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A cladistic analysis of the Trichostrongyloidea (Nematoda).

A morphologically based cladistic analysis of 40 genera included within the Trichostrongyloidea (Amidostomatidae, Dromaeostrongylidae and Trichostrongylidae) is proposed. Two genera were used as outgroups, one from the Strongylina and the other from the Ancylostomatina. Seven genera do not appear in the matrix because some significant morphological characters remain unknown for these genera. Nonetheless, except for Moguranema which is excluded as incertae sedis, a likely systematic position could be assigned to them based on the morphological characters that are known. The classification which best fits the consensus tree is composed of three families. In adding the genera not included in the tree, we obtain: (i) Trichostrongylidae with three sub-families, Amidostomatinae (four genera), Filarinematinae (three genera) and Trichostrongylinae (five genera); (ii) Haemonchidae with two sub-families: Ostertagiinae (eight genera) and Haemonchinae (five genera); (iii) Cooperiidae with three sub-families: Libyostrongylinae (five genera), Obeliscoidinae n. subfam. (five genera) and Cooperiinae (ten genera). Dromaeostrongylus and Ortleppstrongylus, whose females have a caudal spine, are excluded from the Trichostrongyloidea and are placed in the Molineoidea. The hypotheses relating to the evolutionary history of the Trichostrongyloidea are: the origin of the superfamily could have occurred during the upper Cretaceous period. The two most ancient sub-families (Amidostomatinae and Filarinematinae) would be of Gwondwanan origin and evolved during the Paleocene period within Neotropical aquatic birds and within the Australian marsupials. The Trichostrongylinae would have arisen during the Eocene period within birds and then adapted to diverse archaic mammals in the Neotropical region on one hand and in the Nearctic region, on the other hand and lastly adapted to the Lagomorpha and subsequently to the Ruminantia. In both families originating from the Trichostrongylidae, the adaptation to the Lagomorpha may have taken place during the Oligocene but in a different way. In the Haemonchidae, the Ostertagiinae may have passed directly from the Neartic region to Europe. In the Cooperiidae, the adaptation to Lagomorpha may have occurred either within the Libyostrongylinae which may have remained in the Ethiopian region since the Paleocene, or, more likely, by the passage of the Obeliscoidinae from the Nearctic region to the Asian, through the Bering strait. In all cases, the adaptation of the Trichostrongyloidea of Lagomorpha to Ruminants apparently took place during the Miocene, mainly in the Palearctic and the Ethiopian regions.

Adaptation, Physiological↗

[Phylogeny of Trichostrongyloidea nematodes as seen through some of their vertebrate hosts].

Reconstruction of the phylogenetic history of a parasitic group is clearly difficult due to a lack of fossil forms. Among the nematodes parasites of vertebrates, the superfamily Trichostrongyloidea is one of the richest groups in terms of number of species (more than 1000 described) and genera (175) and they therefore offer excellent opportunities to reconstruct their phylogeny. Trichostrongyles occur in the gut and less commonly in the stomach and the lungs of all classes of terrestrial vertebrates except the Perissodactyls and the Proboscidian. They have a world wide distribution and direct life cycle. The criteria used in the classification of the Trichostrongyloidea are essentially morphological. Morphological characters are numerous and an attempt has been made to distinguish their relative value. Information concerning the synlophe, which is the apparatus of locomotion and attachment of the worms in the gut of the hosts, has been found essential in order to construct a classification of the superfamily and understand its evolution. This analysis was greatly facilitated by information from two sources: 1) the morphology of free-living rhaditids, which are the ancestors of the trichostrongyles; 2) the ontogeny of the synlophe. The classification established above takes into account the different morphological characters, but only concerns extant animals. Data provided by the hosts permit us to date the appearance of different lineages and to follow their evolution in time and space. These data come from extant hosts and from paleobiogeography. 1) The evidence from extant host species includes the host distribution of the trichostrongyles in relation to the geographic distribution of the hosts. 2) The evidence from extinct forms includes the date of appearance of the host in the geological record as well as mammalian migration in geologic time periods: a) The date of the appearance of the hosts in the geological records permits us to date the origin of the different families of parasites. b) The manner in which host migration interacts with the parasite is very complex and a few examples will be considered, mainly from the caviomorphs, the sciuromorphs and the myomorphs. The combination of evidence from the morphology of the worms and evidence from the paleobiogeography of the hosts allows us not only to explain the present day host and geographic distributions of the parasites but also to reconstruct their evolutionary history. The phylogenetic tree of the Trichostrongyloidea proposed with Chabaud consists of three main branches which are morphologically clearly defined.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

[Trichostrongyloidea nematodes, parasites of Microchiroptera].

1. a) List of Nematodes collected by Professor Aellen in european Microchiroptera. Additionnal morphological data to the study of Molinostrongylus alatus, M. panousei, M. skrjabini. Description of M. aelleni n. sp. b) Description of M. richardae n. sp., M. benexae n. sp. et M. bauchoti n. sp., parasites of malagasian Molossidae. c) Description of M. colleyi n. sp. and M. owyangi n. sp., parasites of Malaysian Vespertilioninae, and of Allintoschius dunni n. sp., discovered in Myotis mystacinus from Malaysia and Pipistrellus nanus from Africa. 2. Taking into account the characteristics of the synlophe, the 17 species of the genus Molinostrongylus may be divided into five groups, each one being reasonably well characteristic of the genus of their Chiropteran host. 3. The composition of the Trichostrongyloidea fauna of Chiroptera and its relationship with Trichostrongyloidea from other Mammals (Tupaiidae, Pholidotes, Primates, Sciuridés) are analysed. Six groups are separated and divided into two well defined lines: 1) genus Strongylacantha, and 2) 12 genera stemming more or less directly from the Molineinae, 4. The three conical outgrowths at the tip of the female tail which differenciate presently the Anoplostrogylinae from the Molineinae appear to be an unreliable characteristic. The two subfamilies form a complex group which will be better understood if the evolution of the synlophe and that of the caudal bursa of the males are taken into account.

Animals↗

[Nematodirinae (Nematoda:Trichostrongyloidea) of Antilocapra and Ovis in Alberta, Canada].

In this first note, the Trichostrongyloidea belonging to the Nematodirinae parasites of Ruminants from Alberta are identified. New morphological data were given on Nematodirella antilocaprae (Price, 1927) parasite of Antilocapra americana. Nematodirus andersoni n. sp. is described from Ovis canadensis. Because of the deep indentation of the dorsal lobe, the species is similar to N. bioccai Rossi, 1983 parasite of Ovis aries in Iran. However, it is different in the reduced number of cuticular ridges of the males and by the small indentation between the dorsal lobe and the lateral lobes. In our opinion, the species of nematodirus identified as N. archari from Ovis canadensis in North America, are probably N. andersoni.

Animals↗

Tapironema coronatum n. gen., n. sp. (Trichostrongyloidea-Cooperiidae-Obeliscoidinae), a parasite of Holochilus brasiliensis and Tapirus terrestris.

In this paper we provide a description of Tapironema coronatum n. gen. n. sp. (Trichostrongyloidea, Obeliscoidinae) from the cricetid Holochilus brasiliensis or "water rat" in Argentina (Type material) and from Tapirus terrestris in French Guyana (voucher material in poor condition). The new genus is characterized by a corona radiata, an oesophageal tooth, a bilaterally synlophe with about 73 (male), 122 (female) cuticular ridges, a caudal bursa pattern 2-1-2 with rays 5 and 6 parallel and close together and rays 5 longer than rays 3. The most closely related genus is the monospecific Teporingonema Harris, 1985, from a Mexican lagomorph, Romerolagus. The cephalic extremity of this parasite is redescribed after the type-material. The systematic position of Teporingonema amongst the Obeliscoidinae is defined and the hypotheses concerning the origin of this sub family are provided.

Animals↗

A new host record of Camelostrongylus mentulatus (Nematoda; Trichostrongyloidea) from abomasum of a giraffe at a zoo in Japan.

Camelostrongylus mentulatus (Railliet et Henry, 1909) Orloff, 1933 (Nematoda; Trichostrongyloidea) was found from the abomasum of a three-year-old female cape giraffe, Giraffa camelopardalis giraffa, born and died in a zoo park in Yamaguchi prefecture, Japan. This is the new host record from Giraffidae and geographical distribution of C. mentulatus. Present case of C. mentulatus might be infected from other ruminants, e.g., camels, antelopes and goats, kept at a same paddock in the zoo. Risk of imported parasitic diseases by the zoo animals from outside of Japan is discussed.

Abomasum↗

Ovejector structure in the Haemonchinae (Nematoda:Trichostrongyloidea) of ruminants.

The use of ovejector structure in the phylogeny of the Haemonchinae is hampered by differences among nematologists in the application of terminology and the recognition of homologous parts. Some workers recognize a sphincter with 2 parts, but others recognize only the proximal, rounded part of the sphincter and include the distal cylindrical part of the sphincter with the vestibule. The results of this study demonstrate that all sphincters of Haemonchinae of ruminants have 2 parts. To encourage the application of a uniform terminology to homologous parts of the ovejector, we propose the use of the terms "sphincter 1" for the rounded part and "sphincter 2" for the cylindrical part. It is hoped that clarification of the terminology for ovejectors of the Haemonchinae will provide a model useful for improving descriptions of ovejectors throughout the Trichostrongyloidea.

Animals↗

Nematodes of the family Heligmonellidae (Trichostrongyloidea) collected from rodents of the Ryukyu archipelago and Taiwan.

Six species of the family Heligmonellidae (Nematoda: Trichostrongyloidea), including 3 new species, are recorded from rodents of the subfamily Murinae in the Ryukyu Archipelago and Taiwan. Heligmonoides ikeharai n. sp. from Tokudaia osimensis muenninki on Mt. Yonaha, Okinawa Island, is characterized by extremely long spicules and hypertrophied ridges in the prevulval region. Heligmonoides taiwanensis n. sp. from Apodemus draco on Mt. Alishan, Taiwan, is distinguished from other members of the genus in having a markedly asymmetrical bursa and stout bursal rays. Heligmonoides alishanensis n. sp. from Niviventer confucianus on Mt. Alishan differs from the allied forms in lacking hypertrophied ridges at the level of the middle of the spicules and in having longer spicules and a smaller body. Nippostrongylus sp. from N. confucianus on Mt. Alishan resembles Nippostrongylus brasiliensis but is distinguishable in that the externolateral ray is almost the same length as the lateroventral ray in the left lobe, and the fused tips of the spicules are thin and straight. Heligmonoides ryukyensis from Mus caroli and Orientostrongylus tenorai from Bandicota indica are first recorded from Taiwan. Heligmonellid nematodes parasitic in wild rodents in these areas are considered to have been introduced with their hosts from the mainlands of China and Japan through land connections in the Pleistocene.

Animals↗

[Tentative classification of the nematodes Trichostrongyloidea (author's transl)].

Taking into consideration the evolution of different characters, and mainly the evolutionary pattern of the "synlophe", we propose to divide the Trichostrongyloidea into 8 families: Ollulanidae, Dictyocaulidae, Strongylacanthidae, Amidostomatidae, Molineidae, Heligmosomidae, Heligmonellidae and Trichostrongylidae.

Animals↗

[Presence of a new Trichostrongyloidea (Nematoda) Nochtia (Nochtia) atelerixi n. sp. in an Erinacoidea (Insectivora) in Togo].

Nochtia (Nochtia) atelerixi n. sp. (Nematoda, Trichostrongyloidea) of Ethiopian Erinacoidea. - Nochtia (Nochtia) atelerixi n. sp. is found in the intestine of the hedgehog Atelerix albiventris in Togo. It is distinguished from the two other congeneric species parasite of asiatic Primates, N. (N.) nochti Travassos et Vogelsang, 1929 and N. (N.) tani Le Van Hoa, 1966, mainly by the highest number of cuticular crests and spicules with six tips.

Animals↗

[A new classification of Trichostrongyloidea nematodes (author's transl)].

The preliminary classification that we proposed in 1977 grouped provisionally in the family Amidostomatidae all the primitive Trichostrongyloidea (with the exception of the Strongylacanthidae). The study of several nematodes from reptiles and amphibians, and above all parasites of Australian monotremes and marsupials, lead us to split up this family. We believe that two families, Amidostomatidae and Strongylacanthidae evolved from ancestors close to the Ancylostomatoïdea. All the others, however, appear to have evolved from the Strongyloïdea. The morphological characters considered permit the elucidation of three principal evolutionary lines, recognized for the primitive forms by the anatomy of the caudal bursa and for specialized forms by the structure of the synlophe. The principal evolution of the group seems to have occurred during the Secondary Era in Gondwanaland. The families and sub-families remain very numerous in the Neotropical and Australian zones.

Amphibians↗

Cuticular Architecture of Hassalstrongylus epsilon (Nematoda: Trichostrongyloidea).

Hassalstrongylus epsilon is a small nematode, whose adult forms are found among the intestinal microvilli of the water rat Nectomys squamipes, Brants 1827 (Rodentia: Muridae). The external appearance of the cuticle, which presents transversal striations and longitudinal ridges, is described using scanning electron microscopy. Transmission electron microscopy of thin sections and replicas of quick-frozen, freeze-fractured, deep-etched and rotatory shadowed samples showed the presence in the cuticle of struts that arise from the fluid median layer, extending outward to the epicuticle. The cuticle showed the presence of five layers: epicuticle, cortical, fibril-rich, fluid median and fibrous. The cuticle layers were made of an assemblage of fibers that create compartments, which were larger in the fluid region than in the fibril-rich median layer.

Animals↗

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.

Animals↗

Parasitic gastritis in a llama (Lama glama) associated with inhibited larval Teladorsagia spp. (Nematoda: Trichostrongyloidea).

A 7-year-old female llama was experimentally infected with 500 Fasciola hepatica metacercariae then turned out onto pasture also grazed by domestic sheep. The llama was necropsied approximately 22 weeks post infection. Routine examination of the gastrointestinal tract revealed diffusely coalescing, umbilicated nodules covering the caudal one-fifth of the third stomach compartment. Microscopically, the mucosa was irregularly thickened. Numerous nematode larvae were present in glandular lumens, often extending to the base of the glands. Where nematodes were most numerous, decreased numbers of parietal cells, attenuation of glandular epithelium and increased collagen within the lamina propria were evident. Of a total of 6510 adult nematodes in the third stomach compartment, 51% were Teladorsagia spp., 47% were Trichostrongylus axei, and 2% were Camelostrongylus mentulatus. Except for C. mentulatus, three sheep that had grazed with the llama and were necropsied at approximately the same time, harbored the same nematode species. The total number of larvae in the llama was 56,710, with more than 97% being ostertagiine early fourth-stage larvae. Ostertagiine larvae were also present in the sheep. Based on the composition of the adult populations in the llama and sheep, the larvae were considered to be species of Teladorsagia. Additionally, the larger numbers of early fourth-stage larvae indicated that the development of Teladorsagia spp. in the llama is inhibited.

Animals↗

Cross antigenicity among ovine trichostrongyloidea. Preliminary report.

A preliminary trial on the extent of cross-antigenicity among the sheep strongylids Haemonchus contortus, Trichostrongylus colubriformis, Teladorsagia circumcincta and Nematodirus battus in 2.5- to 4-month-old lambs has been carried out using ELISA and Western blotting (WB). Cross antigenicity was tested using soluble extracts from adult and third stage larvae (L3) of H. contortus as antigenic source probed with sera from lambs with monospecific heterologous infections. There was cross-antigenicity between L3 of H. contortus and Trichostrongylus colubriformis in ELISA and WB. Immunodetection results with adult H. contortus antigen showed a closer relationship to Teladorsagia circumcincta. Certain heterologous sera reacted with H. contortus antigens more strongly than the homologous one, but sera from the H. contortus-infected animals had reactivity around the 25 kDa region from adult antigens which could have potential diagnostic use.

Animals↗