Systematic Parasitology 54: 2531, 2003. 2003 Kluwer Academic Publishers. Printed in the Netherlands.

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Dactylogyridae (Monogenea) from the gills of Iheringichthys labrosus (Osteichthyes: Pimelodidae) from the upper Paran a River oodplain, Brazil, with the proposal of Pseudovancleaveus n. g.
Jakeline G. Frana, Andr eia Isaac, Gilberto C. Pavanelli & Ricardo M. Takemoto
Universidade Estadual de Maring a, Nup elia, Bloco G-90, Av. Colombo, 5790, 87020-900, Maring a, State of Paran a, Brazil
Accepted for publication 28th May, 2002

Abstract Two new species of Demidospermus Suriano, 1983 and a new genus, Pseudovancleaveus, and species are described from the gills of Iheringichthys labrosus (Pimelodidae). The shes were collected from the upper Paran River oodplain, Brazil. D. mandi n. sp. is characterised by the distal region of the copulatoty organ having an expanded bulb and D. labrosi n. sp. by the copulatory organ having a funnel-shaped proximal extremity. The latter species also has an accessory piece comprising a variable sheath enclosing the distal shaft of the copulatory organ and two anterolateral structures resembling irregular spheres. Pseudovancleaveus paranaensis. n. g., n. sp. is characterised by a sinistral vagina, overlapping gonads, a copulatory ligament, anchors without a fold on the base and hooks with slightly inated shanks. A new combination, Pseudovancleaveus platensis, is proposed for Vancleaveus platensis Suriano & Incorvaia, 1995.

Introduction Demidospermus was proposed by Suriano (1983) for D. anus Suriano, 1983 from the gills of Loricaria anus (Loricariidae) in Argentina. Gutirrez & Suriano (1992) amended the generic characters of this genus and Kritsky & Gutirrez (1998) its diagnosis, indicating that it includes species having: tandem gonads; a counterclockwise coiled male copulatory organ; a sinistral vaginal aperture; U-, W- or V-shaped haptoral bars; subspherical eye-spots; and a sheath-like accessory piece serving as a guide for the male copulatory organ. Species of Demidospermus parasitise the gills of Neotropical siluriform shes (Loricariidae, Pimelodidae and Auchenipteridae).

Materials and methods Thirty-two specimens of Iheringichthys labrosus (Pimelodidae) were caught using gill-nets in the Paran River, Baa River and Patos Lagoon, all located in the Upper Paran River oodplain, between

March, 1992 and February, 1995. The gills of these hosts were removed and placed in vials containing a 1:4,000 solution of formalin. After 1 hour, the vials were vigorously shaken and additional formalin was added to increase the concentration to c. 5%. Parasites were removed from gills and some were stained with Gomoris trichrome. Other specimens were mounted unstained in Hoyers medium in order to study the sclerotised structures (see Eiras et al., 2000). Measurements, in micrometres, are expressed as the mean, followed by the range and number of specimens measured in parentheses. Drawings were made with the aid of a drawing tube and a Nikon YS 2 microscope. The holotype and paratypes were deposited in the Helminthological Collection of the Instituto Oswaldo Cruz (CHIOC), Rio de Janeiro, Brazil. The ecological terminology follows Margolis et al. (1982) as modied by Bush et al. (1997) and that of the haptoral sclerites tends to follow Kritsky & Mizelle (1968) and related papers.

26 Demidospermus Suriano, 1983 Demidospermus mandi n. sp. Description (Figure 1) [Based upon 21 specimens.] Body 603 (415-936; n = 11) long, 126 (72-174; n = 16) in greatest width, fusiform. Two cephalic lobes and head organs observed; cephalic glands posterolateral to pharynx. Eye-spots 4; posterior pair larger and closer together than anterior pair. Pharynx spherical, 39 (26-46; n = 7) in diameter; oesophagus short. Peduncle broad, 83 (40-120; n = 11) long, 85 (50-120; n = 13) wide. Haptor subhexagonal, 93 (54-188; n = 13) long, 131 (90-178; n = 13) wide. Ventral anchors 44 (41-47; n = 4) long; base 21 (20-22; n = 4) wide, with short roots; shaft short; extremity of point recurved. Dorsal anchors 36 (34-37; n = 4) long; base 17 (15-19; n = 4) wide, with short roots; shaft short; extremity of point slightly recurved. Bars V-shaped; ventral bar 96 (91-102; n = 4) long, distance between ends 66 (5771; n = 4); dorsal bar 73 (60-79; n = 4) long, distance between ends 51 (40-57; n = 4). Hooks pairs 1, 2 and 7 with recurved point, broad attened thumb and dilate shank comprised of 2 distinct regions; lament (FH) loop c. 1/4 shank length; hook pair 1: 42 (3948; n = 4) long; pair 2: 35 (31-38; n = 4); pair 7: 46 (39-53; n = 4). Hook pairs 3 and 4 with slightly inated shank, erect thumb, lament loop similar to shank in length; pair 3: 16 (15-17; n = 4) long; pair 4: 16 (16-17; n = 4). Hook pairs 5 and 6 with attened thumb, thin shank and lament loop similar to shank in length; pair 5: 19 (17-22; n = 4) long; pair 6: 22 (1824; n = 4). Copulatory organ coil forms incomplete ring, frequently J-shaped, 83 (79-88; n = 3) long; distal region of expanded to form bulb. Accessory piece associated with half of copulatory organ, 46 (43-50; n = 3) long. Gonads intercaecal, tandem. Testis posterior to ovary, 110 (88-152; n = 7) 70 (60-90; n = 7). Prostatic reservoirs oval. Ovary 43 (32-52; n = 6) 31 (22-38; n = 6). Vagina with small vestibule; aperture midway between level of ovary and copulatory organ; vaginal canal opens into small medial seminal receptacle. Vitelline follicles dense. Type-host: Iheringichthys labrosus (Ltken). Type-locality: Upper Paran River oodplain, Brazil (2243 S; 53 10 W). Site: Gills. Infestation: Prevalence: 93.75% (30 of 32 shes examined). Mean intensity: 18.8. Specimens deposited: CHIOC no. 34586a (holotype); nos 34586b, 34589, 34590, 34591, 34592, 34593a and 34593b (paratypes). Etymology: The specic epithet refers to the popular name of the host. Remarks Demidospermus mandi n. sp. is similar to D. leptosynophallus Kritsky & Gutirrez, 1998 from Iheringichthys westermanni (Reinhardt) and D. uncusvalidus Gutirrez & Suriano, 1992 from Pimelodus clarias (Lacpde) in the following features: (1) hook pairs 1, 2 and 7 have expanded shanks comprising two distinct regions; (2) the dorsal and ventral bars are V-shaped; (3) the ventral anchors have the tip of the point recurved; and (4) the dorsal anchor has the tip of the point slightly recurved. D. mandi n. sp. differs from D. leptosynophallus in having the distal region of the copulatory organ with an expansion to form a bulb (in D. leptosynophallus it has two sclerotised aps) and from D. uncusvalidus by lacking a broad posteromedial projection on the V-shaped dorsal bar.

Demidospermus labrosi n. sp. Description (Figure 2) [Based upon 15 specimens.] Body 440 (330-510; n = 7) long, 90 (80-104; n = 10) wide, fusiform. Two cephalic lobes and head organs observed; cephalic glands posterolateral to pharynx. Eye-spots 4; posterior pair larger and closer together than anterior pair. Pharynx spherical, 30 (20-31; n = 10) in diameter; oesophagus short. Peduncle 20 (10-30; n = 6) long, 60 (50-90; n = 8) wide. Haptor 50 (49-70; n = 9) long, 80 (79-100; n = 9) wide. Ventral and dorsal anchors similar in form and size, with poorly differentiated roots, short shaft and elongate point. Ventral anchor 24 (2227; n = 5) long; base 17 (13-23; n = 5) wide; roots moderately developed; shaft curved, elongate; point straight. Dorsal anchor 24 (22-27; n = 5) long; base 14 (11-17; n = 5) wide; supercial root relatively well developed; deep root poorly developed; shaft slightly curved; point elongate. Ventral bar U-shaped, 84 (71106; n = 5) long; distance between ends 53 (43-61; n = 5); internal margin serrate; postero-medial margin with concave notch. Dorsal bar V-shaped, 70 (56-89; n = 5) long; distance between ends 41 (30-59; n = 5). Hook pair 1: 23 (22-26; n = 3) long, larger and stouter than others; pair 2: 11 (10-12; n = 3) long; pair 3: 13

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Figure 1. Demidospermus mandi n. sp. A. ventral view; B. copulatory organ; C. hook 1; D. hook 2; E. hook 3; F. hook 4; G. hook 5; H. hook 6; I. hook 7; J. ventral bar; K. ventral anchor; L. dorsal bar; M. dorsal anchor.

(12-13; n = 5); pair 4: 14 (12-17; n = 3); pair 5: 15 (14-16; n = 3); pair 6: 12 (11-13; n = 3); pair 7: 17 (17; n = 3). Hook pair 1 with recurved point, heavy shaft, attened thumb, inated shank tapering proximally and lament loop c. 3/4 shank length; pair 2 with erect thumb, slightly expanded shank and lament loop similar to shank length; pairs 3, 4 and 7 with erect thumb, slender shank and lament loop of shank length; pairs 5 and 6 with attened thumb, delicate shank and lament loop c. 3/4 shank length. Accessory piece in form of elongate duct, associ-

ated with half of copulatory organ, with 2 irregularly spherical structures anterolaterally, 29 (27-35; n = 3) long. Copulatory organ J-shaped, 64 (59-73; n = 3) long; distal end forms large bulb; proximal extremity funnel-shaped. Testis oval, 44 (26-57; n = 6) 33 (24-49; n = 6); seminal vesicle elongate; prostatic reservoirs oval. Ovary 30 (22-42; n = 6) 23 (1928; n = 6). Vaginal aperture midway between level of ovary and copulatory organ; vaginal vestibule complex with sclerotised ridges; seminal receptacle small.

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Figure 2. Demidospermus labrosi n. sp. A. ventral view; B. copulatory organ; C. hook 1; D. hooks 5 and 6; E. hooks 3, 4 and 7; F. hook 2; G. ventral bar; H. ventral anchor; I. dorsal bar; J. dorsal anchor.

Vitelline follicles dense. Type-host: Iheringichthys labrosus (Ltken). Type-locality: Upper Paran River oodplain, Brazil (2243 S; 53 10 W). Site: Gills. Infestation: Prevalence: 100% (32 of 32 shes examined). Mean intensity: 116.6.

Specimens deposited: CHIOC no. 34594a (holotype) and nos 34587a, 34587b, 34587c, 34594b, 34594c, 34595, 34596 and 34597 (paratypes). Etymology: The specic epithet refers to the specic name of the host.

29 Remarks Demidospermus labrosi n. sp. is similar to D. cornicinus Kritsky & Gutirrez, 1998 from Iheringichthys westermanni (Reinhardt) in the following features: (1) the dorsal bar is V-shaped; (2) the dorsal and ventral anchors have poorly differentiated roots, a short shaft and an elongate point; and (3) the vaginal vestibule complex has sclerotised ridges. These species can be differentiated by their haptor and copulatory organ, in that: (1) the ventral bar in both species is U-shaped, but in the new species its internal margin is serrate and there is a concavity in the posteromedial margin; and (2) the copulatory organ of the new species has a funnel-shaped proximal extremity and its proximal shaft is surrounded by an accessory piece with a variable sheath and two anterolateral structures, absent in D. cornicinus, which resemble irregular spheres. Remarks Features distinguishing the new genus include the combined presence of: (1) a sinistral vagina in the from of a tube opening into the seminal receptacle; (2) overlapping gonads; (3) an elongate seminal vesicle; (4) an oval prostatic reservoir; (5) a copulatory ligament; (6) dorsal and ventral anchors with a developed deep root, an elongate shaft and a short point; and (7) hooks with slightly inated shanks and a protruding thumb. Pseudovancleaveus differs from Vancleaveus Kritsky, Thatcher & Boeger, 1986 mainly in the absence of a fold on the base of the anchors. The new genus is named because of its great resemblance to Vancleaveus. Pseudovancleaveus paranaensis n. sp. Description (Figure 3) [Based on 14 specimen.] Body 287 (207-373; n = 6) long, 83 (66-91; n = 9) in greatest width, divisible into cephalic region, trunk, short peduncle and haptor. Head organs and cephalic lobes present; cephalic glands form 2 symmetrical groups posterolateral to pharynx. Eye-spots absent. Pharynx subovate, 19 (1722; n = 7) in diameter; oesophagus short. Haptor 48 (34-60; n = 7) long, 84 (64-121; n = 7) wide. Anchors similar in form and size; each with long shaft and short point. Ventral anchor 33 (27-38; n = 4) long; base 11 (11-12; n = 4) wide, without fold on base; roots well developed. Dorsal anchor 31 (26-34; n = 4) long; base 10 (9-11; n = 4) wide, without fold on base; supercial root well developed; deep root short. Dorsal bar Ushaped, 55 (51-60; n = 4) long; distance between ends 49 (39-57; n = 3). Ventral bar V-shaped, 54 (40-62; n = 4) long; distance between ends 47 (32-55; n = 3); with large posteromedial projection. Haptor with 7 pairs of hooks, similar in form and size, 22 (21-23; n = 3) long; shaft with recurved point; lament loop c. 1/3 of shank length. Accessory piece 19 (17-22; n = 4) long; coil-like copulatory organ with 3 coils; coils 13 (12-17; n = 4) in diameter; base enlarged proximally to form small bulb. Copulatory ligament present. Testis oval, 20 (17-24; n = 2) 13 (9-17; n = 2) wide. Ovary 44 (42-47; n = 2) 20 (18-23; n = 2), overlaps testis ventrally. Vaginal pore sinistral. Vitelline follicles dense. Type-host: Iheringichthys labrosus (Ltken). Type-locality: Floodplain of the upper Paran River,

Pseudovancleaveus n. g. Diagnosis Dactylogyridae; Ancyrocephalinae. Body divisible into cephalic region, trunk, short peduncle and haptor. Tegument thin, smooth. Head organs and cephalic lobes present; cephalic glands unicellular comprising two bilateral groups posterolateral to pharynx. Eye-spots rudimentary or absent. Mouth subterminal; pharynx subovate; oesophagus short; intestinal caeca two. Gonads overlapping, intercaecal. Vas deferens loops left intestinal caecum; prostatic reservoir oval; seminal vesicle elongate. Copulatory organ coil-like, with three coils; base with enlarged region forming small bulb; accessory piece elongate. Copulatory ligament present. Haptor armed with seven pairs of hooks, dorsal and ventral pairs of anchors, and dorsal and ventral bars. Hooks with inated slightly shanks, protruding thumb, elongate shaft, short, recurved point and lament loop approx. one-third of shank length. Ventral anchor with fully developed deep root, elongate shaft and short point. Dorsal anchor with poorly developed deep root, elongate shaft and short point. Both anchors lack fold on base. Ventral bar V-shaped, with large posteromedial projection. Dorsal bar Ushaped. Ovary elongate. Testis dorsal to ovary. Vaginal pore sinistral; vagina not sclerotised. Vitelline follicles dense. Type-species P. paranaensis n. sp.

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Figure 3. Pseudovancleaveus paranaensis n. sp. A. ventral view; B. copulatory organ; C. hook; D. ventral bar; E. ventral anchor; F. dorsal bar; G. dorsal anchor. Abbreviation: CL, copulatory ligament.

Brazil (22 43 S; 53 10 W). Site: Gills. Infestation: Prevalence: 90.62% (29 of 32 shes examined). Mean intensity: 92.3. Specimens deposited: CHIOC no. 34598a (holotype) and nos 34588a, 34588b, 34588c, 34588d and 34598b (paratypes). Etymology: The specic name is derived from the type-locality.

Remarks Pseudovancleaveus paranaensis n. sp. is similar to Vancleaveus platensis Suriano & Incorvaia, 1995 from Pimelodus clarias maculatus (Lacpde) in many respects. The new species can be distinguished from V. platensis by the following features: (1) the ventral bar in P. paranaensis. is V-shaped with large posteromedial projection, while in V. platensis it is rod-shaped; (2) the dorsal bar is U-shaped in the new

31 species but V-shaped in V. platensis; (3) the deep root of the ventral anchor is less well developed in P. paranaensis; and (4) the supercial root of the dorsal anchor in V. platensis is better developed. The dorsal and ventral bars were incorrectly labelled in the group of gures of V. platensis presented by Suriano & Incorvaia, 1995 in that the ventral bar was identied as the dorsal bar. V. platensis is transferred to the new genus below. References
Bush, A.O., Lafferty, K.D., Lotz, J.M. & Shostak, A.W. (1997) Parasitology meets ecology on its own terms: Margolis et al. revisited. Journal of Parasitology, 83, 575-593. Eiras, J.C., Takemoto, R.M. & Pavanelli, G.C. (2000) Mtodos de estudo e tcnicas laboratoriais em parasitologia de peixes. Maring: Eduem, 171 pp. Gutirrez, P.A. & Suriano, D.M. (1992) Ancyrocephalids of the genus Demidospermus Suriano, 1983 (Monogenea) parasites from siluriform shes in Argentina, with descriptions of three new species. Acta Parasitologica, 37, 169-172. Kritsky, D.C. & Gutirrez, P.A. (1998) Neotropical Monogenoidea. 34. Species of Demidospermus (Dactylogyridae, Ancyrocephalinae) from the gills of pimelodids (Teleostei, Siluriformes) in Argentina. Journal of the Helminthological Society of Washington, 65, 147-159. Kritsky, D.C. & Mizelle, J.D. (1968) Studies on monogenetic trematodes. XXXV. Some new and previously described North American species of Gyrodactylus. The American Midland Naturalist, 79, 205-215. Kritsky, D.C., Thatcher, V.E. & Boeger, W.A. (1986) Neotropical Monogenea. 8. Revision of Urocleidoides (Dactylogyridae, Ancyrocephalinae). Proceedings of the Helminthological Society of Washington, 53, 1-37. Margolis, L., Esch, G.W., Holmes, J.C., Kuris, A.M. & Schad, G.A. (1982) The use of ecological terms in parasitology (Report of an ad hoc committee of the American Society of Parasitologists). Journal of Parasitology, 68, 131-133. Suriano, D.M. (1983) Demidospermus anus gen. nov. sp. nov. (Monogenea: Ancyrocephalinae) parasita branquial de Loricaria (L.) anus Valenciennes, 1840 (Pisces: Loricariidae) de la Laguna de Chascomus-Provincia de Buenos Aires - Republica Argentina. Neotropica, 29, 111-119. Suriano, D.M. & Incorvaia, I.S. (1995) Ancyrocephalid (Monogenea) parasites from siluriform shes from the Paranean-Platean ichthyogeographical province in Argentina. Acta Parasitologica, 40, 113-124.

Pseudovancleaveus platensis (Suriano & Incorvaia, 1995) n. comb. Remarks This species was originally described from Pimelodus clarias maculatus, from the Rio de la Plata at Buenos Aires Harbour, Argentina, as Vancleaveus platensis by Suriano & Incorvaia (1995). It is here transferred to Pseudovancleaveus n. g. due to the absence of a fold on the base of the dorsal anchor; this is a major distinguishing feature of the new genus.

Acknowledgements The authors wish to thank Nuplia/UEM, Nucleus of Research in Limnology, Ichthyology and Aquaculture, for logistic and nancial support. Our thanks are also due to the Editor and an anonymous reviewer for valuable criticism which improved this manuscript greatly, and to Dr Luis Carlos Gomes, Universidade Estadual de Maring, for help with the translation of the manuscript.