Experimental Parasitology 125 (2010) 55–62
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Experimental Parasitology
journal homepage: www.elsevier.com/locate/yexpr
Parabrachiella jarai sp. nov. (Crustacea: Copepoda: Siphonostomatoida) parasitic on
Sillago sihama (Actinopterygii: Perciformes: Sillaginidae)
Wojciech Piasecki a,*, Magdalena Młynarczyk a, Craig J. Hayward b,c
a
Division of Fish Diseases, Faculty of Food Sciences and Fisheries, West Pomeranian University of Technology, ul. Kazimierza Królewicza 4, 71-550 Szczecin, Poland
South Australian Research and Development Institute-Aquatic Sciences and the Australian Seafood Cooperative Research Centre, Lincoln Marine Science Centre,
Hindmarsh Street, Port Lincoln, SA 5606, Australia
c
National Centre for Marine Conservation and Resource Sustainability, University of Tasmania, Locked Bag 1-370, Launceston Tas 7250, Australia
b
a r t i c l e
i n f o
Article history:
Received 10 April 2009
Received in revised form 24 September 2009
Accepted 14 October 2009
Available online 20 October 2009
Keywords:
Parasite
New species
Copepoda
Parabrachiella
Thysanote
Fish
Sillago sihama
Sillaginidae
a b s t r a c t
The presently reported study provides a detailed morphological description of the female and the male of
a new species of the genus Parabrachiella—Parabrachiella jarai sp. nov. The parasites were sampled from
marine fish, silver sillago, Sillago sihama (Perciformes: Sillaginidae), captured in Malaysia in 1994 and
Hong Kong in 1995. The new species bears some resemblance to Parabrachiella lata (Song et Chen,
1976) but differs from it in details of second antenna, mandible, and maxilliped.
The genus Parabrachiella currently covers 67 species including those recently transferred from Neobrachiella Kabata, 1979. An amended generic diagnosis is proposed for Parabrachiella and Thysanote. Some
members of Parabrachiella are herewith transferred to Thysanote and some Thysanote are now placed in
Parabrachiella.
Ó 2009 Elsevier Inc. All rights reserved.
1. Introduction
Lernaeopodid copepods (Copepoda: Lernaeopodidae) are parasites of predominantly marine fishes, but they also infect freshwater and brackish water species. They are one of the largest families
of parasitic copepods (some 250 valid species) and most extensively adapted to parasitism (Kabata, 1986). Lernaeopodids are also
very host specific.
Only recently the genus Parabrachiella regained its senior position. Twenty years ago Kabata (1979) established the genus Neobrachiella, to accommodate selected species of the genus Brachiella,
as well as species previously assigned to other genera such as
Parabrachiella, Probrachiella, Epibrachiella, Branchiellina, Lernaeopoda, and Isobranchia. Boxshall and Halsey (2004) noticed that
the name Parabrachiella Wilson, 1915 should have priority over
the name Neobrachiella Kabata, 1979. Therefore all of the former
Neobrachiella species require a transfer to Parabrachiella.
One of us (CJH) surveyed fish, in the frames of his PhD studies,
and collected a number of parasite specimens, including crustaceans. The presently reported study focuses on one of those para* Corresponding author. Fax: +48 91 449 6657.
E-mail addresses: wojciech.piasecki@zut.edu.pl, editor@aiep.pl (W. Piasecki).
0014-4894/$ - see front matter Ó 2009 Elsevier Inc. All rights reserved.
doi:10.1016/j.exppara.2009.10.001
site species, which turned out to be a new species. One of the
host fishes studied (Sillago sihama) hosted two species of the genus
Parabrachiella. One of them was P. sihama, while the other represented species hitherto unknown to science. The latter species is
described below as Parabrachiella jarai sp. nov.
Another, closely related, genus is Thysanote Krøyer, 1863. Females belonging to this genus have distinct fimbriate posteriorand maxillary processes (Piasecki et al., 2008). The structure of
the male body is the same as in the genus Parabrachiella. Surprisingly, some species without maxillary processes traditionally belonged to Thysanote and vice versa—species with maxillary
processes have been assigned to Neobrachiella. The need for preparing a checklist for species of Parabrachiella prompted us to propose the species transfers, between two above-mentioned genera,
where necessary.
2. Materials and methods
The parasites were sampled from the operculum of a marine
fish, silver sillago, Sillago sihama (Forsskål) (Perciformes: Sillaginidae), captured in Malaysia (40 fish) and Hong Kong (38 fish) in
1994 and 1995, respectively. Examination of specimens of related
sillaginids occurring in South-East Asia (Sillago parvisquamis,
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W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
S. soringa, S. aeolus, S. chondropus, and Sillaginopsis panijus) and
Middle East (Sillago arabica, S. attenuata) did not reveal those copepods, and nor was the species present on S. sihama sampled from
any other localities in South-East Asia, Middle East, eastern Africa,
Australia, and islands in the western Pacific Ocean, or on any of a
total of 25 other sillaginid species examined. The study material
consisted of ovigerous- and non-ovigerous females and males attached to females. The material studied were 10 copepod females
and 3 males from Hong Kong and 1 female from Malaysia. The
copepods were studied under a compound microscope using a
modified ‘‘wooden slide” method of Humes and Gooding (1964).
Details of all appendages were illustrated and described. The types
are deposited in the Museum für Naturkunde in Berlin, Germany.
Terminology of the body parts is consistent with that proposed
by Kabata (1979) (with some exceptions). Appendage abbreviations used throughout the text are as follows: first antenna = A1,
second antenna = A2, mandible = Mdb, first maxilla = Mx1, second
maxilla = Mx2, maxilliped = Mxp.
The siphonostomatoid mandibular dental formula (Siphonostomatoida including Lernaeopodidae) used here was first introduced
by Kurtz (1877). Kabata (1979) proposed its Anglicized version
which has been widely used ever since. The tip of mandible has a
row of denticles on one edge. The anterior section is occupied by
larger, primary teeth (P) with distinct interdenticular spaces.
Sometimes those spaces feature much smaller, secondary (S) teeth.
Posterior denticles are smaller, usually uniform in size and shape
(sometimes diminishing posteriorly) and they are called basal
teeth (B). The dental formula shows the sequence of those teeth
starting from distalmost tooth (e.g., P2, S1, P1, S1, B4.).
Valid scientific names of host fishes were determined using
FishBase (Froese and Pauly, 2009).
3. Results
3.1. Parabrachiella jarai sp. nov. (Figs. 1–22)
FEMALE (Figs. 1–14). Body typically lernaeopodid (Figs. 1–4),
consisting of three distinct parts: cephalosome, genital trunk, and
maxillary ‘‘trunk” (fused Mx2). Cephalosome elongate, 2.11 mm
long (1.96–2.25 mm), regularly cylindrical; on fixed specimens
flexed dorsally. Genital trunk sub-spherical, or drop-shaped,
2.25 mm long (1.88–2.39 mm), 1.5 mm wide (1.37–1.68 mm); its
length slightly exceeding length of cephalosome. Posterior part
outfitted with two pairs of digitiform posterior processes, one of
which possibly representing caudal rami. Relative length of processes variable, approximating trunk width or even length
(Fig. 4). Small but distinct genital process present between posterior processes. Egg sacs multiseriate, twice as long as genital trunk.
Maxillary ‘‘trunk” relatively short. Appendages (except Mx2) situated at distal end of cephalosome. Anteriormost second antennae
followed by first antennae and mouth cone with inserted mandibles and flanked by first maxillae. Second maxillae displaced anteriorly in front of maxillipeds and located closely behind mouth
cone.
First antenna (Fig. 5) sub-cylindrical; three-segmented. Basal
segment almost as long as two other segments combined, and
armed subterminally with single seta (whip). Terminal segment
with well developed terminal armature, consisting of three long
setae and three tubercles.
Second antenna (Figs. 6 and 7) biramous. Robust, cylindrical
sympod unarmed. Bulbous, one-segmented, unarmed exopod distinctly longer and thicker than endopod. Endopod two-segmented
with small, almost straight claw 1, aligned with long axis of endopod, small seta 2, seta 5 and prominent denticulate pad 4 (Figs. 6
and 7).
Mandible (Fig. 8) typically siphonostome with 9 teeth. Dental
formula: P2, S1, P1, S1, B4.
First maxilla (Fig. 9) biramous with small endopod and prominent tripartite exopod. Endopod composed of short digitiform process surmounted with two setae: small terminal and larger
subterminal. Exopod tripartite with two big digitiform processes
and short third one. Processes ending with conical elongate setiferous processes.
Second maxillae (Fig. 10) completely fused together and forming ‘‘maxillary trunk”. In fixed specimens contracted and wrinkled.
Paired openings of maxillary glands visible at base, ventrally.
Small, mushroom-shaped bulla partly hidden inside terminal collar
(Figs. 3, 4, and 10).
Maxilliped (Figs. 11 and 12) subchelate with robust corpus and
long slender subchela. Corpus robust, covered by thick, wrinkled
cuticle. On its medial side single process with seta and denticulate,
semispherical pad. Subchela sub-cylindrical with single ventral
seta. Claw large (constituting almost 1/3 of subchela) with secondary tooth. Large auxiliary seta at base of claw, medially.
Thoracic appendages not observed.
Caudal rami (Figs. 1–4) probably represented by ventral pair of
posterior processes. Genital process (Fig. 13) consisting of two conical parts partly fused together with attached spermatophores
(Fig. 14) in some females examined.
MALE (Figs. 3 and 15–22) distinctly smaller than female
(Fig. 15); representing male structural type A (Kabata, 1979); consisting of two major parts: cephalosome (0.57 mm long) and genital trunk (0.65 mm long). Long axis of genital trunk inclined at
almost right angle to long axis of cephalosome. Appendages arranged in undisturbed order: A1, A2, Mdb, Mx1, Mx2, Mxp.
First antenna (Fig. 16) three-segmented. Basal segment longest
with small seta (whip). Terminal segment with well developed
armature consisting of 4 setae and 1 tubercle.
Second antenna (Fig. 17) biramous, elongate. Sympod cylindrical, unarmed. Bulbous, one-segmented, unarmed exopod distinctly shorter than endopod. Endopod two-segmented with
basal segment with denticulate pad. Terminal segment with well
developed lernaeopodid armature consisting of big, curved claw
1, big seta 2, tubercle 3 (at base of seta 2), denticulate pad 4,
and big seta 5.
Mandible (Fig. 18) typical siphonostome with 9 teeth. Dental
formula: P2, S1, P1, S1, B5 (last basal tooth small).
First maxilla (Fig. 19) similar as in female, although more slender. Endopod terminating with 2 equal small setae.
Second maxilla (Fig. 20) subchelate (made of strong thick cuticle) with robust pyriform corpus and strong subchela. Corpus unarmed. Subchela with well delimited, powerful claw, and slightly
shorter cylindrical shaft. Subchela closing against large bulbous
outgrowth of corpus.
Maxilliped (Fig. 21) subchelate, similar in structure to second
maxilla but stronger in appearance. Subchela very robust with
claw positioned at right angle to shaft. Closed subchela partly hiding tip of claw behind medial outgrowth of corpus.
Thoracic appendages not observed.
Caudal rami (Fig. 22) partly hidden in depression of cuticle;
consisting of paired conical outgrowths with obscure segmentation
and no additional setation.
Type host: Sillago sihama (Forsskål).
Site of infection: Inner surface of operculum.
Type locality: Sai Kung, Hong Kong.
Prevalence and intensity: 40 host fish from Sai Kung, Hong Kong
(December 1995) were infected by 10 females (and 3 males attached to them) from; 38 host fish from Cendering, Malaysia (April
1994) yielded 1 parasite female.
Etymology: The specific name of the new species is intended to
honour Professor Zbigniew J. Jara (1918–2008)—a renowned Polish
W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
57
Figs. 1–9. Parabrachiella jarai sp. nov. female. Fig. 1. Ovigerous specimen; whole. Fig. 2. Nonovigerous specimen; whole. Fig. 3. Ovigerous specimen (holotype); whole with
male (allotype) attached. Fig. 4. Another non-ovigerous specimen; whole. Fig. 5. First antenna. Fig. 6. Second antenna; whole. Fig. 7. Second antenna; endopod. Fig. 8.
Mandible. Fig. 9. First maxilla. Abbreviations: DP – dorsal papilla; M – male; 1, 2, 3, 4, 5 – armament of endopod of second antenna (according to Kabata, 1979). Scale bars in
mm.
fish pathologist and parasitologist (Lonc, 2004; Olech and Piasecki,
2009).
Deposition of types: Holotype, allotype, and 5 paratypes have
been deposited in the Museum für Naturkunde in Berlin (ZMB
27689) (all type specimens under a single number).
4. Discussion
The morphology of the presently described new species bears
the closest resemblance (in overall appearance) to P. lata (Song et
Chen, 1976). The differences, however, in the structure of append-
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W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
Figs. 10–14. Parabrachiella jarai sp. nov.; female; appendages. Fig. 10. Second maxilla. Fig. 11. Maxilliped. Fig. 12. Maxilliped (from another specimen). Fig. 13. Genital process.
Fig. 14. Spermatophore. Abbreviations: B – bulla; MG – maxillary gland. Scale bars in mm.
ages of both males and females are distinct and explicit (Song and
Chen, 1976; Roubal, 1981; Ho et al., 2007) and they cover: (1)
shape of ‘‘torso” (base of Mx2) (plain in P. jarai female vs. distinct
lateral lobes/inflations in P. lata; (2) unarmed A2 exopod in P. jarai
female (vs. denticulate area in P. lata); (3) different dental formulas
of female Mdb (P2, S1, P1, S1, B4 in P. jarai (vs. P1, S1, P1, S1, P1, S1,
B5 in P. lata); (4) single denticulate pad on medial side of Mxp corpus of female in P. jarai (vs. two denticulate pads in P. lata); (5) unarmed exopod of male A2 in P. jarai (vs. two spiniform setae and
denticulate pad in P. lata. (6) different dental formulas of male
Mdb (P2, S1, P1, S1, B5 in P. jarai vs. P4, S1, B5 in P. lata). Moreover,
P. lata was found on ‘‘Sparus latus” = Acanthopagrus latus (Houttuyn,
1782) (family Sparidae), while P. jarai parasitized Sillago sihama
(family Sillaginidae).
Parabrachiella jarai n. sp. is very host specific. Of 26 sillaginid
species examined by one of the authors (CJH) throughout the
Indo-West Pacific, ranging from East Africa to Australia, New Caledonia and Japan, it parasitized only one: Sillago sihama, and was
only found at two locations that this species was examined, both
in South-East Asia.
The number of species of the genus Parabrachiella considered
valid by the present authors is 67 (Table 1). Castro Romero and
Baeza Kuroki (1987) provided a key to females of those species
(at that time members of the genus Neobrachiella). They proposed
a tentative division into three groups, based on the number of posterior processes (including caudal rami): GROUP I (2 pairs), GROUP
II (1 pair), and GROUP III (more than 2 pairs). We concluded that
constructing an updated key for such high number of species
would be impractical and premature. It should be emphasized that
many of the existing species have been inadequately described and
illustrated. Therefore those species have to be re-described before
any reliable key is proposed. Moreover, a number of specimens,
previously re-described under well established names represent
probably new species and we are going to focus on them our future
studies.
The presently described new species has been assigned to
Group 2. This group is the largest, consisting of 35 members
(including the new species): P. albida, P. anisotremi, P. auriculata,
P. bera, P. brevicapita, P. chavesii, P. chevreuxii, P. dentici, P. dispar,
P. elegans, P. fasciata, P. gracilis, P. gulosa, P. hoi, P. hostilis, P. indica,
P. insidiosa, P. jarai sp. nov., P. johnii, P. lata, P. lutiani, P. menticirrhi,
P. merluccii, P. multifimbriata P. oralis, P. otolithi, P. paralichthyos, P.
regia, P. richiardii, P. rotunda, P. sciaenae, P. seriolae, P. sihama, P.
trichiuri, and P. yongxingensis.
The identity of a number of Parabrachiella species should be
reconsidered. Some specimens used for redescriptions, especially
those found on different hosts should be re-examined. Among
them are ‘‘Neobrachiella pillai Kabata et Tareen, 1987” (=Parabrachiella lutiani (Tripathi, 1962)), the ‘‘short form” of ‘‘Neobrachiella
rostrata” found by Rubec (1988) on Greenland halibut, Reinhardtius
hippoglossoides, and the long-armed form of P. annulata (Markewitsch, 1940) sensu Ho (1975).
P. chevreuxii (van Beneden, 1891), described from an unconfirmed and unlikely host (‘‘Squalus sp.”) is certainly not a species
described from sciaenid fishes by Brian (1906) and Kabata (1966,
1979). It has differing structure and body proportions. The sciaenid
W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
59
Figs. 15–22. Parabrachiella jarai sp. nov.; male; whole; lateral. Fig. 15. Whole; lateral. Fig. 16. First antenna. Fig. 17. Second antenna; whole. Fig. 18. Mandible. Fig. 19. First
maxilla. Fig. 20. Second maxilla. Fig. 21. Maxilliped. Fig. 22. Caudal rami. Abbreviations: 1, 2, 3, 4, 5 – armament of endopod of second antenna (according to Kabata, 1979).
Scale bars in mm.
parasite should be referred to as P. sciaenae (Brian, 1906) comb.
nov.
‘‘Neobrachiella sp.”, described by Piasecki (1993) from a California halibut, Paralichthys californicus, from southern California,
probably represents an unknown species. New specimens are
needed, however, to describe missing details of its morphology.
As mentioned earlier, some representatives of the genus Thysanote bear a close resemblance to those of the genus Parabrachiella.
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W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
Table 1
List of all valid species of Parabrachiella with their principal data (host data for original record only).
Valid name
Original name
1 P. albida (Rangnekar, 1956)
2
3
4
5
6
7
Charopinus
albidus
P. amphipacifica (Ho, 1982)
Neobrachiella
amphipacifica
P. anisotremi (Castro Romero et Brachiella
Baeza Kuroki, 1989)
anisotremi
P. annulata (Markewitsch, 1940) Brachiella
annulata
P. anserina (Wilson, 1908)
Brachiella
anserina
P. auriculata (Castro Romero et
Neobrachiella
Baeza Kuroki, 1987)
auriculata
P. bera (Yamaguti, 1939)
Brachiella bera
8 P. bispinosa (von Nordmann,
1832)
9 P. brevibrachiata (Kabata, 1992)
10 P. brevicapita (Ho et Do, 1984)
11 P. chavesii (van Beneden, 1891)
Brachiella
bispinosa
Neobrachiella
brevibrachiata
Neobrachiella
brevicapita
Brachiella
chavesii
12 P. chevreuxii (van Beneden, 1891) Brachiella
chevreuxii
13 P. chlorophthalmi (Kabata, 1992) Neobrachiella
chlorophthalmi
14 P. cirrocauda (Heegaard, 1962)
Brachiella
cirrocauda
15 P. dentici (Nunes Ruivo, 1962)
Brachiella dentici
Type host
Host family
Type locality
‘‘Pseudoscieana albida” = Daysciaena albida (Cuvier,
1830)
Psychrolutes phrictus Stein et Bond, 1978, P. sio
Nelson, 1980
Anisotremus scapularis (Tschudi, 1846)
Sciaenidae
Indian Ocean, Bombay
Psychrolutidae
Pacific N.
Haemulidae
Pacific S., Chile, Antofagasta
Ereunias grallator Jordan et Snyder, 1901
Ereuniidae
Pacific N., Japan, Misaki
‘‘Sebastodes glaucus” = Sebastes glaucus Hilgendorf,
1880
‘‘Sciaena gilberti” = Cilus gilberti (Abbott, 1899) (and
S. deliciosa)
‘‘Halichoeres poecilopterus” = Parajulis poecilepterus
(Temminck et Schlegel, 1845)
‘‘Gadus sp.” (misidentification possible) Trigla sp.
(according to Kabata 1979)
‘‘Trypterophycis intermedius” = Tripterophycis
gilchristi Boulenger, 1902
Nibea albiflora (Richardson, 1846)
Sebastidae
Pacific N., off Bering Island
Sciaenidae
Pacific S., Chile
Labridae
Pacific N., Japan Tarumi
Triglidae
Atlantic N. ???
Moridae
Pacific S., Australia, Port Kembla
NSW
Pacific N., Japan, Kojima Bay
‘‘Ceratopterus sp.” = Manta birostris (Walbaum,
1792) (misidentification possible)
‘‘Squalus sp.” (misidentification possible)
‘‘Chlorophthalmus nigripinnis” = Paraulopus
nigripinnis (Günther, 1878)
‘‘Physiculus barbatus” = Pseudophycis barbata
Günther, 1863
‘‘Dentex filosus” = Dentex gibbosus (Rafinesque,
1810)
‘‘Sciaena gilberti” = Cilus gilberti (Abbott, 1899)
Sciaenidae
Elasmobranchii:
Atlantic N., Azores
Rajiformes:
Myliobatidae
Elasmobranchii:
Atlantic N., Senegal
Squaliformes: Squalidae
Paraulopidae
Pacific S., Australia New South
Wales
Moridae
Pacific S., Australia, Victoria,
Corner Inlet
Sparidae
Atlantic S., Angola
16 P. dispar (Castro Romero et Baeza Neobrachiella
Kuroki, 1987)
dispar
17 P. elegans (Richiardi, 1880)
Brachiella elegans Lichia amia, L. gauca, Seriola lalandi
18 P. exigua (Brian, 1906)
Brachiella exigua Pagellus erythrinus (L.)
Sciaenidae
Pacific S., Chile, Antofagasta
Carangidae
Sparidae
19 P. exilis (Shiino, 1956)
Kyphosidae
Atlantic N. Mediterranean
Atlantic N., Mediterranean Sea,
off Napoli
Pacific N., Japan, Nagaturo
20
Sciaenidae
Pacific S., Chile, Antofagasta
Ophidiidae
Sciaenidae
Sciaenidae
Serranidae
Atlantic S., Namibia, Fort Rock
Point
Pacific N., California S.
Atlantic N., USA East coast
Pacific S., Australia, Heron Island
Ophidiidae
Oplegnathidae
Pacific N., California S.
Pacific N., Japan, Suruga Bay
Sciaenidae
Atlantic N., Mediterranean Sea
Tetraodontidae
Pacific N., Sea of Japan
Kyphosidae
Pacific N., Japan Seto
Sciaenidae
Gadidae
Sciaenidae
Indian Ocean, India, Puri
Atlantic N., Mediterranean Sea,
Adriatic
Atlantic N., Gulf of Mexico
Sillaginidae
Sciaenidae
Pacific N., Hong Kong
Pacific N., East China Sea
Haemulidae
Pacific S., Peru, Chorrillios
Sparidae
Lutjanidae
Pacific N., China, Hainan Dao,
Sanya
Indian Ocean, India
Sciaenidae
Pacific S., Peru, Chorrillios
Merlucciidae
Atlantic N., UK, Plymouth
Scorpaenidae
Pacific S., Australia, New South
Wales
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
Epibrachilla exilis ‘‘Kyphosus lembus” = Kyphosus vaigiensis (Quoy et
Gaimard, 1825)
P. fasciata (Castro Romero et
Neobrachiella
Sciaena fasciata (Tschudi, 1846)
Baeza Kuroki, 1987)
fasciata
P. genypteri (Capart, 1959)
Brachiella
Genypterus capensis (Smith, 1847)
genypteri
P. gracilis (Wilson, 1908)
Brachiella gracilis ‘‘Cynoscion nobilis” Atractoscion nobilis (Ayres, 1860)
P. gulosa (Wilson, 1915)
Brachiella gulosa Sciaenops ocellatus (Linnaeus, 1766)
Epinephelus merra Bloch, 1793
P. gymnobrachiata (Kabata, 1968) Thysanote
gymnobrachiata
P. hoi (Piasecki, 1993)
Neobrachiella hoi Chilara taylori (Girard, 1858)
P. hoplognathi (Yamaguti, 1939) Epibrachiella
‘‘Hoplognathus sp.” = Oplegnathus sp.
hoplognathi
P. hostilis (Heller, 1868)
Anchorella
Umbrina cirrosa (L.)
hostilis
P. hugu (Yamaguti, 1939)
Clavellopsis hugu ‘‘Spheroides rubripes” = Takifugu rubripes (Temminck
et Schlegel, 1850)
P. incurva Shiino, 1956
‘‘Kyphosus lembus” = Kyphosus vaigiensis (Quoy et
Gaimard, 1825)
P. indica (Tripathi, 1962)
Brachiella indica ‘‘Sciaena glauca” = Johnius glaucus (Day, 1876)
Brachiella
‘‘Gadus sp.”
P. insidiosa (Heller, 1868)
insidiosa
P. intermedia (Bere, 1936)
Brachiella
Sciaenops ocellatus (Linnaeus, 1766)
intermedia
P. jarai sp. nov.
Sillago sihama (Forsskål, 1775)
P. johnii (Yamaguti, 1939)
Clavellopsis johni ‘‘Johnius goma” = Protonibea diacanthus (Lacepède,
1802)
P. kabatai (Luque et Farfan, 1991) Neobrachiella
Isacia conceptionis (Cuvier, 1830)
kabatai
P. lata (Song et Chen, 1976)
Brachiella lata
‘‘Sparus latus” = Acanthopagrus latus (Houttuyn,
1782)
P. lutiani (Pillai, 1968)
Brachiella indica ‘‘Lutianus sp.” = Lutjanus sp.
Pillai, 1968
P. menticirrhi (Luque et Farfan,
Neobrachiella
Menticirrhus ophicephalus (Jenyns, 1840)
1990)
menticirrhi
P. merluccii (Bassett-Smith, 1896) Brachiella
‘‘Gadus merluccius” = Merluccius merluccius (L.)
merluccii
P. microdigitata (Kabata, 1992)
Neobrachiella
‘‘Helicolenus papillosus” = Scorpaena papillosa
microdigitata
(Schneider et Forster, 1801)
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W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
Table 1 (continued)
Valid name
41 P. mirifica (Kabata, 1992)
42 P. mitrata (Wilson, 1915)
43 P. mugilis (Kabata, Raibaut et Ben
Hassine, 1971)
44 P. multifimbriata (Basset-Smith,
1898)
45 P. nitida (Wilson, 1915)
46 P. oralis (Castro Romero et Baeza
Kuroki, 1987)
47 P. otolithi (Pillai, 1962)
48 P. paralichthyos (Castro Romero et
Baeza Kuroki, 1986)
49 P. pinguis (Wilson, 1915)
50 P. regia (Lewis, 1967)
51 P. richiardi (Ben Hassine et
Raibaut, 1978)
52 P. robusta (Wilson, 1912)
53 P. rostrata (Krøyer, 1837)
54 P. rotunda (Pearse, 1952)
55 P. sciaenae (Brian, 1906)
56 P. septicauda (Shiino, 1956)
57 P. seriolae (Yamaguti et Yamasu,
1960)
58 P. sihama Song et Chen, 1976
Original name
59 P. spinicephala Ringuelet, 1945
60 P. stellifera (Heegaard, 1962)
61 P. sublobulata (Barnard, 1955)
62 P. superba (Leigh-Sharpe, 1934)
63 P. supplicans (Barnard, 1955)
64 P. tetrici (Kabata, 1968)
65 P. trichiuri (Yamaguti, 1939)
66 P. triglae (Claus, 1860)
67 P. yongxingensis (Song et Chen,
1976)
Type host
Neobrachiella
‘‘Hoplichtys haswelli” = Hoplichthys haswelli
mirifica
McCulloch, 1907
Brachiella mitrata ‘‘Lopholatilus chameleonticeps” = Lopholatilus
chamaeleonticeps Goode et Bean, 1879
Eubrachiella
‘‘Mugil auratus” = Liza aurata (Risso, 1810)
mugilis
Brachiella
Serranus sp.
multifimbriata
Brachiella nitida Albatrossia pectoralis (Gilbert, 1892)
Sciaena deliciosa (Tschudi, 1846)
Neobrachiella
oralis
Brachiella otolithi ‘‘Otolithus argenteus” = Otolithes ruber (Bloch et
Schneider, 1801)
Paralichthys adspersus (Steindachner, 1867)
Neobrachiella
paralichthyos
Brachiella pinguis ‘‘Antimora viola” = Antimora microlepis Bean, 1890
Brachiella regia
‘‘Lampris regius” = Lampris guttatus (Brünnich, 1788)
Brachiella
Umbrina cirrosa (L.)
richiardi
Clavella robusta ‘‘Sebastodes auriculatus” = Sebastes auriculatus
Girard, 1854
Brachiella
Hippoglossus hippoglossus (L.)
rostrata
Brachiella
Prionotus tribulus crassiceps Ginsburg, 1950
rotunda
Clavella sciaenae Sciaena antarctica??? (in 1923 – ‘‘Sciaena aquilla”)
Epibrachiella
Doederleinia berycoides (Hilgendorf, 1879)
septicauda
Brachiella seriolae Seriola quinqueradiata Temminck et Schlegel, 1845
Brachiella
stellifera
Epibrachiella
sublobulata
Brachiella
superba
Brachiella
supplicans
Brachiella tetrici
Host family
Type locality
Hoplichthyidae
Pacific S., Australia, New South
Wales
Atlantic N., USA, Massachusetts
Malacanthidae
Serranidae
Atlantic N., Mediterranean Sea,
Lake of Tunis (lagoon)
Indian Ocean, Oman, Muscat
Macrouridae
Sciaenidae
Pacific N.
Pacific S., Chile
Sciaenidae
Indian Ocean, India, Kerala
Paralichthyidae
Pacific S., Chile
Moridae
Lampridae
Sciaenidae
Sebastidae
Atlantic N., Newfoundland S.
Pacific N., Hawaii
Atlantic N., Mediterranean Sea,
Tunis
Pacific N. Canada, Nanaimo BC
Pleuronectidae
Atlantic N., Kattegat
Triglidae
Atlantic N., Gulf of Mexico, Texas
Sciaenidae
Acropomatidae
Atlantic N. ???
Pacific N., Japan, Mie Pref.
Carangidae
Pacific N., Japan
Sillago sihama (Forsskål, 1775)
Sillaginidae
‘‘Pinguipes fasciatus” = Pinguipes brasilianus Cuvier,
1829
Sphaeroides sp.
Pinguipedidae
Pacific N., China, Hainan Dao,
Sanya
Atlantic S., Mar del Plata
Tetraodontidae
Indian Ocean, Western Australia
Congiopodus torvus (Gronow, 1772)
Congiopodidae
Atlantic S., Africa, S., Table Bay
Malacocephalus laevis (Lowe, 1843)
Macrouridae
Genypterus capensis (Smith, 1847)
Ophidiidae
‘‘Pseudolabrus tetricus” = Notolabrus tetricus
(Richardson, 1840)
Trichiurus japonicus L.
Labridae
Pacific S. (Indian Ocean); Flores
Sea
Atlantic S., South Africa,
Table Bay
Indian Ocean, Australia, Port
Willunga
Pacific N., Japan Toyama Bay
Clavellopsis
trichiuri
Brachiella triglae Trigla sp.
Brachiella
Lethrinus sp.
yongxingensis
The males in both genera look alike. We believe that the existing
diagnosis of the genus Thysanote (cf. Kabata and Tareen, 1981) is
impractical. According to those authors the principal reason for
retaining Thysanote is the pattern of mandibular denticulation,
allegedly more primitive in Thysanote (2–3 secondary teeth in Neobrachiella versus 1–2 in Thysanote). The majority of nominal species
assigned to Thysanote and Parabrachiella have been recorded only
once or twice. Many descriptions contain inadequate (or non-reliable) data on the mandible. Consequently, if the mandibular criterion is preserved, the status of many species will remain
unresolved. We propose, herewith, to disregard the mandibular
criterion and to distinguish between the two genera based on the
presence of posterior- and maxillary processes in Thysanote and
only posterior processes in Parabrachiella. Consequently, P. multifimbriata (Bassett-Smith, 1898) comb. nov. and P. gymnobrachiata
(Kabata, 1968) comb. nov. are herewith transferred from Thysanote. On the other hand, we noticed that ‘‘Branchiellina papillosa
Pearse, 1952” is very similar to Thysanote chalermwati Piasecki,
Ohtsuka et Yoshizaki, 2008 in the size, shape, and distribution of
the processes. Therefore we propose to transfer the earlier men-
Mugilidae
Trichiuridae
Triglidae
Lethrinidae
Atlantic N. ???
Pacific N., South China Sea
Yongxing Dao, Xisha Quandao
tioned species to the genus Thysanote as Thysanote papillosa (Pearse, 1952) comb. nov. Consequently, also Neobrachiella impudica
(von Nordmann, 1832) should become Thysanote impudica (von
Nordmann, 1832) comb. nov.
Many researchers referring to the posterior processes of Parbrachiella tend to treat one of the pairs as caudal rami (‘‘uropods”).
Such reasoning may not always be reliable. According to Ho et al.
(2007), who first studied a complete life cycle of a Parabrachiella,
the posterior processes, not being caudal rami, are not present at
any of the chalimus stages and they are apparently formed after
the last moult.
Copepods of the genus Parabrachiella seem to be highly host
specific, infecting single fish species (or only a few closely related
hosts). They have been recorded from various teleost fishes, representing an extensive set of the families. The most frequently infected family is the Sciaenidae, hosting as many as 16 species,
followed by Sparidae with 3 species. Three Parabrachiella species
have been found in representatives of Moridae, Ophidiidae, and
Triglidae. Two species have been recorded in Carangidae, Haemulidae, Kyphosidae, Macrouridae, Labridae, Sebastidae, Serranidae,
62
W. Piasecki et al. / Experimental Parasitology 125 (2010) 55–62
Sillaginidae, Tetraodontidae, whereas a number of families (such as
Acropomatidae, Congiopodidae, Ereuniidae, Gadidae, Hoplichthyidae, Lampridae, Lethrinidae, Lutjanidae, Malacanthidae, Merlucciidae, Mugilidae, Oplegnathidae, Paralichthyidae, Paraulopidae,
Pinguipedidae, Pleuronectidae, Psychrolutidae, Scorpaenidae, and
Trichiuridae) have been associated with a single Parabrachiella species. Parabrachiella seem to be parasites of exclusively teleost
fishes, and the finding of ‘‘Brachiella chavesii” by van Beneden
(1891) on a ‘‘Ceratopterus sp.” = Manta birostris (Walbaum)
(Elasmobranchii: Rajiformes: Myliobatidae) seems to be an error.
Similarly, P. chevreuxii (van Beneden, 1891) was allegedly found
on a ‘‘Squalus sp.”, which is also doubtful.
The narrow host specificity of Parabrachiella species seems to be
their important characteristic feature. Therefore, cases of finding
(allegedly) the same species of copepod from fishes representing
different families must be treated with caution (Kempter et al.,
2006). For example: Markewitsch (1940) described P. annulata
from the skin of a fish representing family Ereunidae, while Kabata
and Gusev (1966), Noble (1973), and Ho (1975) recorded it from
fishes representing family Macrouridae. Also P. exilis (Shiino,
1956) was originally described from Kyphosus vaigiensis (Kyphosidae) but Castro Romero and Baeza Kuroki (1986) re-described it
from Mugil cephalus (Mugilidae).
The majority of Parabrachiella species were found in the Pacific
Ocean (37 species: 22 in North Pacific and 15 in South Pacific).
Some 23 species come from the Atlantic (18 form the North and
5 from the South), while the Indian Ocean is the type locality for
only 7 species.
Acknowledgments
This material was collected as part of PhD studies by CJH. We
are grateful to Geoff Boxshall for providing some literature that
was difficult to access.
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