Marie-Claude Durette-Desset

Trichostrongyloid nematodes and their vertebrate hosts: reconstruction of the phylogeny of a parasitic group.

Publisher Summary This chapter introduces trichostrongyloid nematodes as a model for discussing the reconstruction of the phylogeny of a parasitic group. Morphological and biological characters are studied and assessed for their relative phylogenetic importance. On the basis of morphological characters and their presumed evolution, trichostrongyloids are divided into 14 families and 24 subfamilies. The evolutionary trends of each character are analyzed and used to construct a classification that recognizes different phylogenetic lines. A parasitic line may become isolated from its ancestral forms by passing from one host group to another; the isolation may be followed by the evolutionary radiations of variable importance. In the Trichostrongyloidea, the most important characters from a phylogenetic point of view are those of the caudal bursa and synlophe. Using these characters, a number of different evolutionary lines become evident and the degree of evolution of the different genera composing each of those lines can be assessed. The chapter also presents a phylogenetic tree of the Trichostrongyloidea.

Geography and host biogeography matter for understanding the phylogeography of a parasite.

The co-evolution between hosts and parasites has long been recognized as a fundamental driver of macro-evolutionary patterns of diversification. The effect of co-differentiation on parasite diversification is, however, often confounded by underlying geographic patterns of host distribution. In order to disentangle the confounding effects of allopatric versus host speciation, the mitochondrial cytochrome b (cyt b) gene was sequenced in seventy individuals of the parasitic nematode genus Heligmosomoides sampled in the six Apodemus mice species common in the western Palearctic region. The nuclear internal transcribed spacers (ITS) 1 and 2 were also sequenced in fifteen parasites to confirm the mitochondrial data. All lineages differentiated according to a geographic pattern and independently from the sampled host species. This suggests that host speciation did not involve concurrent parasite speciation. However, the geographic distribution range of some parasite lineages mirrors that of A. sylvaticus lineages in SW Europe, and that of A. flavicollis lineages in the Balkans and in the Middle East. Thus, regional co-differentiation likely occurred between the parasite and the two sister Apodemus hosts in different parts of their distribution range. We suggest that differences in regional abundances of A. sylvaticus and A. flavicollis are responsible for generating this pattern of regional co-differentiation. This study highlights the importance of integrating both geography and biogeographic information from potential hosts to better understand their parasite phylogeography.

Helminths From Introduced Small Mammals on Kerguelen, Crozet, and Amsterdam Islands (Southern Indian Ocean)

Four monoxenous nematodes and 1 heteroxenous cestode were found in 4 species of introduced small mammals on isolated sub-Antarctic islands of the Indian Ocean. In the Kerguelen Archipelago, Syphacia obvelata, Passalurus ambiguus (Nematoda: Oxyuridae), and Rodentolepis straminea (Cestoda: Cyclophyllidae) were respectively found in the house mouse Mus musculus, the rabbit Oryctolagus cuniculus, and the black rat Rattus rattus. One accidental nematode, Trichostrongylus sp. (Nematoda: Trichostrongylidae), was also found in a black rat on Kerguelen. On Possession Island (Crozet Archipelago), R. straminea was present in the black rat. On Amsterdam Island, the brown rat R. norvegicus harbored 2 species, R. straminea and Nippostrongylus brasiliensis (Nematoda: Heligmonellidae). The small number of founder hosts and the depauperate terrestrial communities on these remote islands explain the low diversity in the helminth communities of these introduced mammals compared with continental populations.

Internal transcribed spacer rDNA can be used to infer the phylogenetic relationships of species within the genus Nematodirus (Nematoda: molineoidea).

Sequences of the first internal transcribed spacer rDNA were characterised for four veterinary important species of gastrointestinal nematodes from the genus Nematodirus. The sequence data were combined with previously published data of the second internal transcribed spacer to determine whether these rDNA regions provided a suitable number of informative characters to determine the phylogenetic relationships of species within the genus. A total of 32 alignment positions of the first internal transcribed spacer data set and 33 characters from the second internal transcribed spacer data set were informative in phylogenetic analyses. Irrespective of whether the data from each spacer were analysed separately or combined, only one most parsimonious tree was produced, with the relationships of the four species fully resolved. In addition, several regions of conservatism in the first internal transcribed spacer sequence among the four Nematodirus species suggests that this rDNA region may also provide phylogenetic information for higher taxonomic levels within the Molineoidea.

EPIZOOTIOLOGY OF SYPHACIA OBVELATA FROM A DOMESTIC MOUSE POPULATION ON THE SUBANTARCTIC KERGUELEN ARCHIPELAGO

The effects of abundance, age, and sex of feral domestic mice Mus musculus domesticus on infections with the nematode parasite Syphacia obvelata were analyzed during a long-term study of the mouse population on Guillou Island (1.45 km2), a part of the subantarctic Kerguelen Archipelago. The population dynamics of the nematode did not follow the variation in host abundance. However, depending on the year, differences in pinworm abundance were found between the age classes and sex. Such patterns suggest that parasitic infections may have been modulated by host-intrinsic factors, e.g., either by the way of innate or adaptive immunity, rather than extrinsic factors, e.g., host abundance.

A cladistic analysis of the genera in the subfamily pudicinae (nematoda, trichostrongyloidea, heligmonellidae)

A parsimony analysis was performed on 37 specific taxa belonging to the subfamily Pudicinae (family Heligmonellidae), which contains parasites mainly from South American caviomorph rodents. Thirteen characters were used from the synlophe (rotation of axis, presence of carene, carene asymmetry, presence of comaretes, single ventral comarete length, ridge discontinuity, ventral ridge numbers, presence of a peculiar posterior synlophe, presence of supernumerary spines) and the male caudal bursa (relative length of rays 9 and 10, caudal bursa type, division of the dorsal ray, divergence of the 10th rays). The cladogram shows a consistency index of 1.0. The subfamily Pudicinae has two synapomorphies. Two suprageneric groups are recognized. Suprageneric group 1 shows one synapomorphy and contains Heligmostrongylus, Fuellebornema, Sciurodendrium and Pseudoheligmosomum; suprageneric group 2 shows two synapomorphies and contains Pudica, Acanthostrongylus, Justinema and Durettestrongylus. Five genera are defined on the basis of synapomorphies. The genera Heligmostrongylus, Sciurodendrium and Pudica which are considered paraphyletic, however, are retained due to lack of knowledge as to their relationships.

Additional molecular evidence to support a sister taxon relationship between Heligmosomoidea and Molineoidea nematodes (Trichostrongylina)

Sequences of the first and second internal transcribed spacers of the ribosomal DNA were used to infer the evolutionary relationships of 19 species of parasitic nematode belonging to three superfamilies, Trichostrongyloidea, Molineoidea and Heligmosomoidea, within the sub-order Trichostrongylina. Analyses using maximum parsimony, maximum likelihood and neighbor-joining methods revealed strong statistical support for monophyly of each superfamily as defined on morphological criteria. Furthermore, in most analyses, there was also strong support for a sister taxon relationship between the Molineoidea and Heligmosomoidea, which supports the findings of a previous study based on partial LSU rDNA sequence data.

SYSTEMATIC POSITION OF SOME NEARCTIC HELIGMOSOMOIDEA (NEMATODA: TRICHOSTRONGYLINA) FROM THE U.S. NATIONAL PARASITE COLLECTION AND THEIR DESCRIPTION

The systematic position of some heligmosomoid nematodes from rodents, deposited in the U.S. National Parasite Collection (USNPC), is revised, mainly through the study of their synlophe, which in all cases was unknown or insufficiently described. The material was registered as different species of Longistriata Schulz, 1926, a genus whose representatives are only parasitic in Holarctic insectivores. Longistriata norvegica Dikmans, 1935, parasitic in Rattus sp. becomes a synonym of Hassalstrongylus aduncus (Chandler, 1932). Specimens registered as Longistriata dalrymplei Dikmans, 1935, from Ondatra zibethicus, are confirmed to belong to Carolinensis (Travassos, 1937). Specimens registered as Longistriata noviberiae Dikmans, 1935, parasitic in Sylvilagus floridanus alacer, were found to belong to Vexillata, and Vexillata noviberiae n. comb. is here proposed. This is the first record of a species of Vexillata in a lagomorph. Other specimens registered as Longistriata norvegica, parasitic in Geomys floridanus austrinus, were also found to be an undescribed species of Vexillata (Hall, 1916), which is named Vexillata chitwoodi n. sp. This is similar to Vexillata chabaudi Yoyotte-Vado, 1972, Vexillata petteri Durette-Desset, 1970, Vexillata scorzai Guerrero, 1984 and Vexillata tejerai Guerrero, 1984, all having the same number of cuticular ridges (4 dorsal, 5 ventral) and the division of the dorsal ray at its apex. The most related species is V. chabaudi, which is differentiated from the new species by rays 4 not curved distally, by thick rays 8, and by a dorsal ray enlarged at the level of the arising of rays 8.

A NEW GENUS OF NIPPOSTRONGYLINAE (NEMATODA: HELIGMONELLIDAE) FROM THE WATER RAT SCAPTEROMYS AQUATICUS (SIGMODONTINAE) IN ARGENTINA

A new genus of Nippostrongylinae, Malvinema n. gen., with 3 coparasitic species M. frederici n. sp., M. carolinae n. sp., and M. victoriae n. sp. from the intestine of the water rat, Scapteromys aquaticus Thomas (Rodentia: Muridae), from the northeast of Buenos Aires Province, Argentina, is proposed in this study. The new genus shows similarities to 2 Neotropical Nippostrongylinae: Carolinensis (Travassos, 1937) by some characters of the synlophe and Stilestrongylus Freitas, Lent and Almeida, 1937, by the pattern of the caudal bursa. It is characterized by a synlophe with triple or quadruple gradient of size of the ridges, lateromedian, decreasing from the largest left and right ridges. The gradient situated in the right ventral quadrant is always present. The caudal bursa shows a pattern of type 1-4. Malvinema frederici possesses a synlophe with 17 ridges and an axis of orientation inclined at 45° from the sagittal axis; M. carolinae possesses a synlophe with 22–24 ridges and an axis of orientation almost merged with the sagittal axis. Both species have a caudal bursa with the right lobe enlarged transversally. Malvinema victoriae possesses a synlophe with 22–24 ridges, an axis of orientation inclined at 45° from the sagittal axis, and a caudal bursa with the right lobe enlarged vertically.

THE LIFE CYCLE OF NEMATODIROIDES ZEMBRAE (NEMATODA, TRICHOSTRONGYLINA) IN THE RABBIT

The morphogenesis and the chronology of the life cycle of Nematodiroides zembrae (Bernard, 1965), a parasite of Oryctolagus cuniculus from Spain, were studied in detail in its natural host. For each experiment a morphological description of the different stages of the life cycle is provided. The free-living larvae developed in eggs until infective stage. First hatching began at 10 days. Twenty-one, worm-free rabbits were each infected per os with N. zembrae larvae and killed 3 days after infection (DAI) and every day from 4 to 22 DAI. By 3 DAI all recovered larvae were exsheathed and present in the small intestine. The third moult occurred between 6 and 7 DAI. The last moult occurred between 11 and 19 DAI. The prepatent period lasted for 21–22 days. The distribution of N. zembrae along the small intestine of the rabbit is described. Significantly different distributions of the parasite along the small intestine indicated that migrations occurred during the development of N. zembrae in the rabbit. The life cycle of N. zembrae is compared with the 5 known life cycles of Nematodirus spp. in ruminants. The biological data are very similar in both groups except for the prepatent period.