M.C. Durette-Desset

The origins and evolutionary expansion of the Strongylida (Nematoda)

The Strongylida are thought to have arisen from free-living rhabditoid nematodes, but the relationships between the major groupings within the Strongylida, the Strongylina, the Metastrongylina, Trichostrongylina and the Ancylostomatina are far from clear in spite of the abundance of morphological data now available for analysis. Evolutionary mechanisms including co-evolution, host switching, host dispersal, use of intermediate hosts, various sites of localisation within the definitive host and modifications of life-cycle strategies appear to have been utilised in the expansion of the Strongylida, with different mechanisms predominating in different families or superfamilies. Co-evolution appears to have been a major mode of evolution in the Strongylina, in contrast to the Trichostrongylina, which have used host dispersal and host-switching to great advantage. The phylogeny of the Ancylostomatina shows little association with host evolution, but does match the feeding preferences of the hosts. The Metastrongylina have utilised intermediate hosts and life cycle modifications including a shift to extra-intestinal sites as major means of diversification, in contrast to the other sub-orders. The review, while indicating much progress in our understanding of the phylogeny of the Strongylida, also reveals that enormous gaps still exist, and emphasises the tentative nature of many of the phylogenetic hypotheses tendered to date.

Les Nématodes Trichostrongylina parasites d'Amphibiens et de Reptiles: problèmes taxonomiques, phylétiques et biogéographiques

Except for four species of the Mertensinematinae (Molineoidea) not considered in this work, the Trichostrongylina parasitic in amphibians and reptiles, comprising 15 genera and 105 species, are reviewed. The morphological characteristics of each species are analysed. The most important characteristics are provided by the disposition of the caudal bursal rays, the morphology of the synlophe (at the oesophago-intestinal junction and mid-body level) and by the anatomy of the spicules. The species are classified into seven groups:Group 1: “relict” species with six lips and well-developed buccal capsule (six genera and 16 species).Group 2: “ancient” species with one or two primitive characters (eight genera and 15 species).Seventy-four species, without primitive characters, comprise the genus Oswaldocruzia which is subdivided into five groups.Group 3: Oriento-Ethiopian species with non-“idiomorphic” spicules with two to three tips (10 species).Group 4: Neo-Ethiopian species with non-“idiomorphic” spicules with numerous tips (11 species).Group 5: Holarctic species with “idiomorphic” spicules with the spicular fork divided above the distal third of the spicule length (24 species).Group 6: Continental Neotropical species with “idiomorphic” spicules and spicular fork divided within the distal third of the spicule length (21 species).Group 7: Caribbean Neotropical species with modified “idiomorphic” spicules, the three main branches of which are each divided into numerous tips (eight species).The geographical distribution of the species appears to be of greater significance than the host spectrum and suggests the following biogeographical hypotheses: Group 1, entirely Gondwanan, diversified during the Cretaceous era prior to the separation of the southern continents. The expansion of Group 2, which is represented by Gondwanan and Oriental species mainly from India, Malaysia and Indochina, could have occurred throughout South-East Asia at the end of the Cretaceous era when India collided with Eurasia. Group 3 could be interpreted either as a migration from Asia to Africa during the upper Eocene, or more likely during the Miocene, or, by a dispersion due to the migrations of Bufo melanostictus. Group 4 may result from the expansion of the former group in the Ethiopian region. Group 5 could be interpreted as a colonisation of western Europe and the Nearctic from Asia during the Tertiary. Group 6 could have arisen after the migration of the Neartic species to the Neotropical region during the Pliocene period and Group 7 by the expansion of the former group in the Caribbean.Amongst the Trichostrongylina, the “relict” and “ancient” genera parasitic in amphibians and reptiles can only be compared with the genera parasitic in birds and mammals, probably dating from the Palaeocene period (mainly marsupials and primitive insectivores). On the other hand, the genus Oswaldocruzia can be compared with other members of the Molineoidea parasitic in fissipeds, Pholidota and Chiroptera dating from a later period (Eocene) (see Durette-Desset, 1971). Thus, the Trichostrongylina of amphibians and reptiles are distinguished by the persistence of very old species, the evolution and the geographical distribution of which was arrested for millions of years, and by the existence (in some regions) of groups undergoing full evolutionary expansion.The new taxa proposed in the paper are: Bakeria (Moravec & Sey, 1986) status emend. (= Bakeria Moravec & Sey sub. g.), Ragenema n. g., Ragenema robustum (Baker, 1982) n. comb. (= Oswaldocruzia robusta), Typhlopsia quentini (Durette-Desset, 1980) n. comb. (= Trichoskrjabinia quentini), T. secundus (Pinnell & Schmidt, 1977) n. comb. (= Trichoskrjabinia secundus), T. gansi (Crusz & Ching, 1975) n. comb. (= Oswaldocruzia gansi), T. limnodynastes (Johnston & Simpson, 1942) n. comb. (= Oswaldocruzia limnodynastes), T. legendrei (Chabaud & Brygoo, 1962) n. comb. (= Oswaldocruzia legendrei).

The caudal bursa in the Heligmonellidae (Nematoda: Trichostrongylina). Characterization and hypothesis on its evolution

The different patterns of the caudal bursa of the Heligmonellidae (Nematoda) are redefined, taking into account the grouping of rays 2-6 and the sequence of origin of these rays from their common trunk. The type of symmetry of the caudal bursa is also redefined. The following patterns were observed and characterized: the basic patterns: types 2-3, 2-2-1, 1-3-1 and 1-4 and the intermediary patterns: type 2-3 tending to type 2-2-1, type 2-2-1 tending to type 1-3-1, type 1-3-1 tending to type 1-4 and type 2-2-1 tending to type 1-4. An evolutionary interpretation of the patterns is attempted and seems to follow the direction: 2-3 to 2-2-1 to 1-3-1 to 1-4. Seven atypical patterns are described. The caudal bursae were classified based on their symmetry: subsymmetrical, dissymmetrical and asymmetrical. Independently of the type of symmetry, the two latero-ventral lobes may have the same or different patterns. The type of symmetry, the ratio between the two latero-ventral lobes and a characteristic pattern were utilized to characterize the caudal bursae at the level of the genus and the subfamily. The combination of the right/left ratio and the type of symmetry gives heterogeneous results, with no real association between these characters. The most conspicuous asymmetries and dissymmetries were found among the Nippostrongylinae. The most frequent pattern in the Heligmonellidae is the basic type 2-2-1; types 1-3-1 and 1-4 are less frequent but are characteristic of several genera; type 1-4 is absent from the Heligmonellinae. Whatever the pattern, in the Heligmonellidae rays 4 and 5 are the last to diverge from the common trunk of rays 2-6.

Intestinal migrations of Trichostrongylus retortaeformis (Trichostrongylina, Trichostrongylidae) in the rabbit.

Observations were made on histological sections of the stomach and small intestine of seven rabbits infected with Trichostrongylus retortaeformis and from one uninfected control rabbit. At 12h post-infection, larvae were found in the small intestine. At first, only a few larvae were observed entering the mucosa through capillaries of the stroma of villi; the majority of larvae remained in the intestinal lumen, within mucus of the crypts. We consider that the presence of the worms in the stroma is the result of a larval migration. From a phyletic point of view, this migration is interpreted as an ancestral memory of the pulmonary migration seen in the primitive Strongylida.

The systematic position of some nippostrongyline nematodes (Trichostrongylina: Heligmosomoidea) parasitic in Argentinean sigmodontine rodents

The systematic position of two nippostrongyline nematodes described from Argentinean sigmodontine rodents is clarified. The first species, Hassalstrongylus multiovatus Suriano & Navone, 1992, parasitic in Akodon simulator Thomas from the province of Tucumán, was studied on the basis of type and voucher material. H. multiovatus is proposed as a junior synonym of Trichofreitasia lenti Sutton & Durette-Desset, 1991, a parasite described from Oligoryzomys flavescens (Waterhouse) in the province of Buenos Aires. The holotype and three of seven paratypes deposited as H. multiovatus were identified as T. lenti. One male paratype was identified as Guerrerostrongylusuruguayensis Sutton & Durette-Desset, 1991, a parasite described from O. flavescens in Uruguay. Three female paratypes were identified as Guerrerostrongylus sp. The second species, Stilestrongylus scapteromys Suriano & Navone, 1996, parasitic in Scapteromysaquaticus Thomas from the province of Buenos Aires, was studied on voucher material. Stilestrongylus scapteromys and Malvinema frederici Digiani, Sutton & Durette-Desset, 2003, the type-species of Malvinema Digiani, Sutton & Durette-Desset, 2003, were described from the same host and geographical region. As they are considered to refer to one and the same taxon, the new combination Malvinema scapteromys n. comb. is proposed for this species.

Four new species of Oswaldocruzia (Nematoda: Trichostrongylina, Molineoidea) Parasitizing amphibians and lizards from Ecuador

Description of four new species of Oswaldocruzia parasitizing Iguanidae and Leptodactylidae from Ecuador, demonstrate that they are morphologically close to each other. Like most of the other neotropical and holarctic Oswaldocruzia , they are characterized by spicules with three main branches: blade, shoe and fork; the division of the fork within the distal third of the spicule length appears to be characteristic of the neotropical species. - Oswaldocruzia bainae n. sp. parasitizing Anolis chrysolepis and Anolis fuscoauratus possesses a synlophe visible only on transversal sections of the body. It is composed of rounded and not pointed ridges. - Oswaldocruzia tcheprakovae n. sp. parasitizing Eleutherodactylus altamazonicus is closely related to O. bainae , but the synlophe is present only in the anterior and posterior extremities of the body. - Oswaldocruzia cassonei n. sp. parasitizing Eleutherodactylus lanthanites is closely related to O. taranchoni, Ben Slimane and Durette-Desset, 1995, a parasite of Bufo marinus from Brazil. It is differentiated by the synlophe and the measurements. - Oswaldocruzia petterae n. sp. parasitizing Leptodactylus pentadactylus is closely related to O. chambrieri, Ben Slimane and Durette-Desset, 1993, parasitizing Bufo and Eleutherodactylus in the same region. It is differentiated since, for an equivalent length of the body, the ridges are almost two times fewer and the spicules smaller.

A description ofGraphidioides kravetzi n. sp. and the revision ofGraphidioides Cameron, 1923 (Nematoda Trichostrongyloidea), parasites of Neotropical rodents

Graphidioides kravetzi n. sp., from the cricetidHolochilus brasiliensis in Uruguay, is characterised by relatively long rays 2 in relation to the length of rays 3 and by the characters of the synlophe: the same number of ridges at mid-body and at the level of oesophago-intestinal junction; and in the anterior part of the body ridges facing the lateral fields more developed than the others at the same level. Redescriptions of three species belonging to the genusGraphidioides Cameron, 1923 (Trichostrongylinae) from Argentina are presented:G. rudicaudatus (Railliet & Henry, 1909) Cameron, 1925 fromLagostomus maximus; G. mazzai Lent & Freitas, 1935 fromCavia aperea pamparum; andG. taglei Babero & Cattan, 1975 fromOctodon bridgesi. Complementary data are provided concerning the type-species,G. affinis (Megnin, 1895), fromDolichotis patagonum. An amended definition of the genusGraphidioides is proposed, mainly based on a knowledge of the synlophe, with ridges orientated perpendicularly to the body surface and lacking facing the lateral cords, and the pattern of the caudal bursa. A dichotomous key to the species of the genus is provided.

Suttonema delta n. g., n. sp. (Nematoda: Trichostrongylina: Heligmosomoidea) from Oxymycterus rufus (Rodentia: Sigmodontinae) in Argentina

A new nippostrongyline, Suttonema delta n. g., n. sp., is described from the intestine of Oxymycterus rufus (Rodentia: Sigmodontinae) from Argentina, in a host from which trichostrongylid nematodes were hitherto unknown. The new genus is very similar to Stilestrongylus Freitas, Lent & Almeida, 1937 and Malvinema Digiani, Sutton & Durette-Desset, 2003, both parasites of Neotropical sigmodontines, in the features of the caudal bursa (with a pattern of type 1-4, asymmetrical with hypertophied right lobe) and the presence of cephalic structures resembling cuticular cordons. The larval synlophe is also identical to that of Stilestrongylus freitasi Durette-Desset, 1968. The new genus is differentiated by an adult synlophe with few ridges (9-12 at mid-body) of two different types: small, rounded ridges without cuticular support on the dorsal side, and pointed ridges of unequal size on the ventral side and in lateral fields. It is also characterised by the presence of comaretes on the left ventral and ventral fields of the synlophe.

Fauna Europaea : Helminths (Animal Parasitic)

Abstract Fauna Europaea provides a public web-service with an index of scientific names (including important synonyms) of all living European land and freshwater animals, their geographical distribution at country level (up to the Urals, excluding the Caucasus region), and some additional information. The Fauna Europaea project covers about 230,000 taxonomic names, including 130,000 accepted species and 14,000 accepted subspecies, which is much more than the originally projected number of 100,000 species. This represents a huge effort by more than 400 contributing specialists throughout Europe and is a unique (standard) reference suitable for many users in science, government, industry, nature conservation and education. Helminths parasitic in animals represent a large assemblage of worms, representing three phyla, with more than 200 families and almost 4,000 species of parasites from all major vertebrate and many invertebrate groups. A general introduction is given for each of the major groups of parasitic worms, i.e. the Acanthocephala, Monogenea, Trematoda (Aspidogastrea and Digenea), Cestoda and Nematoda. Basic information for each group includes its size, host-range, distribution, morphological features, life-cycle, classification, identification and recent key-works. Tabulations include a complete list of families dealt with, the number of species in each and the name of the specialist responsible for data acquisition, a list of additional specialists who helped with particular groups, and a list of higher taxa dealt with down to the family level. A compilation of useful references is appended.

A new species of Heligmosomoides (Nematoda, Heligmosomidae) parasitic in Peromyscus maniculatus (Rodentia, Cricetidae) from Pennsylvania, USA

Heligmosomoides vandegrifti sp. nov. (Nematoda, Heligmosomidae) is described from Peromyscus maniculatus (Rodentia, Cricetidae) from Pennsylvania, USA. It differs from its closest congener, H. douglasi, in the number of cuticular ridges (35 vs. 32 in male, 36 vs. 41 in female at mid-body), longer bursal rays 2 in relation to rays 3, and in having smaller spicules (635–740 μm long vs. 1 mm). It is proposed that both H. douglasi and H. vandegrifti sp. nov. are parasites of capture from species in North American arvicoline rodents.