Redescription of Corallobothrium solidum (Cestoda: Proteocephalidea) and erection of a new genus, Essexiella, for tapeworms from channel catfishes (Ictaluridae)

The proteocephalidean tapeworm, Corallobothrium solidum, type species of the genus, is redescribed on the basis of the examination of its type specimens and extensive material recently collected from Malapterurus electricus (type host). Some morphological characteristics of taxonomic importance are reported for the first time, such as the presence of semispherical (U-shaped) sphincters on the external (outer) margin of the suckers, a vaginal sphincter, a well-developed seminal receptacle, and a unique morphology of the eggs. Corallobothrium solidum differs from the 2 remaining species of the genus, both parasitic in channel catfishes (Ictaluridae), in its scolex shape, morphology of its suckers, presence of longitudinal and transverse grooves on the body surface, dense network of excretory canals in the apical part of the scolex, morphology of the eggs, and uterine development. The non-monophyletic nature of Corallobothrium is further supported by molecular data (partial sequences of the 28S rRNA gene) because C. solidum and the 2 remaining species from ictalurids do not form a monophyletic assemblage. Therefore, Essexiella n. gen. is proposed to accommodate Essexiella fimbriatum new comb. (type and only species; syn. Corallobothrium fimbriatum) from channel catfish. Essexiella n. gen. differs from Corallobothrium, Megathylacoides, and Megathylacus by the absence of a sphincter in the suckers, from Corallotaenia by the shape of the scolex and the number and shape of proglottids, and from Paraproteocephalus by the structure of the uterus. The diagnosis of Corallobothrium, which becomes monotypic and restricted to electric catfishes in Africa, is emended. The remaining species of Corallobothrium, Corallobothrium parafimbriatum, is tentatively transferred to Corallotaenia as Corallotaenia parafimbriata n. comb., based on molecular data, small size of the strobila, and shape of the scolex.     

The evolution of the Proteocephalidea (Platyhelminthes,Eucestoda) based on an enlarged molecular phylogeny,with comments on their uterine development

We present a molecular phylogeny of the Proteocephalidea based on 28S rDNA sequence data that is a follow-up to the paper by Zehnder & Mariaux (1999). Twenty-three new sequences, including three outgroups are added in our new data-set. The Gangesiinae Mola, 1929 and the Acanthotaeniinae Freze, 1963 appear to be the most primitive clades. They are followed by a robust clade comprising the Palaearctic Proteocephalinae Mola, 1929 from freshwater fishes. The structure of the more derived clades, comprising most Neotropical and Nearctic species, is less resolved. At the nomenclatural level, we erect a new genus, Glanitaenia n. g. for G. osculata (Goeze, 1782) n. comb., previously Proteocephalus osculatus, and define an aggregate for the Palaearctic Proteocephalus Weinland, 1858. After a re-examination of all of the studied taxa, we identify two types of uterine development and show the importance of this character for the systematics of the order. Our phylogeny does not support the classical view of a Neotropical origin for the Proteocephalidea but rather favours an Old World origin of the group either in saurians or Palaearctic Siluriformes.     

Scyphophyllidium uruguayense n. sp. (Eucestoda: Tetraphyllidea) in Mustelus mento (Cope, 1877) (Chondrichthyes: Carcharhiniformes: Triakidae) from La Paloma, Uruguay.

A new species of Scyphophyllidium inhabits Mustelus mento near La Paloma, Uruguay. It resembles Scyphophyllidium giganteum from the Atlantic Ocean and specimens identified as S. giganteum from California by having anapolytic strobilae 155-258 mm long, 250-300 craspedote proglottids, scoleces 1.2-1.4 mm wide, necks 34-41 mm long, immature and mature proglottids wider than long, gravid proglottids wider than long to longer than wide, genital pores averaging 28% of proglottid length from the anterior end, relatively flat ovaries with digitiform lobes reaching the lateralmost extent of the testicular field, vitellaria in 2 fields converging toward the proglottid midline, straight and short cirrus sacs, and postvaginal vas deferens. The bothridia of the new species have accessory bothridial suckers that are smaller than those of California specimens; European specimens reportedly lack accessory bothridial suckers. The new species possesses a uterine duct that joins the uterus at the level of the genital atrium and ventral osmoregulatory ducts medial rather than lateral to the dorsal ducts, an arrangement described for Californian but not European specimens. It differs from both European and Californian specimens by having longer cirri, more testes per proglottid, prominent scales covering the neck, and vaginae and uterine ducts coiled immediately preovarially. Pithophorus, Marsupiobothrium, and Scyphophyllidium may form a clade.     

Molecular systematic analysis of the order Proteocephalidea (Eucestoda) based on mitochondrial and nuclear rDNA sequences

Two ribosomal DNA sequences were used to infer phylogenetic relationships among the Eucestoda order Proteocephalidea. A 437 bp portion of the 16S mitochondrial and a 1149 bp 5' portion of the nuclear large sub-unit rRNA molecule were sequenced for 53 proteocephalidean cestodes (representing nine subfamilies and 22 genera) and for one outgroup species. Parsimony and distance-based analyses of the two databases, alone and combined, failed to support the monophyly of the two traditionally accepted families, of numerous subfamilies (with the exception of the Rudolphielliinae and Othinoscolescinae which were validated in our analysis) and of various genera, including the genus Nomimoscolex (Woodland), Ophiotaenia (La Rue) as well as the type genus Proteocephalus (Weinland). Palaearctic Proteocephalus species nevertheless constituted a well-defined clade. The two genes globally yielded compatible results; however, the nuclear ribosomal gene provided a better resolution of relations among Proteocephalidea.     

Polymorphism and polytypy for pericentric inversions in 38-chromosome Mastomys (Rodentia, Murinae) and possible taxonomic implications.

Chromosome banding analysis (R- and C-bands) of two 38-chromosome Mastomys specimens originating from the Ivory Coast and Uganda revealed different numbers of autosome arms (NFa), equal to 51 and 60, respectively. Comparison of their chromosome banding patterns with those of Mastomys specimens from the Sudan (NFa = 41) and Senegal (NFa = 51-54), studied previously, showed that variation of the NFa from 40 to 60 throughout the species distribution is the result of a pericentric inversion polymorphism involving 3-12 chromosome pairs. At the population level, this variation is much narrower and never results from more than two chromosome pairs involved in inversion polymorphism. Taking into account that the NFa values recorded to date form a well-defined discontinuous row, we presume that introgressive hybridization between populations differing from each other by 3-5 to 11-12 pericentric inversions is interrupted. From there, the hypothesis of the existence of at least three cryptic species (designated provisionally as MER-1, MER-2, and MER-3) within 38-chromosome Mastomys populations previously assigned to M. erythroleucus can be made. It looks likely that one of them, possessing a karyotype with an NFa = 50-56, is widely distributed throughout sub-Saharan Africa and includes karyotyped populations from Senegal, the Ivory Coast, Mali, Benin, Cameroon, Zaire, and the Sudan. The second species (MER-2) includes the specimens karyotyped (NFa = 40-41) from Chad and the Sudan. Finally, a third tentative species (MER-3) corresponds to specimens with NFa = 59-60 found in East Zaire and Uganda, as well as possibly Mali and Chad.     

Palaearctic migrants in the central Sudan

A brief account is given of the topography and climate of the Sudan and a more detailed survey is made of the vegetation and habitats afforded to wintering Palaearctic migrants in the Sahel and Soudan savannah zones in the central Sudan. Climatic conditions during the migration seasons are mentioned and the utilization of habitats by wintering migrants is broadly surveyed.
The status of 196 Palaearctic migrant species is treated briefly. The basic observational data are tabulated to summarize observed patterns of migration in terms of date, relative abundance and region (for which purpose the central Sudan is divided into four from west to east). Hitherto unpublished field notes of the authors and their collaborators are used, together with dated observations (including specimens collected) of earlier authors, to provide the fullest data on each species.
Moreau's (1967) paradox is discussed as it relates to the Sahel and Soudan savannah zones in the central Sudan, together with the degree of competition with resident African species in those zones. Some disparities between the spring and autumn abundance and d. istribution of Palaearctic migrants are discussed in relation to 40 species ana some tentative views are advanced on the possible reasons for such disparities. Evidence is adduced which points to Lanius minor and Coraczus garrulus (and possibly Circus pygargus , partially at least) having loop migrations, south through the Sudan and north through Arabia, and Falco wespertinus south through the Sudan and north through West Africa. An explanation is offered for the greater volume of water birds observed migrating down-Nile in spring compared with their volume up-Nile in autumn, and for the reversed situation observed in the region west of the Nile for these same species.
Shelduck Tadorna tadorna is included in the Sudan list for the first time.     

Morphological differences and molecular phylogenetic relationship of two tapeworm species,Moniezia expansa and Moniezia benedeni,collected from domestic ruminants in northern Vietnam

Moniezia expansa and M. benedeni are two common tapeworm species of domestic ruminants over the world. However, their morphological and molecular data are available for limited specimens from a few countries. In the present study, we compared morphological characteristics of these two species collected from goats and cattle in northern Vietnam and analyzed their phylogenetic relationship based on the 5.8S and second internal transcribed spacer (ITS2) of nuclear ribosomal DNA and the mitochondrial cytochrome c oxidase subunit I gene (cox1). The two species are clearly distinguishable from one another not only in the morphological appearance of the interproglottidal glands but also in the gross appearance of mature and gravid proglottids. Molecular analyses revealed that the 5.8S-ITS2 sequences of Vietnamese M. expansa were highly similar (99.7%) to the sequences from Japan and India, and made a common clade, which was clearly distinct from M. benedeni of Vietnam. For cox1 sequences, Vietnamese M. expansa showed a high similarity to and were grouped with the sequences from Ethiopia and some sequences from Senegal and China to make a common clade, which was separated from the remaining clades of Senegal and China. The cox1 sequences of M. benedeni from China, Vietnam, and Senegal were far distant (10.0–15.9%) from each other. The results of this study suggest that more sequence data of Moniezia species with details of morphological features from various geographical locations should be obtained to clarify the taxonomic status of Moniezia species.     

Mathevolepis junlanae sp. n. (Cyclophyllidea: Hymenolepididae: Ditestolepidini), a new cestode species from shrews of Far East

Mathevolepis junlanae sp. n. from shrews (Sorex spp.) of Far East is described given. The new species occupies an intermediate position between the Palaearctic species M. petrotschenkoi Spassky, 1948 (total number of proglottids--4) and M. skrjabini (Sadovskaya, 1965) (10-12 segments in a series) by the number of synchronously developing proglottids (2-4). The new species differs from M. larbi Karpenko, 1982 by the structure and measurements of copulative apparatus in proglottids. In M. larbi, vagina S-shaped, vagina length 0.147-0.154 mm, cirrus length 0.146-0.149 mm, and cirrus bursa reaching aporal excretory vessels. M. junlanae sp. n. has cirrus bursa slightly crossing center line of sexually mature proglottids, cirrus about half as long (0.084-0.092 mm), and smaller sac-shaped vagina (0.075-0.10 mm). The new species differs from the Nearctic representative M. macyi (Locker and Rausch, 1952) by lesser measurements of cirrus bursa (0.12-0.14 mm versus 0.4 mm) and a fewer number of proglottids in series (2-4 versus 5-7).     

A new genus and species of proteocephalidean (Cestoda) from Clarias catfishes (Siluriformes: Clariidae) in Africa

A new proteocephalidean cestode is described from 2 catfishes, Clarias gariepinus (type host) and C. cf. anguillaris (Siluriformes: Clariidae), from Ethiopia (type locality), Sudan, Tanzania, and Zimbabwe, and a new genus, Barsonella, is proposed to accommodate it. The genus belongs to the Proteocephalinae because its genital organs (testes, ovary, vitellarium, and uterus) are situated in the medulla. Barsonella lafoni, the type and only species of the new genus, is characterized mainly by the possession of an additional opening of each sucker; circular musculature on the anterior margin of suckers, serving as a sphincter; a small thin-walled glandular apical organ; absence of well-developed osmoregulatory canals in mature, pregravid, and gravid proglottids; and a large strobila, up to 173 mm long and 3.2 mm wide. Species of Marsypocephalus Wedl, 1861 (Marsypocephalinae), other large-sized proteocephalidean tapeworms occurring sympatrically in African catfishes (Clarias and Heterobranchus) and also possessing a sphincter-like, circular musculature on the anterior part of suckers, differ from B. lafoni in the absence of an additional sucker opening and glandular apical organ, the cortical position of the testes, well-developed osmoregulatory canals throughout the strobila, and a large cirrus sac. Proteocephalus glanduligerus (Janicki, 1928), another cestode parasitic in Clarias spp. in Africa, is much smaller than B. lafoni (maximum length 15 mm), has suckers without additional opening and circular musculature on the suckers, a large-sized glandular organ, much larger than suckers, and well-developed osmoregulatory canals. Comparison of partial sequences of the 28S rRNA gene for 7 samples of B. lafoni from 2 different hosts and 4 localities in Ethiopia, Sudan, and Tanzania has shown a very low genetic variability. In a limited phylogenetic analysis, B. lafoni formed a clade with Corallobothrium solidum Fritsch, 1886 (Proteocephalidae: Corallobothriinae), an African electric catfish parasite. This clade was the sister group of almost all Neotropical taxa from pimelodid and other catfishes.     

Thaumasioscolex didelphidis n. gen., n. sp. (Eucestoda: Proteocephalidae) from the black-eared opossum Didelphis marsupialis from Mexico, the first proteocephalidean tapeworm from a mammal.

Thaumasioscolex didelphidis n. gen., n. sp. is described from the intestine of the black-eared opossum Didelphis marsupialis L. (Marsupialia: Didelphidae) from Los Tuxtlas, Veracruz, Mexico. The new genus differs from all proteocephalidean genera in the morphology of the scolex that is formed by 4 well separated lobes each containing 1 noncircular sucker opening laterally inside the exterolateral cavity, a large-sized body (length up to 1 m), a large number of testes, the shape of gravid proglottids that are inversely craspedote (the anterior border of a proglottid overlaps the posterior border of a preceding proglottid), eggs in groups mostly of 4-6 eggs each, and an embryophore bearing digitiform projections on its external surface. This is the first tapeworm of the Proteocephalidea, the members of which were previously reported exclusively from poikilotherm vertebrates (freshwater fishes, amphibians, and reptiles), found in a homoiotherm vertebrate.     

Crossing barriers of zoogeographical regions: Molecular evidence of vicariance of the only cestode parasite of loaches (Cobitidae) in the Indomalayan region,Paracaryophyllaeus lepidocephali (Cestoda: Caryophyllidea), and its redescription

Tapeworms of the genus Paracaryophyllaeus Kulakovskaya, 1961 (Cestoda: Caryophyllidea) are specific parasites of loaches (Cypriniformes: Cobitoidea) and occur almost exclusively in the Palaearctic region. The only exception and example of vicariance over the borders of two zoogeographical regions is Paracaryophyllaeus lepidocephali (Kundu, 1985), an insufficiently known species described from the Indomalayan region, with uncertain generic allocation. In the present paper, the species is redescribed based on new material collected from the type host, Lepidocephalichthys guntea, in West Bengal, India. Molecular data reveal this species as a member of Paracaryophyllaeus, within which it is the most closely related to P. cf. gotoi from Misgurnus anguillicaudatus in China and Japan on the basis of large subunit of ribosomal nuclear DNA (28S rDNA). Generic position of P. lepidocephali examined herein is also confirmed by morphology including cross sections, in particular, by a small, cylindrical body, medullary testes with testicular fields crossing the anterior margin of the cirrus sac, medullary vitelline follicles, with some follicles paramuscular, a shallow common genital atrium, short vagina and uterus not extending anteriorly beyond the cirrus sac. It differs from all Palaearctic congeners but Paracaryophyllaeus vladkae Scholz, Oros and Aydogdu, 2014 by the testicular field crossing the anterior margin of the cirrus sac. It differs from P. vladkae by more anterior position of the first vitelline follicles compared to the first testes. This species is a unique example of a fish tapeworm crossing the borders of the Palaearctic and Indomalayan zoogeographical regions.     

Cestodes of cestodes of Peruvian freshwater stingrays

Adult tetraphyllidean tapeworms (Platyhelminthes: Eucestoda) from the spiral intestines of 3 species of potamotrygonid stingrays (Paratrygon aiereba, Potamotrygon castexi, and Portamotrygon motoro) in the Madre de Dios river in Peru were found to host numerous cysts embedded in their parenchymal tissues. Histological sections of the cysts revealed the presence of a scolex bearing 4 suckers and an unarmed apical organ consistent with larval stages of both Cyclophyllidea and Proteocephalidea. To further elucidate their identities, partial 28S ribosomal DNA (rDNA) sequences were characterized from 3 cysts and 4 adult Rhinebothrium spp. 'host' worms and screened against all available cestode 28S rDNA data. Initial BLAST screening and subsequent alignment ruled out the possibility that the cysts were cyclophyllidean, and the cyst and adult sequences were thus aligned together with all available lecanicephalidean, litobothriidean, proteocephalidean, and tetraphyllidean sequences. Sequences from all 3 cysts were identical, and phylogenetic analysis clearly placed them among derived members of the Proteocephalidea, although no exact match was found. Sequences from the adult host worms formed 2 identical pairs and grouped together with other tetraphyllidean species from rays. These results are compared with records of hyperparasites of South American catfish cestodes. This is the first confirmed record of a proteocephalidean cestode parasitizing a tetraphyllidean cestode.     

Parabothriocephalus psenopsis n. sp. (Eucestoda: Pseudophyllidea) in Psenopsis anomala from the Taiwan Strait, China

ABSTRACT: Palabothriocephalus psenopsis n. sp. (Eucestoda: Pseudophyllidea) is described from the gastrointestinal tract of Psenopsis anomala caught off the coast of Xiamen, China. The new species most closely resembles, but differs from, Parabothriocephalus segmentatus in its possession of muscular globular appendages on the posterior margin of the proglottids, a limited proglottid number (9-13), and the shorter strobila (7.6-13.2 mm vs. 165 mm). In addition, the uterus of P. psenopsis is strongly coiled, whereas that of P. segmentatus is S shaped; P. segmentatus has a spherical expansion in the middle of the vagina, whereas that of P. psenopsis does not. Finally, P. psenopsis differs from Parabothriocephalus gracilis. Parabothriocephalus sagitticeps, and Parabothriocephalus macruri by the posterolateral expansion of the proglottids.     

Description of a new species of Proteocephalus pronini sp.n. (Cestoda: proteocephalidae)--a parasite of grayling from lake Hovsgol

A new cestode species, Proteocephalus pronini sp. n., is described from the Lake Hovsgol Grayling Thymallus arcticus nigrescens (Mongolia, Hovsgol Lake). The new species differs from P. thymalli from the Baikal Lake by the length of strobile, number of proglottids, form of mature proglottids, and number of diverticles of uterus.     

Sobolevitaenia japonensis n. sp. (Dilepididae: Dilepidinae) from a dusky thrush, Turdus naumanni eunomus Temminck, in Oita Prefecture, Japan

Sobolevitaenia japonensis n. sp. (Cestoda: Dilepididae) is described from eight specimens from the small intestine of a dusky thrush, Turdus naumanni eunomus Temminck, collected at Beppu City, Oita Prefecture, Japan. S. japonensis most closely resembles S. moldavica among the 11 species now assigned to the genus Sobolevitaenia, from which it is distinguished by a smaller number of proglottids, a larger scolex and a smaller number of testes of larger size, and a larger vitelline gland.     

On the phylogenetic positions of the Caryophyllidea, Pseudophyllidea and Proteocephalidea (Eucestoda) inferred from 18S rRNA

A phylogenetic analysis of tapeworms (Eucestoda) based on complete sequences of the 18S rRNA genes of 43 taxa (including new sequences of 12 species) was carried out, with the emphasis on the groups parasitising teleost fish and reptiles. Spathebothriidea and Trypanorhyncha (the latter group being paraphyletic) appeared as basal groups of the Eucestoda but their position was not stable. The tetrafossate orders (Litobothriidea, Lecanicephalidea, Tetraphyllidea, Proteocephalidea, Nippotaeniidea, Tetrabothriidea and Cyclophyllidea) were well separated from the remaining groups. Results supported polyphyly of the Pseudophyllidea formed by two distinct clades: one with diphyllobothriids (Diphyllobothrium, Schistocephalus, Spirometra and Duthiersia) and another including Abothrium, Probothriocephalus, Eubothrium and Bothriocephalus. The former pseudophyllidean clade formed a separate branch with the Caryophyllidea (Khawia and Hunterella) and Haplobothriidea (Haplobothrium), the latter taxon being closely related to either caryophyllideans or diphyllobothriids in different analyses. Proteocephalideans formed a monophyletic group in all analyses and constituted a clade within the Tetraphyllidea thus rendered paraphyletic. Within the Proteocephalidea, the Acanthotaeniinae (Acanthotaenia from reptiles in Africa) and Gangesiinae (Gangesia and Silurotaenia from silurid fish in the Palearctic Region) were separated from parasites of freshwater fish and mammals. The family Proteocephalidae was found to be paraphyletic due to the placement of a monticelliid species, Monticellia sp., in a clade within the former family. The genus Proteocephalus appeared as an artificial assemblage of unrelated taxa which is congruent with previous molecular analyses.     

Two new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) in the H. (Rhipistoma) spinulosa subgroup,parasites of carnivores and hedgehogs in Africa

Haemaphysalis (Rhipistoma) princeps n. sp. (Acari: Ixodidae) and H. (R.) camicasi n. sp. are described based on adults. Adults of H. princeps n. sp. were mostly collected from various carnivores (Carnivora: Canidae, Felidae, Herpestidae, Mustelidae, Viverridae) and hedgehogs (Erinaceomorpha: Erinaceidae); few specimens were found on hare (Lagomorpha: Leporidae), various rodents (Rodentia: Muridae) and an antelope (Artiodactyla: Bovidae) in Benin, Burkina Faso, Cameroon, Chad, Côte d’Ivoire, Ghana, Mali, Nigeria, Senegal, South Sudan and Sudan. Few adults of H. princeps n. sp. were reared from nymphs collected on rodents (Rodentia: Muridae) and shrew (Soricomorpha: Soricidae). Adults of H. camicasi n. sp. were also mostly collected from various carnivores (Carnivora: Canidae, Felidae, Herpestidae, Mustelidae, Viverridae) and hedgehogs (Erinaceomorpha: Erinaceidae); few specimens were found on different rodents (Rodentia: Muridae, Nesomyidae, Sciuridae), hare (Lagomorpha: Leporidae) and bushbaby (Primates: Galagidae) in Benin, Burkina Faso, Cameroon, Central African Republic, Chad, Côte d’Ivoire, Nigeria, Senegal, South Sudan and Togo. Males and females of both species can be differentiated from each other and other H. spinulosa-like ticks by their size, pattern of punctations on conscutum/scutum, size of setae, shape of genital structures, size and shape of spiracular plates, dorsal cornua, posterodorsal and posteroventral spurs on palpal segments II and spur on coxae.

     

Integrative taxonomy of a poorly known Sahelian rodent, Gerbillus nancillus (Muridae,Gerbillinae)

Aridification processes that affected the Sahelian area of West Africa during the last decades have induced significant changes in plant and animal communities of this region. In rodents, the genus Gerbillus characteristic of North African and Asian arid habitats has been affected by this climatic and environmental trends. Several species of this genus showed a southward range expansion in recent years into the Sahelian bioclimatic zone. Recent sampling in several localities of West Africa (Mali, Niger and Senegal) enabled us to collect numerous specimens of small gerbils. An integrative study of these samples using molecular, morphological and cytogenetical methods revealed that many of them were attributable to Gerbillus nancillus, a secretive and poorly known species. Gerbillus nancillus appears characterized by a well differentiated karyotype with 2n = 56 chromosomes, and to represent a unique genetic lineage within this genus. Body and skull measurements of G. nancillus were compared with those of the morphologically similar Gerbillus henleyi, which provided diagnostic clues between them. These new data significantly expand the distribution area of G. nancillus, which now ranges from Sudan in the East to Senegal in the West. G. nancillus is here reported from numerous new localities in Niger and Mali, and for the first time in Senegal, which raises questions about the origin of its presence and the colonization routes it followed to get there. We also show that G. henleyi and G. nancillus are sympatric and apparently often syntopic in the sub-Saharan part of the distribution of G. henleyi.