Download a review on the biology and parasites of the big

Survey
yes no Was this document useful for you?
   Thank you for your participation!

* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project

Document related concepts
no text concepts found
Transcript
Journal of Multidisciplinary Scientific Research, 2014,2(5):03-08
ISSN: 2307-6976
Available Online: http://jmsr.rstpublishers.com/
A REVIEW ON THE BIOLOGY AND PARASITES OF THE BIG-EYE SCAD, SELAR
CRUMENOPHTHALMUS (BLOCH, 1793).
Francis Albert T. Argente1*,Charina I. Narido2, Herminie P. Palla3 and Milagros A. Celedonio4
1)Department of Fisheries Science, Pangasinan State University – Binmaley Campus, Binmaley, Pangasinan, Philippines.
2)Mathematics and Science Department, Holy Name University, Tagbilaran City, Bohol, Philippines.
3)College of Fisheries and Aquatic Science, Western Philippines University – Puerto Princesa Campus, Palawan,Philippines.
4)Natural Science Department, Dr. Emilio B. Espinosa Sr. Memorial State College of Agriculture and Technology, Masbate, Philippines.
E-mail:[email protected]
Received:02 ,June,2014
Accepted:07, Aug,2014
__________
Abstract
This paper presents a review on the biology and the parasites of Selar crumenophthalmus, a finfish utilized for human food consumption. The
presence of parasites on this species poses a threat to its human consumers. The biology of S. crumenophthalmus must be understood to
explain the prevalence of its parasites. Available literatures were summarized to provide useful information to human consumers.
Keywords: Parasitism, Selar crumenophthalmus, monogenean, Norileca indica, Philippines.
INTRODUCTION
The big-eye scad, Selar crumenophthalmus is a small pelagic
species that is commercially important in the Philippines. This fish is
primarily utilized for human consumption and together with other
small pelagics, it is considered as a cheap source of protein for low
income families in the country (Trinidad et al., 1993). Previous report
showed that 5% of the total value of top species caught from the
marine waters is composed of big-eye scads (BAS, 2011). Gears
that are used to catch this fish include handlines, ringnets, purse
seines and trawls (Trinidad et al., 1993).
Fish are important hosts of parasites in aquatic ecosystems,
harboring wide variety of adult and immature forms and acting either
as sole host or as one in a series of hosts (Barber and Poulin, 2002).
Some parasites are responsible for outbreaks of diseases to fish
populations (Barber and Poulin, 2002) and few (anisakid nematodes)
are capable of infecting humans (Petersen et al., 1993; Adams et al.
1997). Economically, parasites can devaluate fish market value as it
can cause spoilage and it is transmittable to humans (Adams et al.,
1997; Barber and Poulin, 2002). On the other hand, prevalence of
parasites in one habitat is indicative of the health of an ecosystem
(Adams et al., 1997; Hudson et al., 2006) and determinant of status
of fish stocks (Barber & Poulin, 2002). Parasites have also been
used as biological tag to investigate population structure of
commercially important species of fish (Mckenzie, 2002; McKenzie et
al., 2008), as indicator of historical dispersal through migration
(Rohde, 2002), as pollution indicator (Aloo et al., 2004) and as
important driver of biodiversity (Hudson et al., 2006).
S. crumenopthalmus is not exempted from parasite infestation as
reported in several studies (Rokicki, 1982; Work et al., 2008;
Nagasawa and Petchsupa, 2009). The presence of isopods in the
branchial cavities of this carangid (Rokicki, 1982; Nagasawa and
Petchsupa, 2009) indicated its vulnerability to parasitic attack. Hence,
it is a perennial candidate for the study of parasite infestation or
infection.
Parasitism plays a significant role in fish biology and ecology
(Hudson et al., 2006; Luque and Poulin, 2008). Parasites infection on
fish is a usual ecological event, because parasites are natural
component of the aquatic environment and fish forms an important
part of their life cycle (Barber and Poulin, 2002). Parasites influence
individual host survival and reproduction, they can alter fish
behaviour and migration patterns, and they can even regulate fish
populations and affect fish community structure (Garnick and
Margolis, 1990; Barber and Poulin, 2002).
The study of parasitic organisms becomes important because they
can be an indicator species to detect the water quality and altered
environmental condition such as eutrophication (Palm and
Dobberstein, 1999) and to determine the ecology of the host. Fish
parasites can also be used as biological indicators for fish stock
separation (McKenzie, 1983; Santos et al., 2009) and fish feeding
ecology (Campbell et al., 1980; Palm, 1999).
This paper reviews the biology and the parasites of S.
crumenophthalmus. This is particularly important because this
species is consumed as food. Summarized information based on
available literature are necessary to understand the presence of
harmful parasites and its natural infestation patterns in this
economically-valuable carangid. Thus, potential threat to human
consumers will be known and avoided.
Biology of big-eye scad, Selar crumenophthalmus
The big-eye scad belongs to a large group of fishes known as
family Carangidae. The presence of enlarged, thickened scutes in
the straight of the lateral line and a detached two anal fin spines from
the rest of the ventral fin delineate this family from other group of
fishes (Smith-Vaniz, 1999). The genus Selar has only two species
listed that includes the big-eye scad and the ox-eye scad, S. boops.
The former is characterized by an elongated moderately compressed
body, shorter number of scutes and a deep indentation on the lower
margin of the operculum (Rau and Rau, 1980; Schroeder, 1984).
The distribution of S. crumenopthalmus is circumtropical
(Honebrink, 2000) extending throughout the Indo-West Pacific where
occurrence is known from Japan to Australia and eastward to Hawaii.
This fish is found in small to large schools at a water depth not
exceeding 200 m (Smith-Vaniz, 1999) and is largely migratory where
movement ranges from shallow murky inshore reefs and sandflats to
clear pelagic water (Schroeder, 1984). However, sub-adult S.
crumenopthalmus from Hawaii formed large schools in shallow water
Francis Albert T. Argente et al.
from about July through December and ended up joining adult
schools, which apparently do not make any large scale movements,
even around the same island (Honebrink, 2000). Feeding biology of
this species showed ontogenic shifts where juveniles feed primarily
on euphausiids and decapods while adults feed on small fishes,
crustaceans and gastropods (Roux and Conand, 2000; Schroeder,
1984).
S. crumenophthalmus is gonochoristic; that they have separate
sexes and for the most part there is no apparent difference between
the sexes. However, Clarke and Privitera (1995) distinguished a dark
black color in the soft portion of the anal fin of male S.
crumenophthalmus during spawning season, while females of the
same species possessed a white shade. As to the species fecundity
and spawning period, Roos et al. (2007) reported that there was no
difference in the sex ratio between male and female S.
crumenopthalmus. Sex ratio remained constant every month of
sample even in the larger size classes. The big-eye scad started to
mature at the month of April and the number of mature fish increased
as months passed until spawning commenced in the month of
November. It is notable that after reproduction, S.
crumenophthalmus experienced massive mortality and only few
individuals survived. On the other hand, an earlier study (Clarke and
Privitera, 1995) in Hawaii revealed that spawning season of S.
crumenophthalmus was from April through September or October
which is not far from the study of Roos et al., (2007) in Reunion
Island, Southwest Indian Ocean. In addition, Clarke and Privitera
(1995) showed that big-eye scads appeared to spawn between dawn
and dusk. The frequency of postovulatory follicles of big-eye scad
indicated that females spawned every three days and batch cost
averaged about 3% of body weight while in terms of fecundity
estimates of big-eye scad, mean batch fecundities were 92, 000
eggs.
Parasites of big-eye scad, Selar crumenophthalmus
The parasites of S. crumenophthalmus can be categorized into
nine groups, namely, monogeneans (Jianying et al., 2003; Arthur
and Te, 2006; Kohn et al., 2006), digeneans (Vassiliades, 1982;
Dyer et al., 1986; Arthur and Te, 2006; Liu et al., 2010; Madhavi,
2011), nematodes (Deardorff et al., 1982; Arthur and Mayo, 1997;
Aloo et al., 2004; Arthur and Te, 2006), cestodes (Arthur and Te,
2006; Palm et al., 2009), acanthocephalans (Arthur and Te, 2006;
Verweyen, et al., 2011); copepods (Boxshall and Huys, 2007);
isopods (Rokicki, 1982; Nagasawa and Petchsupa, 2009),
myxozoans (Work et al., 2008) and fungi (Petersen et al., 1993;
Arthur and Mayo, 1997). Among these groups of parasites,
digeneans were the most commonly reported for S.
crumenophthalmus (Vassiliades, 1982; Dyer et al., 1986; Arthur and
Te, 2006; Liu et al., 2010; Madhavi, 2011).
Monogenean parasites reported for S. crumenopthalmus were
commonly infesting the gills of the fish (Arthur and Te, 2006; Kohn et
al., 2006). The sites of infestation of digenean parasites of S.
crumenophthalmus were the stomach and the intestine (Dyer et al.,
1986; Arthur and Te, 2006). Nematodes were reported to infect the
body cavity, mesenteries, musculature, viscera, gonads, skin,
stomach, intestine, pyloric caeca, kidney and liver of the big-eye scad
(Arthur and Mayo, 1997; Aloo et al., 2004; Arthur and Te, 2006).
Parasitic cestodes of the fish are found in its body cavity, gonads,
intestine, stomach, gall bladder, kidney and liver (Arthur and Te,
2006; Palm et al., 2009). Arthur and Te (2006) reported the
infestation of the acanthocephalan Gorgorhynchus medius in the
body cavity and inner organs of the S. crumenophthalmus. The
isopod, Norelica indica was accounted as a common parasite of S.
crumenophthalmus infesting its branchial cavities (Rokicki, 1982;
Nagasawa and Petchsupa, 2009). Microsporic fungi were reported to
infect the external surface of the intestine and musculature (Petersen
et al., 1993; Arthur and Mayo, 1997).
Work et al. (2008) reported a new species of myxozoan that
infected the bulbous arteriosus of S. crumenophthalmus from Hawaii.
The newly discovered parasite belongs to the genus Henneguya,
under the myxosporean group which has a complex life cycle. Boyce
et al. (1985) stated that Henneguya changes its structure in the
course of its life probably passing through several developmental
stages and the best known stage is the spore. Henneguya infection
are said to be common on freshwater fishes and parasitic stages of
this parasite mostly invaded the gills, skin, kidneys, musculoskeletal
system, or gastrointestinal tract (Eiras, 2002) though Henneguya sp.
infecting the bulbus arteriosus are rare (Yokoyama et al., 2005).
According to Work et al. (2008), the host response to the parasite has
no profound effect and gross lesions were not evident hence, it has
minimal measurable detrimental consequence on the host. However,
based on the microscopic observation from 17% of infected fish, the
host response was mild and characterized by accumulations of
eosinophilic fibrillar material around spores and occasional
mononuclear infiltrates in the adventitia.
In the study conducted by Aloo et al. (2004), they discovered
nematodes, Camallanus sp. in the liver, below the ovary and testes
as well as under the skin of S. crumenophthalmus and some
mackerels. The prevalence of the parasite was 59.1% for S.
crumenopthlamus indicating that Camallanus sp. infestation was
quite high in this species considering that more than half of the
samples were infested. Although, the author stated that the parasites
were not affecting the health of the host, it is possible though, that it
can affect the physiological functioning of the organs such as the
liver and gonads. The extraction of nematodes from the gonads and
liver suggested that these roundworms preferred to stay from these
organs and perhaps gain nutrients for their survival. According to
Reichenbach-Klinke and Landolt (1973), parasites infecting the
intestines of the host hamper the digestive activity and in some way
prevent vitamin and sugar metabolism as well as growth. Moreover,
parasites that embedded the liver upset glycogen metabolism while
parasites in the gonads and coelomic cavity could possibly lead to
absolute castration and fecundity reduction.
Table 1 shows the summarized list of parasites, infested body parts
and its geographic distribution as reported by several authors..
Parasite
Infested
Body Parts
Distribution
Source
Monogenea
Choricotyle
caulolatili
gills
Jalisco,
Mexico
Kohn et
2006
al.,
Gastrocotyle
trachuri
gills
Gulf
of
Tonkin; South
China Sea
Arthur
and
Te, 2006
Journal of Multidisciplinary Scientific Research , 2014,2(5):03-08
Hexteracine
heterocerca
gills;
pharyngeal
cavity
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Ectenurus
trachuri
Jaliscia
caballeroi
Metacamopia
chormemi
gills
Jalisco,
Kohn et al.,
Mexico
2006
South China Jianying et
Sea
al., 2003
Ectenurus
trachuri
Ectenurus
virgulus
Lecithocladium
excisum
Pseudaxinoides
vietnamensis
Pseudaxinoides
vietnamensis
Pseudomazocra
es monsivaeisae
Vallisia
chorinemi
Digenea
Aponurus
carangis
Aponurus
laguncula
Aponurus
laguncula
Bucephalus
varicus
South
Sea
gills
gills
gills
stomach;
intestine
stomach;
intestine
stomach;
intestine
Bucephalus
varicus
Didymozoidae
Dinurus selari
Dinurus selari
Ectenurus selari
stomach;
intestine
et
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Oaxaca,
Mexico
Kohn et al.,
2006
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
South China
Sea
Gulf
of
Tonkin; South
China Sea
South
Sea
body
cavity;
eye
sockets;
gills, intestine,
kidney; liver
stomach;
intestine
China Jianying
al., 2003
Liu et al.,
2010
Arthur
and
Te, 2006
China Liu et
2010
al.,
Gulf
of
Arthur
and
Tonkin; South
Te, 2006
China Sea
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
South China
Sea
Gulf
of
Tonkin; South
China Sea
Liu et al.,
2010
Arthur
and
Te, 2006
stomach
stomach
Lecithocladium
excisum
Lecithocladium
harpodontis
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
South China Liu et al.,
Sea
2010
Senegal
Vassiliades,
1982
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
South
Sea
stomach
Lecithocladium
harpodontis
stomach;
intestine
Parasite
Infested
Body Parts
al.,
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
South
Sea
Lecithocladium
seriolellae
China Liu et
2010
China Liu et
2010
al.,
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Distribution
Source
Lecithocladium
seriolellae
South
Sea
China Liu et
2010
al.,
Lecithochirium
ghanense
Senegal
Vassiliades,
1982
Dyer et
1986
Lecithochirium
microcercus
intestine
Puerto Rico
Lecithochirium
montecillii
stomach
South
Sea
China Arthur
and
Te, 2006
Senegal
Vassiliades,
1982
China Liu et al.,
2010
Monascus
typicus
Neoprosorhynch
us xishaensis
South
Sea
Visakhapatna
m Coast; Bay
of Bengal
Paramonorcheid
es selaris
Proctotrema sp.
stomach;
intestine
al.,
Madhavi,
2011
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Francis Albert T. Argente et al.
Proctotrema sp.
Tergestia
laticollis
intestine
Tergestia
laticollis
South China
Sea
Gulf
of
Tonkin; South
China Sea
South
Sea
Liu et al.,
2010
Arthur
and
Te, 2006
China Liu et
2010
Anisakis sp.
Anisakis sp.
Camallanus sp.
Camallanus
carangis
Contracaecum
sp.
Porrocaecum sp.
Hawaiian
Islands
below
ovary/testis;
within
liver;
Kenyan coast
under the skin
intestine;
pyloric caeca
body
cavity;
kidney;
liver;
outer wall of
stomach;
intestine;
stomach
Parasite
Scolex
pleuronectis
New
Caledonia
Boxshall
and
Huys,
2007
branchial
cavities
branchial
cavities
Southeast
Asia
Pattani Bay,
Thailand
Rokicki,
1982
Nagasawa
and
Petchsupa,
2009
Henneguya
akule
bulbus
arteriosus
Oahu, Hawaii
Work et al.,
2008
Fungi
Microspora
flesh
Visayan Sea, Petersen
Philippines
al., 1993
Deardorff et
al., 1982
Isopoda
Norileca indica
Norileca indica
Aloo et
2004
al.,
Gulf
of
Thailand; Gulf
Arthur
and
of
Tonkin;
Te, 2006
South China
Sea
Distribution
Verweyen,
et al., 2011
Naobranchia
spinosa
Deardorff et
al., 1982
body
cavity;
gonads;
Gulf
of Arthur
intestine,
and
Tonkin;
South
Te, 2006
kidney;
liver;
China Sea
stomach
Infested
Body Parts
Arthur
and
Te, 2006
Oahu, Hawaii
Arthur
and
Mayo, 1997
Cestoda
Nybelinia sp.
Palm et al.,
2009
Copepoda
Gulf
of Arthur
and
Tonkin; South Te, 2006
China Sea
Hawaiian
Islands
Hawaii
body
cavity; Gulf of Tonkin
inner organs
Rhadinorhynchu
s lintoni
Gulf
of
Thailand; Gulf
body
cavity; of
Tonkin; Arthur
and
intestine
South China Te, 2006
Sea
Terranova sp.
Acanthocephal
a
Gorgorhynchus
medius
body cavity
al.,
Nematoda
body
cavity;
mesenteries;
Luzon;
musculature;
Visayan
viscera
Islands
Tentacularia
coryphaenae
Source
body
cavity;
gall bladder;
Gulf
of
intestine;
Arthur
and
Tonkin; South
Te, 2006
stomach
China Sea
Myxosporea
(Cnidaria)
Microsporidia
external
surface of the
Panay;
intestine;
Visayan
musculature
Islands
et
Arthur
and
Mayo, 1997
REMARKS
Records of big-eye scad parasites are relatively fewer compared
with other marine species especially those species that possess a
benthic lifestyle. The pelagic nature of S. crumenopthalmus has
likely attributed to this condition because they are not typically
associated with benthic invertebrates that usually act as intermediate
hosts for known parasites such as digeneans, nematodes,
acanthocephalans and, to a lesser extent, cestodes. However, the
migratory behavior of big-eye scads is a major factor that enables
this marine fish to harbor free living parasites in the water gradient.
In addition, feeding preferences of big-eye scads included various
shrimps and crab larvae (Roux and Conand, 2000) as well as
gastropods (Schroeder, 1984) which further increases the possibility
of acquiring parasites as crustaceans are the most common
intermediate hosts in parasite life cycles while digeneans are known
to require mollusks as obligate first intermediate host (Marcogliese,
2002). Conversely, in the event of parasite invasion, the schooling
behavior of bigeye scads allows dilution of potential parasites and
therefore reduces the risk of being targeted for an attack by any
Journal of Multidisciplinary Scientific Research , 2014,2(4):23-28
given parasite (Poulin and FitzGerald, 1989) or in the same way;
schooling effect could be the same reason for the rapid transmission
of parasite infection of a fish stock. In the end, it is always the
capacity of fishes or any other animals to avoid or defend
themselves from the discomfort and possible diseases brought by
parasites. On the other hand, reports of parasite infection of S.
crumenopthalmus have not seriously damaged the organism or
certain stock hence, its fishing industry is not seriously affected
References
1. Adams A.M., K.D. Murrell and J.H. Cross. 1997. Parasites of fish
and risks to public health. Office International des Epizooties,
Scientific and Technical Review 16(2):652-660.
2. Aloo, P.A., R.O. Anam and J.N. Mwang. 2004. Metazoan
parasites of some commercially important fish along the Kenyan
Coast. Western Indian Ocean Journal of Marine Science 3(1): 71-78.
3. Arthur, J.R. and S.L. Mayo. 1997. Checklist of the parasites of
fishes of the Philippines. FAO Fisheries Technical Paper No. 369.
102 p.
4. Arthur, J.R. and B.Q. Te. 2006. Checklist of the parasites of fishes
of Viet Nam. FAO Fisheries Technical Paper No. 369/2. 133 p.
5. Barber, I. and R. Poulin. 2002. Interactions between fish, parasites
and disease. In: Hart P.J.B. and J.D. Reynolds (eds.). Handbook of
fish biology and fisheries. Oxford: Blackwell Publishing. pp. 359–
389.
13. Honebrink, R. 2001. A review of the biology of the family
Carangidae, with emphasis on species found in Hawaiian waters.
DAR Technical Report. 43 p.
14. Hudson P.J., A.P. Dobson and K.D. Lafferty. 2006. Is a healthy
ecosystem one that is rich in parasites? Trends in Ecology and
Evolution 21(7): 381-385.
15. Jianying, Z., Y. Tingbao, L. Lin and D. Xuejuan. 2003. A list of
monogeneans from Chinese marine fishes. Systematic Parasitology
54: 111-130.
16. Kohn, A., S.C. Cohen. G.S. Maldonado. 2006. Checklist of
monogenea parasites of freshwater and marine fishes, amphibians
and reptiles from Mexico, Central America and Caribbean. Zootaxa
1289. 114 p.
17. Luque, J.L. and R. Poulin. 2008. Linking ecology with parasite
diversity in Neotropical fishes. Journal of Fish Biology 72: 189–204.
18. Liu, S., W. Peng, P. Gao, M. Fu, H. Wu, M. Lu, J. Gao and J.
Xiao. 2010. Digenean parasites of Chinese marine fishes: a list of
species, hosts and geographical distribution. Systematic
Parasitology 75: 1-52.
19. Marcogliese, D.J. 2002. Food webs and the transmission of
parasites to marine fish. Parasitology 124: 83-99.
20. McKenzie, K. 1983. Parasites as biological tags in fish population
studies. Advances in Applied Biology 7: 251-331.
6. Bureau of Agricultural Statistics (BAS). 2011. Fisheries Statistics
of the Philippines, 2008-2010. 421p.
Boxshall, G.A. and R. Huys. 2007. Copepoda of New Caledonia. In.
Payri, C.E. and B. Richer de Forges (eds.), Compedium of marine
species from New Caledonia. Institut de Recherche pour le
Developpement. 435 p.
21. McKenzie K. 2002. Parasites as biological tags in population
studies of marine organisms: an update. Parasitology 124: 153–163.
7. Campbell, R.C., R.L. Haedrich and T.A. Munroe. 1980. Parasitism
and ecological relationships among deep-sea benthic fishes. Marine
Biology 57: 301-313.
23. Madhavi, R. 2011. Checklist of digenean trematodes reported
from Indian marine fishes. Systematic Parasitology 78: 163-232.
8. Clarke, T.A. and L.A. Privitera. 1995. Reproductive biology of two
Hawaiian pelagic carangid fishes, the bigeye scad, Selar
crumenophthalmus, and the round scad, Decapterus macarellus.
Bulletin of Marine Science 56(10): 33-47.
9. Deardorff, T.L., M.M. Kliks, M.E. Rosenfeld, R.A. Rychlinski and
R.S. Desowitz. 1982. Larval ascaridoid nematodes from fishes near
the Hawaiian Islands, with comments on pathogenicity experiments.
Pacific Science 36(2): 187-201.
10. Dyer, W.G., E.H. Williams, Jr, and L.B. Williams. 1986. Some
trematodes of marine fishes of Southwestern and Northwestern
Puerto Rico. Transactions of the Illinois Academy of Science 79(1-2):
141-143.
11. Eiras, J. 2002. Synopsis of the species of the genus Henneguya
Thelohan, 1892 (Myxozoa: Myxosporea: Myxobolidae). Systematic
Parasitology 52: 43–54.
12. Garnick, E. and L. Margolis. 1990. Influence of four species of
helminth parasites on orientation of seaward migrating sockeye
salmon (Onchorhynchus nerka) smolts. Canadian Journal of
Fisheries and Aquatic Science 47: 2380–2389.
22. Mckenzie, K., N. Campbell, S. Matticuccib, P. Ramos P,
A.L.Pintod and Abaunza P. 2008. Parasites as biological tags for
stock identification of Atlantic horse mackerel Trachurus trachurus L.
Fisheries Research 89(2): 136-145.
24. Nagasawa, K. and N. Petchsupa. 2009. Norileca indica (Isopoda,
Cymothoidae) parasitic on big eye scad Selar crumenophthalmus in
Thailand. Biogeography 11: 131-133.
25. Palm, H.W. 1999. Ecology of Pseudoterranova decipiens
(Krabbe, 1878) (Nematoda: Anisakidae) from Antarctic waters.
Parasitology Research 85: 638-646.
26. Palm, H.W. and R.C. Dobberstein. 1999. Occurrence of
trichidinid ciliates (Peritrichia: Urceolariidae) in the Kiel Fjord, Baltic
Sea, and its possible use as a biological indicator. Parasitology
Research 85: 726-732.
27. Palm, H.W., A. Waeschenbach, P.D. Olson and D.T.J. Littlewood.
2009. Molecular phylogeny and evolution of the Trypanorhyncha
Diesing, 1863 (Platyhelminthes: Cestoda). Molecular Phylogenetics
and Evolution 52: 351-367.
28. Petersen F., H. Palm, H. Möller and M.A. Cuzi. 1993. Flesh
parasites of fish from central Philippine waters. Diseases of Aquatic
Organisms 15: 81-86.
29. Poulin, R., and FitzGerald, G.J. 1989. Shoaling as
an anti-ectoparasite mechanism in juvenile sticklebacks
(Gasterosteus spp.). Behavioural Ecology and Sociobiology 24: 251255.
Francis Albert T. Argente et al.
30. Rau , N. and Rau, A., 1980. Fishes of Central Philippines. USCGTZ Publication. 627 p.
Sulu Sea. Bureau of Fisheries and Aquatic Resources and Ministry
of Natural Resources. National Media Production Center, Manila.
266 p.
31. Reichenbach-Klinke, H.H. and M.L. Landolt. 1973. Fish
pathology: a guide to the recognition and treatment of diseases and
injuries of fishes, with emphasis on environmental and pollution
problems. T.F.H. Publications. 512 p.
38. Smith-Vaniz, W. F. 1999. Bony fishes part 2 (Mugilidae to
Carangidae). In: Carpenter, K.E. and V.H. Niem (eds.). The living
marine resources of the Western Central Pacific. Volume 4. FAO
species identification guide for fishery purposes. pp. 2689-2756.
32. Rohde, K. 2002. Ecology and biogeography of marine parasites.
Advances in Marine Biology 43: 1-86.
39. Trinidad, A.C., R.S. Pomeroy, P.V. Corpuz and M. Aguero. 1993.
Bioeconomics of the Philippine small pelagics fishery. ICLARM
Technical Report 38. 74 p.
33. Rokicki, J. 1982. Lironeca indica Edwards 1840 (Crustacea,
Isopoda) from Selar crumenophthalmus (Bloch). Wiadomości
Parazytologiczne 28: 205-206.
40. Vassiliades, G. 1982. Helminthes parasites des Poissons de mer
des côtes du Sénégal. Bulletin de PI.F.A.N. 44(1-2): 78-99.
34. Roos, D., O. Roux and F. Conand. 2007. Notes on the biology of
the bigeye scad, Selar crumenophthalmus (Carangidae) around
Reunion Island, southwest Indian Ocean. Scientia Marina 71(1): 137144.
41. Verweyen, L., S. Klimpel and H.W. Palm. 2001. Molecular
phylogeny of the Acanthocephala (Class Palaeacanthocephala) with
a paraphyletic assemblage of the Orders Polymorphida and
Echinorhynchida. PLoS ONE 6(12): e28285.
35. Roux, O. and F. Conand. 2000. Feeding habits of the bigeye
scad, Selar crumenophthalmus (Carangidae) in La Reunion Island
waters (South-western Indian Ocean). Cybium 24(2): 173-179.
42. Work, T.M., G. Takata, C.M. Whipps and M.L. Kent. 2008. A new
species of Henneguya (Myxozoa) in the big-eye scad (Selar
crumenophthalmus) from Hawaii. Journal of Parasitology 94(2): 524529.
36. Santos, M.J., A. Saraiva, C. Cruz, J. C. Eiras, M. Hermida, C.
Ventura and J. P. Soares. 2009. Use of parasites as biological tags
in stock identification of the black scabbardfish, Aphanopus carbo
Lowe, 1839 (Osteichthyes: Trichiuridae) from Portuguese waters.
Scientia Marina 73(2): 55-62.
37. Schroeder, R.E. 1984. Philippine shore fishes of the Western
43. Yokohama, H., N. Itoh, and S. Tanaka. 2005. Henneguya pagri n.
sp. (Myxozoa: Myxosporea) causing
cardiac henneguyosis in red sea bream, Pagrus major (Temminck &
Schlegel). Journal of Fish Diseases 28: 479–487.