 2 setae on

posteroventral side of genital complex.

Remarks

The new species belongs to a group of congeners

sharing possession of a three-segmented leg 4 which

carries a single distal spine on the first exopodal

segment plus three apical spines on the distal margin

of the compound distal exopodal segment. They lack

the outer spine derived from the ancestral second

exopodal segment. This form of leg 4 is characteristic

of the group of species within Caligus referred to as

the macarovi-group by Boxshall & Gurney (1980).

Other characteristics shared by members of this group

include a one-segmented abdomen in the female, a

two-segmented abdomen in the male and the presence

of a myxal process opposing the tip of the subchela on

the male maxilliped. The new species does not share

the remaining two typical characteristics of that group

Table 1 Species of Caligus characterised by the possession of

a three-segmented leg 4, with one spine on the first exopodal

segment and three distal spines only on the second exopodal

segment

1 Caligus absens Ho, Lin & Chen, 2000

2 Caligus aduncus Shen & Li, 1959

3 Caligus alatus Heegaard, 1943

4 Caligus amblygenitalis Pillai, 1961

5 Caligus antennatus Boxshall & Gurney, 1980

6 Caligus brevicaudatus A.Scott, 1901

7 Caligus brevis Shiino, 1954

8 Caligus callyodoni Prabha & Pillai, 1986

9 Caligus calotomi Shiino, 1954

10 Caligus dampieri Byrnes, 1987

11 Caligus dasyaticus Ragnekar, 1957

12 Caligus eventilis Leigh-Sharpe, 1934

13 Caligus fistulariae Yamaguti, 1936

14 Caligus flexispina Lewis, 1964

15 Caligus hamruri Pillai, 1964

16 Caligus itacurussensis Luque & Cezar, 2000

17 Caligus kalumai Lewis,1964

18 Caligus klawei Shiino, 1959

19 Caligus littoralis Luque & Cezar, 2000

20 Caligus longiabdominalis Shiino, 1959

21 Caligus longicaudatus Brady, 1899

22 Caligus longipedis Bassett-Smith, 1889

23 Caligus longispinosus (Heegaard, 1962)

24 Caligus macarovi Gussev, 1951

25 Caligus nolani Longshaw, 1997

26 Caligus orientalis Gussev, 1951

27 Caligus oviceps Shiino, 1952

28 Caligus pampi Lin & Ho, 2002

29 Caligus patulus Wilson, 1937

30 Caligus planktonis Pillai, 1985

31 Caligus polycanthi Gnanamuthu, 1950

32 Caligus praecinctorius Hayes, Justine & Boxshall, 2012

33 Caligus pseudokalumai Lewis, 1968

34 Caligus punctatus Shiino, 1955

35 Caligus rogercresseyi Boxshall & Bravo, 2000

36 Caligus rugosus Shiino, 1959

37 Caligus scabei Gnanamuthu, 1950

38 Caligus sclerotinosus Roubal, Armitage & Rohde, 1983

39 Caligus sensilis Kabata &Gussev, 1966

40 Caligus sibogae Boxshall & Gurney, 1980

41 Caligus stokesi Byrnes, 1987

42 Caligus tenuicaudatus Shiino, 1959

43 Caligus thyrsitae Kazachenko, Korotaeva & Kurochkin, 1972

44 Caligus triangularis Shiino, 1954

45 Caligus wilsoni Delamare Deboutteville & Nunes-Ruivo, 1958

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as it has two sensillae on each of the three papillae

associated with the postantennal process (see Fig. 1E,

black arrows) (compared to a single sensilla typical of

other members of the macarovi-group) and it also

lacks any marginal serrations or denticles on the

maxillary brachium. There were originally 28 species

listed in the macarovi-group (Boxshall & Gurney,

1980) but numerous others (see Table 1) have been

added since 1980, including commercially important

species such as C. rogercresseyi Boxshall & Bravo,

2000 (see Boxshall & Bravo, 2000).

Considering the length to width proportions of the

female genital complex and of the abdomen (length about

equal to, or just greater than width), and the relative length

of the female genital complex and abdomen into account

(abdomen c.60% length of genital complex), a total of ten

species are comparable with the new species, as follows:

C. dasyaticus Rangnekar, 1957, C. latus Byrnes, 1987, C.

macarovi Gusev, 1951, C. nolani Longshaw, 1997, C.

oviceps Shiino, 1952, C. pampi Ho & Lin, 2002, C.

stokesi Byrnes, 1987, C. thyrsitae Kazachenko, Korota-

eva & Kurochkin, 1972, C. triangularis Shiino, 1954, and

C. wilsoni Delamare Deboutteville & Nunes-Ruivo,

1958. One of the other species from the macarovi-group,

C. longicaudatus Brady, 1899, is known only from the

male. As re-described by Parker (1968), the male of C.

longicaudatus differs from C. solea n. sp. in the

possession of a serrated margin on the brachium of the

maxilla (margin smooth in the new species), in the shape

of the sternal furca, in having seta 4 on the apex of leg 1

exopod about equal in length to spines 2 and 3 (compared

to more than twice as long in the new species), and in the

relatively longer distal spines on leg 4.

Two of the ten species, C. dasyaticus and C. pampi,

lack a posterior process on the first segment of the

antenna in the female. The new species differs from

both in having a well-developed posterior spinous

process on the antenna (Fig. 1D). Caligus dasyaticus

has an unusual armature at the tip of the exopod of

leg1, where the spines 1 to 3 are enlarged, almost as

long as the segment, and about twice as long as seta 4.

This is a unique configuration amongst species of the

macarovi-group. Caligus pampi has a highly modified

sternal furca, with both tines lying in close proximity,

adjacent to each other (Ho & Lin, 2002). This

configuration is unique amongst members of the

macarovi-group. The only other Caligus with a similar

sternal furca is C. bocki Heegaard, 1943, a member of

the productus-group (see Boxshall & El-Rashidy,

2009).

The new species can be distinguished from each of

the remaining eight species as follows:

• C. latus: The female genital complex of C. latus

has irregular lateral margins rather than weakly

convex margins as in the new species; the tines of

the sternal furca are slightly incurved and rounded

at the tip (vs straight and tapering in C. solea n.

sp.); the papillate sensillae associated with the

postantennal process are single rather than double,

as in the C. solea n. sp. The female abdomen is less

than half the length of the genital complex (vs

slightly more than half in the new species); and the

caudal rami are about half as long as the abdomen

(vs less than 25% in the new species).

• C. macarovi: The abdomen of the female is relatively

longer (c.85% of the length of the genital complex) in

C. macarovi, compared to only c.60% in the new

species; the papillate sensillae associated with the

postantennal process are single rather than double as

in C. solea n. sp.; the margin of the brachium of the

maxilla is serrated distally (vs margin smooth in the

new species), and a postoral process is present in

female C. macarovi but absent in the new species.

• C. nolani: The female abdomen represents less

than half the length of the genital complex (vs

slightly more than half in the new species); the

papillate sensillae associated with the postantennal

process are single rather than double as in C. solea

n. sp. The tines of the sternal furca are incurved

distally (vs straight in the new species); the second

abdominal segment of the male is about twice as

long as the first in C. nolani compared to 2.8 times

longer in C. solea n. sp., and the myxal process on

the male maxilliped is much larger in C. nolani

than in the new species.

• C. oviceps: The genital complex of the female is

very broad in C. oviceps, c.1.4 times wider than

long, compared to longer than wide in the new

species; the papillate sensillae associated with the

postantennal process are single rather than double

as in C. solea n. sp.; the margin of the brachium of

the maxilla is serrated distally (vs margin smooth

in the new species), and the caudal rami are almost

half as long as the abdomen (vs less than 25% in

the new species).

30 Syst Parasitol (2014) 89:23–32

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• C. stokesi: The genital complex of the female is

subtriangular in shape rather than subrectangular

as in the new species; the tines of the sternal furca

are spatulate and the box is narrower than in C.

solea n. sp.; the papillate sensillae associated with

the postantennal process are single rather than

double as in the new species; the female abdomen

is less than half the length of the genital complex

(vs slightly more than half); and the margin of the

brachium of the maxilla is serrated distally (vs

margin smooth in the new species),

• C. thyrsitae: The abdomen of the female is

relatively longer (c.85% of the length of the

genital complex) in C. thyrsitae, compared to only

c.60% in the new species; the papillate sensillae

associated with the postantennal process are single

rather than double as in the new species; and the

margin of the brachium of the maxilla is serrated

distally (vs margin smooth in C. solea n. sp.).

• C. triangularis: The genital complex of the female

is subtriangular in C. triangularis rather than being

subrectangular without posterolateral lobes as in

the new species; the female abdomen is less than

half the length of the genital complex (vs slightly

more than half); and the myxal process of the male

maxilliped is strongly bifurcated and located

proximally on the segment in C. triangularis

compared to the small conical process at mid-

length of the myxal margin in the new species.

In terms of body proportions of the adult female, the

new species most closely resembles C. wilsoni as

redescribed by Cressey (1991). This parasite was

originally reported by Wilson (1905) under the name

of C. belones Krøyer, 1863, based on material from

Coryphaena equiselis Linnaeus collected from off

Woods Hole. Delamare Deboutteville & Nunes-Ruivo

(1958) based their designation of C. wilsoni on Wilson’s

1905 description of C. belones, having ascertained that it

differed from C. belones Krøyer, 1863 in the form of leg

4. Cressey (1991) redescribed C. wilsoni based on new

material collected from Lutjanus griseus (Linnaeus)

taken off the coast of Florida. It can be distinguished

from C. wilsoni by a combination of characters. The

genital complex of the female is subtriangular and has

slightly lobate posterolateral corners in C. wilsoni rather

than being subrectangular without posterolateral lobes

as in the new species; the papillate sensillae associated

with the postantennal process are single rather than

double as C. solea n. sp. and the outer spine on the first

exopod segment of leg 4 reaches almost to the pecten at

the base of the outermost distal spine in C. wilsoni but

reaches only just beyond the mid-point of the margin in

the new species.

Discussion

Solea solea serves as host to a rich diversity of parasitic

copepods, 13 species in total, belonging to seven

genera. These species are: Acanthocondria soleae

(Krøyer, 1838), Acanthocolax exilipes (Wilson C.B.,

1911), Bomolochus solea Claus, 1864, Caligus apodus,

C. brevicaudatus, C. diaphanus, C. elongatus, Ergasi-

lus lizae Krøyer, 1863, Lepeophtheirus pectoralis

(Muller O.F., 1776), L. thompsoni Baird, 1850, Lern-

aeocera branchialis (Linnaeus, 1767), L. lusci (Bas-

sett-Smith, 1896) and Sphyrion lumpi (Krøyer, 1845).

Of the four previously reported species of Caligus, the

first, C. apodus, can be distinguished by the absence of

the fourth leg in the adult female (Ozak et al., 2013).

Caligus brevicaudatus can readily be distinguished

from the new species by its very short abdomen.

Caligus diaphanus differs from the new species in

having two-segmented abdomen (vs one-segmented)

in the adult female and C. elongatus can be distin-

guished from the new species by the possession of a

total of five spines (vs four) on the exopodal segments

of leg 4. Each species of Caligus reported from S. solea

shows irregular distribution on all over the upper body

surface. Like all other congeners, these previously

reported species also have single small spine (vs one

spine and a conspicuous spinule in C. solea n. sp.) at the

outer distal corner of the first exopodal segment of leg

1. The discovery of Caligus solea n. sp. in the

Mediterranean brings to 28 the total number of valid

species of Caligus reported in the region.

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