FOLIA PARASITOLOGICA 53: 287–296, 2006
Camallanus cotti (Nematoda: Camallanidae), an introduced parasite
of fishes in New Caledonia
František Moravec1 and Jean-Lou Justine2
Institute of Parasitology, Biology Centre, Academy of Sciences of the Czech Republic, Branišovská 31, 370 05 České
Budějovice, Czech Republic;
1
2
Équipe Biogéographie Marine Tropicale, Unité Systématique, Adaptation, Évolution (UPMC, CNRS, MNHN, IRD), Institut de
Recherche pour le Développement, BP A5, 98848 Nouméa Cedex, New Caledonia
Key words: parasitic nematode, Camallanus cotti, introduced species, Awaous, Kuhlia, New Caledonia, South
Pacific
Abstract. A pathogenic Asian nematode species of Camallanus, C. cotti Fujita, 1927, was found in New Caledonia, South Pacific, for the first time; it was recorded from two native fishes, Awaous guamensis (Valenciennes) (Gobiidae) (prevalence 51%,
intensity 1–25) and Kuhlia marginata (Cuvier) (Kuhliidae) (a single specimen found), of the La Foa River, about 100 km north
of Nouméa; the latter represents a new host record. Morphological data on C. cotti based on New Caledonian specimens and
those previously collected from aquarium-kept Misgurnus anguillicaudatus (Cantor) in Canada have been provided. The SEM
examination of C. cotti, applied for the first time in this species, made it possible to study some of its morphological details; firststage larvae from the female’s uterus were found to possess several digit-like processes on the tail tip, not previously reported for
any species of the Camallanidae. Camallanus moraveci Petter, Cassone et France, 1974 is considered a junior synonym of C.
cotti. A list of hitherto recorded hosts of C. cotti is provided. Camallanus cotti is assumed to be introduced into New Caledonia
along with the introduction of the exotic poeciliid fishes, which are known to be among the most common hosts of C. cotti in
aquarium cultures worldwide.
New Caledonia is an interesting area from the zoogeographical point of view, with a high degree of endemism. Despite this fact, almost no attention has been
paid to the parasites of native fishes in freshwater habitats. Regarding parasitic nematodes, the only paper is
that recently published by Moravec et al. (2006a), dealing with a new species of Procamallanus Baylis, 1923
from Pacific eels (Anguilla spp.) in freshwater bodies,
including those in northern New Caledonia.
Gargominy et al. (1996) mentioned a total of 800
flowering plants, at least 400 invertebrates and 36 vertebrates now established in the wild in New Caledonia.
This includes several species of freshwater fishes (Marquet et al. 2003) introduced into New Caledonian
streams from different continents. However, along with
the introduction of fishes, also their parasites are often
introduced, which may subsequently endanger the native fish populations, as pointed out, e.g., by Font and
Tate (1994) and Font (1997a, b) in Hawaii. Therefore,
studies of parasites of New Caledonian freshwater fishes
and their inventories are highly needed.
In September of 2004, a sample of fishes was collected from the La Foa River near La Foa, northern New
Caledonia, where two native fish species were found to
be infected with the apparently introduced nematode
Camallanus cotti Fujita, 1927, a pathogenic parasite of
many fish species introduced from East Asia to many
countries of different continents. Although C. cotti has
several times been described, there are still some problems concerning its morphology, taxonomy and life
cycle. Moreover, this species has not previously been
studied using scanning electron microscopy (SEM). The
purpose of this paper is to treat this parasite on the basis
of New Caledonian material, with the use of previously
unpublished results by the senior author (F.M.) based on
specimens from aquarium-kept Misgurnus anguillicaudatus (Cantor) in Canada.
MATERIALS AND METHODS
Fishes were caught by electrofishing in the La Foa River
near La Foa, northern New Caledonia on 23 September 2004.
They were transported alive to the IRD laboratory in Nouméa
and immediately examined for the presence of helminth parasites. The following fish species were collected and examined
(the introduced species are marked with an asterisk): Anguillidae: giant mottled eel Anguilla marmorata Quoy et Gairmard
(n = 2) and speckled longfin eel Anguilla reinhardtii Steindachner (n = 11); Cichlidae: Mozambique tilapia *Oreochromis mossambicus (Peters) (n = 1); Gobiidae: Awaous
guamensis (Valenciennes) (n = 35), speckled goby Redigobius
bikolanus (Herre) (n = 1) and bichique Sicyopterus lagocephalus (Pallas) (n = 1); Kuhliidae: dark-margined flagtail Kuhlia
marginata (Cuvier) (n = 2) and Kuhlia rupestris (Lacépède) (n
= 2); Mugilidae: flathead mullet Mugil cephalus Linnaeus (n =
4) and (?) otomebora mullet Liza melinoptera (Valenciennes)
(n = 1); Poeciliidae: green swordtail *Xiphophorus helleri
Heckel (n = 3). Of them, only A. guamensis and K. marginata
Address for correspondence: F. Moravec, Institute of Parasitology, Biology Centre, Academy of Sciences of the Czech Republic, Branišovská 31,
370 05 České Budějovice, Czech Republic. Phone: ++420 387 775 432; Fax: ++420 385 310 388; E-mail: moravec@paru.cas.cz
287
harboured the nematodes. The nematodes for morphological
studies were fixed in hot 4% formaldehyde solution. For light
microscopical examination, nematodes were cleared with
glycerol. Drawings were made with the aid of a Zeiss microscope drawing attachment. Specimens used for SEM were
postfixed in 1% osmium tetroxide, dehydrated through a
graded ethanol series, critical point dried and sputter-coated
with gold; they were examined using a JEOL JSM-6300 scanning electron microscope at an accelerating voltage of 15 kV.
Caudal ends of the nematode first-stage larvae were studied in
broken female specimens. Specimens of Camallanus cotti
from aquarium-kept Misgurnus anguillicaudatus (Cobitidae)
in Canada (Nanaimo, British Columbia), provided by Dr. N.P.
Boyce, were examined during the senior author’s (F.M.) stay
at the Pacific Biological Station in Nanaimo, BC in the spring
1979 and were deposited there in the parasite collection. It is
unknown where the aquarium-kept M. anguillicaudatus was
caught and which fish species it may have been kept with. For
comparative purposes, voucher specimens of C. cotti from the
material of Rigby et al. (1997) collected from Awaous
guamensis, Poecilia reticulata, Xiphophorus helleri and Lentipes concolor of the Hakalau Stream, Hawaii and from an
aquarium-kept Potamotrygon sp., now deposited in the US
National Parasite Collection (USNPC Nos. 86994, 86995,
86997, 87002, 87004, 87005, 87062), were examined. All
measurements are in micrometres (µm) unless otherwise
stated. Fish names follow FishBase (Froese and Pauly 2006).
RESULTS
Camallanus cotti Fujita, 1927
Figs. 1–5
Syns.: Camallanus zacconis Li, 1941; C. fotedari Raina
et Dhar, 1972; C. moraveci Petter, Cassone et France,
1974.
Description (based on specimens from Awaous
guamensis): Medium-sized nematodes with finely
transversely striated cuticle and large orange-brown
buccal capsule typical of genus. Mouth aperture slitshaped, surrounded by four submedian cephalic papillae
and two lateral amphids. Valves of capsule roughly
pentagonal in lateral view, internally bearing smooth
longitudinal ridges, some incomplete. Anterior outer
surface of each valve with two longitudinally elongate
sclerotized plates. Narrow, sclerotized ring present at
bottom of capsule. Tridents large, only moderately surpassing posterior border of buccal capsule. Excretory
pore somewhat posterior to level of nerve ring. Deirids
small, slightly asymmetrical, located at about two thirds
of muscular oesophagus.
Male (5 specimens): Length of body 2.60–3.63 mm,
maximum width 163–231. Buccal capsule including
basal ring 96–105 long, maximum width 87–93; size of
basal ring 12 × 57–66; length of tridents 66–81. Each
valve of capsule internally with 14–18 longitudinal
ridges, 3–8 incomplete. Muscular oesophagus 340–381
long, 63–78 wide; glandular oesophagus 367–517 long,
66–87 wide; length ratio of both parts of oesophagus
1:1.08–1.36. Buccal capsule and oesophagus represent-
288
ing 25–31% of total body length. Nerve ring, excretory
pore and deirids 174–195, 222–246 and 291–321, respectively, from anterior extremity. Posterior end of
body with vesiculate caudal alae. Pedunculate papillae:
7 pairs preanal and 6 pairs postanal; papillae of first
preanal pair situated somewhat anterior to anterior ends
of caudal alae (Figs. 1 H,I, 2 D,I), postanal papillae of
first three pairs close together (Figs. 1 K, 2 E). Cloacal
aperture surrounded by 2 pairs of transverse sessile
papillae. Spicules unequal, simple; length of larger
(right) spicule 141–156, of smaller (left), less sclerotized spicule 81–132; length ratio of spicules 1:1.07–
1.93. Length of tail 90–111.
Female (5 gravid specimens; measurements of 1
subgravid specimen in parentheses): Body length of
specimens with larvae in uterus 7.00–9.44 (6.62) mm,
maximum width 367–490 (286). Buccal capsule including basal ring 147–174 (153) long, maximum width
129–150 (141); basal ring 18 (18) long, 81–90 (81)
wide; length of tridents 135–156 (141). Each valve of
capsule with 21–22 (21) ridges, 5–7 (5) incomplete.
Muscular oesophagus 517–653 (530) long, 105–117
(102) wide; glandular oesophagus 571–666 (571) long,
99–111 (108) wide; length ratio of both parts of oesophagus 1:1.09–1.29 (1:1.08). Buccal capsule and
oesophagus representing 15–19 (19) % of total body
length. Nerve ring, excretory pore and deirids 249–286
(252), 297–375 (291) and 411–557 (402), respectively,
from anterior extremity. Tail very elongate, 1.27–1.39
(1.12) mm long, representing 12–18 (17) % of body
length, with rounded tip without any processes. Posterior lip of anus with characteristic dotted sculpture (Fig.
2 J). Vulva slightly postequatorial (preequatorial), 3.60–
5.15 (3.17) mm from anterior end of body [at 51–55
(48) % of body length], with somewhat elevated lips.
Vagina muscular, directed posteriorly from vulva.
Uterus extending posteriorly into tail, filled with numerous larvae.
Female fourth-stage larva (1 specimen from Kuhlia
marginata): Body length 1.55 mm, maximum width
117. Four cephalic papillae present. Buccal capsule
including basal ring 69 long, 57 maximum wide; basal
ring 6 long, 39 wide; tridents absent. Each valve of
capsule externally with small median transverse sclerotized plate (anlage of two cephalic sclerotized plates
in adults) on anterior end and internally with 12 longitudinal ridges, 3 incomplete. Muscular oesophagus 285
long, 45 wide; glandular oesophagus 264 long, 45 wide;
length ratio of both parts of oesophagus 1:0.93. Buccal
capsule and oesophagus representing 40% of total body
length. Nerve ring and excretory pore 135 and 165,
respectively, from anterior extremity. Deirids not found.
Small, oval genital primordium situated ventrally 1.01
mm from anterior extremity (at 65% of body length).
Tail conical, 162 long, with 3 small conical cuticular
spikes on tip; spikes 3 long.
Moravec, Justine: Camallanus cotti in New Caledonia
Fig. 1. Camallanus cotti Fujita, 1927, adults from Awaous guamensis, New Caledonia. A, B – anterior end of gravid female,
lateral and dorsoventral views; C, D – buccal capsule, lateral and dorsoventral views; E – tail of gravid female, lateral view; F –
deirid, lateral view; G – vulva, lateral view; H, I – posterior end of male, ventral and lateral views; J, K – tail of male, lateral and
ventral views. Scale bars: A, B, E = 200 µm; C, D, G–I = 100 µm; F = 20 µm; J = 50 µm; K = 30 µm.
H o s t s : Awaous guamensis (Gobiidae) (body length 66–145
mm) [adults and fourth-stage larvae] and dark-margined
flagtail Kuhlia marginata (Kuhliidae) (body length 62 mm)
(both Perciformes) [fourth-stage larva].
S i t e o f i n f e c t i o n : Intestine.
L o c a l i t y : La Foa River, La Foa, northern New Caledonia
(collected 23 September 2004).
P r e v a l e n c e a n d i n t e n s i t y : A. guamensis: 51%
(18 fish infected / 35 fish examined); 1–25 (mean 5) nematodes per fish. K. marginata: 1 / 1; 1.
D e p o s i t i o n o f v o u c h e r s p e c i m e n s : Muséum
National d’Histoire Naturelle, Paris (Cat. Nos. JNC1308–
1330) and Institute of Parasitology, Biology Centre, ASCR,
České Budějovice (Cat. No. N-11).
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Fig. 2. Camallanus cotti Fujita, 1927 from Awaous guamensis, scanning electron micrographs of adults. A, B – cephalic end,
dorsoventral and apical views; C – deirid; D – papilla of first preanal pair; E, F – tail of male, ventral and lateral views (arrowheads show circumcloacal papillae); G – phasmid on female tail; H – detail of cloacal aperture and circumcloacal papillae
(arrowheads); I – posterior end of male, lateral view (arrowhead shows first preanal papilla situated anterior to caudal ala);
J – anal opening of female (note structure of posterior anal lip). Abbreviations: c – cephalic papilla; p – sclerotized plate of buccal capsule; t – trident. Scale bars: C, H = 1 µm; D = 5 µm; F = 10 µm.
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Moravec, Justine: Camallanus cotti in New Caledonia
4.01–4.40 mm from anterior extremity (at 47–56% of
body length). Tail 1.32–1.61 mm (294–902) long, forming 17–18 (15–21) % of body length.
Fourth-stage larva (2 specimens): Body length 0.90–
1.94 mm, maximum width 65–93. Buccal capsule including basal ring 41–64 long, 44–53 wide. Basal ring
5–7 long, 30–37 wide. Each valve of capsule with 15–
17 ridges. Tridents absent. Length of muscular oesophagus 152–230, of glandular oesophagus 138–276.
Nerve ring and excretory pore 87–136 and 145–207,
respectively, from anterior extremity. Deirids not found.
Tail 124–207 long, forming 11–14% of body length,
with three minute spikes on tip.
DISCUSSION
Fig. 3. Camallanus cotti Fujita, 1927, fourth-stage larva from
Kuhlia marginata, New Caledonia. A – general view; B –
anterior end, lateral view; C – tail, lateral view. Scale bars: A
= 200 µm; B, C = 30 µm.
Measurements of Camallanus cotti from aquariumkept Misgurnus anguillicaudatus in Nanaimo, British
Columbia, Canada
Male (7 specimens): Body length 1.84–3.04 mm,
maximum width 93–167. Buccal capsule including
basal ring 69–87 long, 69–81 maximum wide. Basal
ring 7–9 long, 46–51 wide. Each valve of capsule with
15–17 ridges. Length of tridents 69–83. Length of muscular oesophagus 235–294, of glandular oesophagus
290–460. Nerve ring and excretory pore 140–184 and
161–317, respectively from anterior extremity. Deirids
not located. Pedunculate subventral papillae: 7 pairs
preanal and 6 pairs postanal; 2 pairs of sessile ventral
papillae surrounding cloacal aperture. Larger (right)
spicule 115–150 long, smaller (left), less sclerotized
spicule 92–110 long. Length of tail 92–108.
Female (3 gravid specimens; measurements of 3 juvenile specimens in parentheses): Body length of larvigerous specimens 7.92–8.84 mm (juvenile specimens
2.02–4.32), maximum width 279–326 (112–186). Buccal capsule including basal ring 120–143 (81–122),
133–138 (94–115) maximum wide. Basal ring 12–14
(7–12) long, 64–76 (57–81 wide. Each valve of capsule
with 16–18 (15–17) ridges. Length of tridents 97–108
(76–108). Length of muscular oesophagus 428–437
(253–469), of glandular oesophagus 644–736 (258–
391). Nerve ring 230–235 (184–267) from anterior
extremity; excretory pore and deirids not located. Vulva
The morphology and measurements of New Caledonian specimens correspond, more or less, to C. cotti, as
it has been redescribed by Moravec and Nagasawa
(1989) from the type locality in Japan and by Moravec
and Sey (1988) and Moravec et al. (2003) from fishes in
Vietnam and China, respectively. Unlike the abovementioned authors and previous data by Fujita (1927a,b)
and Yamaguti (1935, 1941), Rigby et al. (1997), in C.
cotti specimens from Poecilia reticulata (Peters) introduced into Hawaii, illustrated the tridents of the buccal
capsule reaching to the level of the nerve ring, caudal
alae joining anteriorly, and the larger spicule as the left
one (generally in camallanids the longer spicule is on
the right side); according to Moravec et al. (2003), it
could not be excluded that, in fact, they dealt with a
different species. However, a recent reexamination of
their voucher specimens showed that the morphology of
the Hawaii specimens does not differ from that of C.
cotti in other regions. The SEM examination, applied in
this species for the first time, confirmed the presence of
small deirids (Fig. 2 C) and caudal alae not joining
anteriorly (Fig. 2 I). It showed that, in contrast to some
other Camallanus spp. studied by SEM (e.g., Moravec
et al. 2006b), the female posterior anal lip had a characteristic sculpture (Fig. 2 J) and that the first pair of preanal papillae in male may be situated anterior to caudal
alae (Fig. 2 D, I).
A SEM examination of the first-stage larvae from the
C. cotti female’s uterus revealed that their tail tip was
provided with a crown of several minute digit-like processes (Fig. 5), not previously reported for any species of
Camallanus. They are almost invisible under the light
microscope. Although the first-stage larva of Camallanus oxycephalus Ward et Magath, 1916 was studied
by SEM (Kelly et al. 1989), no such caudal processes
were found; on the contrary, similar processes were
found by SEM in the larvae of this stage in Camallanus
lacustris (Zoega, 1776) (Drábek and Moravec, unpublished) (Fig. 6). Apparently, these caudal processes
serve the larva to the better attachment by its tail to the
bottom, after the larvae are released into the water; the
larvae attach themselves to the bottom by their tails and
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Fig. 4. Camallanus cotti Fujita, 1927, adults and fourth-stage larvae from aquarium-reared Misgurnus anguillicaudatus, Canada.
A, B – buccal capsule of gravid female, lateral and dorsoventral views; C–F – development of tail in female (C – young specimen with body length 2.02 mm, D – with body length 2.84 mm, E – with body length 4.32 mm, F – gravid specimen 7.92 mm
long); G, H – posterior end of male, ventral and lateral views; I – vulva, lateral view; J, K – buccal capsule of fourth-stage larva,
lateral and dorsoventral views; L, M – tail tip of fourth-stage larva, lateral and ventral views; N – tail tip of specimen undergoing
last (fourth) moult, ventral view; O – tail of female fourth-stage larva, lateral view; P – tail of male fourth-stage larva, lateral
view. Scale bars: A, B, G–I, O, P = 100 µm; C–F = 300 µm; J, K = 50 µm; L–N = 20 µm.
292
Moravec, Justine: Camallanus cotti in New Caledonia
Fig. 6. Camallanus lacustris (Zoega, 1776), scanning electron
micrographs of the tail end of the first-stage larva (specimens
from Perca fluviatilis of the Mácha Lake fishpond system,
Czech Republic, collected in June 1997). Scale bar = 1 µm.
Fig. 5. Camallanus cotti Fujita, 1927, scanning electron micrographs of the tail end of the first-stage larva. A, B – subapical and lateral views, respectively. Scale bars: A, B = 100
nm.
coil and discoil their bodies, attracting thus by these
movements the copepod intermediate hosts. The presence/absence and the character of caudal processes in
first-stage larvae might be an important taxonomic feature for different Camallanus spp.
The fourth-stage larva of C. cotti was described from
experimentally infected fishes by Levsen and Berland
(2002a). They reported only two mucrons on the tail tip
of this larval stage, but there were always three small
spikes or mucrons present in the fourth-stage larvae of
C. cotti from both Canada (Fig. 4 L–N) and New Caledonia (Fig. 3 C), of which the dorsal one was larger than
both subventral. In having three caudal spikes, C. cotti
fourth-stage larvae resemble those of other Camallanus
spp. [e.g., C. carangis Olsen, 1952, C. lacustris, C.
oxycephalus, C. truncatus (Rudolphi, 1814) – see Moravec 1969, 1971, Stromberg and Crites 1974, Moravec
et al. 2006b]. The buccal capsule of C. cotti fourth-stage
larvae is similar to that in adults, but the tridents are
absent. The fourth-stage larva of C. carangis, a species
parasitizing marine fishes, has recently been found to
have the buccal capsule provided with two sclerotized
rod-like structures (Moravec et al. 2006b); these are,
however, absent from the capsule of the larvae of C.
cotti, by which this species resembles some other Camallanus spp. (C. lacustris, C. oxycephalus, C. truncatus).
A characteristic feature of C. cotti is the unusually
long tail of the gravid female, which is an important
taxonomic feature distinguishing C. cotti from most
other congeneric species (Moravec 1973). Fig. 3 N
shows that the three small caudal spikes, which are
present in fourth-stage larvae, are lost during the last
(fourth) moult. However, a considerable increase in tail
length occurs after the moult (Fig. 3 C–F), along with
the body growth and development of female reproductive organs, as also demonstrated by Levsen and Berland 2002b). Therefore, only gravid (i.e., larvigerous)
females of Camallanus spp. should be compared, as it is
used in philometrid nematodes (Moravec 2004). By the
conspicuously long tail in gravid female, C. cotti resembles C. oxycephalus from North American fishes and C.
kirandensis Baylis, 1928 and C. longicaudatus Moravec, 1973 from cyprinids in Africa; however, these are
distinctly larger than C. cotti and differ in some other
characters.
Camallanus cotti was originally described by Fujita
(1927a,b) from four fish species of Lake Biwa and Lake
Tazawa in Japan; later Yamaguti (1935, 1941) and Moravec and Nagasawa (1989) redescribed it from the
specimens newly collected from different Japanese
fishes. Moravec (1973) and Moravec and Sey (1988)
synonymized Camallanus zacconis Li, 1941 and C.
fotedari Raina et Dhar, 1972, two species described
from Korea and India, respectively (Li 1941, Raina and
Dhar 1972), with C. cotti. However, it is necessary to
synonymize also Camallanus moraveci Petter, Cassone
et France, 1974 described from Mollienesia latipinna [=
Poecilia latipinna (Lesueur)] and Xiphophorus helleri
Heckel imported from Singapore to France (Petter et al.
1974). This species should differ from C. cotti only in
the allegedly absence of the left spicule; however, the
left spicule is always present in Camallanus spp., but
due to its poor sclerotization and a certain intraspecific
variability, sometimes this spicule may seem to be ab-
293
sent (see Moravec et al. 2006b). On the other hand, the
hosts of C. moraveci belong to the most frequently
recorded hosts of C. cotti in aquarium conditions
worldwide. Probably identical with C. cotti is also
Camallanus maculatus Martins, Piazza et Garcia, 2004
reported from aquarium-reared Xiphophorus maculatus
(Günther) in Brazil (Martins et al. 2004), but since this
species has not yet been formally described (per. comm.
of M.L. Martins), C. maculatus should be considered a
nomen nudum.
Camallanus cotti has been reported from a number of
fish species belonging to different families and orders. It
seems it was originally distributed in eastern, southeastern and southern Asia, from where it has been recorded
from Japan, China, eastern Russia (Amur R. basin),
Korea, Vietnam, Singapore and India (e.g., Fujita
1927a,b, Yamaguti 1935, 1941, Li 1941, Dogiel and
Akhmerov 1959, Roytman 1963, Raina and Dhar 1972,
Chen 1973, Petter et al. 1974, Wang et al. 1979, 1997,
Wu 1984, Vismanis et al. 1987, Moravec and Sey 1988,
Moravec and Nagasawa 1989, Wu et al. 1991, Kim et
al. 2002, Moravec et al. 2003). It also occurs in the
cultures of aquarium fishes in Europe, North America
(Canada), South America (Brazil), West Indies (Puerto
Rico) and Australia, where this pathogenic parasite is
being brought in along with exotic fishes, imported
mainly from Singapore (e.g., Schubert 1972, Petter et al.
1974, Stumpp 1975, Campana-Rouget et al. 1976,
Bunkley-Williams and Williams 1994, Font and Tate
1994, Alves et al. 2000, Evans and Lester 2001, Levsen
2001, Levsen and Berland 2002a,b, Levsen and Jakobsen 2002, Menezes et al. 2006). Camallanus cotti
has also been reported from native and introduced fishes
in some streams in Hawaii, where it was apparently
brought in along with the introduction of poeciliids
(Font and Tate 1994, Font 1997a, b, 1998, 2003, Vincent and Font 2003a, b).
In New Caledonia (La Foa River), adults and a few
larvae of C. cotti were recorded only from the gobiid
Awaous guamensis, with a relatively high prevalence
(51%) and intensity (up to 25), whereas only a single
fourth-stage larva was found in the dark-margined flagtail Kuhlia marginata. Other native fishes examined,
including gobiids (R. bikolanus and S. lagocephalus)
and another Kuhlia species, as well as the introduced
fishes, were not infected (see the list in Material and
Methods). It is remarkable that neither the introduced
Xiphophorus helleri was infected, which is one of the
most frequently reported hosts of C. cotti in aquarium
cultures and in the streams in Hawaii (Rigby et al. 1997,
Vincent and Font 2003a); but only three specimens of
this fish were examined. Awaous guamensis, a species
widespread in Oceania (Froese and Pauly 2006), was
found to be infected with C. cotti also in Hawaii (Rigby
et al. 1997, Font 2003). It is highly probable that, similarly as in Hawaii, C. cotti was introduced into New
Caledonia along with infected poeciliids. This is the
294
first record of C. cotti from New Caledonia and its finding in Kuhlia marginata represents the first host record.
It can be expected that more New Caledonian fish species will be found as hosts of this nematode parasite.
The host specificity of C. cotti is low and, as far as
we could find, it has been recorded from the following
fish species (a = in aquarium culture; b = an introduced
species; c = an autochtonous species parasitized with the
introduced C. cotti; d = in both natural waters and aquarium culture): Rajiformes: Potamotrygonidae: aPotamotrygon sp.; Acipenseriformes: Acipenseridae: Huso
dauricus (Georgi); Cypriniformes: Balitoridae: Triplophysa kashmirensis (Hora); Cobitidae: aChromobotia
macracanthus (Bleeker), dMisgurnus anguillicaudatus
(Cantor); Cyprinidae: Acanthorhodeus asmussii (Dybowski), Acheilognathus taenianalis (Günther), Culter
erythropterus Basilewsky, Culter mongolicus Basilewsky, aDanio rerio (Hamilton), Elopichthys bambusa (Richardson), Gobio cynocephalus Dybowski, Hemibarbus
labeo (Pallas), Hemiculter leucisculus (Basilewsky),
Leuciscus waleckii (Dybowski), Megalobrama terminalis (Richardson), Ochetobius elongatus (Kner), Opsariichthys uncirostris (Temminck et Schlegel), Parabramis pekinensis (Basilewsky), Phoxinus lagowskii
Dybowski, Pseudaspius leptocephalus (Pallas), Squaliobarbus curriculus (Richardson), Tribolodon hakonensis
(Günther), Zacco platypus (Temminck et Schlegel),
Zacco temmincki (Temminck et Schlegel); Siluriformes:
Bagridae: Pelteobagrus fulvidraco (Richardson); Callichthyidae: aCorydoras sp.; Siluridae: Parasilurus
asotus (Linnaeus) (type host); Sisoridae: Bagarius bagarius (Hamilton); Salmoniformes: Salmonidae: Oncorhynchus masou (Brevoort); Cyprinodontiformes:
Aplocheilidae: aFundulopanchax gardneri (Boulenger);
Poeciliidae: bGambusia affinis (Baird et Girard),
b
Poecilia mexicana Steindachner, a,bPoecilia reticulata
Peters, a,bXiphophorus helleri Heckel, aXiphophorus
maculatus (Günther); Scorpaeniformes: Cottidae: Cottus
pollux (Günther), Cottus reinii Hilgendorf, Mesocottus
haitej (Dybowski); Perciformes: Channidae: Channa
argus (Cantor); Cichlidae: aGeophagus surinamensis
(Bloch), aPterophyllum sp., aSymphysodon sp.; Eleotridae: cEleotris sandwicensis Vaillant et Sauvage; Gobiidae: cAwaous guamensis (Valenciennes), Chaenogobius
macrognathos (Bleeker), cLentipes concolor (Gill),
Rhinogobius lindbergi Berg, Rhinogobius similis Gill,
c
Stenogobius hawaiiensis Watson; Kuhliidae: cKuhlia
marginata (Cuvier); Odontobutidae: Odontobutis obscura (Temminck et Schlegel), Percottus glenii Dybowski; Osphronemidae: aBetta splendens Regan,
Macropodus chinensis (Linnaeus), cMacropodus opercularis (Linnaeus); Percichthyidae: Siniperca chuatsi
(Basilewsky); Percidae: bPerca fluviatilis Linnaeus.
As in other members of the Camallanidae, the life
cycle of C. cotti involves a copepod intermediate host;
Cyclops sp., Macrocyclops albidus (Jurine) and Mesocyclops leuckarti (Claus) were found to serve as its suit-
Moravec, Justine: Camallanus cotti in New Caledonia
able experimental intermediate hosts (Wang and Ling
1975, Campana-Rouget et al. 1976, Levsen and Berland
2002a, b). Some authors (Stumpp 1975, Levsen 2001,
Levsen and Jakobsen 2002) consider this species to be
also capable of a direct (monoxenous) development.
However, results of the experiments performed by
Levsen and Jakobsen (2002) to prove this are not quite
conclusive and the presumed monoxenous development
of C. cotti would need a reliable verification.
Acknowledgements. The authors wish to thank Dr. Christine
Pöllabauer (Erbio, Nouméa, New Caledonia) for organizing an
electrofishing expedition to the La Foa River, her collaboration, fish identification and fruitful discussions. Eric Bureau, a
student, participated in the fishing expedition and parasitological examination of fish. Angelo Di Matteo (IRD, Nou-
méa) is thanked for technical help. Drs. Norbert P. Boyce and
late Leo Margolis (Pacific Biological Station, Nanaimo, British Columbia) kindly provided specimens of C. cotti from
aquarium-kept Misgurnus anguillicaudatus and laboratory
facilities during the senior author’s stay in Canada in 1979.
Thanks are also due to the staff of the Laboratory of Electron
Microscopy of the Institute of Parasitology, ASCR, in České
Budějovice for their technical assistance and Irena Husáková
from the Department of Helminthology of the same Institute
for her help with the preparation of illustrations. Voucher
specimens of Camallanus cotti from Hawaii were kindly lent
by Dr. Patricia Pilitt, US National Parasite Collection, Beltsville, Maryland, USA. This study was partly supported by
grant no. 524/06/0170 from the Grant Agency of the Czech
Republic and by the research projects of the Institute of Parasitology, ASCR (Z60220518 and LC522).
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Received 23 May 2006
Accepted 24 July 2006
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