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Bothriocephalus kupermani n. sp. (Figs. 1–4) Informal synonyms: ‘ Bothriocephalus sp. ’ (PBI-487) of Brabec et al. (2015); ‘Pumpkinseed morph’ of Choudhury and Scholz (2020). Material studied: The description is based on whole, unstained, and unmounted specimens, specimens processed for SEM, and 18 specimens prepared as stained whole mounts. Measurements are from stained whole mounts as follows (number of worms in parentheses): 8 gravid specimens and 1 mature non-gravid specimen from pumpkinseed from Paul Lake, 5 November 2001 (2), Peter Lake, 13 July 2003 (4). Bolger Bog, 12 August 2003 (2) and Caribou Lake (1), and 1 gravid specimen from green sunfish from Duck Creek on 12 July 2009. In addition, 8 stained immature worms from pumpkinseed from Paul Lake, Peter Lake, and Shawano Lake were also studied but not used for measurements. Description: Bothriocephalidea, Bothriocephalidae. Body 21– 43 mm (35 mm) long, maximum width 973–1,595 (1,078) in posterior third. Strobila gradually widening posteriorly, comprising craspedote proglottids (primary and secondary), trapezoidal especially in anterior half, posterior proglottids with rounded edges, often losing craspedote appearance (Figs. 1, 2); proglottids covered with coniform spinitriches (Fig. 4F, G). Except for first few proglottids, immature proglottids including those without obvious anlagen of gonads numerous, wider than long, mature proglottids (with sperm in external sperm duct) very few (1–2), 175–507 (495) long by 781–1,232 (894) wide, length:width ratio 0.11–0.55 (0.55). Gravid proglottids much wider than long (Fig. 1D, E), first gravid proglottid 183–512 long by 750–1530 wide, length:width ratio 0.15–0.55, proglottids often narrower at end of strobila, posterior-most proglottid 252–750 (750) long by 751– 1,208 (751) wide, length:width 0.2–1.0 (1.0). Longitudinal musculature well-developed, formed by numerous muscle fibers. Osmoregulatory canals usually in several (usually 2–3) pairs (Fig. 1E). Ventral osmoregulatory canals wide, sinuous to almost straight (Fig. 1E). Dorsal osmoregulatory canals narrow, thin-walled, slightly sinuous, median to ventral osmoregulatory canals (Fig. 1E). Scolex 414–805 (458) long, with nearly parallel margins through most of its length, dorsoventrally (Figs. 1B, C, 2 A, D, 3A, B) and laterally (Figs. 1A, 2C, 3C, 4B), occasionally slightly wider in middle (Figs. 1A, 3C, 4A), maximum lateral width 151–210, maximum dorsoventral width 183–296 (296). Scolex gently narrowing anteriorly before widening to form slightly convex (almost blunt in dorsoventral view), bilobed apical disc with rounded outer corners (Figs. 1A–C, 3A–C, 4A– C). Apical disc 151–215 wide laterally, 179–234 (234) wide dorsoventrally. Bothria shallow, wide, opening anteriorly to shallow hollow at base of apical disc (Figs. 1B, C, 3A, B, 4A). Lateral surface of scolex may possess medial, shallow groove (Figs. 4 A, B). Neck short, 105–180 (180) wide, almost as wide as scolex and first proglottid (Figs. 1A–C, 2A–D). Scolex with coniform spinitriches, capilliform filitriches, tumuli (Fig. 4A, D), patch/tuft of capilliform filitriches at center of apical disc (Fig. 1C, E). Testes oval to subspherical, variable in size, 36–83 long by 26– 57 wide, medullary, 38–62 (49) per proglottid, ovoid to spherical, in 2 lateral fields on each side; most testes in main external (outer) field between ventral osmoregulatory canals, with only few testes in inner, narrow irregular field, median to internal ventral osmoregulatory canal (Figs. 1E, 3F). Cirrus sac median, anterior to ovary, transversely oval to subspherical in dorsoventral view, 61–74 (74) long by 63–90 (78) wide, leading to common genital pore on mid-dorsal surface of proglottid (Fig. 1E); genital pore postequatorial to almost equatorial, situated at 51–65% of proglottid length; external sperm duct (vas deferens) lateral to cirrus sac (Fig. 1E). Ovary compact, median, near posterior margin of proglottid, irregular in shape (slightly asymmetrical), weakly bilobed, with narrower mid-region (ovarian isthmus), 62–100 (88) long by 218– 368 (248) wide (Figs. 1E, 3E), occupying 23–38% of proglottid width. Vagina tubular, narrower in proximal part, sinuous, widened, thick-walled in distal part, opening to common genital pore on dorsal surface (Fig. 1E). Vitelline follicles very variable in shape, from elongate to almost spherical, 25–61 long by 20–46 wide, cortical, extensive, forming wide lateral fields on each side of proglottid from its lateral margins to beyond dorsal osmoregulatory canals medially, also variably and more sparsely present medially, often confluent by narrow field at level of ovary and uterine pore (Figs. 1D, E, 3D). Uterus tubular, thick-walled, looped in proximal part (anterodorsal to ovary and dorsal to vagina), then turning anterolaterally, circumventing vaginal canal ventrally, thereafter directed anteriorly, winding (Fig. 1E). In gravid proglottids, terminal (distal) part enlarged to form transversely oval, thick-walled uterine sac filled with numerous eggs; width of uterine sac in last proglottids 44–69% of proglottid width (usually about 50%). Uterine sacs open on ventral surface by almost medially situated, slit-like uterine pore situated at variable distance (at 18–37% of proglottid length) from anterior margin of proglottids (Figs. 1D, E). Eggs ovoid, 46–54 long by 32–34 wide (n ¼ 48), tanned when fully developed, operculate, unembryonated in utero. Taxonomic Summary Type host: Lepomis gibbosus (Linnaeus, 1758) (Centrarchiformes: Centrarchidae). Additional hosts: Lepomis cyanellus Rafinesque, 1819 (this study and museum material), L. macrochirus Rafinesque, 1819 (museum material), L. megalotis (Rafinesque, 1820) (museum material) (all Centrarchiformes: Centrarchidae) (see Table I). Site of infection: Intestine. Type locality: Paul Lake (46°15 ′ 04.78 ′′ N, 89°30 ′ 12.94 ′′ W; 516 meters above sea level [m a.s.l.]), University of Notre Dame Environmental Research Center (UNDERC), Michigan (Upper Peninsula) near Land-o-Lakes, Wisconsin. Additional localities: Peter Lake (46°15 ′ 10.17 ′′ N, 89°30 ′ 12.94 ′′ W; 518 m a.s.l.); Bolger Bog (46°13 ′ 47.37 ′′ N, 89°29 ′ 96 ′′ W; 507 m a.s.l.), both UNDERC lakes, Wisconsin; Shawano Lake, Wisconsin (44°47 ′ 11.88 ′′ N, 88°32 ′ 39.60 ′′ W); Caribou Lake, Minnesota (47°42 ′ 59.04 ′′ N, 90°39 ′ 18.18 ′′ W), Duck Creek near Hobart, Wisconsin (44°32 ′ 01.09 ′′ N, 88°07 ′ 44.89 ′′ W), Fox River at Voyageur Park, De Pere, Wisconsin (44°27 ′ 03.39 ′′ N, 088°03 ′ 51.83 ′′ W). Geographical distribution: Canada (Ontario), Mexico (precise locality not known; fish host introduced), United States (Alabama, Michigan, Minnesota, Nebraska, Ohio, Oklahoma, Wisconsin). Infection rate: Mean abundance: 20.6 ± 25.3; 0–112 tapeworms/fish; prevalence: 83% of 30 fish examined from the type locality. Type material (all from L. gibbosus): Holotype: USNM 1593369, Paul Lake, UNDERC, Michigan, 5 November 2001. Paratypes: USNM 1593370–1593373, HWML 216811–216817, IPCAS C-914 /1. Molecular data: The following DNA sequence data are available in GenBank (https://www.ncbi.nlm.nih.gov/genbank/) (the region of the genome is followed by the GenBank accession number: 18S rDNA (ssr DNA): KR780953, 18S rDNA (V 8 region, partial): MT 532462– MT 532463; ITS-1 (partial): MT 532532– MT 532537; ITS-2 (partial): MT 532512– MT 532513; 28S rDNA (lsr DNA): KR780906, 16S rDNA (rrnL): KR780867; cytochrome c oxidase subunit 1 (COI): KR780814. ZooBank registration: urn:lsid:zoobank.org:act: 701C450F-6CE8-4CF8-ADB6-164F49BBAF39. Etymology: The species is named after the late Boris Kuperman, whose sudden demise in 2002 robbed parasitology of a wise, gifted, and gentle practitioner who continued his new life in the United States with the same lifelong passion he always had for our beloved subject, and whose seminal work on the genus Triaenophorus inspired the senior authors (A.C. and T.S.) many years ago. Remarks Bothriocephalus kupermani differs from other species of Bothriocephalus parasitizing freshwater fishes in North America, namely B. claviceps, B. cuspidatus, B. formosus, and B. pearsei, by the following characteristics: (1) the scolex is almost rectangular in dorsoventral and lateral profiles, with nearly parallel margins (vs. arrow-shaped and narrowing towards the apical disc in B. cuspidatus and with maximum width in the anterior third in B. claviceps), with shallow and wide bothria with almost parallel margins (widest in the posterior or middle part in other taxa); (2) small body size (maximum of 43 mm vs. 133 and 137 mm in B. cuspidatus and B. claviceps, respectively; Scholz, 1997); and (3) extensive vitelline follicles that are also present medially, i.e., in the cortex of the ovarian and uterine region (absent anteriorly and never or seldom confluent posteriorly in other species except B. formosus, which has markedly different scolex and tiny strobila compared to the new species; see Scholz, 1997; Choudhury and Scholz, 2020). Bothriocephalus kupermani is also typified by a relatively high number of immature proglottids without obvious anlagen of gonads and fewer testes (38–62) than B. cuspidatus from Walleye (62–101). The validity of B. kupermani is also supported by molecular data of Brabec et al. (2015; isolate called ‘ Bothriocephalus sp. n., ‘PBI-487 ′ in their fig. 2) and Choudhury and Scholz (2020). The specimen from green sunfish, L. cyanellus, is similar in its morphology to specimens from the type host, L. gibbosus, and is considered conspecific., Published as part of Choudhury, Anindo, Scholz, Tomáš & Beuchel, Joseph S., 2022, A New Species Of Bothriocephalus (Cestoda: Bothriocephalidae) From Lepomis Spp. (Actinopterygii: Centrarchidae) In North America, pp. 343-352 in Journal of Parasitology 108 (4) on pages 344-348, DOI: 10.1645/21-70, http://zenodo.org/record/7746757, {"references":["BRABEC J., A. WAESCHENBACH, T. SCHOLZ, D. T. J. LITTLEWOOD, AND R. KUCHTA. 2015. Molecular phylogeny of the Bothriocephalidea (Cestoda): Molecular data challenge morphological classification. International Journal for Parasitology 42: 259 - 267.","BANGHAM, R. V. 1944. Parasites of northern Wisconsin fish. Transactions of the Wisconsin Academy of Sciences, Arts and Letters 36: 291 - 325.","SCHOLZ, T. 1997. A revision of the species of Bothriocephalus Rudolphi, 1808 (Cestoda: Pseudophyllidea) parasitic in American freshwater fishes. Systematic Parasitology 32: 85 - 107."]}

Based on morphological and molecular data, a new species of tapeworm, Bothriocephalus kupermani n. sp., is described from pumpkinseed, Lepomis gibbosus (Linnaeus, 1758) (type host), and green sunfish, L. cyanellus Rafinesque, 1819 (Actinopterygii: Centrarchidae) in the United States. The new species differs from its North American congeners mainly in the shape of its scolex, which is almost rectangular in dorsoventral and lateral views (nearly parallel margins in the bothrial region), with shallow, wide bothria, and by an extensive field of vitelline follicles that are also variably present, albeit more sparsely, medially in the cortex of the ovarian and uterine areas. This tapeworm appears to be typically associated with Lepomis sunfishes. Examination of museum specimens of Bothriocephalus species from Lepomis spp. indicates that the tapeworm also parasitizes bluegill, Lepomis macrochirus, longear sunfish, Lepomis megalotis, and that previous records of Bothriocephalus cuspidatus and Bothriocephalus claviceps in these fishes are of the new species, B. kupermani.

Regobothrium microhamulinum n. gen. and n. sp. (Bothriocephalidea: Bothriocephalidae) is proposed to accommodate a new cestode from flatfish Catathyridium jenynsii (Chabanaud, 1922) (Pleuronectiformes: Acharidae; type host) and another 3 freshwater fishes of the orders Characiformes, Cyprinodontiformes, and Siluriformes in the Neotropical Region. The new genus is placed in the Bothriocephalidae because it possesses medioventral uterine and mediodorsal genital pores and a follicular vitellarium. Regobothrium n. gen. is characterized by possessing a tiny, slightly subovate scolex narrower than the strobila, with an apical disc armed with 2 semicircles of 15-17 tiny hooks in each and an acraspedote strobila. Regobothrium n. gen. differs from all bothriocephalid cestodes that have a scolex armed with hooks by their small size (maximum length less than 20 μm) and a triangular shape with the basal part (handle or basal plate) shorter than the distal coniform part (blade). In the other hooked bothriocephalids, hooks have a longer handle than a blade. Regobothrium microhamulinum n. gen. and n. sp. is the third bothriocephalidean cestode described from freshwater teleosts in South America but the first out of Patagonia. Molecular phylogenetics consider Regobothrium as a member of a lineage consisting of (up to now exclusively) freshwater bothriocephalids from the Ethiopian biogeographic region, thus indicating Gondwanan relationship.

Three parasitic gut worms, one cestode and two acanthocephalan species, have been recorded from two fresh water live fishes (Channa punctatus and Channa striatus) of Channidae family from West Bengal, India for the first time. These parasites were revealed from new host species which were hitherto not reported till now. The cestode is Bothriocephalus acheilognathi (Family Bothriocephalidae) and the acanthocephalans are Pallisentis ophiocephali (family Quadrogyridae) and Neoechinorhynchus sootai (family Neoechinorhynchidae). In this communication, the morphological studies of prevailing cestode and two acanthocephalan species were also incorporated.

Objective of research: To study the activities and properties of nucleoside- diphosphatase (NDPase) in cestode Bothriocephalus scorpii. Materials and methods: Cestodes were homogenized with 10 vol. of extraction medium. NDPase was detected in mitochondria and microsomes with substrates (IDP, GDP, UDP). Inorganic phosphorus was determined by the method of Kochetov (1980). The effects of 10 anthelmintic drugs on the activity of NDPase were studied. Results and discussion: It was found that the mitochondrial and microsomal fractions of cestodes B. scorpii have nucleoside diphosphatase activity. The activity of nucleoside diphosphatase depends on substrates and Mg2+ ions. The impact of various effectors and ions (Ca2+, Mn2+, Zn2+) on enzyme activity was determined. Effects of 10 anthelmintic drugs on activity of nucleoside diphosphatase were studied. The anthelmintics Bitionol and Trichlorophen have been proved effective.

Schyzocotyle acheilognathi is a tapeworm cestode commonly found in native freshwater fishes from Asia. This cestode has low host specificity and for that reason it has been registered parasitizing more than 200 cultured and wild fish species, besides amphibians, reptiles and birds from different regions of the world. With a high pathogenic potential, S. acheilognathi may cause mortalities in highly infected fish. In South America, Schyzocotyle was reported in Cyprinus carpio from a Brazilian fish farm at the municipality of Cornélio Procópio, northern Paraná State and from the natural environment in C. carpio from Neuquen River, Patagonia, Argentina. So far, there has been no report of this parasite in South American native fishes. Herein we report the first occurrence of the invasive tapeworm S. acheilognathi in Rineloricaria pentamaculata (Siluriformes, Loricariidae), a native armored freshwater catfish from southern Brazil.

One of 4 Georgia blind salamanders (25%), Eurycea wallacei from Dougherty County, Georgia, U.S.A., were found to be infected in its small intestine with a Bothriocephalus sp. tapeworm; a single E. wallacei from Jackson County, Florida, U.S.A., was negative. Because the internal anatomy of this worm was immature, species identity was not possible. This is the first definitive report of any parasite from this G2 ranked (globally imperiled) host and a new distribution record for the genus.

A survey of bothriocephalidean tapeworms (Cestoda) parasitizing African freshwater fish is provided. Based on criticalevaluation of type specimens and extensive, newly collected material, only the following seven species, instead of 19 taxalisted in the literature, are considered to be valid and their redescriptions are provided: Bothriocephalus acheilognathi Yamaguti, 1934 (with 3 synonyms from Africa); Bothriocephalus claviceps (Goeze, 1782) (marginally in Africa); Ichthybothrium ichthybori Khalil, 1971; Kirstenella gordoni (Woodland, 1937) n. comb. (1 synonym); Polyonchobothrium polypteri (Leydig, 1853) (4 synonyms); and Tetracampos ciliotheca Wedl, 1861 (4 synonyms). In addition, Tetracampos martinae Kuchta n. sp. is proposed for tapeworms from the catfish Bagrus meridionalis from Lake Malawi. The new speciesdiffers from T. ciliotheca in a much larger body (19 cm versus 3 cm), dorsoventally flattened strobila and numerous (39 versus25–35) and longer apical hooks (up to 98 µm versus less than 50 µm). Kirstenella Kuchta n. gen. is proposed to accommodate Senga gordoni Woodland, 1937 as its type species. The new genus is distinguished from other genera of the Bothriocephalidaeby the presence of an apical disc armed with two lateral semicircles of large hooks, cortical vitelline follicles and large-sizedcirrus-sac. All but one valid species were recollected. Bothriocephalidean cestodes are widely distributed throughout Africa,but only two species, B. acheilognathi and T. ciliotheca, occur in other continents. All but one species (B. acheilognathi) exhibit narrow host specificity, being limited either to one host species (K. gordoni in Heterobranchus bidorsalis and T. martinae in Bagrus meridionalis) or one host genus (I. ichthybori in Ichthyborus spp., P. polypteri in Polypterus spp. and T. ciliotheca in Clarias spp.). Molecular data based on partial sequences of the large subunit rDNA (lsrDNA) show monophyletic position of all African taxa analysed (B. acheilognathi, I. ichthybori, K. gordoni, P. polypteri and T. ciliotheca).

Bothriocephalus claviceps (Goeze, 1782) Rudolphi, 1810 Type locality: Unknown (? Germany). Distribution in Africa: Nile basin – Egypt. Type material: Not known to exist. Material studied: Bothriocephalus claviceps ex Anguilla sp. from Egypt (BMNH 1932.5.31.780– 780). References: Meggitt (1930); Tadros (1967). Remarks: Meggitt (1930) identified immature fragments of tapeworms from unidentified eels [most probably Anguilla anguilla (Linnaeus), the only eel occurring in the region], allegedly found by A. Looss in Egypt, as B. claviceps. Examination of the vouchers confirmed this identification. Bothriocephalus claviceps is a specific parasite of eels in the Holarctic Region (Scholz et al. 2004) and its occurrence in Africa represents the southernmost range of its distribution area, apparently related to occasional migration of eels to the rivers in North Africa including the lower Nile. Based on the distribution of A. anguilla, it is possible that future studies will reveal the presence of B. claviceps also in Maghreb ichthyofaunistic province, which has Palaearctic affinities and includes Morocco, northern Algeria and Tunisia 1. Since the distribution of this Holarctic tapeworm in Africa is restricted to lower Nile and possibly Maghreb province, it is not included into the key below., Published as part of Kuchta, Roman, Burianová, Alena, Jirkú, Miloslav, Chambrier, Alain, Oros, Mikuláš, Brabec, Jan & Scholz, Tomáš, 2012, Bothriocephalidean tapeworms (Cestoda) of freshwater fish in Africa, including erection of Kirstenella n. gen. and description of Tetracampos martinae n. sp., pp. 1-35 in Zootaxa 3309 on page 24, DOI: 10.11646/zootaxa.3309.1.1, http://zenodo.org/record/280992, {"references":["Goeze, J. A. E. (1782) Versuch einer Naturgeschichte der Eingeweidewurmer thierischer Korper. Blankenburg, 471 pp.","Meggitt, F. J. (1930) Report on a collection of cestodes mainly from Egypt. IV. Conclusion. Parasitology, 22, 338 - 345.","Tadros, G. (1967) On a new cestode Bothriocephalus prudhoei sp. nov. from the Nile catfish Clarias anguillaris with some remarks on the genus Clestobothrium June [sic], 1899. Bulletin of the Zoological Society of Egypt, 21, 74 - 88.","Scholz, T., Skerikova, A., Shimazu, T. & Grygier, M. J. (2004) A taxonomic study of species of Bothriocephalus Rudolphi, 1808 (Cestoda: Pseudophyllidea) from eels in Japan: morphological and molecular evidence for the occurrence of B. claviceps (Goeze, 1782) and confirmation of the validity of B. japonicus Yamaguti, 1934. Systematic Parasitology, 57, 87 - 96."]}

Bothriocephalus acheilognathi Yamaguti, 1934 (Figs. 1, 2, 4, 12) Syns. (only taxa reported from Africa; for other synonyms – see Pool & Chubb 1985; Pool 1987; Kuchta & Scholz 2007): Bothriocephalus (Clestobothrium) kivuensis Baer & Fain, 1958; Bothriocephalus aegyptiacus Ryšavý & Moravec, 1975; Bothriocephalus barbus Fahmy, Mandour & El-Naffar, 1978; Bothriocephalus sp. of Yimer (2000), Al-Bassel (2003), and Zekarias & Yimer (2007). Type host: Acheilognathus rhombeus (Temminck & Schlegel) (Cypriniformes: Cyprinidae). Other definitive hosts in Africa (for extensive list of hosts from other continents, see Dove & Fletcher 2000): Barbus altianalis Boulenger, Barbus argenteus Günther, Barbus brevipinnis Jubb, Barbus bynni (Forsskål), Barbus mattozi Guimarães, Barbus paludinosus Peters, Barbus trimaculatus Peters, Cyprinus carpio Linnaeus, Labeobarbus aeneus (Burchell), Labeobarbus kimberleyensis (Gilchrist & Thompson), Labeobarbus marequensis (Smith), Labeobarbus nedgia Rüppell (Cypriniformes: Cyprinidae). Type locality: Lake Ogura, Kyoto Prefecture, Honshu, Japan (35 ° 2 'N, 135 ° 53 'E). Distribution in Africa: Congo basin – Democratic Republic of the Congo; Incomati basin – South Africa; Limpopo basin – South Africa; Maputo basin – South Africa; Nile basin – Egypt, Ethiopia; Orange basin – South Africa. Prevalence and intensity of infection: Precise data are not available, but this cestode seems to be rather infrequent. Despite extensive sampling, the present authors found only a single barbel infected with B. acheilognathi in Ethiopia (out of 56 barbels examined; see Appendix 1). However, Bertasso & Avenant-Oldewage (2005) found prevalence up to 90 % in L. kimberleyensis from South Africa. Type material: MPM 23780 (holotype). Material studied: Type material: holotype (one slide of whole mount and one slide of histological sections) of B. acheilognathi; holotype (one worm on 20 slides) of B. aegyptiacus ex B. bynni from Cairo, Egypt (IPCAS C– 14); three syntypes (3 mounted scoleces and histological sections) of B. kivuensis ex B. altianalis from Lake Kivu, Democratic Republic of the Congo (MHNG 40332) (Fig. 12); vouchers: one specimen ex L. kimberleyensis from Vall Dam, South Africa, collected by M. Barson (MHNG 36429); one specimen of Bothriocephalus sp. ex Hydrocynus sp. (Craciformes: Alestidae) from Bagata, Kwilu, Democratic Republic of the Congo (MHNG 55308); new material: three specimens ex L. nedgia from Beshelo River, near Old Bridge, Ethiopia, collected by Moges Beletew (MHNG 55310) (Figs. 1, 2, 4). Published records from Africa: Baer & Fain (1958, 1960); Ryšavý & Moravec (1975); Amin (1978); Fahmy et al. (1978); Boomker et al. (1980); Brandt et al. (1981); van As et al. (1981); El-Naffar et al. (1984); Basson & van As (1993); Paperna (1996); Schulz & Schoonbee (1999); Al-Bassel (2003); Bertasso & Avenant-Oldewage (2005); Retief et al. (2006, 2007, 2009); Zekarias & Yimer (2007); Degger & Avenant-Oldewage (2009); Degger et al. (2009); Stadtlander et al. (2011). Remarks: Up to date, six species of Bothriocephalus Rudolphi, 1808 have been reported to occur in freshwater fish in Africa, especially in barbels (Cyprinidae: Barbinae) (Baer & Fain 1958; Tadros 1967; Ryšavý & Moravec 1975; Fahmy et al. 1978; Bertasso & Avenant-Oldewage 2005; Stadtlander et al. 2011). However, the present study has shown that apart from marginal occurrence of B. claviceps (see below), only one species, the Asian fish tapeworm, Bothriocephalus acheilognathi, actually parasitizes African fish. Bothriocephalus prudhoei Tadros, 1967 is invalidated (synonym of Kirstenella gordoni – see below) and three remaining species of Bothriocephalus, namely B. (Clestobothrium) kivuensis Baer & Fain, 1958; B. aegyptiacus Ryšavý & Moravec, 1975; and B. barbus Fahmy, Manour & El-Naffar, 1978, are synonyms of B. acheilognathi (see Pool 1987; Kuchta & Scholz 2007). Bothriocephalus acheilognathi has been introduced from its original distribution area in East Asia throughout the world (Scholz et al. 2011 b). It is a pathogen of fry of cultured fish, especially carp and other cyprinids, and has been reported to cause mortalities (Williams & Jones 1994). Numerous bothriocephalidean tapeworms have been synonymized with B. acheilognathi, including African taxa (see Kuchta & Scholz 2007 for list of synonyms). Pool (1987) synonymized B. aegyptiacus and B. kivuensis with B. acheilognathi, whereas B. barbus was considered to be a synonym of B. acheilognathi by Kuchta & Scholz (2007). Molecular data confirm these synonymies (Fig. 62). The host spectrum of B. acheilognathi is extraordinarily wide and includes more than 200 species of unrelated fish (Scholz et al. 2011 b), with cyprinids representing the most suitable definitive hosts. In Africa, it has been recorded in 12 cyprinid species of two native genera, Barbus (7 spp.) and Labeobarbus (4 spp.), as well as in introduced Cyprinus caprio (Stadtlander et al. 2011). Records from the Nile basin, South Africa, as well as the Congo basin suggest that B. acheilognathi is present throughout the continent wherever suitable cyprinid hosts are available. Surprisingly, there are no records from Labeo spp., suggesting that members of this speciose pan-African cyprinid genus constituting a significant component of ichthyofauna of many African basins are not suitable hosts. Such presumed unsuitability of Labeo spp. as hosts for B. acheilognathi might be the reason of its apparent absence in basins in which cyprinids are dominated by Labeo spp. This assumption is supported by the absence of B. acheilognathi in Lake Turkana (this study, Appendix 1), where none of the examined cyprinids, i.e. 42 Labeo spp. and 11 Barbus bynni, was infected (ratio of examined fish does not reflect real ratio in the lake in which two Barbus spp. in very low densities share the lake with huge populations of two Labeo spp.). Findings of B. acheilognathi in clariid catfish in Ethiopia, Nigeria and Zimbabwe (Anosike et al. 1992; Yimer 2000; Moyo et al. 2009; Bichi & Yelwa 2010) may represent accidental findings due to predation, but extraordinarily high values of prevalence (up to 60 %; Moyo et al. 2009) indicate that Clarias Scopoli catfish may harbour this parasite more frequently. Unfortunately, no voucher specimens of these remarkable, but suspicious findings have been preserved, which casts doubts upon correct identification of the worms found. It thus cannot be excluded that the authors misidentified Tetracampos ciliotheca Wedl, 1861, which is a very frequent and abundant intestinal parasite of Clarias catfish in Africa (see below) and the scolex of which somewhat resembles that of B. acheilognathi, especially when hooklets on the apical disc are detached after death of worms. Tapeworms from Tilapia sp. (Characiformes: Cichlidae) and Hydrocynus sp. identified as Bothriocephalus sp. (Khalil & Thurston 1973; present study), may represent incidental infection of B. acheilognathi, but species identification could not be confirmed, because the specimen from Tilapia sp. is not available and the worm from Hydrocynus sp. is without scolex. Bothriocephalus acheilognathi is distributed throughout Africa, but its abundance is considerably lower than in newly colonized regions, especially in Europe and North America (García-Prieto & Osorio-Sarabia 1991; Williams & Jones 1994; Salgado-Maldonado & Pineda-López 2003). The absence of records from West Africa is probably artificial, because great majority of parasitological studies on cyprinids in this region were obviously focused solely on ectoparasitic monogeneans (see Khalil & Polling 1997)., Published as part of Kuchta, Roman, Burianová, Alena, Jirkú, Miloslav, Chambrier, Alain, Oros, Mikuláš, Brabec, Jan & Scholz, Tomáš, 2012, Bothriocephalidean tapeworms (Cestoda) of freshwater fish in Africa, including erection of Kirstenella n. gen. and description of Tetracampos martinae n. sp., pp. 1-35 in Zootaxa 3309 on pages 4-6, DOI: 10.11646/zootaxa.3309.1.1, http://zenodo.org/record/280992, {"references":["Yamaguti, S. (1934) Studies on the helminth fauna of Japan. Part 4. Cestodes of fishes. Japanese Journal of Zoology, 6, 1 - 112.","Pool, D. W. & Chubb, J. C. (1985) A critical scanning electron microscope study of the scolex of Bothriocephalus acheilognathi Yamaguti, 1934, with a review of the taxonomic history of the genus Bothriocephalus parasitizing cyprinid fishes. Systematic Parasitology, 7, 199 - 211.","Pool, D. W. (1987) A note on the synonymy of Bothriocephalus acheilognathi Yamaguti, 1934, B. aegyptiacus Rysavy and Moravec, 1975 and B. kivuensis Baer and Fain, 1958. Parasitology Research, 73, 146 - 150.","Kuchta, R. & Scholz, T. (2007) Diversity and distribution of fish tapeworms of the \" Bothriocephalidea \" (Eucestoda). Parassitologia, 49, 129 - 146.","Baer, J. C. & Fain, A. (1958) Bothriocephalus (Clestobothrium) kivuensis n. sp., a cestode parasitic on a barbel of Lake Kivu. Annales de la Societe Royale Zoologique de Belgique, 88, 287 - 302.","Rysavy, B. & Moravec, F. (1975) Bothriocephalus aegyptiacus sp. n. (Cestoda: Pseudophyllidea) from Barbus bynni, and its life cycle. V e stnik C eskoslovenske Spole c nosti Zoologicke, 39, 68 - 72.","Fahmy, M. A. M., Mandour, A. M. & El-Naffar, M. K. (1978) On some cestodes of the freshwater fishes in Assiut Province, Egypt. Veterinary Medical Journal, 24, 253 - 262.","Yimer, E. (2000) Preliminary survey of parasites and bacterial pathogens of fish at Lake Ziway. Ethiopian Journal of Science, 23, 25 - 33.","Al-Bassel, D. (2003) A general survey of the helminth parasites of fish from inland waters in the Fayoum Governorate, Egypt. Parasitology Research, 90, 135 - 139.","Zekarias, T. & Yimer, E. (2007) Study on parasites of fish at Lake Awassa, Ethiopia. Bulletin of Animal Health and Production in Africa, 55, 149 - 155.","Dove, A. D. M. & Fletcher, A. S. (2000) The distribution of the introduced tapeworm Bothriocephalus acheilognathi in Australian freshwater fishes. Journal of Helminthology, 74, 121 - 127.","Bertasso, A. & Avenant-Oldewage, A. (2005) Aspects of the ecology of the Asian tapeworm, Bothriocephalus acheilognathi Yamaguti, 1934 in yellowfish in the Vaal Dam, South Africa. Onderstepoort Journal of Veterinary Research, 72, 207 - 217.","Baer, J. C. & Fain, A. (1960) Observations sur le developpement de Bothriocephalus (Clestobothrium) kivuensis Baer et Fain 1958. Acta Tropica, 17, 374 - 377.","Amin, O. M. (1978) Intestinal helminths of some Nile fishes near Cairo, Egypt with descriptions of Camallanus kirandensis Baylis, 1928 (Nematoda) and Bothriocephalus aegyptiacus Rysavy and Moravec, 1975 (Cestoda). Journal of Parasitology, 64, 93 - 101.","Boomker, J., Huchzermeyer, F. W. & Naude, T. W. (1980) Bothriocephalosis in the common carp in the Eastern Transvaal. Journal of the South African Veterinary Association, 51, 263 - 264.","Brandt, F. d. W., van As, J. G., Schoonbee, H. J. & Hamilton-Attwell, V. L. (1981) The occurrence and treatment of bothriocephalosis in the common carp, Cyprinus carpio in fish ponds with notes on its presence in the largemouth yellowfish Barbus kimberleyensis from the Vaal Dam, Transvaal. Water SA, 7, 34 - 42.","van As, J. G., Schoonbee, H. J. & Brandt, F. d. W. (1981) Further records of the occurrence of Bothriocephalus (Cestoda: Pseudophyllidea) in the Transvaal. South African Journal of Science, 77, 343.","El-Naffar, M. K., Saoud, M. F. & Hassan, I. M. (1984) Some cestodes from some fishes of lake Nasser at Asswan, including a new Marsypocephalus, Marsypocephalus aegyptiacus n. sp. Assiut Veterinary Medical Journal, 12, 69 - 76.","Basson, L. & van As, J. G. (1993) 1 st record of the European trichodinids (Ciliophora, Peritrichida), Trichodina acuta Lom, 1961 and T. reticulata Hirschmann et Partsch, 1955 in South Africa. Acta Protozoologica, 32, 101 - 105.","Paperna, I. (1996) Parasites, infections and diseases of fishes in Africa. An update (Vol. 31). CIFA Technical Paper FAO, Rome, 210 pp.","Schulz, G. W. C. & Schoonbee, H. J. (1999) Aspects of the length, mass, fecundity, feeding habits and some parasites of the shortfin minnow, Barbus brevipinnis (Cyprinidae) from the Marite River, Mpumalanga province, South Africa. Water SA, 25, 257 - 264.","Retief, N. R., Avenant-Oldewage, A. & du Preez, H. H. (2006) The use of cestode parasites from the largemouth yellowfish, Labeobarbus kimberleyensis (Gilchrist and Thompson, 1913) in the Vaal Dam, South Africa as indicators of heavy metal bioaccumulation. Physics and Chemistry of the Earth, 31, 840 - 847.","Retief, N. R., Avenant-Oldewage, A. & du Preez, H. H. (2007) Ecological aspects of the occurrence of Asian tapeworm, Bothriocephalus acheilognathi Yamaguti, 1934 infection in the largemouth yellowfish, Labeobarbus kimberleyensis (Gilchrist and Thompson, 1913) in the Vaal Dam, South Africa. Physics and Chemistry of the Earth, 32, 1384 - 1390.","Retief, N. R., Avenant-Oldewage, A. & du Preez, H. H. (2009) Seasonal study on Bothriocephalus as indicator of metal pollution in yellowfish, South Africa. Water SA, 35, 315 - 322.","Degger, N. & Avenant-Oldewage, A. (2009) Metal accumulation analysis within tissue of Bothriocephalus acheilognathi. Journal of the South African Veterinary Association, 80, 127 - 128.","Stadtlander, T., Weyl, O. L. F. & Booth, A. J. (2011) New distribution record for the Asian tapeworm Bothriocephalus acheilognathi Yamaguti, 1934 in the Eastern Cape province, South Africa. African Journal of Aquatic Science, 36, 339 - 343.","Tadros, G. (1967) On a new cestode Bothriocephalus prudhoei sp. nov. from the Nile catfish Clarias anguillaris with some remarks on the genus Clestobothrium June [sic], 1899. Bulletin of the Zoological Society of Egypt, 21, 74 - 88.","Khalil, L. F. (1971 a) Check list of the helminth parasites of African freshwater fish. Commonwealth Agricultural Bureaux, Farnham Royal, England, 80 pp.","Woodland, W. N. F. (1937) Some cestodes from Sierra Leone. II. A new caryophyllaeid, Marsypocephalus, and Polyonchobothrium. Proceedings of the Zoological Society of London B, 107, 189 - 197.","Scholz, T., Kuchta, R. & Williams, C. (2011 b) Chapter 17. Bothriocephalus acheilognathi Yamaguti, 1934. In: P. T. K. Woo & K. Buchmann (Eds), Fish Parasites: Pathobiology and Protection. CABI, Wallingford, pp. 282 - 297.","Williams, H. & Jones, A. (1994) Parasitic worms of fish. Taylor & Francis, London, 593 pp.","Anosike, J. C., Omoregie, E., Ofojekwu, P. C. & Nweke, I. E. (1992) A survey of helminth parasites of Clarias gariepinus in Plateau State, Nigeria. Journal of Aquatic Sciences, 7, 39 - 43.","Moyo, D. Z., Chimbira, C. & Yalala, P. (2009) Observations on the helminth parasites of fish in Insukamini Dam, Zimbabwe. Research Journal of Agriculture and Biological Sciences, 5, 782 - 785.","Bichi, A. H. & Yelwa, S. I. (2010) Incidence of piscine parasites on the gills and gastrointestinal tract of Clarias gariepinus (Teugels) at Bagauda fish farm, Kano. Bayero Journal of Pure and Applied Sciences, 3, 104 - 107.","Wedl, K. (1861) Zur Helminthenfauna Agyptens. III. Nematoda. IV. Cestoda. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften Wien, 44, 463 - 482.","Khalil, L. F. & Thurston, J. P. (1973) Studies on the helminth parasites of freshwater fishes of Uganda including the descriptions of two new species of digeneans. Revue de Zoologie et de Botanique Africaines, 87, 209 - 248.","Garcia-Prieto, L. & Osorio-Sarabia, D. (1991) Distribucion actual de Bothriocephalus acheilognathi en Mexico. Anales del Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Seria Zoologia, 62, 523 - 526.","Salgado-Maldonado, G. & Pineda-Lopez, R. F. (2003) The Asian fish tapeworm Bothriocephalus acheilognathi: a potential threat to native freshwater fish species in Mexico. Biological Invasions, 5, 261 - 268.","Khalil, L. F. & Polling, L. (1997) Check list of the helminth parasites of African freshwater fish. Second edition. University of the North, Pietersburg, 185 pp."]}

Kirstenella gordoni (Woodland, 1937) Kuchta, n. comb. (Figs. 7, 8, 10, 11, 15–22) Syns: Polyonchobothrium gordoni Woodland, 1937; Polyoncobothrium gordoni (Woodland, 1937) Yamaguti, 1959; Bothriocephalus prudhoei Tadros, 1967; Polyonchobothrium sp. of Fagbenro et al. (1993); Senga gordoni (Woodland, 1937) Kuchta & Scholz, 2007. Type host: Heterobranchus bidorsalis Geoffroy Saint-Hilaire (Siluriformes: Clariidae). Other definitive hosts: Clarias anguillaris (Linnaeus) (Siluriformes: Clariidae), Schilbe mystus (Linnaeus) (Siluriformes: Schilbeidae). Both species are considered accidental/atypical hosts – see the Remarks section. Type locality: Waanje River near Pujehun, Sierra Leone. Distribution: Gambia basin – Senegal (Lampsar River, Taoué River, Gambia River); Turkana basin – Ethiopia (lower Omo River) and Kenya (Lake Turkana – restricted to the northernmost freshwater region part of the lake; see Remarks); Upper Guinea – Sierra Leone (Moa River); Niger basin – Mali (Lake Debo), Nigeria (River Ogbase); Nile basin – the Sudan (White Nile, Kostí) and Uganda (Lake Victoria). Prevalence and intensity of infection: Lake Turkana – Omo River delta, Todonyang, Kenya, 60 %, n = 10; intensity 1–6 (present study); Ethiopia – Omo River, Omorate, 50 %, n = 2, intensity 1 (present study); Nile basin – the Sudan, 20 %, n = 5, intensity 1 (present study); Niger basin – Nigeria, 22 %, n = 185 (Fagbenro et al. 1993). Type material: Syntypes (several slides with fragments of worms and their histological sections) (BMNH 1965.2.24.36– 45). Material studied: Type material: several syntypes; holotype and paratypes of Polyonchobothrium gordoni from Waanje River, Sierra Leone (BMNH 1965.2.24.36 – 45); Bothriocephalus prudhoei ex Clarias anguillaris, Malakal, the Sudan (BMNH 1998.10.15.6.7; RVC C. 1262); vouchers: Ptychobothriidae gen. sp. ex H. bidorsalis, Mali, Lake Dabo (MNHNP bd 10); P. clarias ex H. bidorsalis, Taoué River, Senegal (RMCA 34695; D.T.F. Puylaert; 15.iii. 1966); Polyonchobothrium sp. ex H. bidorsalis, Waanje River, Sierra Leone (BMNH 1965.2.24.62– 65; collected by W.N.F. Woodland); Polyonchobothrium sp. ex Schilbe mystus, Lake Victoria, Uganda (BMNH 1957.12.30.34– 38); new material: 1 specimen ex H. bidorsalis, White Nile in Kostí, the Sudan (MHNG 49379; 26.iii. 2006); 1 specimen from Omo River, Omorate, Ethiopia (MHNG 63067; 15.iv. 2006); 15 specimens from Omo River delta, Todonyang, Lake Turkana, Kenya (MHNG 69955); 3 specimens from Niokolo-Koba National Park, River Gambia, Senegal collected by B. Koubková (2006; Sen 272 - 1). The new material is deposited in BMNH (Nos. 2012.3.20.26– 28), IPCAS (No. C- 609), MHNG (Nos. 55339, 63254, 82036, 82038) USNPC (No. 105390) and ZMB (No. 7524). Published records: Woodland (1937); Tadros (1968); Khalil (1973); Fagbenro et al. (1993). Re-description (based on 10 whole mounts and 1 scolex observed by SEM; measurements from Woodland, 1937 in brackets): Bothriocephalidea, Bothriocephalidae. Strobila up to 10 [1.0–1.5] cm long; maximum width 875 [1,350]. External and internal segmentation present; segments wider than long, markedly craspedote, several ridges on surface of segments (Figs. 10, 15). Two pairs of longitudinal osmoregulatory canals; dorsal canals narrow (diameter up to 7); ventral canals wide (diameter up to 14), connected by transverse anastomoses. Inner longitudinal musculature well developed, muscle fibres diffused (Fig. 21). Surface of strobila covered with capilliform filitriches. Scolex oval, narrow (Figs. 7, 15, 17), 926 – 1,480 (1,160 ± 168) [280–380] long by 224–419 (317 ± 65) [310– 516] wide (n = 15). Apical disc weakly developed, 230–300 (265 ± 18) wide by 74–102 (87 ± 10) long, armed with 40–42 (41 ± 1) [36–42] small hooks 16–88 (67 ± 19; n = 103) long. Hooks variable in size, arranged in two semicircles, smallest on periphery above each bothrium and increasing into middle of semicircle with largest hook 83– 88 (86 ± 3; 3) [64] long (Figs. 8, 16, 18). Bothria elongate, shallow, 757 – 1,355 (1,025 ± 230) long by 73–98 (84 ± 11) wide (n = 7) (Figs. 7, 17). Surface of scolex covered with capilliform filitriches and tumuliform globular structures (diameter around 1) (Fig. 5). Neck absent, first segments appearing immediately posterior to scolex (Figs. 7, 15, 17). Immature segments 112–257 (191 ± 43) long by 217–531 (340 ± 101) wide; segment length/width ratio 0.27– 0.94: 1 (n = 15) (Fig. 15). Mature segments wider than long, 134–371 (242 ± 74) long by 283–998 (667 ± 201) wide; segment length/width ratio 0.23–0.66: 1 (n = 24) (Fig. 22). Gravid segments wider than long, 188–573 (326 ± 97) long by 704 – 1,104 (869 ± 114) wide; segment length/width ratio 0.24–0.79: 1 (n = 20) (Fig. 15). Testes medullary, oval, 70–133 (86 ± 20; 10) [under 30] in number per segment, 23–51 (35 ± 9) long by 26–45 (36 ± 6) wide [69 × 44 in sections] (n = 10) in diameter, forming 2 narrow longitudinal bands, 34–72 (47 ± 11) testes per band, confluent between segments, absent medially and near lateral margins (Fig. 22). Cirrus-sac large, thick-walled (thickness of sac wall 7–14), spherical, 92–161 (121 ± 21) long by 91–171 (132 ± 20) wide, length/ width ratio 0.66–1.15: 1 (n = 24), pre-equatorial to postequatorial (at 30–69 % of length of mature segment from anterior margin; n = 10). Internal seminal vesicle present, Vas deferens forms numerous loops posterolateral to cirrus-sac; internal sperm ducts strongly coiled, cirrus unarmed, opening into genital atrium (Figs. 19, 21, 22). Genital pore dorsal, median, pre-equatorial (Fig. 22). Ovary asymmetrical, compact, bilobed, 44–95 (70 ± 16) long by 224–455 (332 ± 7) wide (n = 16) (Figs. 19, 22). Vagina a straight, thin-walled tube, 22–52 in diameter, opens posterior to cirrus-sac into genital atrium; vaginal sphincter absent (Fig. 19). Vitelline follicles numerous, small, spherical, 11–17 (14 ± 2; 8) [51 × 22] in diameter, cortical, form 2 wide longitudinal bands confluent between segments, separated medially, rarely connected by several follicles in postovarian region (Fig. 22). Uterine duct forms numerous tightly coiled loops, filled with eggs, enlarged in gravid segments (Fig. 15). Uterus thick-walled, median, spherical, enlarged in gravid segments, occupies 4–30 % of segment surface (Fig. 15). Uterine pore thick-walled, opens slightly posterior to midlength of uterus (Figs. 19, 22). Eggs oval, thin-walled, operculate, unembryonated, with abopercular knob, 30–43 (38 ± 4) lo ng by 21–33 (28 ± 4) wide [47 × 31] (n = 11) (Figs. 11, 20). Remarks: Polyonchobothrium gordoni Woodland, 1937 was described from Heterobranchus bidorsalis from Sierra Leone. Woodland (1937) mentioned similarity of his new species with Tetracampos ciliotheca Wedl, 1861 in the shape of the body (“great part of strobila tightly coiled”) and differentiated both taxa by the number and size of apical hooks. Tadros (1967) described a new species Bothriocephalus prudhoei Tadros, 1967 based on material collected by K.N. Soliman from Clarias anguillaris in the Sudan in 1958. Only one complete specimen with several pieces of strobila was available, but the apical part of its scolex is missing. Kuchta and Scholz (2007) considered B. prudhoei to be a junior synonym of T. ciliotheca, but re-examination of the type material did not support this synonymy. In fact, strobilar morphology, including the presence of a large cirrus-sac, is identical to that of K. gordoni. Therefore, B. prudhoei is synonymized with this species, even though B. prudhoei was not found in its type host H. bidorsalis, but in Clarias, which is considered herein to be an accidental host. Tadros (1968) compared Polyonchobothrium gordoni with P. cylindraceum forma major Janicki, 1926 and concluded that these species were closely related and could be separated from each other by the number and size of the hooks. Kuchta & Scholz (2007) transferred P. gordoni to Senga because they considered Polyonchobothrium Diesing, 1854 to be monotypic (see Kuchta et al. 2008 b). However, the present study has shown that P. gordoni differs from all species of Senga, which occur exclusively in the Indomalayan Region, in several morphological characteristics considered to be of generic importance. Therefore, the new genus, Kirstenella, is proposed to accommodate P. gordoni (see above). No reliable record of Kirstenella gordoni has been published since the original description of the species, but new material of this tapeworm was collected by the present authors and their collaborators in Ethiopia, Kenya, Senegal and the Sudan (Fig. 61). Tapeworms from H. bidorsalis from Lake Debo, Nigeria, identified as Ptychobothriidae gen. sp. (MNHNP bd 10), and those from the same host from the rivers Lampsar and Taoué in Senegal (RMCA 34695), identified by Khalil (1973) as Polyonchobothrium clarias, are conspecific with K. gordoni. Heterobranchus currently comprises 4 species and represents a sister clade to African species of the genera Clarias and Bathyclarias Jackson (Pouyaud et al. 2009), but only H. bidorsalis is known as the definitive host of K. gordoni. In Turkana basin, the occurrence of K. gordoni most probably reflects distribution of its host, H. bidorsalis, which is restricted to the Omo River and freshwater part of the Turkana Lake, i.e. Omo River delta and adjacent northernmost part of the lake. Interestingly, K. gordoni has not been found in any of 547 members of the genus Clarias (abundant and sympatric with H. bidorsalis) examined in this study (Appendix 1), further supporting the presumed narrow host specificity. Mature tapeworms morphologically indistinguishable from K. gordoni were found in the the catfish Schilbe mystus (BMNH 1957.12.30.34– 38), which might represent atypical host that became infected via predation., Published as part of Kuchta, Roman, Burianová, Alena, Jirkú, Miloslav, Chambrier, Alain, Oros, Mikuláš, Brabec, Jan & Scholz, Tomáš, 2012, Bothriocephalidean tapeworms (Cestoda) of freshwater fish in Africa, including erection of Kirstenella n. gen. and description of Tetracampos martinae n. sp., pp. 1-35 in Zootaxa 3309 on pages 9-12, DOI: 10.11646/zootaxa.3309.1.1, http://zenodo.org/record/280992, {"references":["Woodland, W. N. F. (1937) Some cestodes from Sierra Leone. II. A new caryophyllaeid, Marsypocephalus, and Polyonchobothrium. Proceedings of the Zoological Society of London B, 107, 189 - 197.","Tadros, G. (1967) On a new cestode Bothriocephalus prudhoei sp. nov. from the Nile catfish Clarias anguillaris with some remarks on the genus Clestobothrium June [sic], 1899. Bulletin of the Zoological Society of Egypt, 21, 74 - 88.","Fagbenro, O. A., Adedire, C. O., Owoseeni, E. A. & Ayotunde E. O. (1993) Studies on the biology and aquaculture potential of feral catfish Heterobranchus bidorsalis (Geoffroy St. Hilaire 1809) (Clariidae). Tropical Zoology, 6, 67 - 79.","Kuchta, R. & Scholz, T. (2007) Diversity and distribution of fish tapeworms of the \" Bothriocephalidea \" (Eucestoda). Parassitologia, 49, 129 - 146.","Tadros, G. (1968) A re-description of Polyonchobothrium clarias (Woodland, 1925) Meggitt, 1930 (Bothriocephalidae: Cestode) with a brief review of the genus Polyonchobothrium Diesing, 1854 and the identity of the genera Tetracampos Wedl, 1861, Senga Dollfus, 1935, and Oncobothriocephalus Yamaguti, 1959. Journal of Veterinary Science of the United Arab Republic, 5, 53 - 84.","Khalil, L. F. (1973) Some helminth parasites from African freshwater fishes with the description of two new species. Revue de Zoologie et de Botanique Africaines, 87, 795 - 807.","Wedl, K. (1861) Zur Helminthenfauna Agyptens. III. Nematoda. IV. Cestoda. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften Wien, 44, 463 - 482.","Janicki, C. (1926) Cestodes s. str. aus Fischen u. Amphibien. In: L. A. Jagerskiold (Ed.), Results of the Swedish Zoological Expedition to Egypt and the White Nile 1901. Part V. The Library of the Royal University of Uppsala, 66 pp.","Kuchta, R., Scholz, T. & Bray, R. A. (2008 b) Revision of the order Bothriocephalidea Kuchta, Scholz, Brabec & Bray, 2008 (Eucestoda) with amended generic diagnoses and keys to families and genera. Systematic Parasitology, 71, 81 - 136.","Pouyaud, L., Sudarto & Paradis, E. (2009) The phylogenetic structure of habitat shift and morphological convergence in Asian Clarias (Teleostei, Siluriformes: Clariidae). Journal of Zoological Systematics and Evolutionary Research, 47, 344 - 356."]}

Ichthybothrium ichthybori Khalil, 1971 (Figs. 3, 6, 9, 13, 14) Syns: “immature Bothriocephalus ” of Woodland (1936); “unidentified ptychobothriid cestode” of Khalil (1969, 1971a). Type host: Ichthyborus besse (Joannis) (Characiformes: Ichthyboridae). Other definitive host: Ichthyborus quadrilineatus (Pellegrin) (new host). Type locality: White Nile near Khartoum, the Sudan. Distribution: Upper Guinea – Sierra Leone (Moa River in Mano or Pujehun); Nile basin – the Sudan (Khartoum and Kostí). Prevalence and intensity of infection: Nile, Kostí, the Sudan; 67 %, n = 6, intensity 15–35 (Khalil 1971 b); 50 %, n = 4, intensity 1–8 (present study). Type material: Holotype – incomplete worm on 2 slides (USNPC 71666) and 2 paratypes – 3 worms on 3 slides (USNPC 71667). Material studied: Type material (USNPC 71666, 71667); vouchers: several slides collected by W.N.F. Woodland from I. quadrilineatus in Sierra Leone (BMNH 1977.3.28.147– 179); new material: 9 specimens from Kostí, the Sudan (IPCAS C- 455). Published records: Woodland (1936); Khalil (1969, 1971a, b); Kuchta et al. (2008 b). Re-description (based on 3 mature and 6 immature worms from Kostí; measurements of holotype in brackets): Bothriocephalidea, Bothriocephalidae. Strobila up to 10.0 [4.2] cm long; maximum width 3.7 [1.5] mm. External and internal segmentation weakly developed; segments variable in shape from wider than long to longer than wide, acraspedote (Fig. 13). Two pairs of longitudinal osmoregulatory canals; dorsal canals narrow (diameter up to 10); ventral canals wide (diameter up to 15), connected by transverse anastomoses. Inner longitudinal musculature weakly developed, formed by few muscle fibres. Surface of strobila uniformly covered with capilliform filitriches. Scolex oval to lanceolate, much narrower than anterior part of strobila (Figs. 3, 13), 490–630 [495] long by 340–530 wide [413]. Apical disc absent. Bothria shallow, elongate, 240–360 [360] long by 140–200 [186] wide (Fig. 3). Surface of scolex covered with capilliform filitriches; tumuliform globular structures not observed (Fig. 6). Neck absent, first segments appear immediately posterior to scolex (Fig. 13). Strobila with immature segments very long, representing up to 93 % [58 %] of total length. Mature segments, i.e. with spermatozoa in vas deferens, of two markedly different forms (morphotypes): longer than wide (486–614 by 374–529) [749 – 1,187 by 564–700] with one genital complex per segment or much wider than long (315–557 by 3,370–3,745) with two or three genital complexes per segment; segment length/width ratio 1.08–1.09 [1.19–1.78] or 0.08–0.16 (Figs. 13, 14). Gravid segments longer than wide (736–831 by 374–523) [1,080–1,280] or wider than long (425–621 by 2,550–3,360); segment length/width ratio 1.59–1.96 [1.50–1.74] or 0.13–0.24. Testes medullary, oval, 37–69 [48–57] in number per proglottid (110–120 testes in segments with duplicate or triplicate proglottides), 36–50 long by 22–43 wide [77–96 by 54–69], forming 2 narrow longitudinal bands (20–36 [22–29] testes per band), confluent between segments, absent medially and near lateral margins (Fig. 14). Cirrussac large, thin-walled (thickness of sac wall up to 32 [35]), spherical, 107–143 in diameter [106–135]; equatorial to pre-equatorial (at 42–50 % [41–48 %] of length of mature segment from anterior margin). Vas deferens forms numerous loops anterolateral to cirrus-sac, cirrus unarmed, opening into genital atrium (Fig. 14). Genital pore dorsal, sub-median, pre-equatorial (at 8–12 % [4–10 %] of length of mature segment from anterior margin), enlarging in gravid segments (Fig. 14). Ovary asymmetrical, elongated or V-shaped, lobulate, 41–143 [131–213] long by 135–394 [186–251] wide (Fig. 14). Vagina a straight, thin-walled tube, with sac-like extension in proximal part, 64–83 [44–60] long by 23– 52 [26–27] wide, opens posterior to cirrus-sac into genital atrium; vaginal sphincter absent (Fig. 14). Vitelline follicles numerous, small, spherical, 10–25 [30–45] long by 17–35 [25–30] wide, cortical, form 2 wide longitudinal bands confluent between segments, separated medially, rarely connected by several follicles in postovarian region (Fig. 14). Uterine duct forms numerous tightly coiled loops, filled with eggs, enlarged in gravid segments. Uterine pore thick-walled, opens slightly posterior to midlength of uterus. Eggs oval, thin-walled, operculate, unembryonated, 38–46 [45–50] long by 27–30 [32–33] wide (Fig. 9). Remarks: This tapeworm was described by Khalil (1971 b) from the Sudan and placed in a newly proposed genus Ichthybothrium Khalil, 1971. It is characterized by a lanceolate scolex, which is narrower than the anterior end of the strobila (Figs. 3, 13) and possesses shallow bothria, but no apical disc. The strobila has weakly developed segmentation and its large proportion (more than 90 % in some specimens) consists of immature proglottides without genital complexes (called “pregenital region” by Khalil 1971 b). Specimens recently collected near the type locality differ markedly from the type specimens in the shape of the segments and number of genital complexes per segment, with duplication or even triplication of genital complexes (proglottides) within mature and gravid segments (Kuchta et al. 2008 b). Marked differences in the size of the worms may be influenced by crowding effect, because worms in more heavily infected hosts (intensity of infection 8–35 worms) are smaller (maximum length 6.2 cm) and consist of just a few mature and gravid segments, which are longer than wide (length/width ratio 1.08–1.96: 1) and usually contain only one genital complex (Khalil 1971 b) (Fig. 13). In contrast, tapeworms from mono- or light infections are up to 10 cm long, their strobila is formed by numerous mature and gravid segments, which are much wider than long (length/width ratio 0.08–0.24: 1), and each of them contains two or even three genital complexes (Kuchta et al. 2008 b; present study) (Fig. 14). Khalil (1971 b) correctly reported the cortical position of vitelline follicles in the diagnosis of I. ichthybori, but erroneously described vitelline follicles to be medullary in the diagnosis of the genus. The surface of the scolex and strobila was studied using SEM for the first time and a uniform pattern of capilliform filitriches was observed. Khalil (1971 b) also reported the presence of “minute hair-like processes” on the scolex and “long hair-like processes” around the uterine pore, which may also represent capilliform filitriches (Fig. 6). However, tumuliform globular structures commonly observed in many bothriocephalideans (Kuchta et al. 2008 b), including all other studied species, are missing in I. ichthybori (Fig. 6). Ichthybothrium ichthybori is a rather frequent parasite of Ichthyborus besse, with high prevalence of infection (50–67 %) and a relatively high intensity of infection. However, its fish host is rare and thus only a limited number of records of this tapeworm exist in the literature. Examination of several slides with immature cestodes designated as “immature Bothriocephalus ” from Neoborus quadrilineatus (= Ichthyborus quadrilineatus) collected by W.N.F. Woodland in Sierra Leone (BMNH 1977.3.28.147– 179) (Woodland 1936) has shown that they belong to I. ichthybori., Published as part of Kuchta, Roman, Burianová, Alena, Jirkú, Miloslav, Chambrier, Alain, Oros, Mikuláš, Brabec, Jan & Scholz, Tomáš, 2012, Bothriocephalidean tapeworms (Cestoda) of freshwater fish in Africa, including erection of Kirstenella n. gen. and description of Tetracampos martinae n. sp., pp. 1-35 in Zootaxa 3309 on pages 6-8, DOI: 10.11646/zootaxa.3309.1.1, http://zenodo.org/record/280992, {"references":["Khalil, L. F. (1971 a) Check list of the helminth parasites of African freshwater fish. Commonwealth Agricultural Bureaux, Farnham Royal, England, 80 pp.","Woodland, W. N. F. (1936) Some cestodes from Sierra Leone. I. On Wenyonia longicauda sp. n. and Proteocephalus bivitellatus sp. n. Proceedings of the Zoological Society of London B, 106, 931 - 937.","Khalil, L. F. (1969) Studies on the helminth parasites of freshwater fishes of the Sudan. Journal of Zoology, 158, 143 - 170.","Khalil, L. F. (1971 b) Ichthybothrium ichthybori gen. et sp. nov. (Cestoda: Pseudophyllidea) from the African freshwater fish Ichthyborus besse (Joannis, 1835). Journal of Helminthology, 45, 371 - 379.","Kuchta, R., Scholz, T. & Bray, R. A. (2008 b) Revision of the order Bothriocephalidea Kuchta, Scholz, Brabec & Bray, 2008 (Eucestoda) with amended generic diagnoses and keys to families and genera. Systematic Parasitology, 71, 81 - 136.","Baer, J. C. & Fain, A. (1958) Bothriocephalus (Clestobothrium) kivuensis n. sp., a cestode parasitic on a barbel of Lake Kivu. Annales de la Societe Royale Zoologique de Belgique, 88, 287 - 302.","Yamaguti, S. (1934) Studies on the helminth fauna of Japan. Part 4. Cestodes of fishes. Japanese Journal of Zoology, 6, 1 - 112."]}

Kirstenella Kuchta n. gen. Diagnosis: Bothriocephalidea, Bothriocephalidae. Medium-sized worms. Segmentation present. Segments trapeziform, craspedote. Scolex elongate. Bothria elongate, shallow. Apical disc present, as wide as, or slightly wider, than bothrial part of scolex, armed with large, simple hooks arranged in two lateral semicircles. Neck absent. Testes in two lateral fields, continuous between segments. Cirrus-sac thick-walled, spherical, large, representing as much as 1 / 6 – 2 / 5 of segment width; internal seminal vesicle present; cirrus unarmed. Genital pore dorsal, median. Ovary compact, bilobed, transversely elongate. Vagina posterior to cirrus-sac, enlarged distally. Vitelline follicles cortical, in two lateral fields continuous between segments. Uterine duct sinuous, S-shaped, enlarging in gravid segments. Uterus small, oval. Uterine pore median. Eggs operculate, unembryonated. In freshwater siluriforms. Africa. Etymology: The new genus is named in honour of Kirsten Jensen, University of Kansas at Lawrence, USA, for her outstanding contribution to cestode systematics. Type and only species: Kirstenella gordoni (Woodland, 1937) Kuchta, n. comb. Remarks: The new genus is placed in Bothriocephalidae Lönnberg, 1889 because it possesses the median genital pore, follicular vitellaria, and a ventral uterine pore (Kuchta et al. 2008 a, b). It is typified by combination of the following characteristics: (1) scolex is elongate with a small apical disc armed with two lateral semicircles of hooks; (2) cirrus-sac is large (its width represents as much as 1 / 6 – 2 / 5 of the width of segments) and it is situated perpendicularly; (3) a low number of testes (less than 75 per segment); (4) vagina is enlarged distally; (5) vitellaria are cortical. Kirstenella differs from all but three bothriocephalid genera (Polyonchobothrium Diesing, 1854, Senga Dollfus, 1934 and Tetracampos Wedl, 1861) in the presence of an apical disc armed with two lateral semicircles of large hooks (Kuchta et al. 2008 b). The new genus can be distinguished from Tetracampos by the presence of cortical (instead of medullary) vitelline follicles and presence of operculate, thin-walled (vs. unoperculated, with hyaline membrane) eggs. The new genus can be distinguished from Polyonchobothrium and Senga mainly by the relative size of the cirrus-sac, which is markedly larger (ratio of its width to the width of segments 16–39 %) than that of species of Senga (ratio 4–10 %) and Polyonchobothrium (ratio 5–10 %), and is situated more perpendicularly in Kirstenella, whereas longitudinally or obliquely in the two remaining genera (Figs. 19, 22, 38, 39). Moreover, Polyonchobothrium has a markedly wider apical disc (350–510 µm) and much larger hooks (up to 190 μm) mostly distributed in 4 quadrants, whereas Kirstenella has a slender apical disc (230–300 μm) with smaller apical hooks (shorter than 90 μm) distributed in 2 semicircles. Hooks of Senga have an indistinct blade and handle (data not shown), whereas those of Kirstenella have the blade well differentiated from the handle (Figs. 16, 18)., Published as part of Kuchta, Roman, Burianová, Alena, Jirkú, Miloslav, Chambrier, Alain, Oros, Mikuláš, Brabec, Jan & Scholz, Tomáš, 2012, Bothriocephalidean tapeworms (Cestoda) of freshwater fish in Africa, including erection of Kirstenella n. gen. and description of Tetracampos martinae n. sp., pp. 1-35 in Zootaxa 3309 on pages 8-9, DOI: 10.11646/zootaxa.3309.1.1, http://zenodo.org/record/280992, {"references":["Woodland, W. N. F. (1937) Some cestodes from Sierra Leone. II. A new caryophyllaeid, Marsypocephalus, and Polyonchobothrium. Proceedings of the Zoological Society of London B, 107, 189 - 197.","Kuchta, R., Scholz, T., Brabec, J. & Bray, R. A. (2008 a) Suppression of the tapeworm order Pseudophyllidea (Platyhelminthes: Eucestoda) and the proposal of two new orders, Bothriocephalidea and Diphyllobothriidea. International Journal for Parasitology, 38, 49 - 55.","Kuchta, R., Scholz, T. & Bray, R. A. (2008 b) Revision of the order Bothriocephalidea Kuchta, Scholz, Brabec & Bray, 2008 (Eucestoda) with amended generic diagnoses and keys to families and genera. Systematic Parasitology, 71, 81 - 136.","Wedl, K. (1861) Zur Helminthenfauna Agyptens. III. Nematoda. IV. Cestoda. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften Wien, 44, 463 - 482."]}

Five new species in the tribe Anthocoptini of the Eriophyidae (Acari: Eriophyoidea) from China are described and illustrated. They are Vittacus shaoguanicus sp. nov. on Carmona retusa (Vahl) Masam. (Boraginaceae), Vittacus cornusis sp. nov. on Cornus officinalis Sieb. et Zucc. (Cornaceae), Aculops buddlejaes sp. nov. on Buddleja lindleyana Fort. (Buddlejaceae ), Aculodes tsukushiensis sp. nov. on Elymus tsukushiensis Honda var. transiens (Hack.) Osad (Poaceae) and Tegolophus spathaceae sp. nov. on Fargesia spathacea Franch. (Poaceae, Bambuseae). All the new species described herein are vagrants on the respective host plant.

Tetrapapillocephalus magnus (i) The holotype ex S. undosquamis from the Indian Ocean (Parasitological Institute, Russian Academy of Sciences, Moscow, Russia ��� GELAN): holotype (GELAN 696); (ii) plerocercoid on the slide with the holotype of Partitiotestis berycis Protasova & Parukhin, 1986 (GELAN 986)., Published as part of Kuchta, Roman, Scholz, Tom����, Vl��kov��, Romana, ����ha, Milan, Walter, Thorsten, Yuniar, Asri T. & Palm, Harry W., 2009, Revision of tapeworms (Cestoda: Bothriocephalidea) from lizardfish (Saurida: Synodontidae) from the Indo-Pacific region, pp. 55-67 in Zootaxa 1977 on page 57, DOI: 10.5281/zenodo.185185, {"references":["Protasova, E. N. & Mordvinova, T. N. (1986) A ptychobothriid cestode, Tetrapapillocephalus magnus n. g., n. sp. (Pseudophyllidea, Ptychobothriidae) from marine fish. Parazitologiya, 20, 313 - 316. [In Russian.]"]}

Bothriocephalidean cestodes reported from lizardfish (Saurida Valenciennes in Cuvier & Valenciennes) were revised on the basis of newly collected material from type-hosts and available typeand voucher specimens. Instead of nine taxa of four genera listed in the literature, only Oncodiscus sauridae Yamaguti, 1934 and Penetrocephalus ganapattii (Rao, 1954) are considered valid and thus both genera become monotypic. Both taxa are redescribed, including the first scanning electron micrographs and data on intraspecific variability. Both species are unique among bothriocephalid cestodes in the lateral position of the vagina in relation to the cirrus-sac. Oncodiscus sauridae found in Saurida longimanus Norman, S. nebulosa Valenciennes (new host record), S. tumbil (Bloch) and S. undosquamis (Richardson) differs from P. ganapattii found in S. micropectoralis Shindo & Yamanda and S. tumbil in the following characters: (i) scolex shape (fan-shaped, with crenulated bothrial margins in O. sauridae, replaced by a scolex deformatus in P. ganapattii); (ii) site of scolex attachment (in the intestinal lumen in the former taxon versus penetrating through the intestinal wall, and encysted on pyloric caeca in P. ganapattii); (iii) shape of segments (usually only slightly wider than long in O. sauridae versus very short, much wider than long in P. ganapattii); and (iv) number of testes (50–100 per segment in O. sauridae whereas fewer than 60 in P. ganapattii). Bothriocephalus sauridae Ariola, 1900 is considered a species inquirendum, although conspecificity with O. sauridae is very probable. The present study demonstrates the necessity of using standard procedures, especially adequate fixation methods of freshly collected cestode material (hot 4% formaldehyde solution is strongly recommended for morphological studies), because most previous descriptions were based on poor-quality material, including partly macerated worms missing hooklets on the apical disc of the scolex in O. sauridae.

Oncodiscus sauridae Yamaguti, 1934 Figs. 1���8, 14��� 18, 21, 24 Syns. O. fimbriatus Subhapradha, 1955; O. waltairensis Shinde, 1975; O. maharashtrae Jadhav & Shinde, 1981; Bothriocephalus indicus Ganapati & Rao, 1955; Tetrapapillocephalus magnus Protasova & Mordvinova, 1986 Type host: Saurida argyrophanes (now S. tumbil). Other hosts: S. longimanus, S. nebulosa (new host), S. undosquamis. Site of infection: Scolex in the pyloric caeca or embedded in the intestinal wall, strobila in the intestinal lumen. Type locality: Inland Sea, Tarumi-ku, Kobe, Hyōgo Prefecture, Kyūshū, Japan, Pacific Ocean. Distribution: Indian Ocean (India, Indonesia, Kuwait, Mozambique); Pacific Ocean (Australia, Japan, New Caledonia). Prevalence: In Indonesia, S. longimanus ��� 12 %; S. tumbil ��� 9 %; S. undosquamis ��� 5 %. In New Caledonia, 9 % of S. undosquamis were infected (J.-L. Justine ��� unpublished data). References: Subhapradha (1955); Devi (1975); Khalil & Abu-Hakima (1985); Protasova & Mordvinova (1986). Re-description: Bothriocephalidea, Bothriocephalidae. Medium-sized worms, up to 130 (90) mm long. Strobila with craspedote segments with prominent posterolateral wing-like projections, overlapping following segment (Figs. 5, 16, 17). External segmentation present, but may be incomplete (missing medially). Two pairs of longitudinal osmoregulatory canals present. Longitudinal musculature well developed, formed by numerous bundles of muscle fibres (Fig. 7). Surface covered with small filiform microtriches similar in shape and size. Scolex oval, irregularly-shaped or fan-shaped (Figs. 1, 2, 4, 14, 15), usually compressed laterally, 1,120���3,140 (1,710���4,000) long by 860 ��� 2,380 (1,360���2,190) wide. Bothria well developed, with conspicuously crenulated margins turned laterally (Figs. 1, 2, 4, 14, 15). Apical disc well developed, 310���890 (780 ��� 1,250) wide, 140���400 (160���370) high, consisting of two lateral lobes, armed with up to four irregular rows of tiny hooklets (Figs. 2, 3, 15, 18, 24); hooklets easy to detach, especially in dead specimens (Figs. 1, 4), 21���26 (12���24) long, with largest hooklets in second and third rows, and diminishing in size from top of lateral lobes to junction of lobes, where rows may be interrupted. Neck absent, first segments starting immediately posterior to scolex (Figs. 1, 15). Immature segments much wider than long, 25���110 (45���165) long by 400 ��� 1,510 (230���850) wide. Mature segments wider than long, 50���810 (80���380) long by 310 ��� 1,715 (755 ��� 1,070) wide (Fig. 5). Gravid segments wider than long to rectangular or, exceptionally, slightly longer than wide, 190���2000 (180���700) long by 615���1865 (985���1520) wide (Figs. 16, 17). Testes medullary, oval, 50���100 (60���100) in number per segment, 22���66 (30���61) long by 11���38 (25���50) wide, in two lateral fields, usually separated medially, continuous between segments (Fig. 5). Cirrus-sac submedian, irregularly alternating, thick-walled, round or slightly oval, 70���215 (85���135) in diameter, just anterodorsal to ovary (Figs. 5, 6, 8). Cirrus armed with filiform microtriches. Cirrus opening to small genital atrium lateral to median line of segment (Fig. 7). Genital pore dorsal, median, slightly postequatorial (Fig. 5). Ovary transverse elongate, bilobed, lobulate, median, near posterior margin of segment, 200���630 (225���460) wide (Figs. 5, 6). Vagina tubular, dorsal to ovarian bridge, arched lateral or anterolateral to cirrussac (Figs. 5, 6). Vitelline follicles numerous, cortical, circumsegmental (Fig. 7), oval, 13���40 (7���28) in diameter, in two lateral fields, often separated medially, reaching to posterolateral projections (Figs. 5, 7). Uterine duct well developed, sinuous, ending in muscular, thick-walled uterine sac situated medially (Figs. 5, 6); sac markedly enlarged in last gravid segments. Uterine pore medioventral, at distance from anterior margin of segment or almost equatorial. In some segments, pore extremely large due to collapse of wall of uterus (Fig. 16). Eggs oval, operculate, unembryonated, 55���64 (60���63) long by 32���45 (39���42) wide, with operculum 11���20 in diameter (Fig. 21). Remarks: Oncodiscus sauridae was described by Yamaguti (1934) from Saurida argyrophanes (now S. tumbil) and redescribed by Khalil & Abu-Hakima (1985), who collected new material and added morphological data to the original description. The total length of 9 mm was given by Yamaguti (1934) for the holotype, but this specimen measures in fact 90 mm (present study). Besides O. sauridae, another three species of Oncodiscus, namely O. fimbriatus, O. waltairensis, and O. maharashtrae, were erected. They were differentiated mainly by the shape of the scolex, the presence or absence of a median longitudinal furrow, the presence or absence hooklets on the apical disc, the number of testes and the presence or absence of an operculum in the eggs (Subhapradha 1955; Shinde 1975; Jadhav & Shinde 1981). However, these characters are unsuitable for species differentiation in this group of cestodes. The shape of the scolex and the presence of furrows and hooklets on the apical disc are not stable characters because they depend into a great extent on the state in which worms were fixed and on fixation procedure (Kuchta & Scholz 2007). In dead tapeworms or in worms that have been relaxed for too long in water, the hooklets detach (one of Yamaguti���s paratypes of O. sauridae lacks hooklets on the apical disc due to tissue destruction ��� Fig. 1), the scolex shape changes and tegumental furrows may appear as a consequence of deformation of the scolex, especially when fixation by pressure between two glass slides is used (Khalil & Abu- Hakima 1985; present study). The number of testes is variable in O. sauridae and may considerably differ even between segments of the same specimen. The maximum number of testes in O. sauridae was found to reach up to 100, which corresponds to the data from the original description (Yamaguti 1934). Jadhav & Shinde (1981) reported as many as 350 testes in one segment of O. maharashtrae, but it is apparent that the authors confused vitelline follicles with testes. Vitelline follicles enlarge in the posterior part of the strobila and may reach a similar size compared to that of the testes in gravid segments. Another feature which was used for species differentiation, i.e. the presence or absence of an operculum in the eggs, can easily be overlooked unless scanning electron microscopy is used (Bray et al. 1994; Kuchta et al. 2008 a; present study). Yamaguti (1934) described a median longitudinal furrow on the scolex of O. sauridae. On the basis of the absence of that structure, Subhapradha (1955) proposed O. fimbriatus. However, this structure does not represent a species-distinguishing character. Such a furrow is missing in one of the paratypes of O. sauridae and it is much less developed in other specimens of the type series. This indicates that its presence depends on the level of worm contraction or fixation procedure (S. Yamaguti strongly flattened the worms, which may have caused unnatural deformations of the scolex and strobila). Khalil & Abu-Hakima (1985) synonymized O. fimbriatus and O. waltairensis with O. sauridae, but they considered O. maharashtrae to be a species inquirendum. This species was described from Pastinachus (Trygon) sephen (Forssk��l) (Dasyatidae) from off Bombay, India. Unlike Khalil & Abu-Hakima (1985), the present authors (Kuchta & Scholz 2007; Kuchta et al. 2008 b; this study) consider O. maharasthrae to be a synonym of O. sauridae which occurred in an unusual (postcyclic or accidental) host. All three species from India were described on the basis of only a few, apparently macerated and deformed specimens. In addition, the descriptions of the taxa from the Indian Ocean were superficial and no histological sections were made (Ariola 1900; Subhapradha 1955; Shinde 1975; Jadhav & Shinde 1981; Protasova & Mordvinova 1984). Bothriocephalus indicus was described from Saurida tumbil. It is obvious from its morphology that the authors erroneously placed these specimens in the genus Bothriocephalus Rudolphi, 1808, because species of this genus are devoid of a fan-shaped scolex with crenulated bothrial margins (Ganapati & Rao 1955). In fact, B. indicus is identical in its morphology with O. sauridae (the absence of hooklets on an apical disc is apparently due to poor state of specimens studied), parasitizes the same species of fish hosts and occurs in the same region. Therefore, the synonymy of B. indicus with O. sauridae, first proposed by Kuchta & Scholz (2007), is confirmed in the present study. Bothriocephalus sauridae was very briefly described on basis of specimens without a scolex that were collected from Saurida nebulosa off the eastern coast of Africa (Zanzibar) (Ariola 1900). The species was considered to be a species inquirendum (Protasova 1977; Kuchta & Scholz 2007; present paper), but its conspecificity with O. sauridae is very probable. In the same region, O. sauridae (reported to as Tetrapapillocephalus magnus) was found in S. undosquamis by Protasova & Mordvinova (1986). Morphological and morphometrical examination of the specimens of Tetrapapillocephalus magnus (holotype and voucher), which are both in poor condition, shows a high similarity with O. sauridae. The taxa differ only in the absence of tiny hooklets on the apical disc in the former. However, hooklets can be easily lost during handling or when dead tapeworms are collected (Khalil & Abu-Hakima 1985; present study). The scolex of Tetrapapillocephalus magnus (= O. sauridae) illustrated in the original description (Protasova & Mordvinova 1986) belongs to another specimen, not representing the holotype as indicated (Fig. 4). This immature specimen was found in Beryx splendens Lowe; it is mounted onto the same slide with the holotype of Partitiotestis berycis Protasova & Parukhin, 1986 (GELAN 986). Numerous immature tapeworms conspecific with T. magnus were found in this host, but only one specimen was mounted (Protasova ��� pers. com.). Besides this report, no further data exist on the occurrence of O. sauridae in berycid fish (Kuchta & Scholz 2007; Kuchta et al. 2008 b) and it is possible that B. splendens represents an accidental or paratenic host only. It is interesting to note that we examined some unusual material from clupeid fish (Chirocentrus dorab) from the Japan Sea, in the collection of Yamaguti (MPM), collected 19.vi. 1928. These specimens are conspecific with O. sauridae (also identified as Oncodiscus by Yamaguti), but this finding was never published. In addition, no other occurrence of any bothriocephalid tapeworm has been reported from this host. This finding may represent misidentification of the host, or an accidental or paratenic host., Published as part of Kuchta, Roman, Scholz, Tom����, Vl��kov��, Romana, ����ha, Milan, Walter, Thorsten, Yuniar, Asri T. & Palm, Harry W., 2009, Revision of tapeworms (Cestoda: Bothriocephalidea) from lizardfish (Saurida: Synodontidae) from the Indo-Pacific region, pp. 55-67 in Zootaxa 1977 on pages 58-61, DOI: 10.5281/zenodo.185185, {"references":["Yamaguti, S. (1934) Studies on the helminth fauna of Japan. Part 4. Cestodes of fishes. Japanese Journal of Zoology, 6, 1 - 112.","Subhapradha, C. K. (1955) Two new bothriocephalids from marine fish Saurida tumbil (Bl.). Proceedings of the Indian Academy of Science, Series B, 41, 20 - 30.","Shinde, G. B. (1975) On a new species of Oncodiscus Yamaguti, 1934 from marine fish in Andhra Pradesh, India. Marathwada University Journal of Science, 14, 339 - 343.","Jadhav, B. V. & Shinde, G. E. (1981) A new species of Oncodiscus Yamaguti, 1934 (Cestoda: Tetraphyllidea) from India. Proceedings of the Indian Academy of Parasitology, 2, 26 - 27.","Ganapati, P. N. & Rao, K. H. (1955) On Bothriocephalus indicus sp. nov. (Cestoda) from the gut of marine fish, Saurida tumbil (Bloch). Journal of the Zoological Society of India, 7, 177 - 181.","Protasova, E. N. & Mordvinova, T. N. (1986) A ptychobothriid cestode, Tetrapapillocephalus magnus n. g., n. sp. (Pseudophyllidea, Ptychobothriidae) from marine fish. Parazitologiya, 20, 313 - 316. [In Russian.]","Devi, R. P. (1975) Pseudophyllidean cestodes (Bothriocephalidae) from marine fishes of Waltair Coast Bay of Bengal. Rivista di Parassitologia, 36, 279 - 286.","Khalil, L. F. & Abu-Hakima, R. (1985) Oncodiscus sauridae Yamaguti, 1934 from Saurida undosquamis in Kuwait and a revision of the genus Oncodiscus (Cestoda: Bothriocephalidae). Journal of Natural History, 19, 783 - 790.","Kuchta, R. & Scholz, T. (2007) Diversity and distribution of fish tapeworms of the \" Bothriocephalidea \" (Eucestoda). Parassitologia, 49, 21 - 38.","Bray, R. A., Jones, A. & Andersen, K. I. (1994) Order Pseudophyllidea Carus, 1863. In: Khalil, L. F., Jones, A. & Bray, R. A. (Eds.) Keys to the cestode parasites of vertebrates. CAB International, Wallingford, pp. 205 - 247.","Kuchta, R., Scholz, T., Brabec, J., Jirsova, D. & Gustinelli, A. (2008 a) Bothriocephalidean tapeworms (Cestoda) from the blackfish, Centrolophus niger. Folia Parasitologica, 55, 111 - 121.","Kuchta, R., Scholz, T. & Bray, R. A. (2008 b) Revision of the order Bothriocephalidea Kuchta, Scholz, Brabec & Bray 2008 (Eucestoda) with amended generic diagnosis and key to families and genera. Systematic Parasitology, 71, 81 - 136.","Ariola, V. (1900) Revisione della famiglia Bothriocephalidae s. str. Archives de Parasitologie, 3, 369 - 484.","Protasova, E. N. (1977) Cestodes of fish - Bothriocephalata. Principles of cestodology. Vol. 8. Nauka, Moscow, 298 pp. [In Russian.]"]}

Oncodiscus sauridae (i) type material (9 specimens) ex Saurida argyrophanes (now S. tumbil) from the Inland Sea and East China Sea, Japan (Meguro Parasitological Museum, Tokyo, Japan ��� acronym MPM): holotype [MPM 23176 (SY 3042)], paratypes [MPM 23176 (SY 3043), MPM 23177 (SY 3165 - 67)]; (ii) three specimens ex Chirocentrus dorab (Forssk��l) from Toyama Bay, Toyama Prefecture, Honshū, Japan, Japan Sea (collected by S. Yamaguti) [MPM 23178 (SY 5616 - 18)]; (iii) six specimens ex S. tumbil from Pelabuhan Ratu, Java, Indonesia (helminthological collection of the Institute of Parasitology, BC AS CR, Česk�� Budějovice ��� IPCAS C- 456 / 1; Natural History Museum, London, UK ��� BMNH 2008.9.5.3; Invertebrate Collection, Mus��um d'Histoire Naturelle, Gen��ve, Switzerland ��� MHNG INVE 62588); (iii) six specimens ex S. longimanus from Pelabuhan Ratu, Java, Indonesia (IPCAS C- 456 / 2; BMNH 2008.9.5.6 and 2008.11.7.1; Zoologisches Museum Berlin ��� ZMB 7447 and 7448); (iv) one specimen ex S. undosquamis from Pelabuhan Ratu, Java, Indonesia and two specimens from off Noum��a, New Caledonia (IPCAS C- 456 / 3; Mus��um National d'Histoire Naturelle, Paris, France ��� MNHN JNC 502, 2588); (v) one specimens from S. nebulosa from off Noum��a, New Caledonia (MNHN JNC 2474)., Published as part of Kuchta, Roman, Scholz, Tom����, Vl��kov��, Romana, ����ha, Milan, Walter, Thorsten, Yuniar, Asri T. & Palm, Harry W., 2009, Revision of tapeworms (Cestoda: Bothriocephalidea) from lizardfish (Saurida: Synodontidae) from the Indo-Pacific region, pp. 55-67 in Zootaxa 1977 on page 57, DOI: 10.5281/zenodo.185185

Penetrocephalus ganapattii (Rao, 1954) Rao, 1960 Figs. 9���13, 19, 20, 22, 23 Syns. Bothriocephalus ganapattii Rao, 1954; Bothriocephalus penetratus Subhapradha, 1955. Type host: Saurida tumbil. Other hosts: S. micropectoralis, S. undosquamis. Site of infection: The scolex and neck (scolex deformatus) is coiled within a cyst on the wall of pyloric caeca, whereas the strobila lies freely in the intestinal lumen. Type locality: Coast of Waltair, Bay of Bengal, Andhra Pradesh, India, Indian Ocean. Distribution: Indian Ocean (India, Indonesia). Prevalence: In Indonesia, S. micropectoralis ��� 27 %; S. tumbil ��� 5 %. References: Subhapradha (1955); Rao (1960); Devi (1975); Radhakrishnan et al. (1983). Re-description: Bothriocephalidea, Bothriocephalidae. Large worms, up to 125 mm long. Strobila with markedly craspedote segments with prominent lateral wing-like projections (Figs. 10, 19, 23). External segmentation present, may be incomplete (missing medially). Two pairs of longitudinal osmoregulatory canals present. Longitudinal musculature well developed, forming numerous bundles of muscle fibres (Figs. 10, 11). Surface covered with small filiform microtriches similar in shape and size. Scolex unarmed, 700���750 long by 150���200 wide, changed to scolex deformatus (Fig. 9, 12). Apical disc absent. Anterior part of body including neck often degenerated, becoming filiform (Fig. 9), up to 60 mm long, penetrating through intestinal wall, and encysted on pyloric caeca. Segments short, much wider than long (up to 14 times in mature segments). Immature segments 50���620 long by 100 ��� 2,540 wide. Mature segments 40���220 long by 1,080���2,900 wide (Fig. 10). Gravid segments 70���620 long by 890 ��� 3,140 wide (Figs. 19, 23). Testes medullary, oval, 40���60 in number per segment, 10���60 long by 10���40 wide, in two lateral fields separated medially, continuous between segments (Fig. 10). Cirrus-sac submedian (Fig. 11), irregularly alternating, round or slightly oval, 65���195 in diameter, just anterior to ovary (Fig. 10, 13). Cirrus unarmed, covered with long filiform microtriches (Fig. 20), opening to small genital atrium. Genital pore dorsal, slightly submedian (Figs. 11, 23). Ovary transverse elongate, lobulate, median, 240���720 wide (Fig. 10). Vagina tubular, lateral or anterolateral to cirrus-sac. Vitelline follicles cortical, oval, 10���40 in diameter, numerous, in two lateral fields separated medially, reaching to posterolateral projections (Fig. 10, 11). Uterine duct sinuous, leading to thick-walled, median uterus (Figs. 10, 13), dilated ventrally in gravid segments. Uterine pore ventral, median, at distance from anterior margin of segment (Fig. 19). Eggs oval, operculate, unembryonated, 50���74 long by 33���42 wide; operculum 9���18 in diameter (Fig. 22). Remarks: The original description of Bothriocephalus ganapattii was very brief (Rao, 1954), which led Protasova (1977) and Bray et al. (1994) to consider B. ganapattii a nomen nudum. However, the species is clearly recognizable on the basis of its original description because it possesses a scolex deformatus that penetrates the intestine and forms a cyst in the mesentery or in the liver (Rao 1954). Subhapradha (1955), probably not aware of Rao���s (1954) publication, described an apparently conspecific cestode, Bothriocephalus penetratus. Both taxa parasitize the same fish host in the same region and are morphologically indistinguishable. Therefore, they are considered synonymous (Rao 1960; Kuchta & Scholz 2007; Kuchta et al. 2008 b). Rao (1960) proposed a new genus, Penetrocephalus, to accommodate B. ganapattii (the type- and only species was misspelled as P. ganapatii). The genus was invalidated by Protasova (1977) and Bray et al. (1994), but resurrected by Kuchta et al. (2008 b); the validity of the genus is also supported by molecular data (Kuchta 2007, fig. 10)., Published as part of Kuchta, Roman, Scholz, Tom����, Vl��kov��, Romana, ����ha, Milan, Walter, Thorsten, Yuniar, Asri T. & Palm, Harry W., 2009, Revision of tapeworms (Cestoda: Bothriocephalidea) from lizardfish (Saurida: Synodontidae) from the Indo-Pacific region, pp. 55-67 in Zootaxa 1977 on pages 61-64, DOI: 10.5281/zenodo.185185, {"references":["Rao, K. H. (1954) A new bothriocephalid parasite (Cestoda) from the gut of the fish Saurida tumbil (Bloch). Current Science, 23, 333 - 334.","Rao, K. H. (1960) Studies on Penetrocephalus ganapatii, a new genus (Cestoda: Pseudophyllidea) from the marine teleost, Saurida tumbil (Bloch). Parasitology, 50, 155 - 163.","Subhapradha, C. K. (1955) Two new bothriocephalids from marine fish Saurida tumbil (Bl.). Proceedings of the Indian Academy of Science, Series B, 41, 20 - 30.","Devi, R. P. (1975) Pseudophyllidean cestodes (Bothriocephalidae) from marine fishes of Waltair Coast Bay of Bengal. Rivista di Parassitologia, 36, 279 - 286.","Radhakrishnan, S., Nair, N. B. & Balasubramanian, N. K. (1983) Adult cestode infection of the marine teleost fish Saurida tumbil (Bloch). Acta Ichtyologica et Piscatoria, 13, 75 - 97.","Protasova, E. N. (1977) Cestodes of fish - Bothriocephalata. Principles of cestodology. Vol. 8. Nauka, Moscow, 298 pp. [In Russian.]","Bray, R. A., Jones, A. & Andersen, K. I. (1994) Order Pseudophyllidea Carus, 1863. In: Khalil, L. F., Jones, A. & Bray, R. A. (Eds.) Keys to the cestode parasites of vertebrates. CAB International, Wallingford, pp. 205 - 247.","Kuchta, R. & Scholz, T. (2007) Diversity and distribution of fish tapeworms of the \" Bothriocephalidea \" (Eucestoda). Parassitologia, 49, 21 - 38.","Kuchta, R., Scholz, T. & Bray, R. A. (2008 b) Revision of the order Bothriocephalidea Kuchta, Scholz, Brabec & Bray 2008 (Eucestoda) with amended generic diagnosis and key to families and genera. Systematic Parasitology, 71, 81 - 136."]}

Penetrocephalus ganapattii (i) two specimens ex S. tumbil from Pelabuhan Ratu, Java, Indonesia (IPCAS C- 462 / 1; MHNG INVE 62589; ZMB 7449). (ii) three specimens ex S. micropectoralis from Pelabuhan Ratu, Java, Indonesia (IPCAS C- 462 / 2; BMNH 2008.9.5.7��� 8; MHNG INVE 62590; ZMB 7450). Illustrations were made using a drawing attachment on the Olympus BX 51 microscope with Nomarski interference contrast. Types or vouchers of the taxa described by Indian authors were not available and they most probably do not exist or have never been deposited (B. Jadhav ��� personal comm.)., Published as part of Kuchta, Roman, Scholz, Tom����, Vl��kov��, Romana, ����ha, Milan, Walter, Thorsten, Yuniar, Asri T. & Palm, Harry W., 2009, Revision of tapeworms (Cestoda: Bothriocephalidea) from lizardfish (Saurida: Synodontidae) from the Indo-Pacific region, pp. 55-67 in Zootaxa 1977 on page 57, DOI: 10.5281/zenodo.185185

Clestobothrium gibsoni n. sp. is described from the intestine of the bullseye grenadier Bathygadus macrops (Macrouridae) collected in the northeastern Gulf of Mexico. The new species can be distinguished from both C. crassiceps (Rudolphi, 1819), the type-species of the genus, and C. neglectum (Lonnberg, 1893), the second species in the genus, by its unique ovary, which is transversely elongate and tapered on both ends rather than being U-or H-shaped, and by having more testes (60-65 vs 40-50 and 35-45 per proglottid). The new species is also compared to Bothriocephalus acheilognathi Yamaguti, 1934 and B. kivuensis Baer & Fain, 1954, two species with similar scoleces which have previously been assigned to Clestobothrium . The generic diagnosis of Clestobothrium is amended to include species with either anoperculate or operculate eggs. This is the first report of an adult tapeworm from a species of Bathygadus , and the first report of a species of Clestobothrium Luhe, 1899 (Bothriocephalidae) from the Gulf of Mexico.

Oncodiscus sauridae Yamaguti, 1934 is described and figured in detail from specimens collected from Saurida undosquamis (new host) and S. tumbil in the Arabian Gulf and Australian waters, respectively (new geographical localities). O. fimbriatus and O. waltairensis are regarded as synonyms of O. sauridae. O. maharashtrae is regarded as species inquirenda. The diagnosis of the genus Oncodiscus is emended.

Observations have been made on the morphology ofBothriocephalus scorpiifrom littoral fishes in Britain, with particular reference to the reproductive system and the ultrastructure of the tegument. Two different kinds of microtriches, and ciliated sense organs, have been found. The shedding of proglottides by pseudoapolysis is described.

Adult pseudophyllidean tapeworms, Bothriocephalus cuspidatus, were recovered from the intestines of three gizzard shad, Dorosoma cepedianum, collected from two Illinois lakes, Crab Orchard and Carlyle. This represents a new host record.

Anantrum histocephalum sp. n. is described from Synodus lucioceps, California lizardfish. The larger worm size, mushroom-shaped scolex, vitelline follicle distribution, and larger eggs distinguish this species from A. tortum. The tapeworm was observed in 8.5% of 164 lizardfish from the coastal waters of Los Angeles County.

The specimens described in the following paragraphs were collected from material secured by the German Soutlh Pole Expedition of 1901-1903. They were sent to Professor Pintner, head of the department of invertebrate zoology in the Un-iversity of Vienna, who placed the material at my disposition for determination and description. The specimens, though now over a quarter of a century old, are in a remarkable state of preservation. Aside from a slight bleaching and some brittleness due to the action oL -the preservative fluid, the specimens are quite suitable for investigation. Mention must be made here of the procedure follbwed in staining sections. The ordinary paraffin method was used, and the sections cut to a thickness of ten micra. Borax carmine as a staining agent proved unsatisfactory. The best results were obtained by using Ehrlich's hematoxylin preceded by iron alum. The sections were takeut through the alcohol series and then treated with distilled water. They were then immersed in a bath of iron alumr for about twenty minutes, washed in distilled water and stained for about five minutes with Ehrlich's hematoxylin. The stain is washed off under running tap water. Distilled water does not give results. Sections prepared in this way show every detail of structure, especially the nucleus. Up to the present, about fourteen species of Bothriocephalidae have been described. But this number must be reduced to eleven because three specific names are evidently synonyms. The species under consideration is Dibothriocephalus perfoliatus, (Diphyllobothrium clavatum, Railliet.) It was founid in sevent specimens of a species of seal,-Leptony