Parasitic Copepoda of fishes from the collection of the Zoological Institute in Leningrad

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1 J. lint^. SOC. ZOO^.), 46, 309, pp math 176jgures Printed Britain October, 1966 Parasitic Copepoda of fishes from the collection of the Zoological Institute in Leningrad BY Z. ICABATA Marine Laboratory, Aberdan AND A. V. GUSEV Zoological Institute, Leningrad (Accepted for publicution February, 1966) Communicated by the Editorial Xecretary INTRODUCTION Despite considerable research during the last 50 years, the parasitic Copepoda of fishes are still relatively imperfectly known. This is particularly true of those species, whose economic importance has not appeared large enough to stimulate interest. It is not surprising, therefore, that many general conclusions about this group of parasites in the past have been little more than inspired guesses, which have had to bridge extensive gaps in our knowledge. The number of the known species of parasitic copepods is now in excess of 2500, but it is obvious to all, who are interested in this group, that this number is far from final and that numerous species are yet to be discovered and described. Many of those already known have been recorded from single finds and are but poorly known. Even some of the commoner species have suffered from inadequate descriptions. The descriptive work on parasitic Copepoda is, therefore, still needed. The collections of the Zoological Institute in Leningrad (Academy of Sciences of the U.S.S.R.) contain an interesting array of these parasites, to which oceanographical expeditions during the last 20 years have added more material gathered all over the world oceans. Most of this material has not been examined so far, although the work of Markevich (1956) and of Gusev (1951) included some information on the Institute s collections. Recent expeditions of the research ships Vityaz and Ob returned with copepods collrcted in the Antarctic and other localities from the southern hemisphere, the faunistic knowledge of which is still very scanty. It is the purpose of this paper to contribute to the expansion of this knowledge by reporting on some Copepoda belonging to the Institute s collection. The work reported on in this paper was done partly in Leningrad (including Figs 1 to 8, 12 to 14, 16 to 31 and 64 to 67) and partly in Aberdeen. All type specimens of the new species described here are preserved in the collections of the Zoological Institute, Leningrad. The examination of the entire and dissected specimens was conducted under magnifications up to x900. Berlese s fluid was used as clearing agent and mounting medium. No stains were employed, but phase contrast illumination was used throughout.

2 156 Z. KABAT AND A. V. GUSEV The species which are fairly well known and have been adequately described in the past are reported on only briefly, with few or without illustrations. All drawings are freehand, made with the aid of an eye-piece graticule. Some difficulty was experienced in deciding on a nomenclature for the mouth-parts, which would find general acceptance. It was finally decided to use terms applied by Lewis (1964) for the caligoid species and to retain the use of Heegaard (1947) designations for the chondracanthids and lernaeopodids. This apparent vacillation should be attributed to the authors' indecision as to the homology of the copepod mouth-parts; the indecision which reflects t,he disagreement, now current in this field. List of copepods and their hosts (The order in which the families are listed and in which the species are arranged within the families is random and implies no relationships.) Family Caligidae Dana, Caligus quadratus Shiino, Caligus coryphaenae Stp. & Ltk., , Caligus sensilis sp. nov. 4. Sciaenophilus benedeni (Bassett-Smith, 1898) Family Euryphoridae Yamaguti, Euryphorus nymph Stp. & Ltk., Dysgamus sagamiensis Shiino, 1958 Family Pandaridae (Wright, 1877) 7. Pandarus satyrus Dana, Nessipus sp. male Family Anthosomatidae Yamaguti, Lernanthropus gisleri Van Beneden, 1852 Family Eudactylinidae Yamaguti, Kroeyeria carchariaeglauci Hesse, 1879 Family Lernaeoceridae Gurney, Sarcotretes eristaliformis Brian, 1908 Family Chondracanthidae Dana, Chondracanthodes tuberofurcatus sp. nov. 13. Chondracanthopsis dogieli (Gusev, 1951) 14. Acanthochondria solida Gusev, Parapharodes semilunaris sp. nov. 16. Lateracanthus qdripedis gen. et sp. nov. Family Lernaeopodidae Olsson, Brachiella chevreuxii Van Beneden, Brachiella annulata Marlievich, Eubrachiella gaini (Quidor, 1913) 20. Eubrachiella gaini dorsituberculata subsp. nov. 21. Dendrapta cameroni longiclavata subsp. nov. 22. Lernaeopodina paci$m sp. nov. 23. Clavella adunca (Strclm, 1762) Coryphaena hippurus Spheroides inermis Spheroides inermis Johnius hololepidotus Coryphaena hippurus Spheroides inermis Pterolamiops longimanus Prionace glauca Pterolamiops longimanus Johnius hololepidotus Eulamia sp. Bathylagus sp. Macrurus whitsoni Myoxocephlus batrachoides Myoxocephalus brandti Pleurogrammus asonus Lycodes diapterus beringi Macrurus acro1ep.s Johnius hololepidotus Macrurus acrolepis Trematomus scotti Chaenodraco cathleenae Chaenicht h ys rhinoceratus Raja kenojei Raja smirnovi Raja sp. Raja kenojei Macrurus whitsoni Trematomus loennbergi

3 Parasitic Copepoda of $shes f rm Leningrad 157 DESCRIPTION OF SPECIES Family CALIGIDAE Caligus quadratus Shiino, 1954 (Figs 3, 4) Syn. : C. productus Wilson, 1905 C. monacanthi Wilson, 1937 C. coryphaenae Yamaguti, 1936 Record of specimens: Two adult females and several males were collected in western Pacific, east of the Ryukyu Islands, in the vicinity of Tanagashima, by Vityaz on 31.x Host : Coryphaena hippurus. Habitat: Branchial cavity. Previous records: The parasite was first described by Shiino (1954) from Japanese waters, where it was found on the inner surface of the operculum of Parathunnus macropterus. The same author described it subsequently from Coryphaena hippurus, Histiophorus orientalis and Rhinobatus schlegeli in the same area. The male of the species is known from Monacanthus sp. in the West Indies (Wilson, 1937). For discussion of synonymy, see Shiino (1959). Shiino s descriptions, particularly that of 1959, are quite detailed and require no amplification. For comparison, some dimensions of the specimens (in mm) from the present collection are presented below together with those of Shiino s material. Present material Shiino (1959) Total length Carapace length Carapace width Free thorax length Free thorax width Genital segment length Genital segment width Abdomen length First segment length Second segment length Abdomen width First segment width Second segment width Female Male Female Male Shiino s specimens differed from the present collection in that his female had a relatively larger carapace and genital segment. Shiino s male, while smaller than the males here described, differed from them in having, like the female, a relatively larger carapace. The relative dimensions of the posterior part of the body also differed between the two groups of males. These differences, however, were not such as to force one to consider a possibility of specific differences.

4 158 Z. KABAT AND A. V. GUSEV

5 Parasitic Copepoda of $shes from Leningrad 159 Caligus coryphaenae Stp. & Ltk., 1861 Syn. : C. bengoensis Scott, 1894 C. aliuncus Wilson, 1905 C. elongatus Heegaard, 1943 C. tesserifer Shiino, 1952 C. alveolaris Heegaard, 1962 Record of specimens: More than 20 female and male specimens were collected by Vityaz in the northwestern part of the Pacific (east of Volcano Island), on 11.x Host : Spheroides inermis. Habitat : Skin. This species was found together with Caligus sensilis and Dysganaus sagarniensis. Previous records: C. coryphuenae is well represented in the literature. It has been recorded from many localities in the Atlantic, the Pacific and the Indian Oceans, as well as from the Mediterranean and the Adriatic. The range of its hosts is also wide. It has been collected from the following fishes : Coryphaena hippurus (Coryphaenidae); Parathunnus macropterus, P. obesus, Euthynnus sp., E. pelarnis, E. lineatus, E. alleteratus, E. vagans, E. yaito, Xardcr sp., Thunnus thynnus (Scombridae); Seriola dorsalis, Caranx hippos (Carangidae) and Polydactylus opercularis (Polynemidae). Shiino (1957) found it on an elasmobranch host, Isurus glaucus (Lamnidae), while Scott (1894) and several subsequent workers recorded its occurrence on another elasmobranch, Squalus fernandinus (Squalidae) The morphology of this species has been adequately described by several authors. Among recent workers, Shiino (1959a) and Pillai (1962/1963) discussed at length its synonymy and its distinctive features. C. coryphaenae differs from many of its congeners by the presence of a transverse suture line dividing the eucephalic from the postcephalic area of the carapace and in the absence of the post-antennary processes (which are considered to be present, though minute, by some workers). C. coryphaenae is widely distributed throughout the warm seas, where it associates with the fast-swimming and active, pelagic hosts, the most common of which is Coryphaena hippurus. However, as suggested by the list above, it is capable of surviving also on hosts of quite different types, such as Spheroides inerrnis, a member of the slow-moving and cumbersome Tetraodontidae, or even on the elasmobranchs. Abbreviations used in figures 1 a-anus abd-abdomen antl-first antenna ant2-second antenna aux-auxiliary spine b-bulla cl-caudal laminae do-duct orifice dbdorsal tubercle en-endopod es-egg-sac ex-exopod gp-genital process gpl-genital plate 16-leg 6 m-membrane mdb-mandible ms-mandibular sclerite mx-maxilla p-papilla pc-pectinate process pop-postoral process pp-posterior process s-seta SB-fifth segment SG-sixth segment Figs 1 to 4. Brachiella annulata and Caligus quadratus Fig. 1. B. annulata, female, lateral. 2. B. annulata, ventral. 3. C. quadratus, female, dorsal. 4. C. quadratus, male, dorsal.

6 160 Z. KABAT AND A. V. GIJSEV 6 8

7 Parasitic Copepoda of fishes from Leningrad Some dimensions of the specimens (in mm) are given below: Total length Carapace length Carapace width Free thorax length Free thorax width Genital segment length Genital segment width First abdominal segment length First abdominal segment width Second abdominal segment length Second abdominal segment width Female *10-1* * Male Caligus sensilis sp. nov. (Figs 5 to 15) Record of specimens: Six mature and seven immature females and four males were collected by Vityaz in the northwestern part of the Pacific, east of the Volcano Island, on 11.x Host : Spheroides inermis. Habitat: Skin. This species was found together with C. coryphuenae and Dysgamus sagamiensis. C. sensilis belongs to a small group of species, which includes also C. balistae Stp. & Ltk., 1861, C. biaculeatus Brian, 1914, and C. polycanthi Gnanamuthu, All these species have in common a short abdomen and the shape of the genital segment of the female, The genital segment of the male in this group has its posterior margin bisected on each side of the abdomen, although the exact shape of the margin differs from one species to another. One common feature of all these species are the spur-like projections on the ventral surface of the carapace, associated with the base of the maxilla (Fig. 11) in the same way as the sternal furca is with the base of the maxilliped. Another common feature is the presence of two chitinous knobs (Fig. 15) on the ventral surface between the sternal bars of the first and second pairs of thoracic legs. A small orifice can be seen in the centre of each knob. The species most closely related to C. sensilis is C. polycanthi. The diagnosis of C. sensilis, therefore, is very much like that of C. polycanthi, with the following exceptions: The firh leg of the female C. sensilis (Figs 8,9) consists of two groups of setae, one of which contains three (two in C. polycanthi) and the other a single seta. The fifth and the sixth legs of the male C. sensilis (Figs 13,14) are at the apices of processes, which differ in shape from those in C. polycanthi. In the present species, the fifth leg surmounts a shorter, while in C. polycanthi a longer process. The caudal laminae in C. polycanthi, in addition to the armature of their posterior margins, bear dorsally and half-way between the base and the tip, two setae, absent from C. sensilis. The most important discriminating feature, which distinguishes C. sensilis from all the other closely related species of the genus Caligus, is the presence of minute papillae, each of which bears from one to three sensitive hairs. These papillae are particularly evident along the margins of the carapace and the abdominal segment (Fig. 9). They are also Figs 5 to 9. Cnligus sensilis Fig. 5. Young female, dorsal. 6. Adult female, dorsal. 7. Chalimus stage, dorsal. 8. Part of abdomen and posterior corner of genital segment, female, ventral. 9. Posterior corner of genital segment, more highly enlarged, ventral.

8 162 Z. KABAT AND A. V. GUSEV rnm 12 ' 25P ' Figs 10 to 15. Culigus sensilis Fig. 10. Part of carapace margin (marginal membrane omitted), dorsal. 11. Anterior sternal spine and its position in relation to postoral process, and base of maxilla, ventral. 12. Male, dorsal. 13. Male, part of abdomen and posterior corner of genital segment, ventral. 14. Male, posterior corner of genital segment, more highly enlarged, ventral. 15. Sternal knob, ventral. (For explanation of labelling see p. 159.) 15

9 Parasitic Copepoda of $shes from Leningrad 163 present on the dorsal surface of the animal, where they are distributed in a symmetrical pattern; some of them are indicated by black dots in Fig. 5. This figure misses out some of the papillae on the posterior part of the lateral zones of the carapace and some on the posterior part of the genital segment. The details of the papillae are shown in Fig. 10, from which the marginal membrane has been omitted for clarity. Figure 10 shows a marginal papilla which is particularly prominent and shown also in Fig. 5. Anteriorly to this prominent papilla the margin of the carapace bears from 15 to 20 smaller papillae, while posteriorly another 15 to 18 papillae are present. Each lateral margin of the genital segment is provided with about 12 smaller, though similar, papillae. The dimensions of the specimens (in mm) are shown below : Total length Carapace length Carapace width Free thorax length Free thorax width Genital segment length Genital segment width Abdomen length Abdomen width Egg-sac length Egg-sac width Female Male * L It is interesting to note that the workers, who have SO far dealt with the species of Caligus belonging to this group, paid no attention to the presence of the ventral spurs (Fig. 11). These spurs, when examined, bear close resemblance to the structure of the sternal furca. Their position in relation to the nearest segmental limb (maxilla) also resembles that of the sternal furca and the maxilliped. It is quite possible that they are also of similar origin. Should this origin prove to be the intercostal plate (Lang, 1951), a similarity would be established between the maxillae and the maxillipeds, adding weight to the views of those who believe that these limbs should be regarded as the first and the second maxilliped respectively. The matter, however, calls for further critical examination. Sciaenophilus benedeni (Bassett-Smith, 1898) Syn. : Caligus (Sciaenophilus) benedeni Bassett-Smith, 1898 Sciaenophilus benedeni Bere, 1936 Caligus benedeni Capart, 1941 ; 1959 Caligus benedeni Nunes-Ruivo, 1962 Xciaenophilus benedeni Yamaguti, 1963 Records of specimens : Six mature females, three ovigerous and three with remnants of discharged egg-sacs, were collected from the Atlantic coast of Africa in Host : Johnius hololepidotus. Habitat : Gills. Taken together with Lernanthropus gisleri and Brachiella chevreuxii. Previous records : This parasite was first recorded by Bassett-Smith (1898) from Sciaena diacanthus in Bombay. All subsequent records came from the Atlantic. Capart (1941, 1959) recorded it from Johnius hololepidotus (Mauretania) and from Umbrina valida, Sciaena angolensis and S. aquila (Angola). Nunes-Ruivo (1962) found it on Urnbrim ronchw in the latter area. On the American side of the Atlantic, Bere (1936) found it on the gills of Larimus fasciatus off Florida.

10 164 Z. KAEATAND A. V. GUSEV The dimensions (in mm.) of the specimens in this collection are well within the range of sizes given by earlier authors. They are shown below: Total length Carapace length Carapace width Genital segment length Genital segment width Abdomen length Egg-sac length Egg-sac width The definite identity of this species must remain in doubt until its relationship with S. tenuis (Van Beneden, 1852) can be carefully investigated. The problem is beset with difficulties in view of the inadequate original descriptions and the loss of the type material. Family EURYPHORIDAE Euryphorus nympha Stp. & Ltk., 1861 (Figs 16 to 24,27,28) Syn. : E. coryphaenae Kr~yer, 1863 E. nordmanni Kirtisinghe, 1937 Record of specimens : Numerous specimens, both male and female, adult and juvenile, were collected east of the Ryukyu Island, in the vicinity of Tanagashima Island, by Vityaz, on 31.x Host: Coryphaena hippurus. Taken together with Caligus quadratus. Previous records : Originally recorded from the West Indies, where it has been found on two host species : Coryphuena hippurus and C. epuisetis (the latter originally referred to as Lampugus punctulatus), this parasite is now known from almost all the warm seas of the world. It has been found on C. hippurus along the Atlantic coast of America (Causey, 1953, 1953~) 1955), from the Pacific (Yamaguti, 1936a; Shiino, 1954) and from the Indian Ocean (Kirtisinghe, 1937). Shiino (1954a, 1959) found it also on two other hosts, Parathunnus macropterus and Thunnus alalunga, off Japan and in the Indian Ocean respectively. The morphology of this species has been described in detail by the earlier authors. It is of interest, however, to note the extensive changes which this parasite undergoes in the course of its progress towards maturity. In particular, the development of the dorsal plates, the lateral alae of the genital segment in the female and the lateral alae of the first abdominal segment in both sexes, progresses with age and causes well-marked differences in the general appearance of the parasite. These differences are shown in Figs 16 to 22. Figure 16 shows the posterior part of the young male, Fig. 17 an older male with larger abdominal alae, Fig. 18 an adult female, with fully developed lateral structures, while Figs 19 to 22 show their gradual expansion. Among other changes with age is that of the shape of the sternal furca, which alters quite markedly in the female during the course of development. An interesting fact brought to attention by the study of the juvenile stages (both sexes) of E. nymph, is their great resemblance to the males of the genus Dysgamus. When fkst examined in this collection, the juveniles of E. nymph were mistaken for Dysgamus. The overall similarity of these two forms can be seen by comparing Fig. 23 with Fig. 25. The clues to the generic identity can be found only in the details of the appendages. E. nympha, which at this stage is still devoid of the typical dorsal plates (Fig. 23), can be distinguished mainly by the presence of an appendage referred to as secondary furca and associated with the sternal bar of the first pair of legs (Fig. 24). This is absent from Dysgamus. Some

11 Parasitic Copepoda of $shes from Leningrad 165 Figs 16 to 22. Euryphoms nympha Fig. 16. Posterior part of developing male, dorsal. 17. Adult male, dorsal. 18. Adult female, dorsal. 19 to 22. Dorsal views of posterior part of female at increasingly later stages of development.

12 166 Z. KABAT AND A. V. GUSEV 26 2QQp, Figs 23 to 28. Euryphorus nympha and Dysgamus sagamiensis Fig. 23. E. nympha, young male, dorsal. 24. E. nympha, first leg and secondary furca, ventral. 25. D. sagamiensis, male, dorsal. 26. D. sagamiensis, sternal furca, ventral. 21. E, nympha, young male, maxilliped, lateral. 28. E. nympha, sternal furca, ventral. (For explanation of labelling see p. 159.)

13 Parasitic Copepoda of fishes from Leiiingrad 167 differences are also present in the sternal furca, which in Euryphorus is associated with a flap of cuticle (m, Fig. 28), not observed in Dysgamus. Pectinate processes ofthe maxilliped (pc, Fig. 27) are another distinguishing feature of Euryphorus. The first to describe a juvenile E. nympha was Shiino (1959). He is now aware (pers. commun.) of the resemblance between this form and the genus Dysgamus and is of the opinion that further work might prove more alleged males of Dysgamus to be members of Euryphorus or other genera. Some dimensions of specimens in this collection (in mm) are shown below: Total length Carapace length Carapace width Free thorax length Free thorax width Dorsal plates length Dorsal plates width Genital segment length Genital segment width Genital alae length Genital alae width First abdominal segment length First abdominal segment width Abdominal alae length Abdominal alae width Second abdominal segment length Second abdominal segment width Juvenile Male Female ;!*1O ' ' ' '1 8 1 '05-1' ' ' Dysgamus sagamiensis Shiino, 1958 (Pigs 25, 26) Record of specimens : Ten males were collected by 'Vityaz' in the northwestern Pacific, east of the Volcano Island, on ll.x Host: Spheroides inermis. Habitat : Skin. Taken together with Caligus sensilis and C. coryphaenae. Previous records: Shiino (1958),found two males on the skin of Caranx uraspsis in Sagami Bay, Japan, on 24.viii The comparison of the specimens taken by 'Vityaz', with the original description shows no great differences between the two lots of specimens. These parasites resemble one another very closely, both in their general appearance and in the details of their appendages. In particular, they have in common the typical shape of the sternal furca (Fig. 26), with its relatively short tines and a square base. The comparison of the dimensions (in mni) of the present material with Shiino's type specimen is given below: Total length Carapace length Carapace width Free thorax length Free thorax width Genital segment length Genital segment width 11 Present material Shiino (1958) '

14 168 Z. KABATAND A. V. Gus~v Present material Shino (1958) First abdominal segment length First abdominal segment width Second abdominal segment length Second abdominal segment width The only discrepancy between Shiino s and the present material is found in the pattern of grooves marking the divisions between the zones of the carapace. Shiino s description shows that the transverse groove does not meet in the centre, but ends on each side in a fork, soon after diverging from the longitudinal grooves inwards. As can be seen in Fig. 25, the present authors have found that each branch of the fork proceeds towards the centre to meet its opposite number and to enclose a small field in the centre, just posterior to the eyes. Shiino (1958) believes the carapace groove pattern to be one of the most important diagnostic features within the genus Dysgamus. This is not the view of the present authors. It is true that the six species of the genus so far described differ in that pattern, but these differences are most probably only apparent and can be attributed to the inaccuracies of observation and to the lack of uniformity in the methods of illustration by various workers. The present authors found the grooves, particularly in the central zone of the carapace, very difficult to trace, especially in undissected specimens and in cleared mounts. A slight change in the depth of focus affects the appearance of the specimen. It is probable that careful re-examination of different species of Dysgamus would, in most instances, show that the differences either do not exist or are negligible. (With the possible exception of D. murrayi (Brady, 1883), in which the position of the eyes in relation to the grooves provides a clear distinguishing feature.) One has also to take into account the possibility that the visibility of the grooves varies with age. Family PANDARIDAE Pandarus satyrus Dana, 1852 Record of specimens: In all, the collection contains 15 females and four males, taken at two localities: (i) northwestern part of the Pacific, east of the northern part of Honshu, Japan, collected by Vityaz on 17.ix.1955; (ii) east of the Volcano Island, collected by Vityaz on 13.x Hosts : Pterolamiops longiinanus (locality (i)) and Prionace glauca (locality (ii)). Habitat: Skin. Previous records: This species has been widely recorded from many hosts and many localities. The host list comprises : Prionace glauca, lsurus glaucus, I. ditropis, Sphyma zygaena, Alopias pelagicus, Makaira mitsukurii, Carcharinus brachyurus, C. gangeticus and C. platyrhynchus, in addition to Pterolamiops longimanus, which is a new host record. Shiino (1960) mentions its occurrence on a black-tipped shark. The localities cover both the Atlantic and the Pacific. The parasite is known from the Gulf of Mexico and the Cape Verde Island in the former and from Hawaii, Japan, Formosa, the coast of Equador and Kermadec Island in the latter. The morphology of this well-known species has been often and well described. Some dimensions (in mm) of the specimens in the present collection are given below : Female Total length Carapace length Carapace width First free thoracic segment, lateral plates length

15 Parasitic Copepoda of fishes from Leningrad 169 Second free thoracic segment, visible part of dorsal plates Third free thoracic segment, visible part of dorsal plates Genital segment, visible part of dorsal plates Sixth segment, dorsal plate length Sixth segment, dorsal plate width Male Total length Carapace length Carapace width First free thoracic segment length Birst free thoracic segment width (without plates) Second free thoracic segment length Second free thoracic segment width Third free thoracic segment length Third free thoracic segment width Genital segment length Genital segment width First abdominal segment length First abdominal segment width Second abdominal segment lenpt 11 Second abdominal segment width * ~ Nessipus sp. male (Figs 29 t,o 35) Record of specimens: Three males were collected in the northwestern Pacific, east of the northern part of Honshu, Japan, by Vityaz, on 17.ix Host : Pterolamiops longimanus. Habitat: Skin. Taken together with Pandarus satyrus. The appearance of the specimens is typical of the males of the genus Nessipus (Fig. 29). They resemble to a considerable extent the males of N. alatus Wilson, 1905, and N. curticaudis (Dana, 1852), but differ from them both in a number of important details. Some of their appendages are shown in Figs 30 to 35. The dimensions of the specimens in the collection (in mm) are as follows : Total length Carapace length (midline) Carapace width First free segment length First free segment width Second free segment length Second free segment width Third free segment length Third free segment width Genital segment length Genital segment widt,h Abdomen length Abdomen width * An interesting feature is the presence of fine spinulation on the dorsal surface of this parasite. Figure 29 shows (black dots) the small spinules of different sizes, arranged symmetrically on the carapace and on the terga of the remaining thoracic segments. The spinules appear to decrease in number from the anterior to the posterior end of the animal.

16 170 Z. KABAT AND A. V. GUSEV Figs 29 to 35. Nessipus sp. male Fig. 29. Entire, dorsal. 30. Postoral process, ventral. 31. Maxilla, ventral. 32. First thoracic leg, ventral. 33. Second thoracic leg, ventral. 34. Third thoracic leg, ventral. 35. Fourth thoracic leg, ventral.

17 Parasitic Copepodn of fishes from Leningrad 171 The Pacific area is now known to be inhabited by several species of Nessipus. Five of these species (N. australis Heegaard, 1962; N. crypturus Heller. 1865; N. gonosaccus Heegaard, 1943; N. incisus Heegaard, 1962 and N. orientalis Heller, 1865) have no known males. In view of this fact and of the pronounced sexual dimorphism prevalent in the genus Nessipus, it is impossible at this stage to decide on the specific identity of the specimens described here. The final identification must await the discovery of the female. Family ANTHUSOMATIDBE Lernanthropus gisleri Van Beneden, 1852 Record of specimens: Three adult females mere collected off the coast of West Africa in Host : Johnius hololepidotus. Habitat : Gills. The parasites were found together with Xciaeiaophilus benecleni and Brachiella chevreuxii. Previous records: This species has been previously recorded from many parts of the Atlantic, the Mediterranean and from the coast of Belgium, where it was first discovered. Yamaguti (1936) recorded it in Japanese waters. The list of the known hosts is as follows: Sciaena sp., S. aquila, Uinbrina cirrhosa, CJ. steindachneri, U. valida, Johnius hololepidotus, Corvina nigra, C. cameronensis, Cynoscion nebulosus (Sciaenidae); Lichia amia (Trachinotidae) ; Centropomus undecimalis (Centroponiiclae); Polyclactylus quadrijilis (Polynemidae) and Paralichthys sp. (Bothidae). The above list shows that the parasite occurs predominantly on the representatives of the family Sciaenidae. It is of interest to note also that its association with the members of this family appears to be most intimate in the area along the west coast of Africa, where all but one of its known hosts belong to Sciaenidae. On the American side of the Atlantic they are about evenly divided between Sciaeiiidae and fishes of other families. It is from that side that Causey (1953, 1953a) recorded it on a flat-fish species, certainly a very atypical host for this parasite. Examining this species, the authors compared the specimens from the collection with some L. gisleri found on the gills of Sciaena aquila in Scottish waters. The comparison showed complete identity of both specimens. Their dimensions (in mm) are shown below. The length of the specimens has been measured from the anterior margin of the cephalothorax to the posterior margin of the dorsal plate, in mid-dorsal line. The length of the third and fourth leg has been measured from the base to the tip (though it must be remembered that the tips might be occasionally broken off). Specimens from Scotland Range Mean Length Cephalothorax length Cephalothorax width Third leg length Fourth leg length Egg-sac length Specimens from Africa Range Mean *05-1* The African specimens were much smaller than those found in the temperate waters. They also had shorter egg-sacs. In addition to their smaller size, the African specimens differed from their Scottish relatives in the size of their appendages. The ratio total length : third leg length was 2.9 : 1 for the Scottish and 3.8 : 1 for the African specimens. Corresponding ratios total length : fourth leg length were 1.5 : 1 and 1.6 : 1. The relative length

18 172 Z. KABATAND A. V. GUSEV of the third leg was clearly greater in the African parasite. It also had relatively longer cephalothorax, the ratio total length: cephalothorax length being 6.0 : 1 for the Scottish and 5.5 : 1 for the African specimens. These interesting differences illustrate the fact that the parasitic copepods of fishes, whose distribution range covers extensive geographical areas, differ in size from one area to another, those inhabiting the tropical and subtropical areas being often smaller than their relatives from the temperate latitudes. Family EUDACTYLINIDAE Kroeyeria carchariaeglauci Hesse, 1879 Sjm. : K. gracilis Wilson, 1922 Record of specimens: Three females and three males were found by the Ob expedition at Station 214 (Indian Ocean, northwest of the COCOS Island), on 3.v Host: Eulamia sp. Habitat; Gills. Previous records: The parasite was first described on Carcharinus glaucus, off the coast of northern France. It was subsequently redescribed from the same host in the Mediterranean (Delamare Deboutteville & Nunes-Ruivo, 1953) and from Prionace glaucu in the western Pacific (Shiino, 1957). Eulamia sp. is a new host record. Some dimensions of the specimens from this collection (in mm) are shown below : Male Female Total length Carapace length Carapace width Free thorax length Genital segment length Genital segment width Abdomen length (less furca) The study of the genus Kroeyeria has always been beset with numerous difficulties, because the descriptions of the species discovered by Hesse have not been accurate enough to permit comparison with later specimens. Of the 13 species recorded in the literature, four (K. acanthimvulgaris Hesse, 1879 ; K. aculeata (Gerstacker, 1854) ; K. galeivulgaris Hesse, 1879; K. scyllicaniculae Hesse, 1879) are not sufficiently known to be considered as definitely established. The description of K. trecai, Delamare Deboutteville & Nunes-Ruivo (1953), has never been published in full. Two attempts have been made to clarify the position. Wilson (1922) produced a key to the species then known. His key, however, was based on several rather variable characters, such as the relative dimensions of the genital segment or of the stylets. These tend to vary with age and are not reliable. Delamare & Nunes-Ruivo (1954) recommended the use of more permanent characters, such as the armature of the thoracic legs. Following their recommendations, the authors attempted to provide a key to the eight species of the genus, which can now be identified with some certainty. The key is given below: 1. No denticulation or papillae on the outer margins of endopod segments Papillae present on outer margins of endopod segments.. K. papillipes - Denticulations present on outer margins of endopod segments Denticulation on second and third endopod segments Denticulation on second endopod segments only.. K. echinata - Denticulation on all endopod segments of third leg and on first endopod segment of fourth leg only.. K.dispar - Denticulation on fist and third endopod segment of first leg only.. K. spatulata

19 Parasitic Copepoda of $shes from Leningrad Long seta absent from inner margin of second endopod segment of second leg. Terminal segment with six setae.. K. sublineata - Long seta present on inner margin of second endopod segment of second leg. Terminal segment with three setae.. K.lineata 4. Armament of caudal laminae consisting of two plumose and one naked terminal setae, as well as three plumose, subterminal setae.. K. carchariaeglauci - Armament of caudal laminae consisting of two plumose setae and one claw-like spine terminally, as well as one plumose seta and one claw-like spine subterrninally. K. sphyrnae Family LERNAEOCERIDAE Sarcotretes eristaliformis (Brian, 1908) (Figs 36 to 46, 105, 106) Syn. : Lernaeenicus eristalijormis Brian, 1908 Record of specimens: Two females wrre taken by the 'Ob' expedition near the shores of the Antarctic. One, a non-ovigerous female, was caught at station 223 (near Lutzow- Holm Bay, depth 3600 m.) on ; the other, a female with damaged egg-strings, was caught at station 169 (Davis Sea, depth 3500 m) on 18.i Ilost : Bathylagus sp. Habitat: Musculature of the dorsum. PrPvious records : The discoverer found this parasite attached to Bathypterois dubius, a very rare abyssal fish, at 39'18'05'' N 31'12' W (depth 1372 m) and on Gastrostonius bairdz, at 36'06'40'' N 10'18' W (depth 0 to 4740 m). Another specimen was taken by Brian near the latter station (depth 0 to StXO m) on an unknomn host. Later, Brian (1914) foundit againong. bairdi,at4l015'n56"25'w (depthoto4000m).theonlyother author to record this species was Leigh-Sharpe (1934), who found it on fiymenocephalus striatissimus in the Madura Sea. Although found on several occasions, this parasite has not been well described so far. Brian's descriptions deal adequately with the general appearance of the species, but leave out the details of the appendages. This is at least partly due to the fact that the appendages of X eristaliformis become very brittle with age (particularly the swimming legs) and are tlifficult to handle without inflicting damage. The general appearance of the younger specimen of this collection is shown in Fig. 36. The dimensions of both specimens (in imi) are given below: Younger Older Total length Length of trunk Width of trunk Width across the holclfast The first antennae of the specimens were damaged and could not be studied in detail. It has been observed, however, that their terminal setae were very long, almost as long as the entire appendage. The second antenna (Fig. 40) is of the usual lernaeocerid pattern and resembles somewhat that of Lernaeenicus in the length of its claw. The structure of the mouth-tube (Fig. 37) is similar to that of other Lernaecoeridae, but differs from Lernaeocera in the absence of the dorsal plaque (Kabata, 1962). The marginal striated membrane of the mouth-tube extends farther inwards, reaching the level of the second ring. The buccal stylets (Fig. 39) were found in their usual place. They were distinguished by their relatively large size and by the length of their terminal spine. The mandibles of the dissected specimen (Fig. 41) were both broken and their distal third was missing. The proximal two-thirds, however, resembled the corresponding part of other lernaeocerid mandibles both in their shape and in the position of their bases near the maxillae. It can be assumed, therefore, that the distal part of the mandible also resembles that of other

20 174 Z. KABAT AND A. V. GUSEV

21 Parasitic Copepodn of $shes from Leningrad 175 Lernaeoceridae and does not form, as suggested by earlier writers, a pointed, toothless spine. The maxilla (Fig. 41) is also typically Icrnaeocerid. No comments are needed to add to the illustration. The maxilliped (Figs 105, 106) consists of three segments, the basal one armed on the outer surface with a prominent outgrowth. The middle segment is rigidly fused with the distal one. At their junction, the middle segment bears a robust seta covered with irregularly scattered hairs. In the specimen examined, the seta was unfortunately broken and its full length could not be determined. The middle segment is armed also with a small tubercle tipped by a sharp spine and covered with very fine hairs (Fig. 106). Just posterior to it is a small patch of fine spinules on the inner margin of the segment. Thc terminal segment is hooked, strong and rimmed with a striated membrane. The parasite has three pairs of swimrning legs. The ovigerous female had legs consisting of lamelliform sympods only, with rami missing. The structure of the first and the second pair is illustrated in Figs 42 to 46. The t liird pair of legs was damaged and could not be studied. The two specimens examined were of different age and showed differences in appearance. In particular, the shape of the cephalothoracic lioldfast was affected, that of the younger specimen being softer and rounder in outline (Fig. 37), while that of the older was more heavily sclerotized and less regular in outlinc (Fig. 38). The present record, in accord with the earlier ones, suggests that S. eristaliformis is a deep-sea parasite and infests only the fishcs of the abyssal depths. It appears to be more catholic in the choice of the host species and its locality. The present record is a new host record and greatly extends the known distribution range of the species. Fa,mily CHONDRACANTHIDAE Chondracanthodes tuberofurcatus sp. nov. (Figs 47 to 63) Record of specimens: Two adult, ovigerous females, one with a male attached, were found by the Ob expedition at station 202 (off Kemp Land, Antarctica), on 2.ii Host : Macrurus whitsoni. Habitat : Branchial cavity. Description of the female The general appearance (Figs 47 to 49) of this species clearly places it in the genus Chondracanthodes. Its cephalothorax is inclined ventrally to the main axis of the body, is wider than long and postero-laterally extended into prominent conical processes (Fig. 48). The trunk is fairly clearly divided into three parts. The anterior part bears dorsally a large tubercle (Figs 47,49) and laterally two processes, which project ventro-laterally and divide distally into two branches (Figs 47,49). The middle and posterior parts are equipped with lateral processes only, those of the posterior part being larger than those of the middle part. Dorsally, these two parts bear only small swellings. Figs 36 to 46. Surcotretes eristaliformis Fig. 36. Female, entire, cephalothorax dorsal, trunk lateral. 37. Younger female, cephalothorax, ventral. 38. Older female, cephalothorax, dorsal. 39. Buccal stylet, lateral. 40. Second antenna, dorsal. 41. Mandible and maxilla (mandible broken at the tip), lateral. 42. Second leg, ventral. 43. First leg, exopod, ventral. 44. Second leg, exopod, ventral. 45. First and second legs, endopod, ventral. 46. First and second legs, tip of serrated spine.

22 176 Z. KABATAND A. V. GUSEV m Figs 41 to 51. Chondrucunthodes tuberofurcatus Fig. 47. Female, dorsal. 48. Female, ventral. 49. Female, lateral. 50. Female, genital segment and abdomen, ventro.latera Male, lateral.

23 Parasitic Copepoda of fishes from Leningrad Figs 52 to 63. Chondracunthodes tuberofurcntus female Fig. 52. First antenna, ventral. 53. Second antenna, ventral. 54. Mandible, ventral. 55. Mandible and both maxillae, diagrammatic, ventral. 56. Second maxilla, ventral. 57. Second maxilla, terminal spines, ventral. 58. First maxilla, ventral. 59. Maxilliped, ventral. 60. Caudal laminae, dorso-lateral. 61. Thoracic leg, entire, ventral. 62. Thoracic leg, rami, ventral. 63. Thoracic leg, duct orifice, ventral. (For explanation of labelling see p. 159.)

24 178 Z. KABAT AND A. V. GUSEV Some dimensions of the two female specimens examined are given below (in mm) : Specimen 1 Specimen 2 Total length Width Egg-sac length Egg-sac width Egg (diameter) The eggs are small and numerous, arranged in about 60 rows, 22 of which are distributed along bhe circumference of the sac. There are some 80 eggs in each row, so that the approximate number of eggs in a single sac is about The appendages: The first antenna (Fig. 52) is relatively large and distinctly twosegmented. The proximal segment is robust and fleshy, with a small conical protrusion on its postero-lateral corner ; the distal is smaller and club-shaped. The second antenna (Fig. 53) is, on the other hand, relatively quite small. It is of the usual hooked shape. The mandible (Figs 54, 55) is also typical of a chondracanthid. Both its margins are provided with about 30 teeth (the exact number could not be ascertained). Its basal segment bears ventrally a small, paddle-like sclerite (ms, Fig. 55), which was first described for Ch. bulbosus Kabata, The first maxilla (Fig. 58), a controversial appendage, believed by many to be the mandibular palp, has a convex outer and concave inner margin. It carries distally a strong spine with another, much smaller one near its base. The second maxilla (Figs 56, 57) is a short and squat limb, consisting of one segment articulating distally with a claw. The claw (Fig. 57) curves slightly upwards and is provided along the inner and outer margins with rows of 10 teeth each (only the outer row can be seen in Fig. 57). There is an auxiliary spine (aux) near its base, with a sharp tip and a number of small denticles near it. The maxilliped (Fig. 59) is devoid of any especially characteristic features. The two pairs of thoracic legs (Figs 61 to 63) resemble those of Ch. bulbosus. Their external margins bulge outwards, while the two rami point inwards and backwards. Both rami have some finely denticulated surfaces (Fig. 62). The endopod has one more prominent terminal spinule, while the exopod has five such spinules. Near the base of the exopod is a flagellate seta, while just medial to the base of the endopod is a small tubercle (do, Figs 61, 63) with a duct opening at its tip. The genital segment (Fig. 50) is a small tubercular structure, with two fairly well delimited ventro-lateral swellings accommodating the openings of the oviducts and providing attachment for the egg-sacs (es). Distally, it bears the abdominal tubercle, ending with caudal laminae (cl). The latter (Fig. 60) are provided with prominent styli at their tips, with smaller denticles near the bases of the latter and with a longer seta on the dorsal aspect. Description of the male Since only one male specimen was present in the collection, it becomes the allotype of the species. It could not be examined in detail. Its general shape is shown in Fig. 51. The posterior part of the male is displaced towards its ventral surface and a prominent bulge is present on the postero-dorsal aspect of the cephalothorax. The posterior part of the body is indistinctly divided into four segments. The usual set of mouth-parts is present, as well as the thoracic legs. The total length of the male, measured in straight line from the anterior to the posterior part of the cephalothorax, is 0.64 mm.

25 Parasitic C opepoda oj$shes from Leningrad 179 Disczcssio?i Although obviously a member of Chondracanthodes, this parasite could not be placed in any of the known species, particularly in view of the presence of the lateral expansions of the cephalothoracic region, with its posteriorly projecting processes and in view of the shape of the first pair of the lateral processes of the trunk. None of the known species of the genus has the characteristic bifurcation at the end of these processes. An interesting feature of Ch. tuberofurcatus is the presence of the paddle-like sclerite associated with the mandible (Fig. 55). When this sclerite was first discovered (Kabata, 1965), it was suggested that its presence between the mandible and the controversial first maxilla precludes the latter from being a mandibular palp. The position of these two structures in relation to the mandible, as shown diagrammatically in Fig. 55, makes it clear, however, that this is not so. The presence of the sclerite obviously cannot affect the argument about the homology of the first maxilla in any way. Chondracanthopsis dogieli (Gusev, 1951) Syn. : Chondracanthus dogieli Gusev, 1951 Chondracanthopsis dogieli Markevich, 1956 Record of specimens: One gravid female with a male attached was collected by Vityaz in Kronotskiy Zaliv (Kamchatka) on 24.v Seven gravid females, all with males attached, were collected off Shikotan Island (Kuril Archipelago, east of Hokkaido) on 7.ix Hosts : Myoxocephalus batrachoides (one female) ; M. brandti (seven females). Habitat : Internal wall of operculum (M. batrachoides) and gills (M. brandti). Previous records: Gusev (1951) found this species for the first time on the gills of M. stelleri and M. decustriensis in the Sea of Japan; Markevich (1956) found it on unspecified Myoxocephalus in the Bering Sea. The present species, closely resembling Ch. irregularis (Fraser, 1920) has been described in detail and calls for no amplification of description here. It might be interesting, however, to compare the size of the specimens, collected from the two new hosts reported here, with that of the type of material. Dimensions (in mm) are given below : Myoxocephalus batrachoides Total length Cephalothorax length 1.89 Trunk width (anterior) 5.04 Trunk width (posterior) 5.35 Posterior processes length 1.26 Egg-sac length Egg-sac width Myoxocephalus brandti Range Mean Paratype The parasite found on M. batrachoides is larger than its relatives from other species of this host genus. In other respects, however, it is indistinguishable from them. Ch. dogieli appears to be specific to the genus Myoxocephalus and has now been reported from its three species and one subspecies. Acanthochondria solida Gusev, 1951 Record of specimens: Three ovigerous females, with males attached, were collected off Vladyvostok, on l.vi.1949.

26 180 Z. KABATAND A. V. GUSEV Host : Pleurogrammus asonus. Habitat: Gills. Previous records: Gusev (1951) found this species on the gills of Hexagrammus octogrammus; in a later report, Shiino (1959) mentioned its occurrence on the gill arches of H. hqocephalus. Both records came from the Sea of Japan. A. solida appears to be closely related to A. deltoidea (Fraser, 1920), which it resembles. The latter species was found on H. decagrammus taken in the waters of British Columbia. The inadequacies of Fraser s description make it rather difficult to make a thorough comparison between the species. One of the possible discriminants between them is the position of the mouth on the ventral surface surface of the cephalothorax. While in Gusev s species the mouth is situated near the posterior margin of that part of the body, with the maxillipeds often extending posteriorly from the buccal cavity and reaching beyond the level of the thoracic appendages, in Fraser s species it is much nearer the centre of the cephalothorax. The dimensions (in mm) of the specimens found off Vladyvostok are compared below with the figures produced by earlier writers: Present material Gusev (1951) Shiino (1959) Total length Cephalothorax length *80-1* Cephalothorax width Trunk width Egg-sac length Egg-sac width Not unexpectedly, some differences occur in the dimensions of parasites from different hosts. The specimens in the present collection are distinguished by greater width of the head. Since, however, they vary in the degree of contraction and since this might affect the curvature and the shape of the carapace, these dimensions must be treated with caution. Parapharodes semilunaris sp. nov. (Figs 64 to 87) Record of specimens: Six females and three males were found by Vityaz in Kronotskiy Zaliv (Kamchatka), on 29.v Host: Lycodes diapterus beringi. Habitat: Skin and fins. Description of the female In its general appearance (Figs 64, 65), the female of this species resembles that of the only other known species of this genus, P. sadoensis Shiino, Somewhat flattened dorso-ventrally, its cephalothorax is covered by a well-defined dorsal carapace. In all but one of the specimens examined, the cephalothorax was flexed ventrally (Fig. 65) to the axis of the body. The trunk is divided into anterior and posterior part by fairly welldefined transverse grooves. The anterior part of the trunk bears two pairs of clavate processes, the first pair being the larger of the two (Fig. 64). The processes project from the lateral margins of the body in ventro-lateral and slightly posterior direction. In one specimen only, a single process of the anterior pair stretched obliquely in the forward direction. The posterior part of the trunk expands postero-laterally in the form of two large and thick processes (Fig. 64), which impart to this part of the body a shape not unlike that of a horse-shoe. These processes are as long as, or longer than, the anterior

27 Parasitic Copepodn of fishes from Leningrad Figs 64 to 74. Parapharodes semilunaris female Fig. 64. Entire, ventral. 65. Entire, lateral. 66. Entire, diagram. 67. Second antenna, ventral. 68. Mandible, ventral. 69. First maxilla, dorsal. 70. Maxilliped, ventral. 71. First thoracic lag, ventro-lateral. 72. Second maxilla, ventral. 73. Genital segment and abdomen, ventral. 74. Caudal laminae, ventral. (For explanation of labelling see p. 159.)

28 182 Z. KABATAND A. V. GUSEV Figs 75 to 87. Parapharodes senailunaris male Fig. 75. Entire, lateral. 76. Posterior part of trunk, ventral. 77. First antenna, entire, ventral. 78. First antenna, tip, ventral. 79. Second antenna, entire, dorsal. 80. Second antenna, tip of exopod, medial. 81. Mandible, ventral. 82. First maxilla, ventral. 83. Second maxilla, ventral. 84. Maxilliped, ventral. 85. First thoracic leg, ventral. 86. Third thoracic leg, lateral. 87. Caudal laminae, dorsal. (For explanation of labelling see p. 159.)

29 Parasitic Copepodrr of fishes from Lcningrad 183 part of the body. In the apex of the horse-shoe lies the abdomen, divided into three subspherical swellings (Fig. 73). The minute caudal laminae (cl, Fig. 73) are found at the apex of the central swelling, slightly to its ventral side. The dimensions of the female specimens were rather difficult to measure, in view of their unusual shape. To give some idea of their size, three dimensions were selected, indicated by letters a, b and c in Fig. 66. They are shown below (in mm) : Sp. 1 Sp. 2 Sp. 3 Sp. 4 Sp. 5 Sp. 6 a 2.80" *92 b C Egg-sac length Egg-sac width *The unusually long distance a in specimen 1 \\as due to the fact that in this specimen the cephalo thorax was not flexed ventrally, but extended in parallel ~tith the long axis. The appendages are very small and difficult to study. Both the first and the second antennae were damaged in most specimens. No details of the first antenna were obtained. The second antenna (Fig. 67) is uniramous and, as in the other species of Parapharodes, expands distally into a hammer-like, transverse structure. The mandible (Fig. 68) does not differ in essence from the usual chondracanthid type, being in the form of a wide-based, narrow, tongue-like segment, with serrations along both margins. The serrations of the ventral side are larger than those of the dorsal side; they are between 20 and 25 in number and do not extend to the tip of the mandible. There are only about 15 dcnticles on the dorsal margin. The first maxilla (Fig. 69) is a flat segment of somewhat spatulate shape, with two small spinules in its lateral corner. The second maxilla (Fig. 72) is distinguished by the relatively small number of denticles on its terminal spine. There are only thee of them on the ventral side, near the tip and only one on the dorsal side. The auxiliary spine (aux) is slender and simple. The maxilliped (Fig. 70) is of the usual type and distinguished by the absence of armature on the middle and distal segments There are two pairs of thoracic legs, small and difficult to observe in detail. They appear to be all of the same structure, shoan in Fig. 71. Their tubercular base carries two subcylindrical rami, one of which is tipped by a fine spinule. Betu-een the rami, at the apex, the basal tubercle is prolonged into a small papilla. The caudal laminae (Fig. 74) are short and cylindrical. They are equipped with stylets which appear to have slight constrictions near their bases. The lateral margins of the laminae bear also two setae each, while a similar, though apparently shorter, seta is present at the base of each stylet. The tips of the stylets appear to be armed with two or three hairs, but it cannot be certain if these arc not simply frayed fibres of the damaged tips Description of the male The male was located attached to the ventral aspect of the genital segment of the female (Fig. 64). It is very small, measuring about 0.5 mm in a straight line drawn from the top of the cephalothorax to the posterior margin of the last segment. The correspondingly small appendages are even more difficult to study than those of the female. The general appearance (Fig. 75) closely resembles that of the male P. sadoensis. Seen in dorsal aspect, the cephalothorax is roughly circular in outline, while the vermiform posterior part of the body gradually tapers towards its extremity. The male is usually somewhat curled ventrally. The divisions between the segments of the posterior part of the body are fairly clear, particularly in cleared specimens. The exception is the first 12

30 184 Z. KABAT AND A. V. GUSEV segment, which is not distinctly divided from the cephalothorax. The abdomen forms a small segment (abd, Fig. 76), pushed into thc last thoracic segment and surrounded laterally by its two folds. The first, second and third free segments all carry thoracic legs. In the corresponding position of the fourth free segment is a minute hair, which is possibly also a vestige of a thoracic leg. On the ventral surface of the fifth segment are two nipplelike orifices (do, Fig. 76) and ducts can be seen leading from them in the anterior direction. The sixth segment carries yet another vestigial pair of thoracic legs (1 6, Fig. 76). The abdomen carries the usual set of caudal laminae. The appendages: The first antenna (Figs 77, 78) is quite clearly divided into three segments. Its total length is about 130 p. The basal segment is the longest, comprising about a half of the length of the limb. The second and the third segments are of about equal length. The outer margin of the basal segment carries six long and flexible setae. There are three similar setae on the middle segment and ten on the distal segment. The distribution of setae on that segment is illustrated in Fig. 78. The second antenna (Figs 79, 80) is biramous. Its endopod (Fig. 79) forms a strong hook, which appears to carry a small tubercle on its outer margin. (This might possibly be an artifact caused by damage.) The exopod is fairly short, not extending beyond the hook of the endopod distally and slightly tapering towards its apex. Its terminal armament is clearly illustrated in Fig. 80. The mandible (Fig. SO), while generally resembling that of the female, is very characteristic in the slenderness of its blade and in size and arrangement of its marginal denticulations. Denticles on both margins are nine, but the outer ones are much larger and more robust than the inner ones, which are not more than mere serrations. The outer ones do not extend to the tip of the limb, while the inner ones do so. The first maxilla (Fig. 82) is semicircular in shape, though there is some doubt about this, since it was not possible to examine it in sufficient detail. It carries a small papilla at its medial corner and two long setae in the lateral corner. The second maxilla (Fig. 83) differs from that of the female only in the absence of denticulations on its claw and in relatively larger auxiliary spine. The maxjlliped (Fig. 84) has a patch of fine denticles on the outer margin of the second segment, near its junction with the terminal claw. Otherwise it does not differ from that of the female. The first two pairs of thoracic legs (Fig. 85) are similar. They have one-segmented sympods, with rami, also one-segmented, at their distal ends. On the lateral margin of the sympod, near the base of the exopod, there is a long seta. The structure of the rami is clearly shown in the figure, though the true shape of the endopod might have been distorted by handling of the delicate specimen. The third leg (Fig. 86) is only a small tubercle with two setae of unequal length, while the only trace of the fourth is a minute hair, seen only in one specimen and not more than 5 p long. There is no fifth leg. The sixth (Fig. 76) is also a vestige, consisting of two small setae. The caudal laminae (Fig. 87) are very much like those of the female, armed with stylets and three setae each. The general resemblance of this species to P. sadoensis made it necessary to compare them very carefully. The fist and the obvious difference between these two species is the position of the fist two pairs of lateral processes. Shiino s specimens all had processes directed obliquely forwards, while the same processes in the present species were either extended laterally or even obliquely posteriorly. This difference, however, is insufficient for a new species to be established upon it. It is possible that the processes possess some degree of movement and can reflect in their positions the differences in the host and the point of attachment. The only possible difference between the appendages of the female of the two species is the armament of the maxillipeds, that of P. sadoensis having, in addition to a spinulated

31 Parasitic Copepoda ofjis1ws j rom Leningrad 185 patch, also a fairly prominent tubercle near the base of the claw. The illustrations of the thoracic legs of P. sadoensis certainly differ to a great extent from those of the present species, but neither was investigated sufficiently to make these differences definite. The same is true of other possible differences (abscnce of denticles on the claw of the second maxilla in P. sudoensis, the four segments of its maxilliped and the number of spines on the caudal laminae). It falls to the male to offer diagnostic fcatiires which definitely allow to distinguish between P. sadoensis and P. semilunuris ant1 to accord to the latter the status of an independent species. The differences bet\\ een the males are tabulated below : First antenna First maxilla Maxilliped First free segment Second free segment Third free segment Fourth free segment Fifth free segment Sixth free segment Parapharodes sadoensis Short, segmentation uncertain No papilla in medial corner Papilla at base of claw First thoracic leg Second thoracic leg Third thoracic leg No appendage Fifth thoracic leg No appendage Parapharodes semilunaris Long, 3-segmented Pa.pilla present in medial corner No papilla at base of claw First thoracic leg Second thoracic leg Third thoracic leg Fine hair (fourth leg?) Duct orifices Sixth thoracic leg The differences between the structures of the thoracic legs of the two species must be suspect, since they might be only apparent. The same applies to the differsnces in the armament of the caudal laminae. The distribution of the thoracic legs on the free segments four to six, however, is clearly sufficient to discriminate between the two species. Lateracanthus gen. nov FPrnale : Chondracanthidae. Cephalothorax with two long lateral processes, which make it many times wider than long. The segnicnts of the free thorax provided with one or tno pairs of similar processes. Thoracic legs either one or two pairs, situated on the bases of the processes. Trunk as in Acanthochondrza, tleroid of all but the two posterior processes. Abdomen with well-developed caudal laminae. Male : Cephalothorax flexed ventrally to the long axis of the body, as wide as long, with the usual set of appendages on its ventral slidace. Two pairs of thoracic legs. Posterior part of the body indistinctly segmented, long and tapering. Abdominal laminae well drveloped. Type species : L. quadripedis sp. nov. Lateracanthus quadripedis sp. nov. (Figs 88 to 104) K~cord of specimens: Two ovigerous females and two males were found by Vityaz in the Aleutian Deep, on 7.vi Host : Macrurus acrolepis. Habitut : Walls of branchial cavity. Descripfptiotz oj the female The general appearance (Figs 88,89). The botlg of the female is easily divisible into four parts. The anterior part is the cephalothorax. It is dorsally covered by a typical carapace with a mid-dorsal suture line, but laterally estcnded into long, posteriorly directed processes, wide at the base and gradually tapering towards the apex. (The upturned tips of these processes in Fig. 88 are an artifact cansed by the conditions of storage.) The second part, the free thorax hich forms the neck of the animal, consists of two segments. J2*

32 186 Z. KABAT AND A. V. GUSEV Figs 88 to 97. Lateruca tithus qucidripedis female Fig. 88. Entire, dorsal. 89. Entire, ventral. 90. Abdomen, dorsal. 91. Tip of first antenna, ventral. 92. Second antenna, ventral. 93. Mandible, ventral. 94. First maxilla, ventral. 95. Second maxilla, ventral. 96. Maxilliped, ventral. 97. Thoracic leg and base of lateral process. (For oxplanation of labelling see p. 159.)

33 Parasitic Copepotllr nf $sties from Leningrad Figs 98 to 106. Laterocanthus qutidripsdis male and Sorcotretes eristoliformis Wig. 98. L. quadripedis, lateral. 99. First and second antenna, anterior First antenna, distal segment, anterior First ma,xilla, ventral Second maxilla, ventral Maxilliped, ventro-lateral Thoracic leg, ventral S. eristcdiformis, maxilliped, lateral S. eristulijormis, tip, lateral. (For esplanation of labelling see p. 159.)

34 188 Z. KABAT AND A. V. GUSEV Each of them is provided with long, simple, ventro-lateral processes with conical ends (Fig. 89); they are wide at their bases and taper distally. It is on the somewhat swollen bases of these processes that the thoracic limbs are situated, a feature unique among Chondracanthidae. The two posterior parts of the body form a typical chondracanthid trunk, subdivided by a transverse constriction into two almost equal parts. The posterior margin of the second of these two parts is produced posteriorly into two processes, which might differ in length from one specimen to another, but which are of considerable length in all of them. Between these processes, the posterior margin is occupied by the genital segment and small abdomen. The appendages: The first antenna (Figs 89, 91) is well developed and consists of two segments, the basal one being longer and more robust than the distal one. Along the anterior margin of the latter there are several setae (exact number not determined). The apical armament consists of eight setae (Fig. 91), four slender and flagelliform and the other four slightly shorter and more robust. The second antenna (Fig. 92) forms prominent hooks, which reach almost half-may down the ventral surface of the cephalothorax. The mandible (Fig. 93) is of chondracanthid type, the distal segment being equipped with about 30 denticles on either margin. The first maxilla (Fig. 94) is rather narrow and nearly oval in shape, its distal end bearing two spines, one large and terminal, the other smaller and subterminal. The second maxilla (Fig. 95) has a terminal claw armed with a row of 11 teeth on the ventral aspect. The inner aspect is unarmed. The auxiliary spine (aux) is robust and tapers to a fine point. The maxilliped (Fig. 96) is distinguished by the presence of a spiny pad on the inner margin of the middle segment. In the middle of the pad a small swelling forms an unusual feature. Its presence, however, should be further investigated. Both thoracic legs (Fig. 97) are of the same structure. They have a base in the form of a papillose swelling, which is surrounded by two rami. The exopod is a flat, paddle-like segment and might be armed distally with some small denticles, which were not definitely observed. On the outer aspect of its base is a fine, hair-like setule. The endopod is much narrower, almost cylindrical but tapering distally. It appears to be unarmed. The abdomen (Fig. 90) is one-segmented. It has a pair of caudal laminae, each equipped with three setae and strong terminal stylet. Some dimensions of the two specimens examined are given below (in mm). Specimen 1 Specimen 2 Total length Cephalothoracic proccss length First thoracic process length 1 * Second thoracic process length Trunk width Egg-sac length Egg-sac width The egg-sacs are slender and slightly curved. The eggs are mainly irregularly oval and are of unequal size, varying in diameter from 110 to 200 p. They are arranged in 10 longitudinal rows, some 90 eggs in one row. Description of the male The general appearance (Fig. 98) has been described in the generic diagnosis. The size of the specimen which was dissected was 550 p, measured in the flexed posture, as shown in the figure.

35 Pcirusitic Copepotln offijislics from LoLiwyrad 189 The first antenna (Figs 99,100) is about 180 p long and composed of two segmcnts, not unlike those of the female. The distal segment (Fig. 100) is armed with a iiuniber of setae. The condition of the specimen did not allow an accurate study of the setation, but it could be seen that the margins of the segment caarried about five setae, while the apical armament consisted of eight setae, much inore slender and longer in relation to the size of the segment than in the female. The second antenna (Fig. 99) is still birariious, the main part of the limb being formed by the powerful, hooked endopod. The exopocl is present as a vestigial tubercle on the outer aspect of the limb s base, tipped ith three short setae. Tlie mandible is an exact replica of that of the female, though smaller and armed with only about 20 teeth on the margins of the distal segment. Tlie first maxilla (Fig. 101), ho~ever, differs from thc female appendage. Its terminal spine is relatively much longer, as is also its smaller subterininal spine. In addition, nearer the base of the limb, its inner margin bears a short tubercle, surmounted by a minute dent icle. The second maxilla (Fig. 102) has a terminal claw devoid of tecth and with a slender auxiliary spine, while the maxilliped does not differ from that of the female, except in size. There are two pairs of thoracic legs (Fig. 104). They are relatively larger than those of the female, though their structure is similar. Tlie base is larger and the rami more prominent. The laminate exopod is armed with five spinules, four apical and one at some distance from the apex along the outer margin. The spine at the base of the exopod is much more prominent and robust than in the female The cndopod is narrower and its armament consists only of one spinule. The condition of the specimen made it impossible to study in detail its caudal laminae, \I hicli arc cylindrical and bear at least tno sctsc each, in addition to the stylets. Comments The characteristic appearance of the fenialc of this genus distinguishes it from all known chondracanthid genera. The only possible rcsemblance between L. quadripedis and any preb iously lino\+ln chondracanthid is v\ it11 Yrotochondracanthus psettodis Kirtisinghe, This species, later transferred by its cliscoverer to the genus Chondracanthus (cf. Kirtisinghe, 1964), also has postero-lateral cephalothoracic processes and well-developed thoracic processes, but no other outgro\+ tlis. The differences between these two forms are, h o ever, ~ quite significant. Lateracanthus has cephalothoracic processes which are in the form of extensions of the cephalothorax, their bases not being delimited from the segment. The thoracic processes of Kirtisinghe s species are rather in the nature of trunk outgrowths, since they arise from the trunk posterior to the level of the last pair of thoracic legs. In Lateraca~ithus, on the other hand, these processes are not only at the level of the thoracic legs, but are closely associated with them, each process bearing one leg directly on its base. This feature is not known in other genera of Chondracanthidae. The male of Kirtisinghe s species can be distinguished from that of Lateracanthus in having only one pair of thoracic legs. The literature contains another record of a chondracanthicl very similar to the present species. It is Chondracanthus macrurus Brady, This parasite was taken by the Challenger expedition on Macrurus sp. at Kermadcc Island. Its description is very brief, but the drawings presented make it quite definite that the two animals are closely related. The original description makes no mention of the thoracic legs. It proved necessary to consult the type material of Brady s species. a task of some difficulty in view of the state of its preservation. It has been found that the structure of the thoracic legs of Ch. macrurus, as well as their position on the bases of the thoracic processes, is the same as in Lateracanthus. In fact, the only difference between these two animals was the number of the thoracic legs. While L. quadripedis has tu o pairs of thoracic legs (and processes), Brady s species has only one pair of each.

36 190 Z. KABAT AND A. V. GUSEV In view of the great similarity between these two species, the authors propose to transfer Chondracanthus macrurus to the newly established genus, to be known as Lateracanthus mc~u~us (Brady, 1883). Family LERNAEOPODIDAE Brachiella chevreuxii Van Beneden, 1891 Syn. : Clavella sciaenae Brian, 1906 Brachiella sciaenae Wilson, 1915 Brachiella macrura Wilson, 1920 Record cf specimens: Five ovigerous females \\ere collected along the Atlantic coast of Africa in Host : Johnius hololepidotus. Habitat : Gills. Found together with Sciaenophilus benedeni and Lernanthropus gisleri. Previous records : This parasite has been reported, under its various synonyms, on many previous occasions. The discoverer gives its host as un squale, which is probably an incorrect record, since no Brachiella has ever been found on an elasmobranch host. The host range of B. chevreuxii, as recorded in the literature reports, is as follows: Xciaena aquila, Larimus fasciatus, Pogonias cromis, Atractoscion equidens, Otolithes sp., 0. senegalensis, 0. brachygnathus, Umbrina valida, Johnius cameronensis (Sciaenidae) and Lutjanus griseus (Lutjanidae). Its main area of distribution is the southern Atlantic, particularly along the west coast of tropical Africa and to the north of that area (Mediterranean, Atlantic seaboard of Europe). It is also prescnt along the shores of eastern America (Gulf of Mexico, Texas coast). This fairly common parasitc has been adcyuately describcd in several previous publications, the most recent of which is that of Kabata (19G6). Speculations have been made on its possible synonymy with Heller s (1865) B. sciaenophila. The original description of Heller s species, however, shows that it is devoid of a long genital process and that, therefore, it cannot be identical with the present species. Its possible identity with Clavellopsis johni Yamaguti, 1939, can also be ruled out, since the latter is distinguished by a complete fusion of its arms, which are separate in Brachiella, except for their tips. Kabata (1988) considered the possibility of specific identity of B. chevreuxii with B. sciaenae. He found that these two species were almost identical, but that the specimens of B. sciaenae at his disposal had their arms, though separate, covered by a common sheath of cuticle. This prompted him to support the distiiictncss of the two species. Since that time, however, several new specimens of B. chevreuxii have been examined and it has become clear that some of them have their arms completely separate, while others have a single cuticular sheath covering both arms. There were also several instances of intermediate positions. It is clear now that the swelling of the cuticle, so common in preserved specimens of Lernaeopodidae, might obliterate the gap between the amis and form a common sheath, within which the two arms are clearly distinguishable from each other. Since this observation removes the only obstacle to the relegation of B. sciaenae to synonymy with B. chevreuxii, it should be duly noted. The five females found in the present collection were relatively small. Their trunk length varied from 1.92 to 2.88 mm., or about two-thirds of the usual length of their relatives from the temperate waters. Brachiella annulata Markevich, 1940 (Figs 1,2) Record of specimens: Three ovigerous females were collected by Vityaz in Tuscarora Deep, east of Japan, on 17.v Host: Macrurus acrolepis.

37 Purusitic Copcporlri.ffislt PS from Lrningrud 191 Habitat : Buccal cavity. Previous record: The only known recortl of this parasite was made by its discoverer, who described it from the skin of a rare, tleepwa fish, Ereunias grallator, taken at Rlisaki (Japan). This species has been fairly well descril)etl. It is easy to diagnose and calls for no comments on its morphology. The dimensions of tlic specimens in this collection are given below (in mm) : Cephalothorax length Cephalothorax width Trunk length Trunk width Arms length Posterior processes length Egg-sac length Egg-sac width Present material Markevich, i " ' Eubrachiella gaini (Quidor, 1913) (Pigs 107 to 120, 124 to 133) Syn.. Brachiella phi Quidor, 1913 Eubrachidlu pitri Wilson, 1915 Rccord of sperimens; (i) One sample of eight fcmales and two males and one of 16 females and two males mas taken by the 'Oh' expedition at station 185 (near Princess Elizabeth Lantl, Antarctica), on 27.i.1957 ; (11) the same expedition collected four fcwiales in the fiord of Bunger Oasis, Antarctica, on ll.i.1957; (iii) at station 204 (near Enderby Lantl, 'Intarctica), 'Ob' collected nine females and three males. Hosts: (i) Trematomus scotti; (ii) T. hn~mti; (111) Chaeizodmco cathleenae. IZabitat: (i) fins, skin and eye-lids; (ii) gills; (iii) buccal cavity. Pwvioz~s records: Quidor (1913) disco\ crm1 this species on tlie gills and in the branchial cavity of l'rematomus sp. at Port Lockroy, Antarctica. It has been later recorded (Kabata, 1965) from T. scotti at 66'46' S 75'00' E. The abundant material in this collection greatly increases our knon ledge of E. gaini which previously was known from only a feu specimens. Some comments on the morphology of tlic female and detailed description of the male, not previously described, are given below. Comments on the female The female of E. gnini n as described in (let ail by Kabata (1965). Careful re-examination revealed three points which require anientlment of that description. One tubercle has bcen lcft out in Figs 58 and 60 of that paper, as can be seen by comparing them nith Figs 125 and 126 of this paper. The figures illustrate the first antenna of the male, but apply equally to the female of the species. The second point concerns the second antenna of the female. The apparent double tubercle, carritd on the endopod (Fig. 57, Kabata, 1965) is, in fact, a single one, accompanied by a roliiist spine (Figs 127, 128). The original diagnosis of the genus Euhmchiella included the absence of posterior processes as a characteristic feature of tlic qeiius Kabata (1965) believed that the posterior processes are present in the species, in the Eorni of small tubercles, flanking the genital process on the posterior margin of the trinili The presence in this collection of specimens at various stages of development, makes it possible now to determine with some accuracy

38 192 Z. KABAT AND A. V. GUSEV, 25011, h 109 Figs 107 to 114. Eubrnchielln gaini Fig Young female, lateral Young female, posterior tip of trunk, dorsal. 109 Young female, caudal lamma, dorsal Feinale oldor than 107, cephalothorax lateral, trunk dorsal Female older than 107, posterior tip of trunlr Adult female, lateral Adult female, posterior tip of trunk Male, lateral (For explanation of labcllirig see )

39 Parusitic Copepodn of fishes from Lenixyrad 193 the nature of various structures on that margin. Figure 107 shows a juvenile female, with its trunk 0.83 mm long. The posterior margin of its trunk (Fig. 108) is equipped with wellrecognizable caudal laminae, resembling tubercles with four spines at their tips (Fig. 109). With the development of the parasite, the caudal laminae become gradually dwarfed. By the time the female's trunk reaches a length of 1.41 mm (Fig. 110), the lateral expansion of the body, due to the development of the reproductive system, begins to obscure the laminae (Fig. lll), which still appear to rctain some setae at their tips. By the time the trunk reaches a length of 2.28 mm. (Fig. 119). the caudal laminae have been reduced to small tubercles, hardly discernible and with no armament left (Fig. 113). Kabata's earlier interpretation of these structures as posterior processes must, therefore, be amended. (It is, of course, not impossible that at least sonie of the posterior processes are, in fact, modified caudal laminae.) With more than one host of E. gaini no\\ known, it is interesting to determine whether the diffcrences in host species and in the habitat of the parasite on the host affect the shape of the parasite. Some measurements for coniparison are given below (in mm) : Cephalothorax length Cephalothorax width Trunk length Trunk width Arms length Egg-sac length Egg-sac width Eggs diameter Trernatom 119 Tremutomus Chaenodraco scotti hansoni cathleenae ' a ' The above figures show that the length :width ratio of the trunk and the relative length of the cephalothorax (the latter difficult to measure due to the pronounced ventral flexure) differ widely in parasites both on the same and on different host species. The differences, however, can be attributed to environmental factors and find parallel in similar variety of shape occurring in Clavella adunca, studied in European waters (Nunes- Ruivo, 1957; Kabata, 1963). Description of the male The general appearance (Fig. 114) of this male outwardly resembles that of the male of E. antarctica. A waist-like construction is present between the cephalothorax and the trunk, the two parts being inclined to each other at the angle of about 90". The appendages are, in general, similar to those of the female, which have been previously described in detail by Kabata (1965), with some amendments mentioned above. The first and the second antenna are shown in Figs 124 to 128. It should be noted that the position at the tip of the first antenna, which in Lernaeopodidae of other genera is occupied by tubercle 2 (Fig. 126), in Eubrachiella houses two separate tubercles (2, Fig. 125). Unlike the female, the male has no small spinule on the penultimate segment of this limb (Fig. 124). The mandible (Fig. 129) differs from that of the female in having only two secondary teeth and a very small first primary one, but the general pattern of dentition is not dissimilar. The third spine on the first maxilla (Fig. 130), present in the female at the base of the dorsal papilla, was not observed in the male. The slender first maxilliped (Fig. 131) of the usual male lernaeopodid structure offers no special features, while the second maxilliped (Fig. 132) is distinguished by a small spinulated tubercle on the margin of the cavity housing the tip of the claw. There are no thoracic legs. The posterior tip of the trunk is equipped with caudal laminae (Fig. 133), each armed with three subterminal spines of different lengths and with prominent terminal stylet.

40 194 Z. KABAT AND A. V. GUSEV 115 I I mm 120 SP m 121 Imm 122, 1 mm, I23 c------(

41 Parasitic Copepodu of fishes from Leningrad Eubrachiella gaini dorsituberculata subsp. nov (Figs 121 to 123) Syn. : Eubrachiella gaini Kabata, 1965 (in part) Record of specimens: Two females were taken by the Ob expedition at Herd Island. on 2.iv Host : Chaenichthys rhinoceratus. aabitat : Buccal cavity. Previous report: Kabata (1965) recorded this parasite from the same host and habitat, collected at Murray Island (Kerguelen). This parasite can be distinguished from E. gaini by the shape of the posterior margin of the trunk, being otherwise identical. While the dorsal aspect of the posterior part of the trunk has in E. gaini a dimple-like, transversely elongated depression (Figs 115 to 118), in the new subspecies the same part of the trunk is characterized by the presence of a very prominent dorsal tubercle (dt, Fig. 122). The postero-lateral corners of the trunk are fairly distinctly subdivided to form two lobes (Figs 122, 123). These do not occur in E. gaini (Figs 116, 119). Finally, the genital process of the new subspecies is distinctly longer than in E. gaini (gp, Figs 122, 123). As far as the authors can ascertain, there are no other differences between the two subspecies. The dimensions of the specimens of the new subspecies in this collection fall within the range of those reported on by Kabata (1965). The differences between these two obviously very closely related forms, are sufficient to warrant the status of distinct subspecies. In particular, the presence of the dorsal tubercle, which is a distinct qualitative difference, distinguishes E. gaini dorsituberculata from its relatives on other host fishes. The decision of Kabata (1965) to place the specimens found on Chaenichthys rhinoceratus in E. guini, was influenced by the imperfect state of preservation of these specimens. The differences in the shape of the posterior margin were then attributed to inadequate preservation. It is interesting to note that the changes, reflecting the differences between the two subspecies, are further accentuated in E. sublobulata, described by Barnard (1955) from Congiopodus torvus of South Africa. It appears that the evolution of the genus progresses from the simple shape of the trunk towards the ever-increasing complexity. Seen in this light, one would place E. gaini at the beginning of the line, which through E. gaini dorsituberculata leads to E. sublobulata. The smooth margins of the trunk, through the development of a dorsal tubercle and subdivision of the postero-lateral corners, pass on to the presence of many tubercles, both on the posterior and the lateral margins of the trunk. E. sublobulata was found on a scorpaenid fish, rather specialized and systematically removed from the hosts of the two subspecies of E. gaini. Dendrapta cameroni longiclavata subsp. nov. (Figs 134 t.o 145) Record of specimens: (i) Four females aid one male were found by Vityaz at station 3259 (Kronotskiy Zaliv, Kamchatka), on 22.v.1955; Figs 116 to 133. Eubrachiella guini and E.g. dorsituberculata Figs 11 5 to E. gaini from Chaenodraco cnthleenae, posterior tip of trunk, dorsal, lateral and ventral respectively. 118 to 120. E. guini from Trematomus scotti, posterior tip of trunk, dorsal, lateral and ventral respectively. 121 to 123. E.g. dorduberculata, posterior tip of trunk, dorsal, lateral and ventral respectively. 124 to 133. E. goini, male, appendages First antenna, lateral First antenna, tip, ventral First antenna, diagram of apical armature Second antenna, lat,eral Second antenna, endopod, diagram of apical armature Mandible, tip, lateral Maxilla, lateral First maxilliped, lateral Second maxilliped, lateral Caudal lamina, ventral. (For explanation of labelling see p. 159.)

42 196 Z. KABAT AND A. V. GUSEV (ii) Three females were found by the vessel Toporok in Zaliv Terpenya (Sakhalin), on 26.iv Hosts: (i) Raja sp. ;(ii) Raja kenojei and R. smirnovi. Habitat: Skin. Previous records: The original species, D. cameroni, was discovered by Heller (1949) on the skin of Raja scabrata in the Baie de Chaleur region, on the Atlantic coast of Canada. It was again recorded from the same host and locality by Kabata (1964). Description of the female A detailed description of the female of D. cameroni was published by Kabata (1964). The new subspecies differs from it only in the proportions of the trunk and, significantly, in the length of its posterior processes (Fig. 134). The data below show the differences between some dimensions of the two forms (in mm) : D. cameroni D. cameroni longiclavata Trunk length Trunk width Genital process length These average measurements were obtained from seven adult females of D. cameroni and four of the new subspecies. They show that the trunk of D. cameroni is much wider than long and its posterior processes are relatively short. In contrast, the trunk of the new subspecies is only slightly wider than long and its posterior processes more than twice as long as the trunk. It should be added, however, that the specimens of the new subspecies show some transitional stages, their trunks being much more slender at an early age and becoming wider with the progress towards maturity. The length of the posterior processes also increases with age. This can be seen by comparing the dimensions of the trunk, as illustrated by the ratios below : D. cameroni D. cameroni D.c. longiclavata D.c. longiclavata (juvenile) (adult) (juvenile) (adult) Trunk length : Trunk width 1:l 1 : :0*6 1 : 1.1 Trunk length : Genital process length 1 : :0*6 1 : :2*4 The only other difference between the subspecies is the mode of branching of the attachment organ, which in the new subspecies is much more profuse and more finely divided than in the original C. cameroni. The comparison of the structure of the appendages shows no differences between the two subspecies. The re-examination of the material of D. cameroni showed that Kabata (1964) failed to observe a small patch of denticles at the base of the distal papillae of the maxilla (Fig. 140), as well as a possible presence of a third small spine on the exopod of this limb. Description of the male The general appearance of the male (Fig. 135) brings it close to the usual type for the so-called Charopinus-group of Lernaeopodidae. The total length of the specimen examined Figs 134 to 145. Dendrapta eameroni longiclavata Fig Female, lateral Male, lateral. 136 to 145. Male appendages First antenna, lateral First antenna, tip, ventro-lateral Second antenna, lateral Mandible, tip, lateral Maxilla, lateral First maxilliped, distal segment, lateral Second maxilliped, medial Mediative process, almost lateral Caudal lamina, ventral Caudal lamina, tip, lateral. (For explanation of labelling see p. 159.)

43

44 198 Z. KABAT AND A. V. GUSEV was 2.07 mm, more than a half of it constituting the cephalothorax, dorsally covered by a carapace, slightly inclined to the axis of the trunk and separated from it by a distinct waist. The prominent mouth-cone is laterally flanked by the two pairs of antennae and ventro-laterally by the first maxillae. Near to the border with the trunk, the ventral surface of the cephalothorax is occupied by the two pairs of the maxillipeds, the second pair being situated in front of the first. A bifid mediative process (Fig. 143) is found between the maxillipeds. The trunk is ovoid, not obviously segmented in preserved specimens. Posteriorly it ends in a pair of caudal laminae, flanking the anal orifice. The appendages: The first antenna (Figs 136, 137) is four-segmented. Its basal segment bears a small spine (111, Fig. 136) on its dorso-lateral aspect. In the corresponding place of the second segment two spines are found (I, 11, Fig. 136), a small spine I1 being present in addition to the usual lernaeopodid spine I. The third segment is also equipped with a small spine (IV, Fig. 136). The apical armament (Fig. 137) consists of three long spines and three tubercles, numbered in the figure in accordance with the practice adopted by Kabata in his earlier publications (1963, 1964). The second antenna (Fig. 138) has the same structure as that of the female and needs no comments. The only possible difference is the relatively larger endopod with a somewhat stronger terminal hook. The mandible (Fig. 139) is a narrow blade, about 200 p long, while its dentiferous marginisarmed witha set of teeth (formula: Hl,Zl, Hl,Zl, Hl,Zl, N5). The teeth of the N series decrease in size towards the base of the limb. The maxilla (Fig. 140) differs from that of the female in size only. The peculiar feature of the maxilla of this species is, as mentioned above, a patch of small denticles at the base of the terminal papillae. The first maxilliped (Fig. 141) is also of a typical shape. The characteristic feature of this limb is the presence of a group of small denticles near the base of the claw and two spinules of the claw itself. The second maxillipeds (Fig. 142) are linked together by means of a medial membrane (m). The unarmed claw of the second maxilliped closes into a depression in a small protuberance based on the ventral aspect of the limb. There is also another, more prominent swelling, on the dorsal aspect. It appears to house the orifice of a duct, which can be seen running through it in cleared specimens. The caudal laminae (Figs 144, 145) are indistinctly segmented and appear to consist of about three segments. Their complicated structure is illustrated in Fig It is interesting to note that even the terminal stylet appears to be armed with minute denticles. There are two pairs of thoracic legs, both in the shape of single small setae. The first pair (Fig. 135) is situated near the waist region of the male, the second at some distance behind the first. Taking into account the nature of differences between the two subspecies, one is inclined to think that their formation must have been caused by recent geographical isolation of their closely related hosts of the genus Raja. None of these differences is of truly qualitative character and the authors are not inclined to recognize them as sufficient grounds for placing the two forms in two independent species. Lernaeopodina pacifica sp. nov. (Figs 146 to 166) Record of specimens: Two samples were collected by the vessel Toporok in Zaliv Terpenya (Sakhalin), on 27.ix The first sample consisted of seven females, the second of 12 females and two males. Host : Raja kenojei. Habitat : Gills. The second sample was found together with a single female specimen of Dendrapta cameroni longiclavata.

45 Parmitic Copepocla of $shes from Leningrad 199 en C 156 Figs 146 to 156. Lernaeopodina pncijca female Fig Young female, lateral Posterior tip of trunk, dorsal Posterior tip of trunk, ventral Tip of posterior process First antenna, dorso-lateral First antenna, tip, ventro-lateral First antenna, diagram of apical armature Second antenna, medial Second antenna, endopod, medial Second antenna, endopod, diagram of apical armature Mandible, tip, lateral. (For explanation of labelling see 1'. 159.)

46 200 Z. KABAT AND A. V. GUSEV I59 Figs 157 to 166. Lernueopodina pacijca, male and female Fig Female maxilla, lateral, slight,ly dorsal Maxilliped, lateral Maxilliped, tip of claw, lateral Male, lateral Male, vestigial thoracic leg Mediative process, lateral Posterior extremity, ventral Mandible, tip, 1at)eral First maxilliped, lateral Second maxilliped, ventral. (For explanation of labelling see p. 159.)

47 Parasitic Copepodu of fishes from Leningrad 20 1 Description of the female The general appearance (Fig. 146): The short cephalothorax, covered by a dorsal carapace, is inclined ventrally to the long axis of the trunk. The trunk has anterior narrow and posterior wide parts, which together give it roughly pyriform shape. Just anterior to a shallow constriction, separating the cephalothorax from the trunk, arise very elongated first maxillipeds, which form the usual lernaeopodid arms. The arms are separate, except for the tips, and of more or less uniform diameter throughout. At the tips they expand into disc-like pads, which join together and partly enclose the bulla (b, Fig. 146). This organ of attachment is circular and flat, devoid of manubrium. In the centre of the posterior margin of the trunk a prominent tubercle provides attachment to the egg-sacs (es, Figs 147, 148). Dorsal to the egg-sacs arid on the lateral aspects of the anal orifice are the postcrior processes (pp, Figs 147, 148), short, cylindrical and ending in short, soft seta each (Fig. 149). The dimensions (in mm), based on measurements of five females, are given below: Trunk, entire length Trunk, anterior part length Trunk width : anterior part posterior part Arms length Posterior processes length Egg-sac length Egg-sac width The appendages: The first antenna (Figs 150 to 152) is four-segmented, with a rather inflated and robust basal segment. The second segment carries a single spine on its dorsomedial aspect. The authors were unable to ascertain whether the third segment also carried a spine in the corresponding position (?, Fig. 150). The apical armament of the limb consists of three spines and three shorter structures, which are numbered from one to six and whose position on the apex are indicated in Fig The character of each spine and their distribution are in agreement with the position in other Lernaeopodidae. The peculiar feature of the species is the shape of the structures 1 to 3, which usually are more tubercular and shorter. The second antenna (Figs 153 to 155) is of the usual type. Its endopod is particularly well developed and provided, in addition to the usual two spines, with fairly prominent and widely scattered spinules, which cover its anterior and dorsal margins. The endopod is two-segmented, with spinulated pads on the ventral aspects of both segments (Fig. 153). Its distal segment, with apical armament, is illustrated in Figs 154 to 155. The twosegmented sympod has another spinulated pad near the base of the endopod. The mandible (Fig. 156) has the formula H1, Z1, H1, Z1, H1, Z1, N4-5, the last tooth of the N series being overlapped by the ventral blade of the appendage. The authors did not observe the presence of the dark streaks on the mandible (Kabata, 1964). The second maxilla is illustrated in Fig. 157 and requires 110 comments. The second maxilliped (Figs 158, 159) is distinguished by the sparsity of the spinulation on its proximal segment (Fig. 158). The same is true of the claw, the inner margin of which (Fig. 159) has only some six or seven denticles near the base and has a small swelling on its inner margin. Description of the male The general appearance (Fig. 160) resembles that of the other known males of the genus. The male is short (0.8 mm), with extensive dorsal carapace covering about half of its

48 202 Z. KABAT AND A. V. Gus~v dorsum. The posterior part of the body is relatively short. The cephalic region and the posterior part are almost parallel to each other. The appendages: The antennae of the male are in general the same as those of the female and need no description. The mandible (Fig. 164) has the same formula as, and greatly resembles, the female appendage. The same is true of the second maxilla. The main difference in the structure of the appendages of the male and the female is noted in the maxillipeds. The first maxilliped (Fig. 165) is equipped with robust subchela, its tip closing into a shallow pit bordered by a sharp-tipped tubercle. It has the usual lateral spine about half-way between the base and the tip. The second maxilliped (Fig. 166) is linked with its opposite number by means of a membrane. Its claw appears to be unarmed and the pit which receives its end is associated with two spinulated tubercles. Near the tubercles, a small orifice can be seen, with a part of the duct leading from it towards the base of the limb. Between the maxillipeds is situated a bifid mediative process (Fig. 162), with a small depression at the base of one of its lobes. There are two pairs of thoracic legs, just posterior to the base of the first maxillipeds and close to each other. Each leg consists of a single seta (Fig. 161) with a slightly inflated base. The postero-lateral corners of the posterior extremity are provided with a pair of genital plates (gpl, Fig. 163). The anal orifice is flanked by caudal laminae, each equipped with four subterminal setae of subequal lengths, and with a short, robust terminal seta. Comments The first record of the occurrence of Lernaeopodina in the Pacific, this species greatly resembles L. longibrachia (Brian, 1912). In fact, the authors have found it very difficult to decide on the diagnosis of the new species. L. longibrachia has been found on only two occasions, a single specimen each time (one with abnormal arms). The available description is quite inadequate to make a critical comparison between L. longibrachia and L. paci$ca. In particular, no comparison of appendages is possible. Such differences, as are evident from the comparison of the description of L. Eongibrachia with the present material, include the dimensions of the trunk, the posterior processes, the relative length of the arms. The anterior part of the trunk of L. longibruchia is relatively wider and the posterior processes and arms longer. There are also some differences in the shape of the posterior margin of the trunk and that of the bulla. The latter, in contrast to the bulla of L. longibruchia, has no manubrium and is much wider. The two animals differ also in their overall size. L. longibrachia, without its posterior processes, is 11.5 mm long, its arms reaching the length of 31 to 32 mm. This can be contrasted with the arms of not more than 8.4 mm. long in the present species. (It should be noted that in this instance the difference in size appears in the manner opposite to the one usually observed. While normally the parasites of the lower latitudes tend to be smaller than their relatives from the colder climatic zones, here the Sakhalin species is much smaller than L. longibrachia, which was found at the low latitudes of the central Atlantic. Indirectly, this also suggests absence of close links between the tn o forms.) One has also to take into account the differences in host species and in habitat, particularly the latter. Although one does find among Lernaeopodidae some species able to parasitize both skin, fins and gills of its hosts, such ability is usually characteristic of members of the higher genera of the family; it is quite rare, among the members of the Charopinus group, to which Lernaeopodina belongs. L. longibruchia was found on both occasions on the skin of its hosts (Brian, 1913), while the present species parasitizes the gills of its host. Any doubts one might still harbour about the possible synonymy of L. pacijica with L. longibrachia can be resolved only by examination of the type material of the older species. The authors, unfortunately, were denied the opportunity to do so. They decided,

49 Parasitic Copepoda of fishes from Leningrad 203 therefore, to accept this accumulation of small differences as sufficient to erect a new species for the Pacific parasite. Clavella adunca (Str~m, 1762) (Figs 167 to 176) Syn. : C. urzciiiata (Miiller, 1776) Record of specimens: (i) Three female specimens were collected by the Ob expedition at station 202 (coast of Kemp Land, Antarctica), on 2.ii One of these females was not ovigerous and carried three males ; (ii) Two ovigerous females were collected by the vessel Ivan Nosenko near the coast of East Antarctica in Hosts : (i) Nacrurus whitsoni ; (ii) Treinutonzus loennberyi. Habitat: (i) buccal cavity; (ii) gills. Previous records: This species is very common and has been recorded by many authors from as many as 30 species of host fishes, mainly belonging to the family Gadidae, but also to others. It has been found in numerous localities in the waters of the northern hemisphere, but its previous southernmost records, as far as the authors are aware, was that of Wilson (1923) who found it on the gills of Doydixodonfasciatum, off the Pacific coast of South America. According to the literature, the genus Chvella inhabits predominantly the waters of the northern hemisphere and its records south of the Equator are comparatively few. It is very interesting, therefore, to find C. adunca at the locality, which brings its distribution range to the shores of the Antarctic and makes it it truly cosmopolitan species. The morphology of C. adunca is very well known and needs no comment here. Some salient features of the appendages and the general appearance of the specimens from this collection are illustrated in Figs 167 to 176. The dimensions of the specimens in this collection are shown below (in mm) : Cephalothorax length Cephalothorax width Trunk length Trunk width Genital process length Arms length Egg-sac length Egg-sac width Eggs diameter Trematomus 7- I L M Macrurus In determining the specific identity of these specimens, the authors took into account the great range of variability, exhibited by C. adunca and commented on by several authors (Poulsen, 1939; Nunes-Ruivo, 1957 ; Kabata, 1963). Both sets of specimens were rather smaller than is usual for C. adunca from the gadoid hosts of the northern hemisphere. They also differed in the proportions of the body (relative length of the cephalothorax, the trunk length :width ratio, the length of the genital process). In some respects, the specimens from Trematomus resembled C. perjida Wilson, 1915 more than they did a typical C. adunca. Neither set, however, showed any differences in the structure of the appendages from those of C. adunca, when compared with the European material of that species. As regards the similarity with C. perfida, the authors believe that a critical examination of that species will show it to be synonymous with C. adunca. No known description of

50 204 Z. KABAT AND A. V. GUSEV 70 Figs 1G7 to 176. Clavella adunca Fig Female from Trematomus Zoennbergi, ventral Female from T. Zoennbergi, lateral Female from T. Zoennbergi, mandible tip, lateral Female from T. Zoennbesgi, first antenna, tip, lateral Female from T. Zoennbergi, first antenna, diagram of apical armature Male from T. Zoennbergi, lateral Female from Macrurus whitsoni, lateral Female from M. ruhitsoni, ventral Female from M. whitsoni, mandible, tip, lateral Female from M. whitsoni, maxilla, lateral.

51 Parasitic Copepoda offishu from Leningrad 205 C. perfla shows distinctive features (save perhaps for the shape of the bulla) which would merit for it a status of an independent species. ACEN OWLEDGEMEXTS The authors wish to acknowledge their indebtedness to Prof. B. E. Bykhovsky (Academy of Sciences, U.S.S.R.), who collected a considerable proportion of the specimens described in this paper. They are also grateful to Dr J. P. Harding (British Museum, London), who kindly placed at their disposal type material of Chondracanthus macrurus and to Dr J. R. Grindley (South African Museim, Cape Town), who made available to them the type material of Eubrachiella sublobulatn. The authors benefited greatly from advice anti assistance given to them by Dr S. M. Shiino (University of Mie, Japan) and by Prof. C. Delamare Deboutteville (Museum Nacioiial d Histoire Naturelle, Brunoy, France). YUhIMARY This paper describes some para5itic copepods of marine fishes from- the collection of the Zoological Institute of the Academy of Scienccs, Leningrad, U.S.S.R. The descriptions and data are given of 21 species and two subspecies of copepods, belonging to 17 genera. The descriptions include one genus (Laterncanthue), five species (Ca,ligus sensilis, Chondra,canthodes tuberofurcatus, Parapharodes semilunaris, Lateracanthus quadripedis a,nd Lenaaeopodina pacifica) and two subspccies (Eubrachiella gaiizi dorsituberculah and Deiidrapta cameroni longiclasata) new tjo science. - REFEREXCES BaRNARD, I<. H., South Afcican para,sitic Copepoda. A12)Z ,fr. Jflis., 41 : BASSETT-SMITE, P. W., 1x98. Sohe new parasitic copepods found on fish at Bombay. Ann. Jlog. irut. Hist. (ser. 7), 1 : BENEDEN, v.4~ P. J., Note sur un notivetin genre de crustace parasite de la faniille des PeltocPphalides. Bull. Accrd. r. Bely. CC. Sci., 19: 4(L-467. BFNEDEIU, VAN P. J., Deus lernkopodiens nouvertux recuillis, l un aux Aqores, l autre sur les c6tes du Senegal. Bull. Accrd. r. Bely. CI. Sci., 22 ; BEIZE, R., Parasitic copepods from Gulf of Rlexico fish. Am. MicZl. fiat., 17: BRADY, G. S., Report 011 the Copepocla collected by H.M.S. Challenger. Challenger Rep., 8(1): BRIAN, A,, C opepodi para dei pcschi tl ltrtlicc, 1S7 pp. Genova. BRI~N, A,, Note prdliininaire sur les COpdpodeS parasites des poissons provenant des campagnes scientifiques de S.A.S. le Prince Albert ler do RIonnco ou ddposks dam les collections du Muske ochnographique. Bull. Inst. oce unogr. Mo?ucco, No. 110 : 1-19 BRIAN, a., Copkpodes parasites des poissons et echinides provenant des carnpagnes scientifiques de S.A.S. le Prince Albert Ier de Monaco ( ). IZe suet. Cump. scient. Prince Albert I, 38: BRI~N, A,, Sur un cas d anomalie present6 pkw un specimen dc Lernaeopoda Zongibrachia Brian. Bull. Inst. ocianogr. Monaco, No. 259: 1-5. BRIAN, A., CopBpodes parasites provenant des rpcentes campagnes scientifiques de S.A.S. le Prince Albert Ier de Monaco ou dkpos6s dans lcs collections du MusBe OcBanographique. Bull. Inst. ocdanogr. Monaco, NO. 286: CAPART, A,, Copepoda paraskica. V. RAdtnts scientifiques des croisieres du navire ecole Belge Mercator. Mdm. Mus. r. Hist. not. Bely. (2 ser.), 21: CAPART, A,, CopBpodes parasites. Re s. sci. Exp&Z. oc&anogr. bely. Enux c6t. afr. Atl. sud., 3(3): CAUSEY, D., Parasitic Copepoda. of Texas coastal fishes. Publs Inst. mar. Sci. Univ. Ten., 3 : CAUSEY, D., 1953 a. Parasitic Copepoda from Grand Isle, La. Occ. Pap. mar. Lab. La St. Univ., No. 7: CAUSEY, D., Parasitic Copepoda from Gulf of Mexico fish. Occ. Pap. mar. Lab. La St. Univ., NO. 9: DANA, J. D., United States ExpZoring Expedition during the years , under the command of Charles Wilkes, U.S.N., 13, pts. I and 11.

52 206 Z. KABAT AND A. V. GUSEV DELAMARE DEBOUTTEVILLE, C. & NUNES-RUIVO, L., Copepodes parasites des poissons Mediterraneens (ser. 3) (I). VieMillieu, 4: FRASER, C. hl., Copepods parasitic on fish from the Vancouver Island region. Proc. Trans. R. Soc. Can. (ser. 3), 13 (5): GNANAMUTHU, C. P., Sex differences in t,he chalinius and adult forms of CaZigus polycanthi n. sp. (Crustacea: Copepoda), parasitic on Balistes mnculatzts from Madras. Rec. Indian Mus., 47: GUSEV, A. V., Parasitic copepods of some marine fishes (In Russian). Parazit. Sb., 13: HEEGAARD, P., Parasitic copepods mainly from tropical and Antarctic seas. Ark. Zool., 34 (A18): HEEOAARD, P., Contribution to the phylogeny of the arthropods, Copepoda. Spolia zool. Mus. haun., 8 : HEEGAARD, P., Parasitic Copepoda from Australian waters. Rec. Aust. MPLS., 25 (9): HELLER, A. F., Parasites of cod and other marine fishes from the Baie de Chaleur region. Can. J. Res., D, 27: HELLER, C., Crustaceen. Reise der Oesterreichischeii Fregnttc Xovara um die Erde, 2 (3): HESSE, E., Description des crustacbs rapes ou nouveaux des c8tes de France. 29. Description de 10 nouveaux crustaces, dont 7 appartiennent au genre Cycrius de Kroyer et 3 au genre Kreyeria de Van Beneden. AnnlsSci. nut., b, (ser. G), 8 (11): KABATA, Z., The mouth and the mouth-parts of Lernaeocera brunchialis (L.), a parasitic copepod. Crustaceann, 3 (4): KABATA, Z., Clavelb (Copepoda) parasitic on British Gadidae : one species or several? Crustaceann, 5 (I): KABATA, Z., Revision of the genus Charopinus Kroyer, 1863 (Copopoda: Lernaeopodidae). Vidensk. Meddr dansk naturh. Foren., 127 : KABATA, Z., Parasitic Copepoda of fishes. Rep. B.A.hr.Z. antarct. Res. Exped., 8 (6): KABATA, Z., Bruchiella chevreuzii Van Beneden, 1891 (Copepoda: Lemaeopodidae) ; a taxonomic problem. Crustaceana, 10: KIRTISINGHE, P., Parasitic copepods of fish from Ceylon. 11. Porasitology, 29: KIRTISINGHE, P., Parasitic copepods of fish from Ceylon. III. Parasitology, 40: KIRTISINQHE, P., A review of the parasitic copepods of fish recorded from Ceylon, with description of additional forms. Bull. Fish. Res. Stn Ceylon, 17 (1): KRQYER, H., Bidrag ti1 kundskab om snylteltrehsene. Naturhist. Tiddskr. (ser. 3). 2: Lma, K., On the nature of the so-called sternal furca in the Caligidae. Ark. Zool. (ser. 2), 1: LEIGH-SHARPE, W. H., The Copepoda of the Sihoga Expedition. Part TI: Commensal and parasitic Copepoda. Monogr. Siboga-Exped., 29b (No. 123) : LEWIS, A. G., Caligoid copepods of the Hawaiian Islands. 1. Parasitic on fishes of the family Acanthuridae. Proc. US. natn. Mus.. 115: MARPEVICH, A. P., New species of parasit,ic copepods (In Russian). Dopov. Akad. Nauk URSR, 11: MARPEVICH, A. P., Parasitic copepods of$shes of the U.S.S.R. (In Russian), 258 pp. Kiev. NUNES-RUIVO, L., Contribution B 1 6tude des variations morphologiques de Clavella adu?lca (H. Strom), copepode parasite de Gadus callarias L. Considbrations sup quelques Cluvella parasites des Gadidae. Revta Fae. Citric. Univ. Li.sb., 2 ser. C, 5 (2): NUNES-RUIVO, L., Copepodes parasitas de peixes das costas de Angola (Lists faunistica). Notas mimeop-. Cent. Biol. pise., No. 33: PILLAI, K. N., 1962 (1963). Observations on the synonymy of Caligus coryphaenae. Ann. Mag. nat. Hist. (ser. 13), 5: POULSEN, E. M., Investigations on CZavella uncinata from cod in Danish waters. Vidensk. Mecldr dansk naturh. Poren., 102: QUIDOR, A., Cop6podes parasites. Deuzi2m.e Expdd. antaret. fr. ( ). Sci. nat. : SCOTT, T., Report on Entomostraca from the Gulf of Guinea. Trans. Linn. Soc. Load., 6 (2001.): SHIINO, S. M., Copepods parasitic on Japanese fishes. I. On the species of Caligus and Lepeophtheirus. Rep. Fac. Fish. prefect. Uni u. Mie, 1 : SHIINO, S. M., Note on Caligus quadratus sp. nov., a copepod parasitic on the fish Neothynnus macropterus (T. and S.). Bull. Jap. Soc. scient. Fish., 20: SHIINO, S. M., 1954a. Copepods parasitic on Japanese fishes. 4. The family Euryphoridae. Rep. Pac. Fish. prefect. Univ. Mie, 1 : SHIINO, S. M., Copepods parasitic on Japanese fishes. 13. Parasitic copepods collected off Kessennuma, Miyagi Prefecture. Rep. Fac. Fish. prefect. Univ. Mie, 2: SHIINO, S. M., Ueber eine neue Art der schmarotzenden Copepodengattung Dysgamus aus der Bucht von Sagami. Annotnes zool. Jap., 31 (3): SRIINO, S. M., Sammlung der parasitischen Copepoden in der Prafecturuniversitit von Mie. Rep. Fac. Fish. prefect. Univ. Mie, 3 :

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