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First record of Hamaticolax scutigerulus (Copepoda: Bomolochidae) in Brazil, ectoparasite of the spotted goatfish Pseudupeneus maculatus (Actinopterygii: Mullidae)/Primeiro registro de Hamaticolax scutigerulus (Copepoda: Bomolochidae), parasito de saramunete Pseudupeneus maculatus (Actinopterygii: Mullidae), no Brasil.

Introduction

Copepods comprise the second most frequent parasites of marine fishes in the Neotropical region (Luque & Poulin, 2007). The family Bomolochidae Claus, 1875 has at least 20 genera and the species are characterized by specialized structures on the body for attachment and feeding on their hosts, generally on the gills, branchial cavity and noses (Kim & Moon, 2013, Maran, Moon, Adday, Khamees, & Myoung, 2014, Walter & Boxshall, 2015).

The genus Hamaticolax Ho and Lin (2006) was proposed together with Cresseyus Ho and Lin (2006), to accommodate some orphan species previously allocated to the genus Holobomolochus Vervoort (1969). Hamaticolax differs from the genera Cresseyus and Holobomolochus in having a pair of rostral hooks (Ho & Lin, 2006). To make a taxonomic identification of Hamaticolax, several characteristics need to be taken into consideration. However, the leg armor (Le), i.e. leg composition in terms of setae and spines, are the feature that has been most characterized in previous descriptions (Kabata, 1971).

The spotted goatfish Pseudupeneus maculatus (Bloch, 1793) is a demersal marine fish that is frequently found in shallow coral reef waters, along the entire western Atlantic seaboard from New Jersey, USA, to Santa Catarina, Brazil (Lessa & Nobrega, 2000, Hostim-Silva et al., 2006). This species is caught mainly by means of trap fishing on the coast of the state of Pernambuco, Brazil, and its meat has been appreciated in both the national and the export market. Hence, this fish is economically valuable (Lessa & Nobrega, 2000).

The aims of this study were to provide the first report of occurrences of the parasite Hamaticolax scutigerulus (Wilson, 1935) in Pseudupeneus maculatus caught in the state of Pernambuco, northeastern Brazil, and to present information on the leg armor of this parasite, in comparison with 10 described species.

Material and methods

A total of 120 fish were obtained for parasite evaluation between October 2012 and September 2013. They were caught by an artisanal fisherman at a mean depth of 20 m, in an area comprising the following coordinates: site 1 (7[degrees] 37' 28.43" S; 34[degrees] 1' 10.24" W), offshore from the municipality of Goiana; site 2 (7[degrees] 37' 50.34" S; 34[degrees] 43' 41.73" W), in the metropolitan region of Recife; and site 3 (8[degrees] 54' 41.79" S; 33[degrees] 57' 23.45" W), offshore from the municipality of Sao Jose da Coroa Grande (Figure 1).

The fish were stored in thermal boxes with ice and were transported to the Marine Fish Farm Laboratory of the Fisheries and Aquaculture Department of the Universidade Federal Rural de Pernambuco (UFRPE) for necropsy. The gills were collected and fixed in 70% alcohol, and were slightly agitated to detach the parasites (Eiras, Takemoto, & Pavanelli, 2006, Jeronimo, Martins, Ishikawa, Ventura, & Tavares-Dias, 2011) for subsequent analysis.

The copepods were clarified in lactic acid (Humes & Gooding, 1964), and then dissected and observed on compressed slides under a DIC Iamger.A2 microscope equipped with Axio Cam MRc camera. The adult specimens were identified as described by Vervoort (1969), Ho and Lin (2006) and Morales-Serna and Gomez (2010). The composition of the setae and spines of the leg armor was compared with previous studies on 10 described species in the genus Hamaticolax. Prevalence, mean intensity of infestation and mean abundance were calculated as described by Bush, Lafferty, Lotz, and Shostak (1997).

Results and discussion

A total of 80 copepods were collected, comprising 77 females and three males in hosts of both sexes. Nine gravid females and three males were measured (Table 1) and identified as Hamaticolax scutigerulus (Wilson, 1935) (Figure 2).

The total length was 1480[+ or -]70 pm for females and 549[+ or -]32 pm for males, and the greatest width was about 993 [+ or -]67 pm for females and 290[+ or -]20 pm for males.

The measurements and characteristics of the setae and spines on the legs of the present material were compared with those of another species (Table 2). The prevalence of Hamaticolax scutigerulus in the fish examined here was 35%; its mean intensity was 1.9[+ or -]1.3 and its mean abundance was 0.7[+ or -]1.2.

The present specimens were similar to the specimens of H. scutigerulus that were observed parasitizing P. maculatus in Piscadera Bay, Curasao, by Vervoot (1969). On the other hand, our specimens and those reported by Vervoot (1969) showed larger measurements (total length, greatest width, length and width of somites and furcal rami, and length of longest furcal seta) than those found by Cressey (1983).

However, some differences in comparison with other species of the genus Hamaticolax need to be emphasized (Table 1). Females of H. scutigerulus showed shorter total body length than those of H. attenuatus and H. paralabracis, longer total body length than those of H. embiotocae and similar total body length to those of H. galeichthyos, H. occultus, H. spinulus, H. unisagittatus and H. prolixus.

The body width of the specimens collected in Pernambuco was similar to the width in H. attenuatus, H. paralabracis and H. spinulus and greater than in H. galeichthyos, H. unisagittatus and H. prolixus. The long seta length was shorter than in H. attenuatus, H. galeichthyos, H. occultus and H. spinulus and longer than in H. paralabracis and H. unisagittatus. The male specimens in the present study had shorter body length than in H. galeichthyos and H. prolixus, but their length resembled the body length of H. embiotocae. However, the body width was shorter than that found in H. galeichthyos.

The legs of copepods in the genus Hamaticolax have distinct composition in terms of the number of segments (proximal, medial and distal articulations) that form the endopodite and exopodite rami. The number of setae and spines of these rami differ among Hamaticolax species and the findings from the present study are in agreement with those from previous studies (Table 2). Vervoort (1969) argued that the exopodite of the first legs (Le1) in females of H. scutigerulus possess one segment with six setae and five spines, thus differing from the other species. Moreover, male specimens present distal segments of the Le1 exopodite (three setae and three spines) that differ from those of other species.

In contrast to that related by Vervoort (1969), the textual information and drawings from Cressey (1983) showed an Le4 endopodite with one seta and no spines, both in proximal and medial segments, and three setae and no spines in the distal segment. Also, Le5 had four setae and no spines in the distal segment was reported in H. scutigerulus females.

In the present study, it was found that the medial segment of the Le4 endopodite of the H. scutigerulus females had one element, which was probably a spine and not a seta as proposed by Cressey (1983), and the distal segment of Le5 had four setae, as also observed by Cressey (1983). In addition, for the males, the distal segment of the Le4 endopodite had one seta and one spine, according to Cressey (1983). In the present study, it was not possible to discern whether there were two or three elements in the distal segment of the Le4 endopodite for the males.

The parasitological indices of H. scutigerulus in P. maculatus examined in Brazil were lower than those found for Bomolochus soleae Claus, 1864, in Soleasolea in Portugal (Durieux, Marques, Sasal, Begout, & Cabral, 2007). Similarly, Tavares and Luque (2003) described Hamaticolaxunisagittatus (Tavares & Luque, 2003) (= Acantholochus unisagittatus sp. nov.) in Centropomus undecimalis in Rio de Janeiro, with high prevalence and mean abundance. Pseudupeneus maculatus has also been found to be parasitized by two other species of Bomolochidae: Orbitacolax analogus Vervoort (1969) and Orbitacolax hapalogenyos (Yamaguti & Yamasu, 1959-Maran et al., 2014) on the gills of fish in Belize (Cressey & Cressey, 1989). On the other hand, all of the present material examined was H. scutigerulus.

The ventral body structures and hooks of the antennae in the family Bomolochidae are adapted so as to allow the parasite to become attached to the host on the gills, internal wall of the operculum, fins and integument, and around the eyes (Radhakrishnan & Nair, 1983, Boxshall, 2005).

Conclusion

In conclusion, some morphometric differences in the composition of the setae and spines of the leg armor can be found in the genus Hamaticolax. Nonetheless, these constitute the most valid characteristic for specific identification.

Doi:10.4025/actascibiolsci.v39i2.34100

Acknowledgements

The authors thank Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq) for research grant to M. L. Martins (305869/ 2014-0).

References

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Luque, J. L., & Bruno, M. (1990). Two new species of Acantholochus Cressey, 1984 (Copepoda: Bomolochidae) parasitic on Peruvian marine fishes. Journal of Natural History, 24(1), 241-249.

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Received on November 1, 2016.

Accepted on March 21, 2017.

Lucas Cardoso (1) *, Mauricio Laterga Martins (1), Ana Carolina Figueiredo Lacerda (2), Douglas Ismael Cadorin (1), Carolina Nunes Costa Bonfim (3) and Ricardo Luis Mendes de Oliveira (3)

(1) Laboratorio de Sanidade de Organismos Aquaticos, Departamento de Aquicultura, Universidade Federal de Santa Catarina, Rodovia Admar Gonzaga, 1346, 88040-900, Florianopolis, Santa Catarina, Brazil. (2) Departamento de Ecologia e Sistemas, Universidade Federal da Paralba, Joao Pessoa, Paralba, Brazil. (3) Departamento de Engenharia Pesqueira e Aquicultura, Centro de Ciencias Agrarias Aplicadas, Universidade Federal de Sergipe, Sao Cristovao, Sergipe, Brazil. * Author for correspondence. E-mail: lucas.cardoso@ufsc.br

Caption: Figure 1. Location of the three sites (1, 2 and 3) of capture of 120 specimens of Pseudupeneus maculatus used in this study.

Caption: Figure 2. Hamaticolax scutigerulus, ventral view: (A) female and its (B) anterior portion, (C) egg sacs and (D) uropods; (E) male.
Table 1. Comparative values (mean values  [+ or -]  standard
deviation, in [micro]m) of measurements of Hamaticolax species,
especially Hamaticolax scutigerulus from the gills of Pseudupeneus
maculatus captured in the Coast of Pern ambuco State, Brazil, and
other regions. Length (L), Width (W), and minimum and maximum values
between parentheses.

Species               Hamaticolax           Hamaticolax
                     scutigerulus (a)      scutigerulus (b)

Author               Vervoort (1969)       Cressey (1983)

Measurements          Female      Male     Female    Male

Total length           1640       611       1116      --
Greatestwidth          1000       286        750      --
L/W of cephalic      419/999     226/286     -/-      -/-
  somite
L/W of second        338/850     61/231      -/-      -/-
  thoracic somite
L/W of third         284/601     50/198      -/-      -/-
  thoracic somite
L/W of fourth        176/338     44/138      -/-      -/-
  thoracic somite
L/W of fifth         135/243     55/126      -/-      -/-
  thoracic somite
L/W of genital       216/473     121/132    83/130   -/-
  complex
L/W of third          95/243       -/-      33/71    -/-
  abdominal somite
L/W of fourth         68/216     39/72 2    21/59    -/-
  abdominal somite
L/W of fifth          95/176     22/55 5    56/53    -/-
  abdominal somite
L/W of furcal rami    68/81 2     22/19      -/-      -/-

Length of longest      210 2      200        150      --
  furcal seta
L/W of ovisac        837/270       -/-       -/-      -/-

Species                  Hamaticolax scutigerulus (c)

Author                           Current study

Measurements              Female                Male

Total length          1480 [+ or -] 70     549 [+ or -] 32
Greatestwidth        993 [+ or -] 67      290 [+ or -] 20
L/W of cephalic      488 [+ or -] 67/     211 [+ or -] 14/
  somite              992 [+ or -] 70     286 [+ or -] 13
L/W of second        279 [+ or -] 59/      65 [+ or -] 3/
  thoracic somite     883 [+ or -] 65     238 [+ or -] 14
L/W of third         217 [+ or -] 32/      49 [+ or -] 8/
  thoracic somite     618 [+ or -] 62     203 [+ or -] 18
L/W of fourth        164 [+ or -] 31/      50 [+ or -] 5/
  thoracic somite     355 [+ or -] 34     143 [+ or -] 10
L/W of fifth         130 [+ or -] 14/     55 [+ or -] 10/
  thoracic somite     267 [+ or -] 41      112 [+ or -] 2
L/W of genital       174 [+ or -] 43/     119 [+ or -] 20/
  complex            349 [+ or -] 106      143 [+ or -] 4
L/W of third           93 [+ or -] 8/            -/-
  abdominal somite   205 [+ or -] 60
L/W of fourth          76 [+ or -] 6/      36 [+ or -] 9/
  abdominal somite   189 [+ or -] 69       74 [+ or -] 3
L/W of fifth          75 [+ or -] 14/      28 [+ or -] 7/
  abdominal somite    137 [+ or -] 35      63 [+ or -] 11
L/W of furcal rami     74 [+ or -] 8/      29 [+ or -] 11/
                       76 [+ or -] 5       20 [+ or -] 3
Length of longest     196 [+ or -] 15     225 [+ or -] 10 6
  furcal seta
L/W ofovisac         768 [+ or -] 124/           -/-
                      296 [+ or -] 61

Species               Hamaticolax
                       attenuatus

Author               Vervoort (1969)

Measurements          Female     Male

Total length           2280       --
Greatestwidth           860       --
L/W of cephalic       500/860     -/-
  somite
L/W of second         176/675     -/-
  thoracic somite
L/W of third          242/567     -/-
  thoracic somite
L/W of fourth         135/324     -/-
  thoracic somite
L/W of fifth          149/243     -/-
  thoracic somite
L/W of genital        257/257     -/-
  complex             257/176     -/-
L/W of third
  abdominal somite
L/W of fourth         230/149     -/-
  abdominal somite
L/W of fifth          162/135     -/-
  abdominal somite
L/W of furcal rami     -/122      -/-

Length of longest       608       --
  furcal seta
L/W ofovisac         2200/160     -/-

Parasite of Pseudupeneus maculatus from Curacao (a), Belize
(b) and Brazil (c).

                                                 Hamaticolax
Species              Hamaticolax galeichthyos    galeichthyos

                                                   Luque and
Author              Luque and Bruno (1990)       Bruno (1990)

Measurements                  Female                 Male

Total length             1542 (1425-1750)       1025 (1000-1075
Greatestwidth             517 (425-575)          342 (325-350)
L/W of cephalic                -/-                    -/-
  Somite
L/W of second                  -/-                    -/-
  thoracic somite
L/W of third                   -/-                    -/-
  thoracic somite
L/W of fourth                  -/-                    -/-
  thoracic somite              -/-                    -/-
L/W of fifth
  thoracic somite         157 (140-190)/              -/-
L/W of genital            190 (150-210)
  complex                 107 (100-120)/          60 (50-70)/
L/W of third              120 (110-140)               70
  abdominal somite
L/W of fourth              90 (70-110)/           47 (40-50)/
  abdominal somite        107 (100-120)           57 (50-60)
L/W of fifth               67 (60-80)/                -/-
  abdominal somite         97 (90-110)
L/W of furcal rami         67 (60-80)/                -/-
                           50 (40-60)
Length of longest         390 (340-430)               --
  furcal seta
L/W ofovisac                                          --

Species                Hamaticolax     Hamaticolax
                         maleus         occultus

Author                   Oldewage        Kabata
                          (1994)         (1971)

Measurements           Female   Male     Female

Total length             --      --     1390-1920
Greatestwidth            --      --        --
L/W of cephalic         -/-     -/-      508/894
  somite
L/W of second           -/-     -/-      127/834
  thoracic somite
L/W of third            -/-     -/-      179/597
  thoracic somite
L/W of fourth           -/-     -/-      127/241
  thoracic somite       -/-     -/-      89/236
L/W of fifth
  thoracic somite       -/-     -/-      165/241
L/W of genital
  complex               -/-     -/-      76/165
L/W of third
  abdominal somite
L/W of fourth           -/-     -/-      63/140
  abdominal somite
L/W of fifth            -/-     -/-      63/114
  abdominal somite
L/W of furcal rami      -/-     -/-       51/38

Length of longest        --      --        394
  furcal seta
L/W ofovisac            -/-     -/-     1079/240

Species              Hamaticolax   Hamaticolax paralabracis
                      occultus

Author                 Kabata      Luque and Bruno (1990)
                       (1971)
Measurements            Male           Female        Male

Total length             --        2525 (2375-2725    --
Greatestwidth            --         925 (875-975)     --
L/W of cephalic          -/-             -/-         -/-
  somite
L/W of second            -/-             -/-         -/-
  thoracic somite
L/W of third             -/-             -/-         -/-
  thoracic somite
L/W of fourth            -/-             -/-         -/-
  thoracic somite        -/-             -/-         -/-
L/W of fifth
  thoracic somite        -/-       207 (200-220)/    -/-
L/W of genital                     217 (210-230)
  complex                -/-       166 (120-190)/    -/-
L/W of third                       135 (130-150)
  abdominal somite
L/W of fourth            -/-       160 (150-170)/    -/-
  abdominal somite                 120 (110-130)
L/W of fifth             -/-       130 (120-150)/    -/-
  abdominal somite                  103 (90-120)
L/W of furcal rami       -/-       113 (100-130)/    -/-
                                     43 (40-50)
Length of longest        --          50 (45-55)       --
  furcal seta
L/W ofovisac             -/-             --          -/-

Species              Hamaticolax      Hamaticolax unisagittatus
                      spinulus
Author               Cressey (1969)   Tavares and Luque (2003)

Measurements         Female    Male         Female          Male

Total length          1490      --      1240 (1020-1380)     --
Greatestwidth          870      --    614.2 (540.6-690.1)    --
L/W of cephalic      -/870     -/-            -/-           -/-
  somite
L/W of second          -/-     -/-            -/-           -/-
  thoracic somite
L/W of third           -/-     -/-            -/-           -/-
  thoracic somite
L/W of fourth          -/-     -/-            -/-           -/-
  thoracic somite      -/-     -/-            -/-           -/-
L/W of fifth
  thoracic somite    200/200   -/-    132.8 (90.7-172.5)/   -/-
L/W of genital                        182.4 (130.4-198.4)
  complex             224/-    -/-      63.6 (51-73.7)/     -/-
L/W of third                           86.4 (73.7-90.7)
  abdominal somite
L/W of fourth         200/-    -/-      54.5 (51-56.7)/     -/-
  abdominal somite                     79.4 (62.3-96.3)
L/W of fifth          112/-    -/-      62.3 (51-90.7)/     -/-
  abdominal somite                      75.1 (68-90.7)
L/W of furcal rami   142/60    -/-       73.4 (68-85)/      -/-
                                        34.7 (34-39.7)
Length of longest      768      --      88.5 (68-107.7)      --
  furcal seta
L/W ofovisac         950/ -    -/-            -/-           -/-

Species              Hamaticolax prolixus

Author               Cressey (1969) and
                          Ho (1972)

Measurements          Female      Male

Total length         1600-1870   710-780
Greatestwidth           580        --
L/W of cephalic         -/-        --
  somite
L/W of second           -/-        --
  thoracic somite
L/W of third            -/-        --
  thoracic somite
L/W of fourth           -/-        --
  thoracic somite       -/-        --
L/W of fifth
  thoracic somite     132/103      --
L/W of genital
  complex              118/-       --
L/W of third
  abdominal somite
L/W of fourth          94/-        --
  abdominal somite
L/W of fifth           118/-       --
  abdominal somite
L/W of furcal rami     100/65     38/23

Length of longest       --         --
  furcal seta
L/W ofovisac            -/-        -/-

Species                   Hamaticolax embiotocae

Author                         Hanan (1976)

Measurements              Female             Male

Total length         938 [+ or -] 125   507 [+ or -] 59
Greatestwidth               --                --
L/W of cephalic            -/-                -/-
  somite
L/W of second              -/-                -/-
  thoracic somite
L/W of third               -/-                -/-
  thoracic somite
L/W of fourth              -/-                -/-
  thoracic somite          -/-                -/-
L/W of fifth
  thoracic somite          -/-                -/-
L/W of genital
  complex                  -/-                -/-
L/W of third
  abdominal somite
L/W of fourth              -/-                -/-
  abdominal somite
L/W of fifth               -/-                -/-
  abdominal somite
L/W of furcal rami         -/-                -/-

Length of longest           --                --
  furcal seta
L/W ofovisac               -/-                -/-

Table 2. Setation of legs (Le) of the Hamaticolax species (setae in
arabic, spine sin roman numerals).

Species                  Hamaticolaxprolixus          Hamaticola-
                             [Female]                  xprolixus
                                                        [Male]

Author                      Cressey (1969)             Ho (1972)

Host                    Pleuronichthycoenosus         Paralichthys
                              Giard, 1854             californicus
                                                       Ayres, 1859

Site                          gillcavity               gillcavity

Locality                  La Jolla, California         Anaheim Bay,
                                                        California

                                           Segments

Le            Rami       1         2          3         1         2
3

Le1        exopodite    I-0       I-1       III-5      I-0       I-1
70
           endopodite   0-1       0-1        0-5       0-1       0-1
50
Le2        exopodite    I-0       I-1        IV-5      I-0       I-1
I-III-5
           endopodite   0-1       0-2        II-3      0-1       0-2
II-3
Le3        exopodite    I-0       I-1       III-5      I-0       I-1
III-4I
           endopodite   0-1       0-2        II-2      0-1       0-2
II-2
Le4        exopodite    I-0       I-1       III-5      I-0       0-1
III-4I
           endopodite   0-1       0-1       I-1-I      0-1      I-1-I
I-1-I0
Le5            --       0-1       * -1        --       0-1       0-2
Le6            --       Three long setae at area of       Lacking
                        ovisac attachment

Species                 Hamaticolaxprolixus       Hamaticolax
                              [Male]          scutigerulus [female]

Author                       Ho (1972)          Vervoort (1969)

Host                       Paralichthys       Pseudupeneus maculatus
                           californicus
                            Ayres, 1859

Site                        gillcavity                gill

Locality                   Anaheim Bay,       Piscadera Bay, Curacao
                            California

                                          Segments

Le            Rami               3              1      2      3

Le1        exopodite          II-I-4           V-6    --     --
           endopodite           I-5            0-1    0-1    50
Le2        exopodite          II-I-5           I-0    I-1   III-5
           endopodite          II-3            0-1    0-2   II-3
Le3        exopodite          II-I-5           I-0    I-1   III-5
           endopodite          II-2            0-1    0-2   II-2
Le4        exopodite          II-I-4           I-0    I-1   III-5
           endopodite           --             0-1    0-0   I-1-
Le5            --               --             0-1    0-3    --
Le6            --            Lacking          Three naked setae at
                                              area ovisac attachment

Species                    Hamaticolax        Hamaticolax galeichthyos
                        scutigerulus [male]          [female]

Author                  Vervoort (1969)       Luque and Bruno (1990)

Host                      Pseudupeneus          Galeichthys peruvianus
                            maculatus              Lutken, 1874

Site                          gill                      gill

Locality                 Piscadera Bay,        Cura Chorrillos, Peru
                             Curacao

                                        Segments

Le            Rami       1      2       3      1        2         3

Le1        exopodite    I-0    I-1    III-3   I-0     III-6      -I
           endopodite   0-1    0-1     I-5    0-10     0-10     0-50
Le2        exopodite    I-0    I-1    III-4   I-0I     I-1I     IV-5
           endopodite   0-1    0-1    II-3    0-1      0-2I     II-3
Le3        exopodite    I-0    0-1    III-4   I-0I     I-1I     III-5
           endopodite   0-1    0-1    II-2    0-1      0-2I     II-2
Le4        exopodite    I-0    0-1    III-4   I-0I     I-1I     III-4
           endopodite   0-1   I-1-I    --     0-10     0-10     0-20
Le5            --       0-1    0-2     --     0-1    II-I-I-0    --
Le6            --             Lacking          Three long setae at
                                                 genital complex

Species                 Hamaticolax galeichthyos    Hamaticolax
                                [male]               embiotocae
                                                     [female]

Author                  Luque and Bruno (1990)     Hanan (1976)

Host                    leichthys peruvianus       Cymatogaster
                           Lutken, 1874              aggregata
                                                   Gibbons, 1854

Site                             gill                nasalcavity

Locality                    Chorrillos, Peru       Anaheim Bay and
                                                      Huntington
                                                       Harbor,
                                                     California
                                          Segments

Le            Rami         1         2       3        1      2

Le1        exopodite      I-0      III-6    --      I-0I   I-17
           endopodite     0-1       0-1     0-5     0-10   0-15
Le2        exopodite      I-0       I-1    III-6    I-0I   I-1I
           endopodite     0-1       0-2    II-3      0-1   0-2I
Le3        exopodite      I-0       I-1    III-5    I-0I   I-1I
           endopodite     0-1       0-2    II-2      0-1   0-2I
Le4        exopodite      I-0       0-1    III-4    I-0I   I-1I
           endopodite     0-1       0-2     --      0-10   0-1I
Le5            --         I-1       --      --       0-1  1-I-III
Le6            --                Lacking            Three long setae
                                                      at genital
                                                        complex

Species                    Hamaticolax       Hamaticolax embiotocae
                            embiotocae                [male]
                             [female]

Author                     Hanan (1976)            Hanan (1976)

Host                       Cymatogaster      Cymatogaster aggregata
                            aggregata             Gibbons, 1854
                          Gibbons, 1854

Site                       nasalcavity             nasalcavity

Locality                 Anaheim Bay and     Anaheim Bay and Huntington
                            Huntington           Harbor, California
                             Harbor,
                            California

                                            Segments

Le            Rami              3              1        2        3

Le1        exopodite            --           0-1 **    I-1     III-4
           endopodite          0-5            0-1      0-1     1-5 **
Le2        exopodite          III-6           I-0      I-1     III-5
           endopodite          II-3           0-1      0-2      II-3
Le3        exopodite          III-5           I-0      I-1     III-4
           endopodite          II-2           0-1      0-2      II-2
Le4        exopodite          III-4           I-0     I-II-3     --
           endopodite           --            0-1     I-1-I      --
Le5            --               --            0-1     I-1 **     --
Le6            --        Three long setae             Lacking
                        at genital complex

Species                  Hamaticolax         Hamaticolax
                           attenuat         unisagittatus
                           [female]            [female]

Author                  Vervoort (1969)      Tavares and
                                             Luque (2003)

Host                   Scorpaena plumieri
                          Bloch, 1789         Centropomus
                                            undecimalis Bloch,
                                                 1792

Site                    gill and inner            gill
                           operculum

Locality                  Montego Bay,       Angra dos Reis,
                            Jamaica              Rio de
                         Piscadera Bay,      Janeiro, Brazi
                            Curacao

                                      Segments

Le            Rami       1     2      3      1     2      3

Le1        exopodite    I-0   I-0   III-6   0-3   0-4    --
           endopodite   0-1   0-1    0-5    0-1   0-1    0-5
Le2        exopodite    I-0   I-1   IV-5    I-0   I-1   IV-3
           endopodite   0-1   0-2   II-3    0-1   0-1    0-5
Le3        exopodite    I-0   I-1   III-5   I-0   I-1   III-4
           endopodite   0-1   0-2   II-2    0-1   0-1    0-4
Le4        exopodite    I-0   I-1   III-5   I-0   I-1   III-3
           endopodite   0-1   0-1   I-1-I   0-1   0-1    0-3
Le5            --       0-1   I-3    --     0-1   0-4    --
Le6            --        Three setae at       Three pinnate
                           area ovisac        setae at area
                           attachment       ovisac attachment

Species               Hamaticolax maleus         Hamaticolax occultus
                          [female]                     [female]

Author                 Oldewage (1994)              Kabata (1971)

Host                 Malacocephalus laevis         Lyopsettaexilis,
                          Lowe, 1843             Jordan and Gilbert;
                                                   Hippoglossoides
                                                elassodon, Jordan and
                                                       Gilbert

Site                     Gill and inner         Gill and gill cavity
                            operculum

Locality                South Africa Coast      West Coast Vancouver
                                                Island; Chatham Sound
                                                 (Border of British
                                                Columbia and Alaska)

                                         Segments

Le            Rami       1       2        3       1      2      3
                                         --
Le1        exopodite    I-0    IV-6      0-5     V-6    --      -
           endopodite   0-1    0-10     IV-5     0-10   0-1    I-5
Le2        exopodite    I-0    I-1I     II-3     I-0    I-1   IV-5
           endopodite   0-1    0-2I     III-5    0-1    0-2   II-3
Le3        exopodite    I-0    I-1I     II-2     I-0    I-1   III-5
           endopodite   0-1    0-2I     II-5     0-1    0-2   II-2
Le4        exopodite    I-0    I-1I      V-3     I-0    I-1   III-5
           endopodite   0-1    0-1V      --      0-1    0-1   I-1-I
Le5            --       0-1   0-4 ***            0-1    III-1  --
Le6            --       Three naked setae        Three pinnate setae
                                                   at area genital
                                                      complex

Species                 Hamaticolax spinulus [female]

Author                         Cressey (1969)

Host                       Scorpaena guttata Giard,
                               Oxylebiuspictus
                          Sebastodesmystinus Jordan
                        and Gilbert 1880, Sebastodesse-
                           rranoides Eigenmann and
                                   Eigenmann.

Site                               Gill cavity

Locality                      La Jolla, California

                                   Segments

Le            Rami         1           2           3
                                     I-1 I
Le1        exopodite     I-0I        0-1 0        II-5
           endopodite     0-1        I-1 I        0-50
Le2        exopodite     I-0I        0-2 I        IV-5
           endopodite     0-1        I-1 I       II-30
Le3        exopodite     I-0I        0-2 I       III-5
           endopodite     0-1        I-1 I       II-20
Le4        exopodite     I-0I        0-1 I       III-5
           endopodite     0-1     I-1-I-1 ****   I-1-I0
Le5            --         0-1
Le6            --          Three setae at area ovisac
                                   attachment

Species                 Hamaticolax paralabracis
                                [female]

Author                   Luque and Bruno (1990)

Host                      Paralabraxhumeralis

Site                    Gill and inner operculum

Locality                    Chorrillos, Peru

                                Segments

Le            Rami        1         2       3

Le1        exopodite     I-0      III-6    --
           endopodite    0-1       0-1     0-5
Le2        exopodite     I-0       I-1    III-6
           endopodite    0-1       0-2    II-3
Le3        exopodite     I-0       I-1    II-6
           endopodite    0-1       0-2    II-2
Le4        exopodite     I-0       I-1    II-6
           endopodite    0-1       0-1     0-3
Le5            --                  0-1     --
Le6            --       Three pinnate setae at
                        area ovisac attachment

* According to Cressey (1969), three terminal spines or setae.

** Textual information of Hanan (1976) is different from his
schematic drawings.

*** According to schematic drawnings of Oldewage (1994).

**** According to schematic drawnings of Cressey (1969), setae
or spines.
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Author:Cardoso, Lucas; Martins, Mauricio Laterca; Lacerda, Ana Carolina Figueiredo; Cadorin, Douglas Ismael
Publication:Acta Scientiarum. Biological Sciences (UEM)
Date:Apr 1, 2017
Words:4442
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