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1 This article was downloaded by: [CINCEL Consortium - Chile] On: 23 April 2009 Access details: Access Details: [subscription number ] Publisher Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Journal of Natural History Publication details, including instructions for authors and subscription information: Four new species of Hargicotyle Mamaev, 1972 (Diclidophoridae) parasites on sciaenid fishes from Peru and Chile Marcelo E. Oliva a ; José L. Luque a a Laboratorio de Parasitología, Fac. Cs. Biológicas, Universidad Ricardo Palma, Lima 18, Perú Online Publication Date: 01 December 1989 To cite this Article Oliva, Marcelo E. and Luque, José L.(1989)'Four new species of Hargicotyle Mamaev, 1972 (Diclidophoridae) parasites on sciaenid fishes from Peru and Chile',Journal of Natural History,23:6, To link to this Article: DOI: / URL: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

2 JOURNAL OF NATURAL HISTORY, 1989, 23, Four new species of Hargicotyle Mamaev, 1972 (Diclidophoridae) parasites on sciaenid fishes from Peru and Chile MARCELO E. OLIVA and JOSl~ L. LUQUE Laboratorio de Parasitologla, Fac. Cs. Biolbgicas Universidad Ricardo Palma, P.O. Box 138, Lima 18, PenJ (Accepted 22 June 1989) Hargicotyle magna sp. nov., Hargicotyle paralonchuri sp. nov., Hargicotyle sciaenae sp. nov., from the gills of the Sciaenids Sciaenafasciata, Paralonchurus peruanus and Sciaena deliciosa, respectively, and Hargicotyle menticirrhi sp. nov. from the gill and mouth of Menticirrhus ophicephalus are described from the northern Chilean and central Peruvian coast. Distinct characteristics of the new species are the distribution of the vitelline follicles; the number and distribution of the testes, the size, shape and number of larval hooks; and the presence of one or two suckers in each damp, KEYWORDS: Marine fish parasites, Monogenea, Hargicotylinae, Sciaenidae, Taxonomy, South Pacific. Introduction The genus Hargicotyle was erected by Mamaev (1972) for those species described as Choricotyle and characterized mainly by the special features of the genital armature, i.e. multitudinous spines without bifid base and arranged in many rows. The species considered to belong in Hargicotyle by Mamaev (1972) were Hargicotyle louisianensis (Hargis, 1955) and Hargicotyle pacifica (Bravo-Hollis, 1966). Later on, Tantalran (1974) described two additional Choricotyle species from the Peruvian coast. These species were considered as members of Hargicotyle by Mamaev and Aleshkina (1984) and Oliva (1987). During a parasitological survey on marine fishes from Antofagasta, Chile and Chorrillos, Per/t, specimens of Hargicotyle magna sp. nov., Hargicotyle paralonchuri sp. nov., Hargicotyle menticirrhi sp. nov. and Hargicotyle sciaenae sp. nov. were collected from the gills and mouth of the sciaenid fishes Sciaenafasciata (Tschudi), Paralonchurus peruanus (Steindachner), Menticirrhus ophicephalus (Jordan) and Sciaena deliciosa (Tschudi), respectively. The new species are described and illustrated below and compared with descriptions of the known species of Hargicotyle Mamaev, Hargicotyle species described by Tantalran, are considered as species inquirenda. As a consequence of this paper, the genus Hargicotyle comprises six species, all parasitic in sciaenid fishes. Materials and methods The fishes examined were obtained fresh from the fish-market in Antofagasta, Chile (23 42' S ' W.) and Chorrillos, Peril (12 30' S ' W.). Monogeneans were removed, washed in saline solution, pressed between slides, fixed with hot AFA /89 $ Taylor & Francis Ltd.

3 1388 M.E. Oliva and J. L. Luque (Alcohol: formalin: acetic acid) and preserved in 70 alcohol. Worms were stained with Grenacher's alcoholic Borax-carmine and mounted in Canada balsam. Drawings were made with the aid of a camera lucida. Measurements, in miuimetres, were taken with a micrometer ocular; the mean is followed by the range in parentheses. The following institutional abbreviations apply: USNMHC: United States National Museum, Helminthological collection. MHNJP: Museo de Historia Natural 'Javier Prado' (Lima, Peril). CHURP: Colecci6n Helmintol6gica, Universidad Ricardo Palma (Lima, Peril). Results Hargicotyle magna sp. nov. (Figs 1-5) Description (Based on 6 specimens stained and mounted.) With the characters of Hargicotyle Mamaev, Body proper long and slender (Fig. 1) 18.7 (16"2-20.8) by 1.10 ( ), distinctly set-off posthaptor. Total length 23-0 ( ), maximum width at ovarian level. Posterior portion of body proper devoid of testes and vitelline follicles. Prohaptor with two well developed suckers (0" 19-0"22 in diameter), located close to oral aperture. Mouth terminal, pharynx immediately posterior to oral suckers. Intestinal caeca confluent posteriorly, branches extent into posthaptor and penetrate peduncles. Posthaptor palmate, long and slender, damps wider than peduncles. First pair of peduncles longer than the remainder. First and second peduncles originated from a common base. Terminal lappet elongate and thin, bearing two pairs of hooks, similar in size and shape, 22.5 micrometers long (Fig. 2). Clamps typical of Hargicotyle (Clamp nomenclature follows that of Euzet and Suriano, 1975). With one sucker in each clamp, and an oval accessory sclerite in the antero-axial quadrant. Anterior portion of M sclerite with a projection that reaches the middle of sclerite As (Fig. 4). Sclerite Al with a slightly sclerotized portion that runs near to Ma. Pal and Pll sderites articulated with As and A1 sclerites, respectively, and do not reach, ventrally, sclerites Ps2 and Pl2. O sclerite reaches the curved portion of Ps2 and P12. Testes , intercaecal, postovarian, about ( x 0.064).13). Vas deferens sinuous, extending anteriorly in midline to genital atrium. Ovary inverted 'U'-shaped, ootype and Mehli's gland located in space between ovarian branches. Oviduct originates from this depression, curved and runs posteriorly, oviduct receives common vitelline duct near curve, then ascends and form uterus (Fig. 5). Transverse vitelline ducts join at level of anterior third of the ovary. Genito-intestinal canal originates posteriorly from the oviduct. Genital atrium armed with numerous sicklelike spines, 22.5 #m long (Fig. 3). Genital pore 0.21 in diameter. Three pairs of frontal glands around the genital atrium. Eggs not observed. Host. Sciaena fasciata (Tschudi). Habitat. Gills. Locality. Chorrillos, Peril. Holotype. MHNJP Paratypes. MHNJP 1031, USNMHC 80536, CHURP 507. Etymology. The specific name refers to the size of the worm, the longest found in this study.

4 New species of Hargicotyle (Diclidophoridae) ! N- i O. O.V. 6. FIGS Hargicotyle magna sp. nov., Paratype, ventral view. 2. Larval hooks 3. Armature of genital atrium. 3 A. Free hand sketch of one hook of the genital armature. 4. Clamp. 5. Reproductive system, S.R.: Seminal receptacle, O: Ovary, O.V.: Oviduct, U: Uterus, M.G.: Mehli's gland, T: Testes, Note: dotted line in Fig 4 show slightly sclerotized piece. Hargicotyle menticirrhi sp. nov. (Figs 6-8) Description (Based on 4 specimens stained and mounted.) With characters of Har#icotyle Mamaev, Body proper (Fig. 6) long and slender, distinctly set-off posthaptor, 7.2 (7"1-7-4). Total length 8-6 ( ), maximum width 1.23 (1-06-1"39) at testicular level. Posterior part of body proper without testes and vitelline follicles. An

5 1390 M.E. Oliva and J. L. Luque 7 6 FIGS 6-8 Hargicotyle menticirrhi sp. nov. 6. Paratype, ventral view. 7. Larval hooks. 8. Clamp. anterior contraction is found at pharynx level. Prohaptor with two well developed suckers 0.11 x0-13 ( x ). Mouth terminal, pharynx posterior to suckers. Intestinal caeca confluent posteriorly, branches penetrate posthaptor. Posthaptor palmate, peduncles long and slender, similar in size, all peduncles join base at same level. Terminal lappet with three pairs of larval hooks, dissimilar in shape and size, 17.5, 15.0 and 8-8 #m long, respectively (Fig. 7). Clamps (Fig. 8) typical of Har#icotyle. With two suckers, each one in the anterior quadrant. An oval accessory sclerite is found in the antero-axial quadrant. Clamps sclerites as in H. magna, but distinct projection of sclerite A1 was found. A hook-like sclerite, namely herein As' sclerite, is present close to sclerite As. Testes 56-63, intercaecal, postovarian, about 0.18 in diameter. Female reproductive system as in H. magna. Genital pore 0-14 in diameter, armature of genital atrium as in H. magna, but genital hooks are smaller, 15.8 #m long. Three pairs of frontal glands are present near the genital atrium. Eggs not observed. 8

6 New species of Haroicotyle (Diclidophoridae) 1391 Host. Menticirrhus ophicephalus. Habitat. Gills and mouth. Locality Chorrillos, Perf. Holotype. MHNJP Paratypes. MHNJP 1033, USNMHC 80534, CHURP 508. Etymology. The specific name refers to the genus of the fish host. Hargicotyle parmonehmi sp. nov. (Figs 9-10) Description (Based on 6 specimens stained and mounted.) With characters of Hargicotyle Mamaev, Body proper long and slender (Fig. 9), 6.3 ( ) by 0.74 ( ), distinctly set-offposthaptor. Total length 8-2 ( ), maximum width at ovarian level. Vitellaria and testes reach the posterior end of the body proper, but do not penetrate the posthaptor. Prophaptor with two well developed suckers, 0.10 x 0.13 ( x ), located close to mouth. Mouth terminal, pharynx immediately posterior to suckers. Intestinal caeca confluent posteriorly, branches penetrate the posthaptor. Posthaptor 10 Fi~s Hargicotyle paralonchuri sp. nov. Paratype, ventral view. 10. Clamp (dotted line as in Fig. 4).

7 1392 M.E. Oliva and J. L. Luque palmate, peduncles long and slender, different in size, the first is the longer, the last pair, originated from a projection of the posthaptor. Terminal lappet with two pairs of hooks, similar in shape and size, 17-5/~m long. Clamps typical of Hargicotyle (Fig 10). With two suckers in each clamp, as in H. menticirrhi. An oval accessory sclerite in the antero-axial quadrant. Sclerite As' with a long slight projection that reaches near the oval accessory sclerite. A1 sclerite also with a long and slight projection. Other pieces as in H. magna. Testes , intercaecal, postovarian, small, about 0-07 in diameter. Female reproductive system as in H. magna. Genital atrium armed with numerous small sicklelike spines, 19/~m long. Three pairs of frontal glands near the genital atrium. Eggs not observed. Host. Paralonchurus peruanus (Steindachner). Habitat. Gills. Locality. ChorriUos, Peril. Holotype. MHNJP Paratypes. MHNJP 1035, USNMHC 80535, CHURP 509. Etymology. The specific name refers to the genus of the fish host. Hargicotyle sciaenae sp. nov. (Figs 11-13) Description (Based on 8 specimens stained and mounted.) With the characters of Hargicotyle Mamaev, Body proper (Fig. 11), long and slender, 6-1 ( ), distinctly set-off posthaptor. Total length 7-9 ( ), maximum width (0.89) at ovarian level. Vitelline follicles invading posthaptor. Prohaptor with two well developed suckers x (0-1784)-187 x ). Mouth terminal, pharynx immediately posterior to the suckers. Intestinal caeca confluent posteriorly, can penetrate the peduncles. Posthaptor palmate, first and second pairs of peduncles originated from a common base and are larger than remainder. Terminal lappet long and thin, armed with two pairs of larval hooks, different in size and shape, the longer is 22.5/~m and the smaller is 10 #m long. The larger one have bifid base and curved tip, the smaller are sickle shaped (Fig. 13). Clamps (Fig. 13) typical of Hargicotyle. With two suckers and an oval accessory sclerite. A strong projection of anterior portion of M sderite reach near the posterior portion of the same piece (M sclerite). Other pieces as in the above described species but without slight projection. Female reproductive system as in the above described species. Genital atrium armed with numerous spines, 30/~m long. Three pairs of frontal glands near the genital atrium. Eggs yellow, with two polar filaments, eggs without filament, 0-22x0.055 (0"214).23 x ).062) Host. Sciaena deliciosa (Tschudi). Habitat. Gills. Locality. Antofagasta, Chile. Holotype. MHNJP Paratypes. MHNJP 1037, USNMHC 80537, CHURP 510. Etymology. The specific name refers to the genus of the fish host.

8 New species of Hargicotyle (Diclidophoridae) 1393 Fms D l 112!I 11. Har#icotyle sciaenae sp. nov. Paratype ventral view. 12. Larval hooks. 13. Clamp (dotted line as in Fig. 4). 15 Discussion According to Mamaev and Aleshkina (1984), the genus Har#icotyle comprises four species, namely H. louisianensis (Hargis, 1955), H. americana (Bravo-Hollis, 1966), H. peruensis (Tantalran, 1974) and H. chimbotensis (Tantalran, 1974). The original description of H. louisianensis was based on an immature worm (Hargis, 1955), but Bravo-Hollis and Arroyo (1962) gave a complete redescription of the parasite. Oliva (1987) claimed that Hargicotyle peruensis is a common parasite of sciaenids from the northern coast of Chile, but a more exhaustive analysis showed that the specimens from Chile are a new species, Hargicotyle sciaenae. For comparative purposes, the authors requested the types ofh. peruensis and H. chimbotensis described originally as Choricotyle peruensis and Choricotyle chimbotensis, respectively (Tantalran 1974), but they were never deposited in the Museo de Historia Natural Javier Prado, as stated in Tantalran's paper. Material from the type locality (Chimbote,

9 1394 M.E. Oliva and J. L. Luque ca. 400 km north of Lima) was not available, but the fish host of Tantalran's material, the sciaenid Polyclemus (= Paralonchurus) peruanus is a common fish in the central coast of Perr. The study of the parasites obtained from this fish species revealed an undescribed species (described herein as Hargicotyle paralonchuri sp. nov.), but not Tantalran species. H. paralonchuri is easy to differentiate of Tantalran's material; the most notorious differences are the testicular number: in H. peruensis, in H. chimbotensis and in H. paralonchuri. H. chimbotensis have a vagina and a small number of genital hooks (14-16 spines). H. paralonchuri have a number of genital hooks and no vagina. H. peruensis harbour only one sucker in each clamp, instead there are two in H. paralonchuri. The absence of type material of Hargicotyle species described by Tantalran and the finding of H. paralonchuri in Paralonchurus peruanus, but not H. peruensis and/or H. chimbotensis, allow us to consider these species as species inquirenda. Thus, only two species of Hargicotyle are considered as valid herein There is only one diclidophorid known that has two suckers in each clamp, Choricotyle australiensis, a species described by Roubal et al. (1983). The known species of Hargicotyle, including those described herein, can be separated into two groups: those with one sucker in each clamp, namely H. louisianensis, H. pacifica and H. magna sp. nov., and those with two suckers in each clamp, namely H. sciaenae sp. nov. H. menticirrhi sp. nov. and H. paralonchuri. H. magna sp. nov. can be differentiated from the remaining species of the first group, by the presence of two pairs of larval hooks on the terminal lappet, instead of three pairs in H. louisianensis and only one in H. pacifica. Moreover, the testicular number is also different, there are around 46 in H. pacifica, in H. louisianensis and in H. magna sp. nov. In the second group, H. sciaenae is easily differentiated because it is the only species in the group with vitelline follicles invading the posthaptor deeply; in H. menticirrhi and H. paralonchuri the viteuine follicles do not penetrate the posthaptor. Moreover, the armature of the terminal lappets are dissimilar: H. sciaenae harbours two pairs of larval hooks different in size and shape, H. menticirrhi has three pairs of larval hooks and H. paralonchuri has two pairs, but similar in size and shape The testicular number is also different, in H. sciaenae it is 72-97, in H. menticirrhi and in H. paralonchuri The following artificial key permits the separation of the known species of Hargicotyle Mamaev, With one sucker in each clamp - With two suckers in each clamp. 2. Three pairs of larval hooks, testes. - Two pairs of larval hooks, testes - One pair of larval hook with bifid base, 46 testes 3 Vitelline follicles invading posthaptor, testes - Vitelline follicles not invading posthaptor 4 With three pairs of larval hooks, testes - Two pairs of larval hooks, testes 2 3 H. louisianensis H. magna H. pacifica H. sciaenae 4 H. "menticirrhi H. paralonchuri A c k n o w l e d g e m e n t s The authors are deeply indebted to the Consejo Nacional de Ciencia y Tecnologia (CONCYTEC-PERU) for the financial support to develop the research project 'Estudio parasitol6gico en peces marinos de importancia comercial de la Familia Sciaenidae'. We also express our thanks to Mr Miguel Marzal for his valuable help with the drawings.

10 New species of Haroicotyle (Diclidophoridae) 1395 References BRAvO-HOLLIS, M., and ARROYO, G., 1962, Helmintos de peces Costarricenses. Anales del Instituto de Biologia. Universidad Nacional Aut6noma de M~xico 37, EUZET, L., and StaUANO, D. M., 1975, Orbocotyle marplatensis n. gen. n. sp. (Diclidophotidae) monogene parasite branchial de teleosteens matins du genre Prionotus (Triglidae) en Argentine. Bulletin du Museum National d'historie Naturalle 3 Ser., Zool. 192, HARGIS, W., 1955, Monogenetic trematodes of Gulf of Mexico fishes. Part IX. The family Diclidophoridae Furhmann, Transactions of the American Microscopical Society 74, MAMAEV, Y. I., 1972, The description of a new monogeneans from the subfamily Choricotylinae with some notes about the genus Choricotyle composition. Trudy Biologo-Pochviennie Instituta Noboya Serie 11 (114), (In Russian). MAMAEV, Y. I., and ALESHKINA, L. D., 1984, Four new species of the higher monogeneans of the tropical part of the Atlantic Ocean. In Mamaev, Y. I. et al. (Eds). Parasites of Animals and Plants. Vladivostock: DVNTs AN SSSR, pp (In Russian). OLIVA, M., 1987, Choricotyle anisotremi n. sp. (Monogenea: Diclidophoridae) parasitic on Anisotremus scapularis (Tschudi) from the northern Chilean coast. Systematic Parasitolo#y 10, ROUBAL, F. R., ARMITAGE, J. and ROHD~ K., 1983, Taxonomy of Metazoan ectoparasites of Snapper, Chrysophrys auratus (Family Sparidae), from Southern Australia, Eastern Australia and New Zealand. Australian Journal of Zoology. Supplementary Series 94, 1~8. TANTAL~AN, M., 1974, Dos nuevas especies de monogeneos parhsitos de peces comerciales del mar Peruano. Biota (Per6) 10,

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