New arrangement of three genera of fish tapeworms (Cestoda: Proteocephalidae) in catfishes (Siluriformes) from the Neotropical Region: taxonomic implications of molecular phylogenetic analyses.

Tapeworms (Cestoda: Proteocephalidae) are the dominant component of communities of intestinal parasites in pimelodid and other catfishes (Siluriformes) from South America. Even though these parasites have been studied intensively over more than one century, molecular taxonomy and phylogenetics have questioned their morphology-based classification, thus raising doubts about the systematic value of traits commonly used to circumscribe individual taxa. In the present study, members of three morphologically well-characterized genera of proteocephalids from pimelodid (Hemisorubim platyrhynchos and Sorubim lima) and auchenipterid (Ageneiosus inermis) catfishes from the Paraná or Amazon River basins were subjected to DNA sequencing of the large subunit nuclear ribosomal RNA (lsrDNA) and complete mitochondrial cytochrome c oxidase subunit I (COI). Phylogenetic analyses revealed the sister relationship between Manaosia bracodemoca and Mariauxiella piscatorum, and among Mariauxiella pimelodi and Ageneiella brevifilis. As a result, Mar. piscatorum and A. brevifilis are transferred to Manaosia and Mariauxiella, respectively, as Manaosia piscatorum n. comb. and Mariauxiella brevifilis n. comb., and the genus Ageneiella is suppressed. Diagnoses of Manaosia and Mariauxiella are amended. In addition, the present study revealed misidentification of tapeworms whose sequences are deposited in the GenBank database.

Description of the first species of Pseudoendorchis (Cestoda: Proteocephalidae) with uniloculate suckers from the pimelodid catfish Megalonema platycephalum, with comments on the taxonomic importance of the terminal vagina.

The proteocephalid genus Pseudoendorchis (Cestoda: Onchoproteocephalidea) has recently been proposed to accommodate seven species/species-level lineages of tapeworm parasites of catfishes (Siluriformes) in the Neotropical region, based on integration of genetic information, and morphological and ecological data. Its members are typified by having a large Mehlis' gland, representing more than 1/5 (usually 1/4-1/2) of proglottid width, and the vagina always anterior to the cirrus-sac. Critical examination of previously unstudied museum cestodes tentatively designated as Pseudoendorchis sp. 3 from Megalonema platycephalum (Pimelodidae) in the Peruvian Amazon made it possible to formally describe this taxon and to differentiate it from all congeneric species. The new species, which is the first parasite of M. platycephalum ever recorded, is characterised mainly by having the scolex bearing four uniloculate suckers (biloculate in all nominal species of the genus), the lowest relative surface of the ovary (ratio of its surface to that of the whole proglottid) among species of the genus (< 9% versus > 11%, usually 15-20%), and an extraordinarily large Mehlis' gland (its diameter represents 41-50% of proglottid width). Morphology of the terminal portion of the vaginal canals in proteocephalids is briefly discussed.

Revision of Acanthotaenia von Linstow, 1903 (Cestoda: Proteocephalidae), parasites of monitors (Varanus spp.), based on morphological and molecular data.

A morphological and molecular phylogenetic study of proteocephalid tapeworms of the genus Acanthotaenia von Linstow, 1903, parasites of monitors (Varanidae), was carried out. The type species, A. shipleyi von Linstow, 1903, which was originally described based on an immature specimen from Sri Lanka, is redescribed based on new material from the type host, Varanus salvator, in Sri Lanka, Malaysia, and Vietnam, and its neotype is designated. In addition, Acanthotaenia susanae n. sp. is described from Varanus nebulosus in Vietnam. The new species differs from congeners by the large size of the scolex, width of the rostellum and the number of testes. New molecular data (sequences of lsrDNA and cox1) revealed Acanthotaenia paraphyletic with the inclusion of Australotaenia bunthangi de Chambrier & Scholz, 2012, a parasite of Enhydris enhydris (Ophidia: Homalopsidae) in Cambodia. Molecular data confirm a wide distribution of A. shipleyi (isolates from Malaysia and Vietnam were almost identical) and indicate a strict host specificity (oioxeny) of individual species of the genus. Type specimens of four species made it possible to supplement their morphological descriptions. A survey of all species of Acanthotaenia recognised as valid is presented and the following taxonomic changes are proposed: Acanthotaenia pythonis Wahid, 1968 described from the green python, Morelia viridis, in a zoo, is transferred to Kapsulotaenia as Kapsulotaenia pythonis (Wahid, 1968) n. comb., because it possesses intrauterine eggs grouped in capsules. Acanthotaenia gracilis (Beddard, 1913) from Varanus varius in Australia is considered to be species inquirenda because its original descriptions did not contain sufficient data for adequate circumscription and differentiation from congeners and type material was not available. Generic diagnosis of Acanthotaenia is amended and a key to its seven species is provided.

Amendment of Rostellotaenia Freze, 1963 (Cestoda: Proteocephalidae) from African monitors (Varanus spp.) and a redescription of its type-species, R. nilotica (Beddard, 1913).

Based on morphological evaluation of type-specimens and new material collected in three species of monitor lizards (Squamata: Varanidae), the diagnosis of the monotypic Rostellotaenia Freze, 1963 (Proteocephalidae: 'Acanthotaeniinae') is amended. The genus differs from Acanthotaenia von Linstow, 1903, with which it was previously synonymised, by the shape of the scolex with a wide, dome-shaped rostellum, the posterior extent of the uterus, which does not overpass the ovarian isthmus posteriorly, well-developed retractor muscles connecting the rostellum with the neck region, absence of a well-developed, ring-like vaginal sphincter, shorter and wider proglottides, and geographical distribution (Ethiopian region only). The type- and only species, R. nilotica (Beddard, 1913), is redescribed and two new definitive hosts, Varanus albigularis Daudin and V. exanthematicus (Bosc), are reported. Host specificity of R. nilotica, which exhibits remarkable morphological variability, is stenoxenous, i.e. it occurs in several congeneric species of African monitor lizards, unlike species of Acanthotaenia, which are strictly specific (oioxenous).

Untangling convoluted taxonomy of Chambriella Rego, Chubb & Pavanelli, 1999 (Cestoda: Proteocephalidae), with erection of Riggenbachiella n. g. and the description of a new species from pimelodid catfishes in the Neotropical Region.

As part of a complex revision of proteocephalid cestodes parasitic in freshwater bony fishes of the Neotropical Region, the genus Chambriella Rego, Chubb & Pavanelli, 1999 is redefined based on detailed examination of type-specimens and newly collected material of both nominal species of the genus. This examination revealed that the type-species C. agostinhoi (Pavanelli & Santos, 1992) from Zungaro jahu (Ihering) (type-host) and Z. zungaro (Humboldt) is indistinguishable from Lenhataenia megacephala (Woodland, 1934) from Sorubimichthys planiceps (Spix & Agassiz) (all hosts Siluriformes: Pimelodidae), the type- and only species of the genus. New molecular data (partial sequences of the large subunit nuclear ribosomal RNA gene) support the conspecificity of these taxa. As a result, Lenhataenia de Chambrier & Scholz, 2008 becomes a junior synonym of Chambriella and its type-species, C. agostinhoi, a junior synonym of C. megacephala (Woodland, 1934) n. comb. The second species of Chambriella, C. paranaensis (Pavanelli & Rego, 1989) from Hemisorubim platyrhynchos (Valenciennes), is transferred to a new genus, Riggenbachiella n. g., as R. paranaense (Pavanelli & Rego, 1989) n. comb. Riggenbachiella amazonense n. sp. (syn. Chambriella sp. of de Chambrier & Scholz, 2008) from S. planiceps (type-host), Phractocephalus hemioliopterus (Bloch & Schneider) and Z. zungaro is described and designated as the type-species of the new genus. Riggenbachiella n. g. is placed in the subfamily Monticelliinae Mola, 1929, because of the cortical position of the testes, ovary, vitelline follicles and uterus, and is mainly characterised by the possession of a sigmoid cirrus-sac with voluminous, chambered internal seminal vesicle, and bi-loculate suckers.

An annotated list of the species of Gangesia Woodland, 1924 (Cestoda: Proteocephalidea), parasites of catfishes in Asia, with new synonyms and a key to their identification.

An annotated list of tapeworms of the genus Gangesia Woodland, 1924 (Cestoda: Proteocephalidea), parasites of siluriform fishes in Asia, is provided. Based on the morphological examination of museum specimens and newly collected material from China, Japan and Russia, as well as the results of a previous revision of the Indomalayan species, only eight of more than 50 nominal taxa are considered to be valid. These are: from India and neighbouring countries, Gangesia bengalensis (Southwell, 1913) (type-species), G. agraensis Verma, 1928, both from Wallago attu (Bloch & Schneider) (Siluridae), G. macrones Woodland, 1924 from Sperata seenghala (Sykes) (Bagridae) and G. vachai (Gupta & Parmar, 1988) from different catfishes (type-host Eutropiichthys vacha (Hamilton); Schilbeidae), and, from the Palaearctic, G. margolisi Shimazu, 1994, a parasite of Silurus biwaensis (Tomoda) (Siluridae) in Japan, G. oligonchis Roitman & Freze, 1964 from Tachysurus fulvidraco (Richardson) (Bagridae) in Russia, and G. parasiluri Yamaguti, 1934 and G. polyonchis Roitman & Freze, 1964, both from Silurus asotus L. (Siluridae) in Japan and Russia, respectively. The poorly known G. oligonchis is redescribed. Seven new synonyms are proposed: G. chauhani Mathur & Srivastav, 2000, G. wallaguae Pradhan, Kulkarni, Kale & Wakle, 2010 and G. shivajiraoi Dhole, Waghmare & Chavan, 2012 are synonymised with G. agraensis; G. striatusii Bhure & Nanaware, 2012 and Silurotaenia govindii Sawarkar, 2013 with G. macrones; G. spasskajae Demshin, 1987 with G. polyonchis; and Silurotaenia spinula Chen, 1984 with Postgangesia orientalis Akhmerov, 1969. Gangesia pseudobagrae Chen, 1962 is considered to be a species inquirenda, whereas G. chauhani Mathur, 1992 and G. dineshei Jaysingpure, 2002 are recognised as unavailable names. An amended generic diagnosis of Gangesia and a key to the identification of its recognised species are also provided.

A new species of Pseudocrepidobothrium (Cestoda: Proteocephalidea) from Pseudoplatystoma reticulatum (Pisces: Siluriformes) in the Paraná River basin (Argentina).

This study describes the proteocephalidean tapeworm Pseudocrepidobothrium chanaorum sp. n. (Proteocephalidae: Proteocephalinae), which was found in the intestine of Pseudoplatystoma reticulatum (Eigenmann et Eigenmann) from the Colastiné River, a tributary of the Paraná River. The new species differs from the two other species of the genus, P. eirasi (Rego et de Chambrier, 1995) and P. ludovici Ruedi et de Chambrier, 2012, parasites of Phractocephalus hemioliopterus (Bloch et Schneider) from the Amazon River in Brazil, in having fewer proglottides (4-8 without ventral appendages vs 7-12 with ventral appendages and 20-36 without ventral appendages, respectively), a smaller scolex (350-450 µm wide vs 495-990 µm and 515-1020 µm wide, respectively), in the total number of testes (21-25 vs 21-51 and 37-79, respectively), a cirrus-sac usually directed anteriorly if the vagina is posterior to the cirrus-sac vs transversely situated in the known species. The study of the tegumental surface of Pseudocrepidobothrium spp. revealed the presence of four types of microtriches: papilliform, acicular and capilliform filitriches, and gladiate spinitriches. The three species have a similar microthrix pattern, with minor differences on the immature proglottis surface. Pseudocrepidobothrium chanaorum sp. n. is the ninth proteocephalid reported from P. reticulatum.

Taxonomic status of Woodland's enigmatic tapeworms (Cestoda: Proteocephalidea) from Amazonian catfishes: back to museum collections.

Poorly known proteocephalidean cestodes of peculiar morphology, described by Woodland (1934) from pimelodid catfishes in Amazonia, Brazil, were studied. Re-examination of their type-specimens and evaluation of newly-collected material from Brazil and Peru made it possible to clarify their taxonomic status. Brayela karuatayi (Woodland, 1934), the type-species of the monotypic Brayela Rego, 1984, which has never been recorded since its original description, is redescribed and its scolex morphology, which has been misinterpreted in the original description, was studied using scanning electron microscopy (SEM). The actual definitive host of B. karuatayi is not a species of Glanidium Lütken (Auchenipteridae), but coroatá, Platynematichthys notatus (Jardine) (Pimelodidae). Peru is a new geographical record for B. karuatayi. The definitive host of other two proteocephalidean cestodes, Megathylacus jandia Woodland, 1934 and Proteocephalus jandia Woodland, 1934, is not a species of Rhamdia Bleeker (family Heptapteridae), but the pimelodid Zungaro zungaro (Humboldt) [syn. Paulicea luetkeni (Steindachner)]. Proteocephalus jandia is in fact conspecific with Travassiella avitellina Rego & Pavanelli, 1987, type-species of Travassiella Rego & Pavanelli, 1987. As a result, a new combination, Travassiella jandia (Woodland, 1934), is proposed. Megathylacus jandia Woodland, 1934 is considered conspecific with M. brooksi Rego & Pavanelli, 1985 described from the congeneric host [Zungaro jahu (Ihering)] from the Paraná River in Brazil; the latter species becomes its new junior synonym. The validity of M. travassosi Pavanelli & Rego, 1992, a parasite of Pseudoplatystoma corruscans (Spix & Agassiz) in the Paraná River basin in Brazil, is confirmed by a study of its type- and voucher specimens. The present account provides strong arguments to always study museum specimens in taxonomic studies; it also represents an evidence of the importance of depositing types and vouchers in internationally recognised collections. Based on the new information provided in this study, the generic diagnoses of Brayela and Megathylacus Woodland, 1934 are amended.

Tapeworms (Cestoda: Proteocephalidae) of the gars (Lepisosteidae), living fossils in America, including proposal of a new genus and a new species.

A new genus, Cordicestus, is proposed to accommodate proteocephalid tapeworms parasitising gars (Lepisosteiformes: Lepisosteidae) in North and Central America that were previously placed in the polyphyletic genus Proteocephalus Weinland, 1858. The new genus differs from other proteocephalid genera by the particular morphology of the scolex, which is small, protrudes apically but has no apical organ, and bears flat, heart-shaped (= cordis) suckers. In addition, the species of the new genus have an elongated cirrus sac with an almost straight internal vas deferens and wide, sinuous ventral osmoregulatory canals with secondary canals directed outwards. The type species of the new genus, Cordicestus singularis (La Rue, 1911) n. comb., is redescribed based on new material from the shortnose gar, Lepisosteus platostomus Rafinesque (type host), and the spotted gar, L. oculatus Winchell, in the United States. Cordicestus rafaeli n. sp. is described from the tropical gar, Atractosteus tropicus Gill, in Mexico. The new species differs from its relatives primarily by the presence of craspedote proglottids (acraspedote in other species) and some biometric features. All species of Cordicestus are revised, including unidentified specimens from A. tropicus and the Cuban gar A. tristoechus (Bloch and Schneider) in Nicaragua and Cuba, respectively, which may be new species, and a key to the identification of these taxa is provided. Molecular data available for two nominal species of the new genus indicate the possible existence of another species of Cordicestus in Lepisosteus in the USA.

A new genus and species of the Monticelliinae (Eucestoda: Proteocephalidea), a parasite of Pseudoplatystoma fasciatum (Pisces: Siluriformes) from the Paraná River basin (Argentina), with comments on microtriches of proteocephalideans.

This paper describes Regoella brevis gen. n. et. sp. n. (Proteocephalidea: Monticelliinae), a parasite of the intestine of the barred sorubim Pseudoplatystoma fasciatum (Linnaeus) from the Paraná River basin. The new genus is placed in the Monticelliinae because of the cortical position of the genital organs. It differs from all known genera included in the Monticelliinae by the following combination of characters: 1) a quadrangular scolex with a truncated conical apex and formed by four lobes separated by grooves; 2) uniloculate suckers of inverted triangular shape possessing a small cone-shaped projection at each corner of the anterior margin; 3) strobila consisting of a low number of proglottides; 4) testes arranged in one dorsal field; 5) a cirrus-sac, which represents more than one half of the proglottis width, cirrus surrounded by conspicuous chromophilic gland cells; 6) a butterfly-shaped and strongly lobulate ovary; and 7) formation of uterus of type 2. The examination of the tegument surface with scanning electron microscopy revealed the occurrence of three types of microtriches: acicular and capilliform filitriches and gladiate spinitriches. The new species is the eighth proteocephalidean reported from P. fasciatum, six of which are commonly found in the Amazon and Paraná River basins.

Macrobothriotaenia ficta (Cestoda: Proteocephalidea), a parasite of sunbeam snake (Xenopeltis unicolor): example of convergent evolution.

The poorly known proteocephalidean cestode Macrobothriotaenia ficta (Meggitt, 1931) from the sunbeam snake Xenopeltis unicolor (Ophidia: Xenopeltidae) is redescribed on the basis of re-examination of its type specimens from Burma (Myanmar), and vouchers from Thailand and Vietnam. The peculiar morphology of the scolex, which is formed by four pedunculate lobe-bearing pincer-shaped suckers, is described for the first time using scanning electron microscopy. In scolex morphology, M. ficta closely resembles phyllobothriidean cestodes, parasites of elasmobranchs. However, this similarity does not reflect phylogenetic relatedness of these cestodes but instead presents an example of convergent morphological evolution of attachment organs of unrelated groups of cestodes that parasitize different groups of vertebrates. Besides scolex morphology, the genus is characterised by the possession of a very large cirrus-sac, which may reach up to the midline of proglottides, few testes (less than 60), vitelline follicles limited to the dorsal side of proglottides, a large vaginal sphincter, and eggs with a three-layered embryophore covered with rounded projections. Numerous errors in the diagnosis of M. ficta, which appeared in the literature as a result of uncritical compilation of data without examination of original material, are corrected. Multilocus phylogenetic analysis of nuclear ribosomal RNA genes ssr- and lsr-DNA and mitochondrial genes rrnL and cox1 place this species among other snake-parasitizing proteocephalideans of the genus Ophiotaenia. The convergent evolution of scolex morphology across distantly related taxa is discussed.

Discovering high species diversity of Ophiotaenia tapeworms (Cestoda: Proteocephalidae) of watersnakes (Colubridae) in North America.

Tapeworms of the genus Ophiotaenia La Rue, 1911 parasitize herptiles (= amphibians and 'reptiles') throughout the world, with about 100 species recognised as valid. In the present work, the North American species found in watersnakes (Colubridae) are reviewed. An examination of the holotype of Ophiotaenia perspicua La Rue, 1911, the type species of the genus, and other specimens from Nerodia rhombifer (Hallowell) revealed that two species were used for the species description. The 'true' O. perspicua has a small scolex and small, round suckers. This species is redescribed based on new material from Oklahoma, USA. The other species from N. rhombifer, Ophiotaenia laruei n. sp., has a larger scolex and larger, almost triangular suckers. Examination of the types of O. variabilis (Brooks, 1978) from N. rhombifer and N. cyclopion (Duméril, Bibron et Duméril) from Louisiana, USA has revealed that it is a mixture of two or more species. Because of poor quality of these specimens, it is not possible to adequately characterise O. variabilis, which is considered a species inquirenda. In addition, two new species are described from Nerodia fasciata confluens (Blanchard). Ophiotaenia currani n. sp. from Mississippi, USA is characterised by elongate, narrow proglottids, few testes, and a relatively long cirrus sac. Ophiotaenia tkachi n. sp. from Louisiana, USA is characterised by relatively short and wide proglottids, more testes, and an unusual terminal part of the vagina with folds. Morphologically similar tapeworms of N. fasciata confluens, N. erythrogaster (Förster), N. sipedon (Linnaeus), and Agkistrodon piscivorus (Lacépède) (Viperidae) from Arkansas and Oklahoma, USA, which are genetically nearly identical, are considered to be conspecific with O. tkachi n. sp. The present data suggest a high, previously undescribed species diversity of proteocephalid tapeworms in watersnakes in North America, and generally strict host specificity of these tapeworms.

A global survey of tapeworms (Cestoda: Proteocephalidae) of 'true' frogs (Amphibia: Ranidae), including a tabulated list of all proteocephalids parasitising amphibians.

Proteocephalid tapeworms of frogs of the family Ranidae ('true' frogs) are reviewed with emphasis on their species diversity, host specificity and geographical distribution. New molecular data (nuclear lsrDNA and mitochondrial COI sequences) are presented for tapeworms of four species of ranid frogs in North America, including the poorly known Ophiotaenia saphena Osler, 1931 of Rana clamitans Latreille and R. catesbeiana (Shaw), which is redescribed using new material from Arkansas, USA. Tapeworms of R. sphenocephala (Cope) and R. pipiens Schreber, the latter previously identified as O. saphena, represent another, putative new species, but are not formally described due to insufficient available material. Proteocephalus papuensis Bursey, Goldberg et Kraus, 2008 from Sylvirana supragrisea (Menzies) is transferred to Ophiotaenia La Rue, 1911 as a new combination. After a critical review of the literature, only nine nominal species of Ophiotaenia are recognised as valid, which is in contrast to the large number of ranid frogs (> 440 spp.). The reasons for this striking disparity are briefly discussed, and a key based on morphology is presented for the identification of all species of Ophiotaenia from the Ranidae. Molecular data are available for only two taxa from North America that form a monophyletic group. The relationships among tapeworms of ranid frogs occurring in other zoogeographical regions are not yet known. The taxonomic status of Batrachotaenia Rudin, 1917, which was erected to accommodate proteocephalids from amphibians, is also discussed. To facilitate future studies, a tabulated summary of all 32 species of proteocephalids belonging to three genera reported from amphibians (frogs and salamanders) is presented, with information on their hosts, distribution, and taxonomically important characters, including key measurements.

TAPEWORMS (CESTODA: OPHIOTAENIA) FROM THE NORTHERN COTTONMOUTH (AGKISTRODON PISCIVORUS).

The northern cottonmouth, Agkistrodon piscivorus (Lacépède, 1789) (Viperidae: Crotalinae), occurs in the southeastern United States and is one of the few semiaquatic vipers in the world. Three proteocephalid tapeworms of the genus OphiotaeniaLa Rue, 1911 (Cestoda), have been described from this venomous snake. A critical evaluation of type specimens and tapeworms recently collected from A. piscivorus has revealed that only 2 species, Ophiotaenia marenzelleri (Barrois, 1898) and Ophiotaenia grandisLa Rue, 1911 (new synonym Ophiotaenia agkistrodontis [Harwood, 1933]), are specific parasites of this crotaline viper; both species are redescribed here. Ophiotaenia grandis was originally described from a mixture of 2 species: 'true' O. grandis, which is indistinguishable from O. agkistrodontis described 22 yr later and thus considered to be a junior synonym of O. grandis, and O. marenzelleri. Ophiotaenia marenzelleri, also reported from the pygmy rattlesnake, Sistrurus miliarius (Linnaeus, 1766), is a larger tapeworm with a massive scolex, a large cirrus sac, a very large, annular vaginal sphincter near the equatorial gonopore, and an oval, 3-layered embryophore surrounded by a nearly square hyaline outer membrane. Ophiotaenia grandis is much smaller and more slender and has a narrower scolex, a smaller cirrus sac and vaginal sphincter, a 2-layered embryophore, and a distinctly pre-equatorial gonopore. In addition to O. marenzelleri and O. grandis, other Ophiotaenia species typical of semiaquatic snakes (Colubridae: Natricinae) may be present in the northern cottonmouth, which serves only as a postcyclic or occasional host. There is also a tabular summary of 18 species of Ophiotaenia from semiaquatic snakes worldwide, with information on their hosts, distribution, and taxonomically important characters, including key measurements.

A new species of Australotaenia (Cestoda: Proteocephalidea) from a snake in Cambodia: host switching or postcyclic parasitism in a distant region?

Australotaenia de Chambrier et de Chambrier, 2010 has been proposed to accommodate two species of proteocephalidean cestodes from hylid frogs (Litoria spp.) in Australia. Recently, apparently congeneric cestode, for which the name A. bunthangi sp. n. is proposed, was found in the homalopsid snake Enhydris enhydris (Schneider) (Serpentes: Homalopsidae) from South-East Asia (Cambodia). This finding indicates a much wider range of definitive hosts of species of this genus, i.e. amphibians and reptiles, which is exceptional among proteocephalideans. Postcyclic parasitism, i.e. predation of the definitive host infected with sexually mature parasites, cannot be excluded but does not seem to be probable. In addition, the occurrence of A. bunthangi in the former Indochina extends the range of the geographical distribution of the genus to another zoogeographical region. The new species differs from both species of Australotaenia in the relative size of an apical organ, the diameter of which equals to that of suckers (versus much smaller in the remaining species, in which the width of the apical organ represents less than 2/3 of the diameter of the suckers), much smaller scolex and suckers (width 150 microm and diameter of suckers 50-55 microm versus 245-420 microm and 100-140 microm, respectively), and longer body (224 mm versus 57-121 mm). In addition, A. bunthangi differs from A. hylae (Johnston, 1912) (type-species of the genus) by the number of testes (46-64 versus 74-106 in A. hylae) and by the ovary width/proglottis width ratio (55-65% versus 68-71% in A. hylae). Australotaenia bunthangi differs from A. grobeli de Chambrier et de Chambrier, 2010 by relative size of the cirrus-sac (its length represents 18-24% of the width of the proglottis versus 27-33% in A. grobeli) and by the diameter of the embryophore (25-27 microm versus 18-23 microm in A. grobeli).

Ophiotaenia bungari n. sp. (Cestoda), a parasite of Bungarus fasciatus (Schneider) (Ophidia: Elapidae) from Vietnam, with comments on relative ovarian size as a new and potentially useful diagnostic character for proteocephalidean tapeworms.

Ophiotaenia bungari n. sp. (Cestoda: Proteocephalidea) is described from the intestine of the banded krait Bungarus fasciatus (Schneider) (Ophidia: Elapidae) in Vietnam. The new species differs from all but three Ophiotaenia species parasitic in Asian reptiles in the possession of a glandular apical organ. It differs from O. andersoni Jensen, Schmidt & Kuntz, 1983 in the position of the vagina in relation to the cirrus-sac (anterior and posterior in O. bungari versus anterior only in the latter species), in the cirrus-sac/proglottis width ratio (29-38 versus 50%) and by having more testes (100-150 versus 42-116 in O. andersoni); from O. chattoraji Srivastava, 1980 in the number of uterine diverticula (50-65 versus 10-26) and in the cirrus-sac/proglottis width ratio (29-38 versus 22%); and from O. rhabdophidis (Burt, 1937) by having more uterine diverticula (50-65 versus 30-45), by the cirrus-sac/proglottis width ratio (29-38 versus 20-25%) and by the width of the scolex (360-420 versus 130-187 µm). The taxonomic importance of the relative size of the ovary (i.e. the ratio of the ovarian size in relation to that of the entire proglottis), a character previously not used in the systematics of proteocephalidean cestodes, is discussed. Comparison of measurements of all of the nominal species of Ophiotaenia La Rue, 1911 and Proteocephalus Weinland, 1858 (c.135 species) has shown that the ovary of species parasitic in snakes in the Americas, Africa, Asia and Australia is not only considerably smaller than that of congeneric species from European hosts, but also smaller than in all species of Proteocephalus parasitic in teleost fishes throughout the world.

A new tapeworm from Compsophis infralineatus (Pseudoxyrhophiidae), an endemic snake of Madagascar: Scratching the surface of undiscovered reptilian parasite diversity.

A new species of proteocephalid cestodes, provisionally assigned to the polyphyletic genus Ophiotaenia La Rue, 1911 (Cestoda: Proteocephalidae), is described from Compsophis infralineatus (Günther) (Serpentes: Pseudoxyrhophiidae) endemic to Madagascar. Ophiotaenia oreae n. sp. differs from all African and Asian species of Ophiotaenia by possessing more uterine diverticula (68-82 on one side). It is also characterised by the absence of an apical organ, the relative sizes of the cirrus-sac and ovary, the almost equatorial position of the gonopore, the diameter of the embryophore, and other biometric characteristics. Phylogenetic relationships of the new species indicate its relatedness to Indomalayan and Australasian proteocephalids from reptiles. Ophiotaenia oreae n. sp. formed a well-supported clade composed of species of Australophiotaenia de Chambrier, Beveridge and Scholz, 2018 from Australian snakes, Macrobothriotaenia ficta (Meggitt, 1931) from Xenopeltis unicolor Reinwardt in Boie from Vietnam, Ophiotaenia sp. from Trimeresurus flavomaculatus (Gray) from the Philippines, and Ophiotaenia bungari de Chambrier, Binh and Scholz, 2012 from Bungarus fasciatus (Schneider) from Vietnam. The only proteocephalid from Madagascan snakes sequenced so far, Ophiotaenia lapata Rambeloson, Ranaivoson and de Chambrier, 2012 from Madagascarophis colubrinus (Schlegel), does not form a monophyletic group with the new species. The actual species diversity of reptilian cestodes in Madagascar is undoubtedly underestimated. Because of the assumed strict (oioxenous) host specificity of reptilian proteocephalids and rich fauna of snakes occurring in Madagascar, it is plausible to expect the existence of dozens new species of proteocephalids on this island.

Ritacestus gen. n. (Cestoda: Proteocephalidea) and redescription of R. ritaii comb. n., a parasite of Rita rita (Siluriformes) in India.

A new genus, Ritacestus, is proposed to accommodate Ritacestus ritaii (Verma, 1926) comb. n. (syn. Proteocephalus ritaii), a parasite of the catfish Rita rita (Hamilton) in India. The new genus, which is placed in the Gangesiinae, is characterized by (i) a small, subspherical scolex formed by four large lobes separated from one another by longitudinal grooves, with a large, widely oval to pyriform rostellum-like apical organ, larger than suckers and possessing an apical hemispherical depression; (ii) paramuscular and cortical position of some vitelline follicles (most follicles are situated medullary); (iii) ventral and dorsal bands of vitelline follicles usually uninterrupted ventral to terminal genitalia and reaching to the posterior margin of proglottides; (iv) the vagina always anterior to the cirrus-sac; (v) a large size of the body (length up to 51 cm); and (vi) development of the uterus of type 2. In its morphology, especially shape of the scolex and apical organ, and paramuscular and cortical position of some vitelline follicles, Ritacestus resembles Postgangesia Akhmerov, 1969, but differs in the presence of a genital atrium (both genital pores of Postgangesia are separate), the anterior position of the vagina (almost always posterior in the latter genus), position of vitelline follicles in cross sections (dorsal and ventral bands in Ritacestus versus only a lateral band in the latter genus), and dorsal excretory canals indistinguishable in mature and gravid proglottides of R. ritaii (well developed in Postgangesia spp.). The type and only species of the genus, R. ritaii, is redescribed on the basis of new material from the type host from the Ganges River basin in India and its neotype is designated.

Ophiotaenia echidis n. sp. (Cestoda: Proteocephalidae) from the saw-scaled viper, Echis carinatus sochureki Stemmler (Ophidia: Viperidae), one of the world's deadliest snakes, from the United Arab Emirates.

Ophiotaenia echidis n. sp. (Cestoda: Proteocephalidae) is described from the intestine of one of the world's deadliest snakes, the saw-scaled viper Echis carinatus sochureki Stemmler (Ophidia: Viperidae) in the United Arab Emirates. The new species differs from other species of the non-monophyletic Ophiotaenia by the position of testes in two longitudinal lines on both sides of the uterus, and by the large size of an embryophore (diameter of 44-55 µm versus less than 40 µm in other species). Phylogenetic reconstructions based on lsrDNA and concatenated lsrDNA + COI datasets place the new species among proteocephalids from unrelated zoogeographical realms but mostly infecting venomous snakes. In all analyses, O. echidis n. sp. exhibited a strongly supported sister relationship with O. lapata Rambeloson, Ranaivoson et de Chambrier, 2012, a parasite of a pseudoxyrhophiid snake endemic to Madagascar. Despite a shared close evolutionary history between these taxa, morphological synapomorphies remain unclear, which impedes the erection of a new genus to accommodate them. A list of the 71 tapeworms of the former, non-monophyletic subfamily Proteocephalinae, parasitising snakes and lizards, including species inquirendae, and the phylogenetically closely related Thaumasioscolex didelphidis from opossum, with selected characteristics, is also provided, together with a checklist of helminth parasites reported from E. carinatus.

Two new genera and two new species of proteocephalidean tapeworms (Eucestoda) from reptiles and amphibians in Australia.

The examination of the type series of Ophiotaenia Gallardi (Johnston, 1911) (syn. Proteocephalus gallardi Johnston, 1911) revealed that it is a mixture of two species of different genera. Lectotype of Ophiotaenia gallardi is designated and the species is redescribed on the basis of it, conspecific paralectotypes and additional materials. The remaining part of the type series belongs to Vandiermenia gen. n. (Acanthotaeniinae), with V Beveridgei sp. n. as the type- and only species. The new genus differs from all other acanthotaeniine genera, i.e. Rostellotaenia Freze, 1963, Acanthotaenia von Linstow, 1903 and Kapsulotaenia Freze, 1963, by the presence of cortical uterine stem and paramuscular vitelline follicles, particular structure of the internal longitudinal musculature (absent laterally and more developed than in the three above-mentioned genera) and testes limited in two fields separated medially. Type series of Ophiotaenia mjobergi (Nybelin, 1917) (syn. Crepidobothrium mjobergi Nybelin, 1917), O. amphiboluri (Nybelin, 1917) (syn. Crepidobothrium amphiboluri Nybelin, 1917), O. striata (Johnston, 1914) (syn. Acanthotaenia striata Johnston, 1914) and O. longmani Johnston, 1916 are revised and compared with Ophiotaenia gallardi. Australotaenia hylae (Johnston, 1912) comb. n. is proposed for Ophiotaenia hylae Johnston, 1912. Australotaenia gen. n. differs from the remaining genera of the subfamily Acanthotaeniinae by (1) the Type 2 of the formation of the uterus (sensu de Chambrier et al. 2004) (all the other acanthotaeniines have the Type 1 of uterine development), (2) the cortical position of the uterine stem (all the other genera have medullary uterine stem) and (3) the morphology of the internal longitudinal musculature, which is composed of few well-developed bundles of fibres (in contrast to the other genera). The new genus also differs from ãby eggs not in clusters, the presence of two testicular fields (versus one in Vandiermenia) and the structure of the longitudinal internal musculature with only 8-10 bundles (versus formed by numerous bundles and with the presence of secondary muscles in Vandiermenia). Ophiotaenia sp. sensu de Chambrier (2004), a parasite of Litoria moorei, is described as Australotaenia grobeli sp. n., which can be distinguished from Australotaenia hylae by the smaller number of testes (46-76 versus 74-106), greater cirrus-sac length/width of proglottis ratio (27-33% versus 17-19%) and the smaller ovary width/proglottis width ratio (55-63% versus 68-71%).