ORIGINAL ARTICLE /ARTÍCULO ORIGINAL
PARASITIC COPEPODS OF THE VERMILION ROCKFISH SEBASTES MINIATUS
(PISCES: SCORPAENIDAE) FROM INSHORE WATERS OF BAJA CALIFORNIA
(EASTERN PACIFIC)
COPÉPODOS PARÁSITOS DEL PEZ VERMILLON SEBASTES MINIATUS (PISCES:
SCORPAENIDAE) EN LAS COSTAS DE BAJA CALIFORNIA (PACÍFICO NORESTE)
1,2 2 3 4,5
*Rodríguez-Santiago Maria Amparo , Gómez Samuel ,Rosales-Casián Jorge A & Grano-Maldonado Mayra I
1 Universidad Autónoma del Carmen (UNACAR), Facultad de Ciencias Naturales, Centro de Investigación de Ciencias
Ambientales (CICA), Av. Laguna de Términos s/n Col. Renovación 2da Sección, C.P. 24155 Ciudad del Carmen, Campeche,
México. (Current address) *Corresponding author e-mail: marodriguezsa@conacyt.mx, amparoshalom@hotmail.com
2 Universidad Nacional Autónoma de México, Instituto de Ciencias del Mar y Limnología, Unidad Académica Mazatlán, Joel
Montes Camarena s/n, Mazatlán 82040, Sinaloa, México.
3Centro de Investigación Científica y de Educación Superior de Ensenada, B. C., División de Oceanología, Departamento de
Ecología Marina, Km 107 carretera Tijuana-Ensenada, Ensenada, B. C., México. C.P. 22800.
4 Departamento de Ecofisiología, Facultad de Ciencias del Mar, Universidad Autónoma de Sinaloa. Paseo Claussen s/n. A.P.
5
610. Mazatlán, Sinaloa. México; Centro de Ci?ncias do Mar, Universidade do Algarve, Campus de Gambelas, 8005-139
Faro, Portugal.
Neotropical Helminthology, 2015, 9(1), jan-jun: 1-12.
ABSTRACT
Keywords: Baja California - Clavellotis sebastidis - Lepeophtheirus rotundipes – Mexico - Neobranchia scorpaenae -
parasitic copepods - Sebastes miniatus.
A qualitative and quantitative survey of the parasitic copepod fauna of the rockfish Sebastes
miniatus from the Pacific coasts of Baja California, Mexico, is presented. The species
composition, temporal changes in abundance, prevalence and intensity of infection of parasitic
copepods were assessed over an annual cycle (2005). Three copepods species were found,
Clavellotis sebastidis and Naobranchia scorpaenae were identified from gill rakers, whereas
Lepeophtheirus rotundipes showed preference for gill arches. Clavellotis sebastidis and L.
rotundipes were the most abundant and most prevalent species. The intensity of infection was
higher for L. rotundipes than for the other species. The average abundance, prevalence and
intensity of infection showed significant variation over the year (p < 0.05). The overall mean
abundance of copepods was highest in summer (August), whereas maximum values of
prevalence were detected in the autumn-winter and the intensity of infection was highest in the
spring. The occurrence of these copepod species on S. miniatus constitutes a new host record and
extends their known geographical distribution. This study is important as it provides new
information and ecological data from copepods parasites whose ecology is poorly studied
worldwide.
1
ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043
INTRODUCTION
2
Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Rodríguez-Santiago et al.
The genus Sebastes from the Pacific coast of
North America is composed of 65 species, and
the greatest diversity (56 species) is found
within the Southern California Bight (Love et
al., 2002). Rockfishes are economically
important for the coastal fisheries of Baja
California (Mexico) (Rodriguez-Medrano,
1993; Hernández-Hernández, 2002; Rosales-
Casián & Gonzalez-Camacho, 2003) and
California (USA) (Eschmeyer et al., 1983;
Love et al., 2003; Stephens et al., 2006). The
vermilion rockfish, Sebastes miniatus (Jordan
& Gilbert 1880), is a common target fish
species in the recreational and commercial
fisheries of Baja California (Rodríguez-
Santiago & Rosales-Casián, 2008), as well as a
highly prized fish for party and private vessel
anglers throughout California, with the
majority of catches occurring at south of
Monterey Bay (Love et al., 2002). This
rockfish distributes from Prince William
Sound, south Alaska to central Baja California,
and inhabits rocky reefs, kelp forests and
canyons at depths of 15–467 m, but is most
commonly found at depths of 50-150 m
(O'Connell et al., 1992; Love et al., 2002).
Given its wide latitudinal distribution, this
species represents a potential intermediate host
for many parasites, and can be prey of final
hosts such as the California sea lion (Lowry et
al., 1991). Parasitic copepods are commonly
found on cultured and wild marine finfish. The
attachment and feeding activities on mucous,
tissues, and blood of these types of parasites
may cause diseases and mortality (Lin et al.,
1994; Pike & Wadsworth, 1999; Ho et al.,
2001). Therefore, knowledge of their
occurrence and abundance on cultured fishes is
of great importance as they may have the
potential to affect growth, fecundity and
survival (Krkosek et al., 2006; Costello, 2006;
Morales-Serna et al., 2011) and in wild
population (Iannacone et al., 2010; Iannacone
et al., 2011; Acacio et al., 2012; Mendoza-
Cruz et al.,2013).
Currently, few parasite studies in commercial
marine fishes from the Mexican Pacific coasts,
RESUMEN
Palabras clave: Baja California - Clavellotis sebastidis - copépodos parásitos - Lepeophtheirus rotundipes - Mexico - Neobranchia scorpaenae -
Sebastes miniatus.
En este trabajo se presenta un estudio cualitativo y cuantitativo de la fauna de copépodos
parásitos del pez vermillon o rockotrojo Sebastes miniatus de las costas del Pacífico de Baja
California, México. Se evaluó la composición de las especies de copépodos parásitos así como la
abundancia, prevalencia e intensidad de infección durante un ciclo anual (2005). Las especies de
copépodos identificados fueron: Lepeophtheirus rotundipes, Naobranchia scorpaenae y
Clavellotis sebastidis. De los cuales N. scorpaenae y C. sebastidis mostraron preferencia por las
branquiespinas, mientras que L. rotundipes mostró preferencia por los arcos branquiales. La
especie más abundante fue C. sebastidis, mientras que la prevalencia fue mayor en L. rotundipes.
La intensidad de infección fue relativamente mayor en L. rotundipes. La abundancia media,
prevalencia e intensidad de infección mostraron variaciones significativas durante el año (p <
0.05). La abundancia promedio total de copépodos fue mayor en el verano (agosto), mientras que
los valores máximos de prevalencia fueron detectados en el periodo otoño-invierno. Por otra
parte, la intensidad de la infección fue mayor en primavera. El registro de estas tres especies de
copépodos en S. miniatus presentadas en este estudio constituye un nuevo registro de hospedero y
amplían su distribución geográfica conocida.
MATERIALS AND METHODS
3
especially from Baja California, are available
(Rodríguez-Santiago & Rosales-Casián, 2008;
Rodriguez-Santiago et al., 2014). Considering
the number of coastal systems along the
Mexican coasts, particularly along the Baja
California peninsula, and the economic
importance of rockfish species, this lack of
parasitological studies is noteworthy. Reports
on the copepod parasitic fauna of fishes from
Mexican coastal systems is scarce and are
either published only sporadically or deal with
the occurrence of certain copepod species or
description of new taxa (Morales-Serna et al.,
2012). Therefore, the aim of the present study
is to assess the intra-annual variability in the
abundance, prevalence and intensity of
infection of parasitic copepods of S. miniatus
from the Pacific coast of Baja California
(Mexico), previous studies (Rodríguez-
Santiago & Rosales-Casián, 2008; Rodriguez-
Santiago et al., 2014) have identified this area
which has potential value for a wide range of
commercial fish, as well as to analyze the
relationship between copepod abundance and
the size and condition factor of the host and
water temperature.
Samples of the red rockfish, Sebastes miniatus
were obtained bimonthly from catches of
recreational fishing at San Quintín, Baja
California, México (30°33'37'' N; 115°56'33''
W) (Figure 1). A total of 210 individuals (210-
610 mm total length) were sampled. Surface
water temperatures (°C) and at a depth range of
50–140 m depth were obtained during 2005
from station 107.32 during the IMECOCAL
cruises (30°27'17.3''N, 116°09'41.8''W),
located close to Isla San Martin (García-
Córdova et al., 2005).
Fish samples were transported in individual
plastic bags to the laboratory in a cool box.
Each fish was examined for the presence of
parasitic copepods on skin, fins, gills and gill
rakers. External examination of copepods on
the body surface of the hosts was performed
under good illumination, and gill arches were
removed from each fish and carefully
inspected in a Petri dish using a
stereomicroscope (LEICA MZ9.5). The plastic
bag contents were also examined for the
presence of detached copepods.
Parasites found on each fish were preserved in
labeled vials with 70% ethanol. Copepods
identification was performed following
Castro-Gonzalez (2005), Dojiri (1981) and
Dojiri (1979). Voucher specimens were
deposited in the fish collection of the
Laboratorio de Ecología Pesquera, of the
Centro de Investigación Científica y de
Educación Superior de Ensenada CICESE,
Ensenada, Baja California, México. The
Fulton's Condition Factor (KLP) was
3
calculated for each fish as K = [W/TL ] 10,000;
where: W = weight (g) and TL = total length
(mm) (Ricker, 1975). This factor is used to
describe the physiological condition of the
individual fish. Weight (g) of specimens was
measured with an analogic balance and the
total length (cm) with an ictiometer.
Prevalence (percent of infected hosts among
all hosts examined), abundance (number of
parasites per host) and intensity (number of
parasites per infected hosts) of parasites were
determined according to Margolis et al.
(1982). To assess significant variations in
copepod abundance, prevalence and intensity
of infection over the year, non-parametric
analyses of variance of Kruskall-Wallis (KW)
were performed (Steel & Torrie, 1986).
Spearman rank correlations were used to
assess relationships between the parasite
abundance and the host size (mm) and weight
(g), Fulton's condition, and seawater
temperature (°C).
Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Copepods parasites in marine fish
4
Water temperature
During 2005, the surface water temperature in
the fishing area showed a mean (± SE) of 16.0
o
± 0.3 C. The highest mean temperature was
o
observed in August (18.2 ± 0.5 C), and the
o
lowest in February (14.9 ± 0.3 C). At the
fishing depth (50-140 m), the water
temperature did not show considerable
variation; the average annual temperature was
10.9 ± 0.09°C with maximum values in
o
October (11.5 ± 0.20 C) and lowest values in
o
June (10.2 ± 0.08 C).
Abundance, prevalence and intensity of
parasitic copepods
Three parasitic copepod species Le
peophtheirus rotundipes Dojiri, 1979,
Naobranchia scorpaenae Dojiri, 1981 and
Clavellotis sebastidis Castro & González,
2005 were identified on from individuals of S.
miniatus the skin, gills and rakers. Specimens
of L. rotundipes were found on the skin and gill
filaments whereas those of N. scorpaenae and
C. sebastidis were found attached to gill
filaments and gill rakers.
The overall mean abundance of parasitic
copepods was of 0.5 ± 0.2 ind/host with
maximum values in August and lowest values
in February (Table. 1). The most abundant
species was C. sebastidis (annual mean
abundance = 0.6 ± 0.2 ind/host), whereas N.
scorpaenae was the least abundant species
(annual mean abundance = 0.3 ± 0.2 ind/host).
In general, the highest proportion of infected
hosts was recorded in October (mean
prevalence = 30 ± 12.3%) and the lowest in
February (4.4 ± 5.4%). The prevalence was
relatively higher in L. rotundipes (16.2 ± 2.8%)
than in C. sebastidis and N. scorpaenae (14.9 ±
7.6 and 14.5 ± 5.6%, respectively) (Fig. 2b).
The intensity of infection showed an annual
average value of 1.97 ± 0.4 and did not show
significant variations over the year. The higher
average value of intensity was in October (3.3
± 1.0). The mean abundance of N. scorpaenae
showed significant changes (Kruskall-Wallis,
H = 11.05, p < 0.05) over time.
The Spearman rank correlation analyses
showed that there were no significant
relationships between abundance and
RESULTS
Figure. 1. Location of the study area at San Quintín Bay, Baja California, Mexico.
Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Rodríguez-Santiago et al.
5
prevalence of parasitic copepods with the size
(r = 0.10, p > 0.05) and weight (r = 0.19, p >
0.05) of the hosts, surface water temperature (r
= -0.10, p > 0.05), bottom temperature (r =
0.19, p > 0.05) and condition of fish (r = -0.01,
p > 0.05). However, the highest (August) and
lowest (February) recorded abundance of
parasitic copepods coincided when surface
water temperatures reached their highest and
lowest values respectively (Figure 5).
Figure. 2. Mean abundance, prevalence and intensity of infection of copepods parasites in Sebastes miniatus collected at San
Quintin, Baja California, during an annual cycle, 2005.
Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Copepods parasites in marine fish
6
Table. 1. Characterization of copepods parasitic infections of Sebastes miniatus (n = 210) from the coasts of San
Quintin, Baja California, Mexico. Number of examined fish (NF), number of copepod species (NS), prevalence
(MP%), mean abundance (MA), mean intensity (MI), mean bottom temperature (BT), mean surface water
temperature (ST), mean condition factor (K), total length in cm (TL), weight in g (W).
Sampling
month
NF
NS
NP
MP %
MA
MI
BT ST K TL W
February 15 1 3
4.4±5.44
0.1±0.08 0.5±0.61
12.4 14.4 0.11 400 1081.6
April 28 3 26 14.0±6.48 0.5±0.24 2.27±0.17 10.6 13.9 0.15 615 629.5
June 62 3 40 14.0±6.49 0.5±0.23 1.39±0.25 10.8 15.0 0.13 520 1448.5
August 35 3 11 7.5±2.16
1.3±0.30 1.33±0.40 10.6 21.8 0.13 485 1128.6
October 29 3 46 29.7±12.3 0.5±0.23 3.33±1.08 12.4 15.2 0.13 550 823.7
December 45 3 54 22.0±4.56 0.2±0.04 3.0±0.0
12.2 15.6 0.14 680 1285.7
1864) and S. proriger (Jordan & Gilbert,
1880), in the Northwest Pacific (Sekerak &
Arai, 1977; Kabata, 1988). Love et al. (1984)
also reported Naobranchia occidentalis C.B.
Wilson, 1915 in Sebastes serranoides
(Eigenmann & Eigenmann, 1890) in central
California. Prior to this study, C. sebastidis had
not been recorded in Mexican waters. The
genus Clavellotis includes nine species
(Kroyer, 1863), seven of which were
transferred from the genus Clavellopsis
(Wilson, 1915) by Kabata (1990) and recently
C. sebastidis female and male were described
for the fish Sebastes oculatus by Castro &
Gonzalez (2005). Clavella parva C. B. Wilson,
1912 has been found in Sebastes aleutinus
(Jordan & Evermann, 1898), S. elongatus
Ayres, 1859, S. bobcocki (Thompson, 1915)
and S. caurinus Richardson, 1844 in the North
Pacific (Sekerak & Arai, 1977), S. alutus
(Gilbert, 1890) and S. auriculatus Girard, 1854
in British Columbia (Wilson, 1915; Sekerak &
Arai, 1977), S. serranoides (Eigenmann &
Eigenmann, 1890) in central California (Love
et al., 1984), S. diploproa in British Columbia
(Kabata, 1970), S. maliger in the North Pacific
(Kabata, 1988) and S. melanops Girard, 1856
in Alaska (Wilson, 1915). In this work, the
genus Clavellotis is reported for the fist time
parasitising S. miniatus, and San Quintin Baja
California in the Mexican Pacific coast is
reported as a new geographic distribution area.
The present report of L. rotundipes, N.
scorpaenae and C. sebastidis in S. miniatus
Despite the economic importance of the
rockfish Sebastes miniatus in the Pacific coasts
of North America (Rosales-Casian &
González-Camacho, 2003) little is known
about the ecology of its parasitic fauna,
particularly of the three parasitic copepods
found in this study (L.rotundipes, N.
scorpaenae and C. sebastidis). The genus
Lepeophtheirus Normann, 1832, has about 133
valid species (Ho & Lin, 2000) from which
only L. longipes Wilson C.B., 1905, L.
parviventris Wilson C.B., 1905, L. paulus
Cressey, 1969 and L. oblitus Kabata, 1973
have been reported in 12 of the 65 Sebastes
species recorded in this region (Love et al.,
2002). It is important to mention that species of
genus Lepeophtheirus have also been reported
in other fish hosts and are very common in
cultured fish (Johnson et al., 2004).
Other species of the genus Naobranchia (i.e.,
N. robusta [Kabata, 1970] and N. occidentalis
Wilson C.B., 1915 [Sekerak & Arai, 1977]) are
known to parasitise other Sebastes species
(Love et al., 1984), such as S. alutus (Gilbert
1890), S. bobcocki (Thompson, 1915), S.
borealis Barsukov 1970, S. brevispinis
(Bean,1884), S. caurinus Richardson, 1844, S.
diploproa (Gilbert, 1890), S. maliger (Jordan
& Gilbert, 1880), S. nigrocinctus Ayres, 1859,
S. paucispinus Ayres, 1854, S. pinniger (Gill,
DISCUSSION
Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Rodríguez-Santiago et al.
7
constitutes a new host record in the Pacific
coasts of Baja California and a new
geographical record. The low overall
prevalence was probably due to loss of
parasites, while accidental transfer among
hosts may also have occurred in this study. In
addition, ecological and environmental
conditions have been shown to influence the
prevalence of ectoparasites (Williams &
MacKenzie, 2003). Copepods embedded in the
tissues of the host were rejected by an immune
reaction of the host tissues (Jonhson &
Albright, 1992; Tsotetsi, 2005) apparently
resulting from the integument's response to
penetration by the parasite. N. scorpaenae and
C. sebastidis showed relatively high
abundances from spring to autumn, whereas its
prevalence and intensity of infection were
highest in spring. Only L. rotundipes showed a
seasonal pattern throughout the year.
Similarly, the prevalence and mean intensity of
N. scorpaenae and C. sebastidis was
significantly higher in October and December
compared to the rest of the year. The
prevalence and mean intensity of L. rotundipes
was significantly higher in April and June
(with the highest water temperature) than in
other months. Seasonality has also been
documented in two other species of
Lepeophtheirus (L. salmonis and L. pectoralis)
from higher latitudes; both of which reached
high infection levels in spring (Schram et al.,
1998; Cavaleiro & Santos, 2009).
The prevalence of L. rotundipes varied
significantly between sampling months, but
did not show a regular pattern. This suggests
that the occurrence of L. rotundipes on S.
miniatus is not affected by water temperature.
However, the mean intensity of this species
was almost constant throughout the one year
period of study, except for a significantly lower
value observed in February. Nagasawa et al.
(1993) found increasing infection levels of the
copepod Lepeophtheirus salmonis (Krøyer,
1837) with host age and size, while Morand et
al. (1999) found that parasite species and
parasite abundance correlated positively with
host body length, and Boxshall (1974) found
that the age of the host influenced the
frequency distribution of the parasitic copepod
Bomolochus confusus Stock, 1953. The
difference between the frequency distributions
of the parasite on the different size classes of
hosts suggests that there is a gradual
accumulation of parasites with increasing size,
and therefore age, of host fishes. Iannacone et
al. (2011) found that the total fish length was
not related to the prevalence and abundance of
helminth infection, except for a negative
relationship with the prevalence of the
copepod Ommatokoita elongata Grant, 1827
in the Pacific Guitar fish Rhinobatos planiceps
(Garman, 1880) in Peru. Subsequent,
Iannacone & Alvariño (2013) reported the total
length and weight of the fish Brama japonica
Hilgendorf, 1878 showed a close relation
between fish sex and the average volumetric
abundance of the copepod parasite Hatschekia
conifera Yamaguti, 1939.
The prevalence of all three parasitic copepod
species (L. rotundipes, N. scorpaenae and C.
sebastidis) was more variable than their mean
intensity. This supports the hypothesis that
intensity of infection is a species characteristic
and that the biological features of parasitic
species can potentially override local
environmental conditions in driving parasite
population dynamics. According to Poulin
(2006), this is possible because prevalence is
determined by encounter rates between
parasites and hosts, which are influenced by
processes external to the fish (e.g., survival of
free-living parasite stages), probably
determined by local factors, whereas intensity
of infection is determined to a large extent by
processes acting within the fish. No significant
association was detected in this study between
the burden of parasitic copepods and fish
condition factor K. Similarly, other studies
failed to indicate that parasites affect the fish
condition in natural systems (Mo & Heuch,
1998; Olivero-Vervel et al., 2005; Jones et al.,
Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Copepods parasites in marine fish
8
2006). In contrast, Lemly & Esch (1984)
provided a clear example of mortality due to
the direct effects of parasitism in a natural
population of fish. Evidence of damage
produced by parasitic copepods is better
documented from aquaculture studies. In a
farm of the southern bluefin tuna Thunnus
maccoyii (Castelnau, 1872), Hayward et al.
(2009) found that the sea louse Caligus chiatos
Lin & Ho, 2003 was strongly associated with
both fish condition and severity of eye damage.
It is possible that the health of S. miniatus was
already reduced by copepod infection because
these crustaceans damage their hosts directly
through their attachment mechanisms and
feeding activities. However, the severity of
damage reported by Hayward et al. (2009) was
likely subtle, and the methods used here waere
unable to detect any significant effect of
copepod parasitism on S. miniatus. The
presence of L. rotundipes (n=67), N. scorpenae
(n=60) and C. sebastidis (n=53), are described
for the first time parasitising the vermilion
rockfish, S. miniatus, in Baja California. It is
evident that both taxonomic and ecological
aspects of fish copedods parasite research are
greatly needed and the knowledge concerning
the biodiversity of parasitic copepods in
Mexico (Morales Serna et al., 2012). Although
C. sebastidis has been previosly reported in
Argentina (Castro & Gonzalez, 2005).
Although some work has been perfomed, still a
lack of knowledge which impede our
understanding of the evolutionary and
biogeographical history of the host-parasite
associations. The ecological studies of
parasitic copepods in commercially important
fish species like the vermilion rockfish S.
miniatus are important due to a high market
value. Also, this fish species is highly
appreciated by its white meat, texture and
flavor. This fish species is abundant in the
commercial fishing and sport-fishing from
Baja California (México) and California
(USA) and is a common target for recreational
and commercial fisheries of Baja California
(Rodríguez-Santiago & Rosales-Casián, 2008;
Rodriguez-Santiago et al., 2014). In
conclusion, in this study the magnitude of
infection is very well represented, also the
importance of analysing the copepod parasitic
fauna of commercially important fish species
has been highlighted and shed some light on
the knowledge of biodiversity and biology of
parasitic copepods. New geographical host
records are important, for the reason that you
can detect distribution patterns, and may
employ other fish to colonise, etc. Studies on
copepod parasitizing other commercial fish
hosts, particularly in Mexico and Baja
California, and the completion of their life
cycle warrant further investigation.
This study was funded by the CICESE project:
Analysis of the recreative sport-Wshing
catches from San Quintín, B.C., Mexico and
by the UCMEXUS-CONACYT grant project:
Baseline study of the nearshore non-reef
Wshes of Bahía de Los Angeles, Baja
California, México, prior to proposed
development. We thank to E. Fajer (CIAD-
Unidad Mazatlan) for providing us with
materials and to J. E. Kalman of the Marine
Aquarium, San Pedro, California, for his help
in obtaining some old articles.
ACKNOWLEDGMENTS
Acacio, M, Varella, AMB & Malta, JCO. 2012.
Boxshall, GA. 1974.
The parasite crustacea ns of
Serrasalmus rhombeus (Linnaeus, 1776)
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Central Amazon, Brazil. Neotropical
Helminthology, vol. 6, pp.179-184.
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of Lepeophtheirus pectoralis (Müller,
1776): seasonal variation in abundance
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