High prevalence of cestodes in Artemia spp. throughout the annual cycle: relationship with abundance of avian final hosts.

Brine shrimp, Artemia spp., act as intermediate hosts for a range of cestode species that use waterbirds as their final hosts. These parasites can have marked influences on shrimp behavior and fecundity, generating the potential for cascading effects in hypersaline food webs. We present the first comprehensive study of the temporal dynamics of cestode parasites in natural populations of brine shrimp throughout the annual cycle. Over a 12-month period, clonal Artemia parthenogenetica were sampled in the Odiel marshes in Huelva, and the sexual Artemia salina was sampled in the Salinas de Cerrillos in Almería. Throughout the year, 4-45 % of A. parthenogenetica were infected with cestodes (mean species richness = 0.26), compared to 27-72 % of A. salina (mean species richness = 0.64). Ten cestode species were recorded. Male and female A. salina showed similar levels of parasitism. The most prevalent and abundant cestodes were those infecting the most abundant final hosts, especially the Greater Flamingo Phoenicopterus ruber. In particular, the flamingo parasite Flamingolepis liguloides had a prevalence of up to 43 % in A. parthenogenetica and 63.5 % in A. salina in a given month. Although there was strong seasonal variation in prevalence, abundance, and intensity of cestode infections, seasonal changes in bird counts were weak predictors of the dynamics of cestode infections. However, infection levels of Confluaria podicipina in A. parthenogenetica were positively correlated with the number of their black-necked grebe Podiceps nigricollis hosts. Similarly, infection levels of Anomotaenia tringae and Anomotaenia microphallos in A. salina were correlated with the number of shorebird hosts present the month before. Correlated seasonal transmission structured the cestode community, leading to more multiple infections than expected by chance.

Spermiogenesis and spermatozoon ultrastructure of the paruterinid cestode Notopentorchis sp. (Cyclophyllidea).

Ultrastructural characters of the spermiogenesis and mature spermatozoon of Notopentorchis sp. (Cestoda, Cyclophyllidea, Paruterinidae), a parasite from Apus affinis (Aves, Apodiformes, Apodidae) from Gabon, are described by means of transmission electron microscopy. Cytochemical analysis for detection of glycogen was applied. Vestigial striated roots associated with the two centrioles are present in the zone of differentiation. The spermiogenesis is characterized by an external growth of free flagellum followed by a proximodistal fusion of the latter with cytoplasmic protrusion, thus, corresponding to the cestode spermiogenesis of the type III pattern described by Bâ and Marchand (Mem. Mus. Natl. Hist. Nat. 166:87-95, 1995). In the final stage of spermiogenesis, a single crested body appears at the base of the forming spermatozoon. The mature spermatozoon of Notopentorchis sp. is filiform and tapering at both extremities. It consists of five regions differing in their ultrastructural characteristics. The anterior extremity of the mature spermatozoon is characterized by the presence of an apical cone and a single crested body. The cytoplasm contains one axoneme of 9 + "1" type of the trepaxonematan Platyhelminthes, a periaxonemal sheath, a layer of twisted cortical microtubules, transverse intracytoplasmic walls, and granules of glycogen. The nucleus is coiled in spiral around the axoneme. The posterior extremity of the spermatozoon is characterized by the presence of electron-dense material. This structural organization corresponds to the morphology of cestode spermatozoon of type VII as defined by Levron et al. (Biol Rev 85: 523-543, 2010). The comparison of the results with those of the two previous studies on paruterinids suggests that several characters of the spermiogenesis and the mature spermatozoon are invariable, i.e. the type III spermiogenesis and the presence of vestigial striated roots, a single crested body, a periaxonemal sheath, and intracytoplasmic walls. The main differences of the sperm cells among members of this family are the lack of dense granules (as in Triaenorhina rectangula) and the presence of electron-dense material in the posterior extremity of the spermatozoon (as in Notopentorchis sp.).

Participation of metanauplii and juvenile individuals of Artemia parthenogenetica (Branchiopoda) in the circulation of avian cestodes.

Adult crustaceans of the genus Artemia (brine shrimps) are intermediate hosts in the life cycle of cestode species parasitic in aquatic birds as their definitive hosts. However, there are no data on the role of larval and juvenile brine shrimps in the transmission of avian helminth parasites. In order to examine the possible role of early developmental stages (nauplii, metanauplii and juveniles) of Artemia for the circulation of avian cestodes, the natural cestode infection in the population of Artemia parthenogenetica from La Mata Lagoon, Mediterranean coast of Spain, was studied. Metacestodes (cysticercoids) of four cestode species were recorded in adult brine shrimps: Flamingolepis liguloides and Flamingolepis flamingo (hymenolepidids parasitic in flamingos), Confluaria podicipina (a hymenolepidid species parasitic in grebes) and Eurycestus avoceti (a dilepidid species parasitic in avocets, stilts, plovers and, to a lesser extent, in flamingos). No cysticercoids were found in nauplii. Two species, F. liguloides and F. flamingo, were found in metanauplii and juvenile brine shrimps. Only 36.3% of the cysticercoids of F. liguloides occurred in adult brine shrimps; the remaining 63.7% were parasitic in metanauplii (39.6%) and juveniles (24.1%). Similarly, the metacestodes of F. flamingo were also distributed among various age groups: in adults (44.4% of cysticercoids), juveniles (27.8%) and metanauplii (27.8%). These results indicate that the early developmental stages of Artemia have an important role for the circulation of certain parasite species. No cysticercoids of C. podicipina and E. avoceti were recorded in larval and juvenile brine shrimps. The selective infestation of larval brine shrimps with flamingo parasites is probably associated with the feeding behaviour of definitive hosts, which are filtering predators; in contrast, grebes and waders pick brine shrimps individually one by one. The possible underlying mechanism for selective infestation of metanauplii and adults by certain cestode species is associated with the size of parasite eggs, allowing only cestode species with small eggs to be ingested by larval brine shrimps.

Ultrastructure of spermiogenesis and mature spermatozoon of Triaenorhina rectangula (Cestoda: Cyclophyllidea: Paruterinidae).

Ultrastructural characters of spermiogenesis and mature spermatozoon of Triaenorhina rectangula (Fuhrmann, 1908) are examined by transmission electron microscopy. Spermiogenesis follows the Bâ and Marchand's Type III spermiogenesis of cestodes. The process begins with the formation of a differentiation zone containing two centrioles and a cytoplasmic protrusion. The centrioles are associated with vestigial striated roots. One of the centrioles develops a free flagellum externally to the cytoplasmic protrusion. After a slight rotation, the free flagellum fuses with the cytoplasmic protrusion. In the final stage of spermiogenesis, a single crested body appears in the anterior part of the differentiating spermatozoon. The anterior extremity of the mature spermatozoon is characterised by an apical cone and a single crested body. The axoneme is of the 9+"1" trepaxonematan type. A periaxonemal sheath and electron-dense rods are described in some parts of the mature spermatozoon. The nucleus is electron-dense and spirally coiled around the axoneme. The cortical microtubules are spirally arranged at an angle of about 40 degrees to the spermatozoon axis. The present results show that the ultrastructural characters of spermiogenesis and mature spermatozoon of T. rectangula resemble most closely those in taeniids and metadilepidids. The comparison of these results with the only previous spermiological description of a paruterinid species reveals differences relative to the occurrence of filamentous rods of electron-dense material versus intracytoplasmic walls in the mature spermatozoon that may reflect the polyphyletic character of the Paruterinidae.

Cestodes from Hector's beaked whale ( Mesoplodon hectori ) and spectacled porpoise ( Phocoena dioptrica ) from Argentinean waters.

Single individuals of 2 little-known cetacean species, Mesoplodon hectori and Phocoena dioptrica , stranded and died on the coast of Argentina (Buenos Aires and Chubut provinces, respectively) and were studied for the presence of helminths. The cestodes found were described and illustrated using light microscopy. The following cestode taxa were recovered: Tetrabothrius ( Tetrabothrius ) hobergi n. sp. (several fragmented specimens, at least 1 gravid) and Tetrabothrius ( s.l. ) sp. 1 (several fragmented immature specimens) from M. hectori , and Tetrabothrius ( s.l. ) sp. 2 (single fragmented immature specimen) and 2 morphotypes of tetraphyllidean larvae from P. dioptrica. Tetrabothrius ( T. ) hobergi n. sp. can be distinguished from Tetrabothrius ( T. ) forsteri by the greater number of testes and larger eggs and oncospheres, from Tetrabothrius ( T. ) curilensis by the smaller testes and vitellarium, the shape and size of the ovary, and the larger oncospheres and longer embryonic hooks, and from Tetrabothrius ( T. ) sp. from Ziphius cavirostris by the narrower strobila, smaller scolex, and smaller number of testes. The generic designations of Tetrabothrius ( s.l. ) sp. 1 and Tetrabothrius ( s.l. ) sp. 2 were based on the scolex morphology. Tetrabothrius ( s.l. ) sp. 1 is closest to Tetrabothrius ( T. ) forsteri and Tetrabothrius ( Biamniculus ) innominatus based on the number of testes, while the scolex size of Tetrabothrius ( Tetrabothrius ) sp. 2 is within the variability range reported for Tetrabothrius ( T. ) forsteri . More definite identification of the 2 species was not possible due to the condition of the available material. The present study provides the first descriptions of cestodes from M. hectori and P. dioptrica , thus enriching the knowledge regarding the helminths of insufficiently studied marine mammals.

Taxonomic revision and phylogenetic analysis of the cestode genus Paraprogynotaenia Rysavy, 1966 (Cyclophyllidea: Progynotaeniidae).

The type-series of Paraprogynotaenia jimenezi Rysavy, 1966 [type-species of Paraprogynotaenia Rysavy, 1966] from Charadrius wilsonia Ord in Cuba and P. charadrii (Yamaguti, 1956) (= Proterogynotaenia charadrii Yamaguti, 1956) from C. alexandrinus L. in Japan are redescribed. Two new species are described: Paraprogynotaenia minuta n. sp. from C. alexandrinus on the Black Sea coast of Bulgaria [type-host and type-locality, based on specimens previously reported as P. charadrii], from the same host species in Tunisia and from "Tringa totanus" (the host identification is uncertain) in France [the latter two samples were previously reported as Progynotaenia odhneri Nybelin, 1916]; and Paraprogynotaenia canarisi n. sp. from C. marginatus Vieillot in the Republic of South Africa [type-host and type-locality] and from C. alexandrinus and Pluvialis dominica (Müller) in Taiwan [specimens from both localities were previously reported as Paraprogynotaenia charadrii]. P. jimenezi can be distinguished from its congeners by having 12 hooks of equal length. Among the remaining species with 18-22 hooks of varying length, P. minuta n. sp. can be distinguished by the smaller number of proglottides: up to 8 vs 15-20 for P. charadrii and P. canarisi n. sp. The latter two species can be differentiated from one another by the blade length/total length ratio of the smallest (lateral) rostellar hooks, which is less than 0.5 for P. canarisi n. sp. and 0.5 or more for P. charadrii. An identification key to the species of Paraprogynotaenia is provided and the generic diagnosis is amended. A phylogenetic analysis based on 17 morphological characters resulted in the following hypothesis for the relationships within the genus: (P. jimenezi (P. minuta (P. charadrii, P. canarisi))).

Spermiogenesis and sperm ultrastructure of Valipora mutabilis Linton, 1927 (Cestoda, Cyclophyllidea, Gryporhynchidae).

This is the first ultrastructural study of the spermiogenesis and the mature spermatozoon of a cyclophyllidean cestode of the family Gryporhynchidae. The spermiogenesis of Valipora mutabilis begins with the formation of a differentiation zone delimited by arching membranes and containing two centrioles. One of the centrioles develops an axoneme that grows directly into the cytoplasmic protrusion. The other centriole remains situated in a cytoplasmic bud and is subsequently aborted (type IV of cestode spermiogenesis). The mature spermatozoon of V. mutabilis is a filiform cell, tapered at both extremities and lacks mitochondria. The anterior extremity is characterised by the presence of an apical cone and a single helicoidal crested body. The axoneme is of 9 + '1' trepaxonematan pattern, with a periaxonemal sheath. The cortical microtubules are twisted at an angle of about 45 degrees to the spermatozoon axis. The nucleus is electron dense and spirally coiled around the axoneme. The cytoplasm is electron lucent and contains numerous granules of electron-dense material. In contrast to a recent opinion for close phylogenetic relationships, these ultrastructural data demonstrate the distant position between gryporhynchids and the family Taeniidae. The most similar pattern in the ultrastructure of the spermiogenesis and the mature spermatozoon has been described for dilepidids, some hymenolepidids and some anoplocephalids.

Taxonomic revision of the cestodes of the family Progynotaeniidae (Cyclophyllidea) parasitising stone curlews (Charadriiformes: Burhinidae).

The type series of Progynotaenia evaginata Fuhrmann, 1909 from Burhinus senegalensis in Sudan, P. foetida Meggitt, 1928 from B. oedicnemus in Egypt and Angularia australis Maplestone, 1921 from B. grallarius in Australia are redescribed. As a comparative material, specimens of P. evaginata from B. oedicnemus in Kazakhstan were studied. The type-series of P. evaginata and P. foetida were found to be heterogeneous due to the presence of scoleces and fragments of cestodes of the genus Stenovaria Spasskii & Borgarenko, 1973 (Dilepididae). For P. foetida, a lectotype is designated. P. foetida is recognised as a synonym of P. evaginata (new synonymy). Angularia australis, previously considered a member of the Dilepididae, is transferred to the Progynotaeniidae as a synonym of P. evaginata (new synonymy). The synonymy of P. indica Johri, 1963 with P. evaginata, proposed by Ryzhikov & Tolkacheva (1981), is supported. The host range and the geographical distribution of P. evaginata are restricted to birds of the genus Burhinus from the Eastern Hemisphere.

New data on the morphology of species of Leptotaenia Cohn, 1901 (Cestoda: Progynotaeniidae).

Leptotaenia ischnorhyncha (Lühe, 1898) is redescribed on the basis of a specimen from Phoenicopterus ruber in Cuba. In contrast to previous descriptions, the presence of two rows of rostellar hooks, a secondary female copulatory organ and four types of cirrus spines are observed. Leptotaenia sp. from P. minor from Kenya, identified as L. ischnorhyncha and figured by Khalil (1994), is described on the basis of poorly preserved specimens. The synonymy of L. skrjabini Shakhtakhtinskaya, 1953 from P. ruber in Azerbaijan with L. ischnorhyncha, proposed by Ryzhikov & Tolkacheva (1981), is questioned and the former is regarded as a species inquirenda. The diagnosis of Leptotaenia Cohn, 1901 is amended.