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Phylogenetic relationships of the soil nematode orders Dorylaimida, Mononchida, Triplonchida and Alaimida, with a revised classification of the subclass Enoplia
... The longidorid nematodes, containing some virus vector nematode genera, were assigned to the superfamily Dorylaimoidea under suborder Dorylaimina of order Dorylaimida. While the trichodorid nematodes were placed in superfamily Trichodoroidea under suborder Diptherophorina of the order Triplonchida (Hunt, 1993) as earlier proposed by Siddiqi (1983) and Jairajpuri and Ahmad (1992). ...
... texanus Cobb, 1913 Latocephalus Patil & Khan, 1982L. smithi (Heyns, 1963 Patil & Khan, 1982 Sindellus Siddiqi, 1983S. sindi Siddiqi, 1983 RashidanemaSiddiqi, 1983 R. cognatum (Siddiqi, 1983) Jairajpuri & Ahmad, 1992 Tylencholaimoidea Filipjev, 1934Tylencholaimidae Filipjev, 1934Tylencholaiminae Filipjev, 1934 Tylencholaimus de Man, 1876 ...
... smithi (Heyns, 1963 Patil & Khan, 1982 Sindellus Siddiqi, 1983S. sindi Siddiqi, 1983 RashidanemaSiddiqi, 1983 R. cognatum (Siddiqi, 1983) Jairajpuri & Ahmad, 1992 Tylencholaimoidea Filipjev, 1934Tylencholaimidae Filipjev, 1934Tylencholaiminae Filipjev, 1934 Tylencholaimus de Man, 1876 ...
... A common feature of all these concepts, except that of Siddiqi (1983), is the presumed close relationship between Dorylaimida and the diphtherophorids and trichodorids, now usually placed in the order Triplonchida Cobb, 1920. Molecular studies employing SSU rDNA have suggested a basal position for the Dorylaimia (clade I in Blaxter et al., 1998) on the overall tree for the phylum. ...
... Diphtherophorina/Triplonchida has not been recovered as a sister taxon to Dorylaimida and this group clusters with other Enoplia Pearse, 1942 based on evaluation of the phylogenetic signal transmitted by SSU sequences. To date there have been no molecular studies which have included any sequences obtained from members of Bathyodontina to investigate Siddiqi's (1983) hypothesis. The genus Bathyodontus was originally placed in Nygolaiminae Thorne, 1935 when described (Fielding, 1950), but subsequently transferred to a separate family, Bathyodontidae (Clark, 1961). ...
... In his original description, Cobb (1920) placed Isolaimium in the order Isolaimia Cobb, 1920, along with numerous other species. Isolaimium was not treated by Clark (1961), but Goodey (1963) Clark (1961), Andrássy (1976), Lorenzen (1981Lorenzen ( , 1983, Maggenti (1983), Siddiqi (1983), and De Ley and Blaxter (2002). Vol. ...
Phylogenetic reconstructions based on 18S rDNA sequence data indicate that Dorylaimida, comprising the suborders Nygolaimina and Dorylaimina, is a monophyletic lineage, but that there is a deep division within Nygolaimina, giving rise to the possibility that Nygolaimina is paraphyletic. A well-supported clade comprising members of the traditional orders Mermithida and Mononchida (including Bathyodontina) forms the sister taxon to the Dorylaimida. Inferred relationships within this clade indicate that Mermithida shares more recent common ancestry with Mononchina than does Bathyodontina. Vertebrate parasites within Dorylaimia (Dioctophymida and Trichinellida) are reconstructed in a sister-taxon relationship with the Mononchida/Dorylaimida lineage. The enigmatic order Isolaimida (represented by Isolaimium) appears to be ancestral to all other Dorylaimia sampled. Expanded taxon sampling for phylogenetic analyses of the subclass raises new possibilities for the reconstruction of hypothetical character states in the common ancestor of Dorylaimia.
... Thus, the evolution of Campydoridae was believed to represent 'an independent development of the pre-bathyodontidstock' (Yeates, 1967); the systematic position of the Campydoridae remained 'uncertain'. This uncertainty was primarily due to the absence of pharyngo-intestinal 'glands' in Campydora; presence of these structures is characteristic of most Nygolaimidae, and they also occur in some aporcelaim taxa, as well as in some members of the presumably ancestral Bathyodontina Siddiqi, 1983(Coomans & Loof, 1970. ...
... This species remains a species inquirenda. Siddiqi (1983) took exception to Maggenti's 1982 classi cation, stating that the homology between the mural tooth of Campydora and that of nygolaim nematodes could not be justi ed 'as Campydora is not a dorylaim'. Other morphological features, including the structure of the pharynx and of the amphids (in Dorylaimida, the amphidial fovea is connected to the sensillum pouch by a duct, the canalis amphidialis; this structure is lacking in Campydora), were employed to support the assignment of Campydorina to the Enoplida (Siddiqi, 1983). ...
... Siddiqi (1983) took exception to Maggenti's 1982 classi cation, stating that the homology between the mural tooth of Campydora and that of nygolaim nematodes could not be justi ed 'as Campydora is not a dorylaim'. Other morphological features, including the structure of the pharynx and of the amphids (in Dorylaimida, the amphidial fovea is connected to the sensillum pouch by a duct, the canalis amphidialis; this structure is lacking in Campydora), were employed to support the assignment of Campydorina to the Enoplida (Siddiqi, 1983). Due to the nature of the stomatal armature, Maggenti (1991) retained Campydoridae in the Nygolaimina. ...
The systematic position of Campydora Cobb, 1920, which possesses many unique morphological features, especially in pharyngeal structure and stomatal armature, has long been a matter of uncertainty with the 'position of the Campydorinae' (containing only Campydora) being questionable. A review of the morphology of C. demonstrans, the only nominal species of Campydora concluded that the species warranted placement as the sole member of a monotypic suborder, Campydorina, in the order Dorylaimida. Others placed Campydorina in the order Enoplida. We conducted phylogenetic analyses, using 18s small subunit ribosomal DNA sequences generated from a number of taxa in the subclasses Enoplia and Dorylaimia, to evaluate these competing hypotheses. Although precise taxonomic placement of the genus Campydora and the identity of its closest living relatives is in need of further investigation, our analyses, under maximum parsimony, distance, and maximum likelihood criteria, unambiguously indicate that Campydora shares a common, more recent, ancestry with genera such as Alaimus, Pontonema, Tripyla and Ironus (Enoplida), rather than with any members of Dorylaimida, Mononchida or Triplonchida.
... Before the advent of molecular techniques, a number of taxonomic classifications attempted to catalogue the substantial morphological diversity observed within the subclass Enoplia. Filipjev [11] was the first author to produce a comprehensive morphological classification of free-living nematodes, and subsequent revisions that included a focus on the Enoplia were completed by Pearse [12], Chitwood and Chitwood [13], Clark [14], De Coninck [15], Andrassy [16], Maggenti [17] , Loren- zen [18], and Siddiqi [19] . Morphological schemes primarily differed in their placement of the Tripyloididae, Alaimidae, Ironidae (all currently grouped in the Enoplida ), and the Mononchoidea (now grouped under the subclass Dorylaimia). ...
... This investigation consistently recovered the Tripyloididae as a sister taxon to the clade containing the Trefusiidae, Tripylina, and Trischistoma. Such a relationship was first suggested by Siddiqi [19] who placed the Tripylidae species and the Tripyloididae together in the order Tripylida based on morphology. The molecular framework outlined by Van Megen et al. [10] recovered a similar association between these taxa, although this relationship demonstrated low support values in their analysis (ML bootstraps <50%). ...
... Morphological evidence has previously suggested a close relationship between the Rhabdolaimidae and Syringolaimus; most classifications grouped Syringolaimus within the Ironidae [11,14,15,18], but some placed this genus within the Rhabdolaimidae in the Ara- eolaimida [16,37] . The genus Campydora was historically classified within the Dorylaimia based on morphology [15,36,38]; only Siddiqi [19] was the only author to suggest a relationship with the Enoplia based on the structure of the pharynx and amphids. The current placement of Syringolaimus clearly denotes the polyphyly of the Ironidae sensu Lorenzen [18], a finding that was originally put forth by Meldal et al. [7] upheld in later phylogenies [10]. ...
The subclass Enoplia (Phylum Nematoda) is purported to be the earliest branching clade amongst all nematode taxa, yet the deep phylogeny of this important lineage remains elusive. Free-living marine species within the order Enoplida play prominent roles in marine ecosystems, but previous molecular phylogenies have provided only the briefest evolutionary insights; this study aimed to firmly resolve internal relationships within the hyper-diverse but poorly understood Enoplida. In addition, we revisited the molecular framework of the Nematoda using a rigorous phylogenetic approach in order to investigate patterns of early splits amongst the oldest lineages (Dorylaimia and Enoplia).
Morphological identifications, nuclear gene sequences (18S and 28S rRNA), and mitochondrial gene sequences (cox1) were obtained from marine Enoplid specimens representing 37 genera. The 18S gene was used to resolve deep splits within the Enoplia and evaluate the branching order of major clades in the nematode tree; multiple phylogenetic methods and rigorous empirical tests were carried out to assess tree topologies under different parameters and combinations of taxa. Significantly increased taxon sampling within the Enoplida resulted in a well-supported, robust phylogenetic topology of this group, although the placement of certain clades was not fully resolved. Our analysis could not unequivocally confirm the earliest splits in the nematode tree, and outgroup choice significantly affected the observed branching order of the Dorylaimia and Enoplia. Both 28S and cox1 were too variable to infer deep phylogeny, but provided additional insight at lower taxonomic levels.
Analysis of internal relationships reveals that the Enoplia is split into two main clades, with groups consisting of terrestrial (Triplonchida) and primarily marine fauna (Enoplida). Five independent lineages were recovered within the Enoplida, containing a mixture of marine and terrestrial species; clade structure suggests that habitat transitions have occurred at least four times within this group. Unfortunately, we were unable to obtain a consistent or well-supported topology amongst early-branching nematode lineages. It appears unlikely that single-gene phylogenies using the conserved 18S gene will be useful for confirming the branching order at the base of the nematode tree-future efforts will require multi-gene analyses or phylogenomic methods.
... The members of this phylum occur in very high abundance within an environment (Lambshead, 2004;Danovaro et al., 2008) and thought to be hyper diverse (Appeltans et al., 2012). The order Enoplida Filipjev, 1929, comprises 7 suborders, amongst which the suborder Ironina Siddiqi, 1983, consist of one superfamily Ironoidea de Man, 1876, which further consists of 3 families; Ironidae de Man, 1876, Leptosomatidae Filipjev, 1916 andOxystominidae Chitwood, 1935. Family Ironidae is further divided into two subfamilies; Ironinae de Man, 1876 andThalassironinae Andrássy, 1976. ...
... The specimens have been deposited in the National Zoological Collections of Nemathelminthes section, Zoological Survey of India, Kolkata. Chen and Guo, 2015 Phylum : Nematoda Cobb, 1932 Class : Enoplea Inglis, 1983 Subclass : Enoplia Pearse, 1942 Order : Enoplida Filipjev, 1929 Suborder : Ironina Siddiqi, 1983 Superfamily ...
Two species of free-living marine nematodes were collected from the intertidal areas of East Coast of India namely, Conilia sinensis Chen and Guo, 2015 and Corononema parvum Nicholas and Stewart, 1995, both species are new records for the Indian fauna. Previously, the genus Conilia Gerlach, 1956 was known from Santos Beach, South Japan, East China Sea and Korea. The genus Corononema Nicholas and Stewart, 1995 was known only from Australia and Thailand and very recently reported from off the coast of Vietnam. Both the genera are being reported for the first time from India. The species are described and photomicrographed.
... In order to take into account this new evidence regarding the placement of the genera Tripylina and Trischistoma, Zhao (2011) proposed an updated classification where these two genera are placed within the family Trischistomatidae Andrássy, 2007 in the suborder Tripyloidina. This placement was considered conservative, as the Tripylidae had previously been classified together with the Tripyloididae by Lorenzen (1981) and Siddiqi (1983), and the Trefusiina was exclusively marine. The classification of the Tripylidae with the Enoplida by Lorenzen (1981) was partly based on his observation of metanemes in Tripylina glomerans Bastian, 1865;Zhao (2011) later also observed metanemes in four Trischistoma species he described from New Zealand. ...
... Caudal glands (when present) lie completely within the tail or position unknown. Lorenzen (1981) erected the order Trefusiida, which was subsequently the lowered to the level of suborder within the Enoplida in the classification of De Ley & Blaxter (2004) based on analyses of 18S rRNA sequences (Rusin et al. 2001) and following Siddiqi (1983). The Trefusiida was not considered monophyletic by Lorenzen (1981) because it is not characterized by any character which is apomorphic for that taxon. ...
We provide a review of the enoplid suborder Trefusiina Siddiqi, 1983, based on morphological considerations and analyses of new and published 18S rDNA sequences. We also describe Halanonchus scintillatulus Leduc sp. nov. from the Hauraki Gulf, northern New Zealand, as well as females of Trefusialaimus idrisi Leduc, 2013 from the continental slope of New Zealand. We show for the first time that the structure of the female reproductive system of Trefusialaimus Riemann, 1974 consists of two opposed and outstretched ovaries, an unusual feature for the Enoplida. The Trefusiina did not form a monophyletic group in the 18S rDNA phylogeny due to the placement of Lauratonema Gerlach, 1953 and Trefusialaimus sequences well away from the main Trefusiina clade. However, due to generally weak Maximum Likelihood support values, we refrain from changing the classification of these taxa until more comprehensive analyses can be conducted. Our phylogenetic analysis supports the inclusion of the Trischistomatidae Andrássy, 2007 within the Trefusiina, meaning that all of the enoplid suborders now include at least some terrestrial/freshwater representatives. The Trefusiina currently comprises five families, 14 genera and 92 valid species.
... The unique and poorly understood morphology of the genus Campydora hindered understanding of its relationships with other nematodes and resulted in the taxonomic inflation from the genus to the suborder level (Jairajpuri 1983). This decision was followed in subsequent publications, although the suborder was transferred from the order Dorylaimida to the order Enoplida based on the morphological and molecular evidence, and its relationships within the order Enoplida remained unresolved (Siddiqi 1983;Winiszewska 2001;Mullin et al. 2003;Zhao et al. 2012;Smol et al. 2014). Subsequent phylogenetic studies using 18S rDNA sequence suggested close affinities between the genera Campydora, Rhabdolaimus de Man, 1880 and Syringolaimus de Man, 1888, placing these three genera as a monophyletic clade within the unresolved order Enoplida (Meldal et al. 2007;van Megen et al. 2009;Bik et al. 2010;Shokoohi et al. 2013;Smythe 2015;Leduc et al. 2018). ...
... This grouping can be further substantiated by at least one morphological character: the presence of a well-developed basal bulb, not found in any other Enoplida (the genera Polygastrophora de Man, 1922 andBelbolla Cobb, 1920 have the posterior part of the pharynx modified into a series of bulb-like swellings that are of independent origin). Placement of the genus Syringolaimus in the suborder Campydorina is further supported by the monorchic male reproductive system similar to the reproductive system found in the species from the genera Campydora and Rogerus (Hoeppli & Chu 1932;Winiszewska 2001;Tchesunov 2017), while it is diorchic in other genera of Ironina Siddiqi, 1983(Shokoohi et al. 2013Chen & Guo 2015). The revised classification of the suborder Campydorina is given below. ...
The new genus and species Campydoroides manautei Holovachov gen. et sp. nov. is placed in the suborder Campydorina and is characterised by a transversely striated cuticle without lateral alae, body pores or epidermal glands; somatic sensilla only on pharyngeal region and on tail; a truncate labial region with papilliform inner labial, outer labial and cephalic sensilla; a stirrup-shaped amphid with transverse slit-like opening; a conoid stoma with strongly cuticularised walls and large protrusible dorsal tooth; a cylindrical pharynx with distinct basal bulb but without valves; a large ovoid cardia; didelphic, amphidelphic female gonads with antidromously reflexed ovaries and without spermatheca; a transverse vulva; a straight vagina without pars refringens vaginae or epiptygmata; an elongate tail with caudal glands and spinneret. The new genus is similar to the genera Campydora Cobb, 1920 and Udonchus Cobb, 1913 in having papilliform labial and cephalic sensilla, a stirrup-shaped amphid with a transverse slit-like opening, a stoma with a well-developed protrusible dorsal tooth, and a muscular pharynx with a strongly developed basal bulb, but can be easily separated from both in details of a stoma morphology. The systematics of the suborder Campydorina is revised. Halirhabdolaimus Siddiqi, 2012 is synonymised with Syringolaimus de Man, 1888.
... The Triplonchida were conWrmed as an order within the Enoplia, consistent with Siddiqi (1983) but contrary to many earlier classiWcations that were based on morphological data alone and placed part of this group among the Dorylaimia (e.g. Thorne, 1939;Clark, 1961;Siddiqi, 1961Siddiqi, , 1973De Coninck, 1965;Coomans and Loof, 1970). ...
... The Tripyloididae were not associated with the Chromadoria, as has been inferred from the presence of spiral amphids (De Coninck and Schuurmans Stekhoven, 1933;Chitwood and Chitwood, 1950;De Coninck, 1965;Andrássy, 1976). The position of Campydora demonstrans as part of the Enoplida has been conWrmed (Siddiqi, 1983;Mullin et al., 2003). This taxon was previously classiWed amongst the Dorylaimia (Thorne, 1939;Jairajpuri and Ahmad, 1992;. ...
... The Triplonchida were conWrmed as an order within the Enoplia, consistent with Siddiqi (1983) but contrary to many earlier classiWcations that were based on morphological data alone and placed part of this group among the Dorylaimia (e.g. Thorne, 1939;Clark, 1961;Siddiqi, 1961Siddiqi, , 1973De Coninck, 1965;Coomans and Loof, 1970). ...
... The Tripyloididae were not associated with the Chromadoria, as has been inferred from the presence of spiral amphids (De Coninck and Schuurmans Stekhoven, 1933;Chitwood and Chitwood, 1950;De Coninck, 1965;Andrássy, 1976). The position of Campydora demonstrans as part of the Enoplida has been conWrmed (Siddiqi, 1983;Mullin et al., 2003). This taxon was previously classiWed amongst the Dorylaimia (Thorne, 1939;Jairajpuri and Ahmad, 1992;. ...
Phylogenetic reconstructions of relations within the phylum Nematoda are inherently difficult but have been advanced with the introduction of large-scale molecular-based techniques. However, the most recent revisions were heavily biased towards terrestrial and parasitic species and greater representation of clades containing marine species (e.g. Araeolaimida, Chromadorida, Desmodorida, Desmoscolecida, Enoplida, and Monhysterida) is needed for accurate coverage of known taxonomic diversity. We now add small subunit ribosomal DNA (SSU rDNA) sequences for 100 previously un-sequenced species of nematodes, including 46 marine taxa. SSU rDNA sequences for >200 taxa have been analysed based on Bayesian inference and LogDet-transformed distances. The resulting phylogenies provide support for (i) the re-classification of the Secernentea as the order Rhabditida that derived from a common ancestor of chromadorean orders Araeolaimida, Chromadorida, Desmodorida, Desmoscolecida, and Monhysterida and (ii) the position of Bunonema close to the Diplogasteroidea in the Rhabditina. Other, previously controversial relationships can now be resolved more clearly: (a) Alaimus, Campydora, and Trischistoma belong in the Enoplida, (b) Isolaimium is placed basally to a big clade containing the Axonolaimidae, Plectidae, and Rhabditida, (c) Xyzzors belongs in the Desmodoridae, (d) Comesomatidae and Cyartonema belongs in the Monhysterida, (e) Globodera belongs in the Hoplolaimidae and (f) Paratylenchus dianeae belongs in the Criconematoidea. However, the SSU gene did not provide significant support for the class Chromadoria or clear evidence for the relationship between the three classes, Enoplia, Dorylaimia, and Chromadoria. Furthermore, across the whole phylum, the phylogenetically informative characters of the SSU gene are not informative in a parsimony analysis, highlighting the short-comings of the parsimony method for large-scale phylogenetic modelling.
... There are six other species/populations of Onchulus described on the base of males. Both O. nolli and O. fl agellatus do not possess cervical sensilla and have different stoma morphology, and Onchulus sp. in Siddiqi (1983) is not described in details enough for comparison. The description of O. oistospiculum (Allgen, 1933) Gerlach & Riemann, 1974 is very incomplete, the only putative dif-ference between O. straticauda and O. oistospiculum is that cervical sensilla were not described in the former species. ...
... On the other hand, O. capriviensis, O. longicauda, O. gracilis Andrássy, 2001, O. dorsalis Andrássy, 2001and O. biannulatus Hodda, Bloemers & Wanless, 2005 are similar in having smaller body size, stoma morphology and presence of subcephalic setae. The male of O. capriviensis is morphologically similar to onchulid males described under the names O. longicauda by Gerlach (1966), O. straticauda by Altherr (1972) and Onchulus sp. by Siddiqi (1983) in the presence of cervical papillae and spicule morphology. O. microdontus Hodda, Bloemers & Wanless, 2005 is insuffi ciently described to be included in the cladistic analysis. ...
... | 1 view, grounded in the pharyngeal and amphid morphology, supported the position of Campydorinae within Enoplida (Siddiqi, 1983), eventually identifying the morphology-based alternatives to place the Campydora genus either within Dorylaimida or Enoplida. The affinity of C. demonstrans within the order Enoplida has been validated in SSU rDNA-based phylogenetic analyses (Mullin et al., 2003;van Megen et al., 2009), including a large-scale Bayesian assay of >200 taxa (Ahmed & Holovachov, 2021). ...
The phylum Nematoda remains very poorly sampled for mtDNA, with a strong bias toward parasitic, economically important or model species of the Chromadoria lineage. Most chromadorian mitogenomes share a specific order of genes encoded on one mtDNA strand. However, the few sequenced representatives of the Dorylaimia lineage exhibit a variable order of mtDNA genes encoded on both strands. While the ancestral arrangement of nematode mitogenome remains undefined, no evidence has been reported for Enoplia, the phylum's third early divergent major lineage. We describe the first mitogenome of an enoplian nematode, Campydora demonstrans, and contend that the complete 37-gene repertoire and both-strand gene encoding are ancestral states preserved in Enoplia and Dorylaimia versus the derived mitogenome arrangement in some Chromadoria. The C. demonstrans mitogenome is 17,018 bp in size and contains a noncoding perfect inverted repeat with 2013 bp-long arms, subdividing the mitogenome into two coding regions. This mtDNA arrangement is very rare among animals and instead resembles that of chloroplast genomes in land plants. Our report broadens mtDNA taxonomic sampling of the phylum Nematoda and adds support to the applicability of cox1 gene as a phylogenetic marker for establishing nematode relationships within higher taxa.
... Considering both the terrestrial and the marine nematodes, about 25% of them are freeliving marine forms (Du et al., 2014). The order Enoplida Filipjev, 1929, consist of 7 suborders, amongst which the suborder Ironina Siddiqi, 1983, consist of one superfamily Ironoidea de Man, 1876, which further comprises of 3 families; Ironidae de Man, 1876, Leptosomatidae Filipjev, 1916 andOxystominidae Chitwood, 1935. The genus Trissonchulus Cobb, 1920 belongs to the family Ironidae de Man, 1876. ...
We report two new records of free-living Marine Nematodes from India, viz. Trissonchulus provulvatus Orselli and Vinciguerra, 1997 and Antomicron quindecimpapillatus Holovachov, 2012. Sediment samples were collected from Coastal regions of Puducherry. Morphological data along with description and distribution for both the species are provided here. Both the species reported here have been recorded previously only from their type locality: T. provulvatus Orselli and Vinciguerra, 1997 from the Mouth of River Simeto, Catania, Italy, while, A. quindecimpapillatus Holovachov, 2012 from Gullmarn Fjord near Fiskebackskil, Sweden.
... Oxystominina Siddiqi, 1983 Suborder Alaimina Clark, 1961 Diagnosis: The lips are provided with papillae or are represented by small pit-like depressions. The cephalic sensillae are papillose. ...
... Ironidae is now the only family of the superfamily Ironoidea de Man, 1876 and the suborder Ironina Siddiqi, 1983 of the order Enoplida. Lorenzen (1981Lorenzen ( , 1994 established the holophyly of the Ironidae based on buccal cavity armature: three (or four if the dorsal tooth is split into two) movable teeth on the anterior edge of the elongate cylindrical buccal cavity; in juvenile nematodes, the teeth replacing those of the following stage are formed behind the functioning teeth. ...
Three nematode species of the genus Syringolaimus (Enoplida, Ironidae) were found in the Angola Basin (south-east Atlantic Ocean) at depths of almost 5500 m. One of them is described as new for science and the other two were initially recorded from the deep sea off the Brazilian coast and are redescribed here. Syringolaimus elegans sp. n. differs from all other species of Syringolaimus by a slimmer body (a=99 vs. 30–73) and from all species except S. ingens (nom. nud.) by its greater body length (2718 μm vs. 760–1750 μm), longer buccal cavity (79 μm vs. 28–58 μm), and longer terminal pharyngeal bulb (70 μm vs. 53 μm). All the Syringolaimus species together make up from 0.75 to 2.82% of all nematode individuals from all samples. Diagnoses of Syringolaimus annae Lima et al., 2009 and S. smolae Lima et al., 2009 are updated. An annotated list of Syringolaimus species is provided; the species annae, elegans, filicaudatus, loofi, magdae, renaudae, smolae, striatocaudatus and taniae are considered as valid, while brevicaudatus, caspersi, lichenii, marisalbi, smarigdus and venustus as species inquirendae, and gladiatus, ingens, nitidus and venustus as nomina nuda. Diagnostic characters and problems of species identification are discussed.
... Oxystominina Siddiqi, 1983 Suborder Alaimina Clark, 1961 Diagnosis: The lips are provided with papillae or are represented by small pit-like depressions. The cephalic sensillae are papillose. ...
Soil Nematodes of Grasslands in Northern China presents research on China's temperate grasslands, providing the findings and results of a large field survey along a transect across the northern temperate grassland. It examines nematode distribution patterns along the transect from trophic group and family, to genus level, also evaluating their relationship with climatic conditions, plant biomass and soil parameters. The book then presents detailed taxonomy information of nematodes to genus or species level, providing keen insights into nematode diversity along the grassland transect in north China. Final sections review the advances and perspectives for the research of soil ecology on soil nematodes in China, including recent major discoveries of soil microbial diversity and eco-function during this field survey. This work will help researchers predict the impact of global change drivers on below ground soil biota and better understand the functioning and services they provide in terrestrial ecosystems. © 2017 Zhejiang University Press. Published by Elsevier Inc. All rights reserved.
... Trefusiidae is now considered as a member of the order Enoplida Filipjev, 1929, which is supported by morphological (Andrássy 1983;De Coninck 1965;Filipjev 1934) and molecular data (Bik et al. 2010;Holterman 2008;Rusin et al. 2001;van Megen et al. 2009). Morphological (Siddiqi 1983) and molecular analyses (Bik et al. 2010) indicate that Trefusiidae is closely related to Tripyloididae Filipjev, 1918, differing from the latter by the cephalic setae widely separated from the six outer labial setae, the two testes, and the lack of teeth in the buccal cavity. ...
A new nematode species, Africanema multipapillatum sp. nov., is described from a sand beach in the East China Sea. The new species is assigned to the family Trefusiidae based on both the morphological and molecular analyses. Africanema multipapillatum sp. nov. is a distinct species characterized by its toothless and spacious buccal cavity, jointed labial setae, non-spiral and elongate groove-shaped amphidial fovea, faintly striated cuticle, and a single posterior ovary. Within the family Trefusiidae, the new species is most similar to the monotypic genus Africanema, but differs distinctly from Africanema interstitiale by the long and curved spicules with gubernaculum apophysis, long and slim sperm cells, and the lack of pharyngeal papillate supplements. Molecular phylogenetic analyses indicate that Africanema multipapillatum sp. nov. and the genera Rhabdocoma and Trefusia always fell within a single clade of the family Trefusiidae, where it is closely related to Rhabdocoma. Thus, we propose assigning the genus Rhabdocoma from the subfamily Trefusiinae to the subfamily Halanonchinae.
... So far, the only morphological feature claimed to be unique for the order Triplonchida is the modification of spicule protractor muscles into two " capsule-like structures that surround the anterior part of each spicule and which appear to squeeze out the spicules rather than simply pulling them backwards " (cited from De Ley & Blaxter 2002, p. 17). This feature was already mentioned by Siddiqi (1983, p. 9), but he proposed a separate ordinal status for Tripylida Siddiqi, 1983 and Triplonchida. Unfortunately, the exact morphology of the male copulatory musculature is unknown in the families Rhabdodemaniidae, Pandolaimidae and Triodontolaimidae, and their placement within the order Triplonchida is provisional. ...
... Adoncholaimus differs from the other three genera in the family mainly by the structure of the Demanian system (Smol & Coomans, 2006), a sperm-storage organ in the female and unique to Oncholaimidae (Rachor, 1969). Siddiqi (1983) established a new order, Oncholaimida, for Oncholaimidae and Enchelidiidae Filipjev, 1918, which had formerly been placed in the Enoplida Filip-western Japan, but there has been no taxonomic study on Japanese Adoncholaimus. In the present study, we describe two new species of Adoncholaimus from Hokkaido, northern Japan, representing the first species in this genus identified from Japan. ...
Two new species of free-living marine nematodes, Adoncholaimus daikokuensis sp. nov. and A. pseudofervidus sp. nov., from the coastal area of northern Japan, are described and illustrated. Adoncholaimus daikokuensis sp. nov. is similar to five congeners, A. derjugini, A. punctatus, A. oxyuroides, A. squalus comb. nov. and A. filicauda comb. nov. in the absence of a gubernaculum, but differs in the absence of a ventral swelling on tail, spicule length and buccal cavity length. Adoncholaimus pseudofervidus sp. nov. is similar to A. fervidus in having large body size, short tail, similar positions of the excretory pore and nerve ring, short spicules, arrangement of subventral setae in posterior region of body in males, and a single pair of terminal pores of the Demanian system surrounded by small gland cells in females. Adoncholaimus pseudofervidus sp. nov. differs from A. fervidus in having smaller, more anteriorly located amphids, longer buccal cavity, absence of gubernaculum in the male, and a different position of the terminal pores in the female. Partial sequences of the mitochondrial cytochrome c oxidase subunit I gene (309-337 bp) of the new species are provided for identification based on the DNA barcoding method. Metoncholaimoides is proposed as a junior synonym of Adoncholaimus. The new diagnosis of Adoncholaimus and a key to species are provided. Adoncholaimus squalus comb. nov., Adoncholaimus filicauda comb. nov., and Admirandus papillatus comb. nov. are proposed.
... A posição sistemática da família Trichodoridae é a seguinte, segundo Siddiqi (1983) Pearse, 1942 Ordem Triplonchida Cobb, 1920Subordem Diphtherophorina Coomans & Loof, 1970 Superfamília Diphtherophoroidea Micoletzky, 1922 Família Trichodoridae Thorne, 1935 Para além da criação recente do gênero Ecuadorus, este grupo de nematóides passou por algumas propostas taxonômicas, embora sem aceitação unânime por parte dos especialistas, referindo-se as principais: em 1974, Siddiqi propôs a divisão do gênero Paratrichodorus em três sub-gêneros: Paratrichodorus, Atlantadorus e Nanidorus. Posteriormente, o mesmo autor elevou estes três sub-gêneros ao nível genérico (Siddiqi, 1980); no entanto os gêneros Atlantadorus e Nanidorus Siddiqi, 1974 foram rejeitados por diversos autores (Decraemer, 1980;Decraemer & De Waele, 1981;Sturhan, 1985;Baujard & Decraemer, 2000), apesar de terem sido repostos mais tarde ao nível de sub-gênero, por Ahmad (1989) e Jairajpuri & Ahmad (1992) Baujard, 1980 T. azorensis Almeida, De Waele, Santos & Sturhan, 1989 T. beirensis Almeida, De Waele, Santos & Sturhan, 1989 T. borai Rahman, Jairajpuri & Ahmad, 1985 T. borneoensis Hooper, 1962 T. californicus Allen, 1957 T. carlingi Bernard, 1992 T. cedarus Yokoo, 1964: sin. ...
... The molecular phylogenetic study of Meldal et al. (2007) confirmed that 1) the Triplonchida is an order within Enoplia, consistent with Siddiqi (1983) but contrary to many earlier classifications that were based on morphological data alone and placed part of this group among the Dorylaimia (Thorne 1939;Clark 1961;Siddiqi 1961Siddiqi , 1973De Coninck 1965;Coomans & Loof 1970); 2) within Triplonchida, the Diphtherophoroidea were well supported as monophyletic; 3) contrary to morphological classifications, Trischistoma monohystera appears to be more closely related to Enoplida than to Triplonchida as the latter order forms a well supported clade excluding T. monohystera. ...
We have made an extensive study of New Zealand representatives of nematodes from the family Tripylidae de Man, 1876. Based on SSU DNA sequence data and phylogenetic analysis, the genera Tripylina Brzeski, 1964 and Trischistoma Cobb, 1913 are not closely related to Tripyla Bastian, 1865, the type genus of the family Tripylidae de Man 1876. The genus Tripylina is sister to Trischistoma and Trefusia de Man, 1893 and is more closely related to Enoplida than to Triplonchida. Our phylogenetic results indicate that Tripylina should be placed in Enoplida.
... Cobb (1913) described the genus Trischistoma and proposed (1920) the Order Triplonchia. The latter was amended to Triplonchida by Siddiqi (1983). Brzeski (1963) erected a new genus Tripylina under Tripylidae and considered Trischistoma as genus inquirendum. ...
The paper deals with description of three new and two known species of Tripylidae Örley, 1880. Two new and a known species of the genus TripylinaBrzeski, 1963 while one new and one known species of Trischistoma are described. Tripylina ymyensis n. sp. is characterised by robust body; long, slender outer labial setae; stoma with thick cuticularised wall bearing a prominent dorsal tooth and two equal-sized posteriorly-placed subventral teeth; large ovoid coelomocytes with conspicuous, globular inclusions; simple vagina without cuticularised pieces and a long, curved tail with a U- or S-shaped turn, uniformly tapering into a narrow terminus. Tripylina valiathani n. sp. is characterised by moderately large body; plump, leaf-shaped outer labial setae; stoma with thick cuticularised dorsal wall bearing a prominent dorsal tooth and two small anteriorly-placed subventral denticles; large fusiform coelomocytes with tapering ends having fine granular inclusions; cuticularised vagina, a distinct prerectum and a stumpy, laterally curved tail with a blunt terminus. T. stramenti (Yeates, 1972) Tsalolikhin, 1983 has been reported for the first time from India. Trischistoma minor n. sp. is characterised by medium-sized body; with dense crystalloids and granular aggregates; low, flattened lip region; plump outer labial setae; slenderer post-labial cephalic setae lying closely posterior to outer labials; a post-uterine sac of one body diameter and a dorsally curved tail with ventral terminal turn. T. pellucidumCobb, 1913 forms the first report from India. The relationships of Tripylina and Trischistoma have been discussed.
... The Tripyloididae were not associated with the Chromadoria, as has been inferred from the presence of spiral amphids (De Coninck and Schuurmans Stekhoven, 1933; Chitwood and Chitwood, 1950; De Coninck, 1965; Andrássy, 1976 ). The position of Campydora demonstrans as part of the Enoplida has been conWrmed (Siddiqi, 1983; Mullin et al., 2003). This taxon was previously classiWed amongst the Dorylaimia (Thorne, 1939; Jairajpuri and Ahmad, 1992;). ...
Phylogenetic reconstructions of relations within the phylum Nematoda are inherently difficult but have been advanced with the introduction of large-scale molecular-based techniques. However, the most recent revisions were heavily biased towards terrestrial and parasitic species and greater representation of clades containing marine species (e.g. Araeolaimida, Chromadorida, Desmodorida, Desmoscolecida, Enoplida, and Monhysterida) is needed for accurate coverage of known taxonomic diversity. We now add small subunit ribosomal DNA (SSU rDNA) sequences for 100 previously un-sequenced species of nematodes, including 46 marine taxa. SSU rDNA sequences for >200 taxa have been analysed based on Bayesian inference and LogDet-transformed distances. The resulting phylogenies provide support for (i) the re-classification of the Secernentea as the order Rhabditida that derived from a common ancestor of chromadorean orders Araeolaimida, Chromadorida, Desmodorida, Desmoscolecida, and Monhysterida and (ii) the position of Bunonema close to the Diplogasteroidea in the Rhabditina. Other, previously controversial relationships can now be resolved more clearly: (a) Alaimus, Campydora, and Trischistoma belong in the Enoplida, (b) Isolaimium is placed basally to a big clade containing the Axonolaimidae, Plectidae, and Rhabditida, (c) Xyzzors belongs in the Desmodoridae, (d) Comesomatidae and Cyartonema belongs in the Monhysterida, (e) Globodera belongs in the Hoplolaimidae and (f) Paratylenchus dianeae belongs in the Criconematoidea. However, the SSU gene did not provide significant support for the class Chromadoria or clear evidence for the relationship between the three classes, Enoplia, Dorylaimia, and Chromadoria. Furthermore, across the whole phylum, the phylogenetically informative characters of the SSU gene are not informative in a parsimony analysis, highlighting the short-comings of the parsimony method for large-scale phylogenetic modelling.
A catalogue is presented of the nematode slide collection of W.L. Nicholas, which is deposited in the National Research Collections Australia at CSIRO. This is the most extensive slide collection of free-living marine and estuarine nematodes from Australia to date, and consists of 553 putative species, collected across a wide range of Australia’s eastern and northern regions over the course of nearly 40 years. The collection contains mostly marine and estuarine free-living nematodes collected on coarse substrate in littoral habitats. The most abundant genera were Desmodora, Theristus, and Onyx. Most taxa were found rarely, being recorded only once, and repeated sampling at several sandy beach sites revealed only a small proportion of the fauna on more than one occasion. A significant proportion of the taxa were also found to be widespread, occurring on more than one occasion at more than one location, with Theristus sp., Onyx sp., and Viscosia sp. occurring in the greatest number of localities. The catalogue adds an additional 90 species and 160 genera to the documented fauna of Australian free-living nematodes verifiable by specimens in permanent collections. It thus provides a better framework for studying nematode biodiversity and biogeography in the region.
The present study provides an updated species list of free-living marine nematodes reported from coastal India (Coasts and Islands) based on the thorough consultation of literature published from 1956 to 2022. This exercise resulted in a total of 617 valid species belonging to 266 genera, 48 families, 21 superfamilies and 9 orders. Class Chromadorea comprises 487 species represented by 205 genera, while class Enoplea includes 130 species belonging to 61 genera. The most common family was Xyalidae, with 76 species and the least common families having a single species each were represented by Aegialoalaimidae, Rhadinematidae, Aphanolaimidae, Rhabditidae, Pandolaimidae and Rhabdodemaniidae. The checklist provides a robust framework for the distribution and biogeography of free-living marine nematodes from the Indian waters and could be used to relate with marine ecosystems of other countries.
Background
Adoncholaiminae is one of the seven subfamilies in the free-living aquatic nematode family Oncholaimidae. Nematodes in Adoncholaiminae are found from various water environment of the world. However, a checklist of all Adoncholaiminae species including full literature, especially information of experimental (not taxonomic) works, has not been updated for more than 40 years.
New information
A revised checklist of the subfamily Adoncholaiminae of the world is provided. It contains 31 valid and 13 invalid species names in four genera with synonyms, collection records, and full literature from 1860's to 2015 for each species. A literature survey of total 477 previous papers was conducted in this work, and 362 of them are newly added to checklist.
In the present article, morphological characteristics with reference to evolutionary patterns of the nematodes inhabiting soil and belonging to the Order Alaimida on the basis of their sensory structures — amphids have been discussed. The alaims are simple nematodes, having a thin slender body and unarmed stoma, more or less gradually widening esophagus single or paired female gonads, single testis, simple and short spicules, few ventromedian supplements and no caudal glands or spinneret.
Depending on the type of amphids two different routes of evolution i.e ‘Alaimus group’ and ‘Amphidelus group’ have been traced. A third line that is, ‘Cristamphidelus group’ has diverged mainly on the basis of numerous prominent longitudinal ridges. Possible evolutionary pattern of thirteen genera of alaims has also been discussed.
‘Alaimus group’ having amphidial apertures which are small, pore-like or indistinct and posteriorly placed, diverges into the genus Alaimus and Cosalaimus respectively.
In ‘Amphidelus group’ the amphidial apertures are very large, transverse slit — like or oval or even longitudinally oval, placed generally close to the head. Ten different genera coming under this group i.e Amphidelus, Caviputa, Etamphidelus, Laxamphidelus, Megamphidelus, Metamp hide lus, Paramphidelus, Postamphidelus, Scleralaimus, Scleramphidelus, have been discussed. These seem to have evolved from one single Amphidelus-like ancestor. Scleramphidelus have large amphids but instead of having a cup-shaped appearance transform to a stirrup-shaped form and also the margins are sclerotized. Probably, later three lines arose such as Paramphidelus slit-like, crescentshaped amphids, Etamphidelus, oval amphids with margins sclerotized and Metamphidelus having small, irregularly oval ones also with the same feature of the amphid margins. The genus Postamphidelus, as the name means posteriorly directed amphids, derive the character of these sensory structures from Paramphidelus. On the other hand, Etamphidelus gave rise to Megamphidelus with very large amphids and Caviputa where only the males have large, oval, kidney-shaped amphids. Lastly Caviputa gave rise to those alaims that have amphids which are oval and bilobed-Scleralaimus and Laxamphidelus with amphids that are kidney-shaped.
‘Cristamphidelus group’ has become a little more advanced in comparison to other alaims in the sense that it has evolved dorylaim characters such as presence of long slender spicules and also a small gubernaculum.
This article presents information related to the biodiversity anddistribution of nematodes belonging to the families Longidoridae andTrichodoridae in different habitats in the countries belonging to the CIS(Commonwealth of Independent States, formerly known as theUSSR).The investigations have been conducted in 13 republics of theformer USSR, and in 83 regions of the Russian Federation. The studiesrevealed 40 species of Longidoridae, including 18 species belonging tothe genus Longidorus, 3 belonging to the genus Paralongidorus, and 19belonging to Xiphinema. The list of species belonging to the familyTrichodoridae is represented by 12 species, including 9 species belongingto the genus Trichodorus, and 3 belonging to Paratrichodorus. Most ofthese nematodes are characteristic for the palearctic fauna. Thecombination of field and laboratory studies allowed us to assess theimportance of various agroecological factors (e.g. host plants,geographical distribution, location, climatic conditions, edaphiccharacteristics/soil properties, etc.) for population abundance and theoccurrence of particular species.
Redescription of Campydora demonstrans Cobb, 1920 is provided. Male is described and illustrated for the first time in the genus; it is characterized by one testis, presence of caudal alae and gubernaculum. Observations on this species are given on the basis of material collected in Poland.
Two new species of the family Enchelidiidae Filipjev, 1918 were collected from marine sediments near Maemul Island in South Korea: a new species of Abelbolla Huang & Zhang, 2004 and a new species of Ledovitia Filipjev, 1927. Abelbolla maemulensis sp. nov. is characterized by its small size (1,493 × 38 µm, body length × maximum body diameter); the presence of a circular amphid; the gubernacular apophysis with swollen distal tip; and the complex structure of the gubernaculum. It is close to Abelbolla huanghaiensis Huang & Zhang, 2004, but differs by the structure of gubernacular apophysis and body length (1,493 vs 2,303 µm). Ledovitia brevis sp. nov. can be separated from its congeners by its small size of body, the length of gubernacular apophysis, and the length of the spicules. It is close to Ledovitia pharetrata Wieser, 1953a, but differs by the length of the body (1,699 vs 2,640 µm) and the spicules (40 vs 100 µm).
The αβ tubulin heterodimer is the structural subunit of
microtubules, which are cytoskeletal elements that are essential for
intracellular transport and cell division in all eukaryotes. Each
tubulin monomer binds a guanine nucleotide, which is non-exchangeable
when it is bound in the α subunit, or N site, and exchangeable
when bound in the β subunit, or E site. The α- and
β-tubulins share 40% amino-acid sequence identity, both exist in
several isotype forms, and both undergo a variety of post-translational
modifications. Limited sequence homology has been found with the
proteins FtsZ and Misato, which are involved in cell division in
bacteria and Drosophila, respectively. Here we present an atomic model
of the αβ tubulin dimer fitted to a 3.7-Å density map
obtained by electron crystallography of zinc-induced tubulin sheets. The
structures of α- and β-tubulin are basically identical: each
monomer is formed by a core of two β-sheets surrounded by
α-helices. The monomer structure is very compact, but can be
divided into three functional domains: the amino-terminal domain
containing the nucleotide-binding region, an intermediate domain
containing the Taxol-binding site, and the carboxy-terminal domain,
which probably constitutes the binding surface for motor proteins.
Syringolaimus is the most abundant and diverse genus of the family Ironidae (Nematoda) found in the Campos Basin, off Rio de Janeiro, Brazil. In this article, four new species of this genus are described. S. annae sp. n. is characterized by a conical-cylindrical tail without a spinneret. S. magdae sp. n. possesses a pair of ejaculatory glands located in the precloacal region, and a spinneret, which comprises 21.2% of the total tail length. In S. smolae sp. n. the spicule is setiform. S. taniae sp. n. has the anterior region of the tail rounded, with its terminal portion cylindrical-filiform.
Soil material collected at Camberene, Senegal yielded two of the three known species of the rare genus Bathyodontus: B. cylindricus Fielding, 1950 and B. indicus (Khan, 1972). A male B. cylindricus is described, having a gubernaculum with lateral guiding pieces. The four females found of B. indicus lack a fully developed anterior reproductive branch. Comparison with collection specimens of B. mirus (Andrássy, 1956) allows a new generic character to be identified: presence of a mononchid-like excretory system. Stoma, pharynx and vagina structure are examined in detail; S.E.M. photographs are included and relationships within the genus and with other genera are discussed.
Observations are reported on the fine structure of the cuticle of Paratrichodorus minor, P. nanus, P. rhodesiensis and Trichodorus eburneus. The ultrastructure of the cuticle was similar in Paratrichodorus and Trichodorus species; six layers were observed, the innermost layer consisting of three multilaminate units. The anatomy of the cuticle in Trichodorids was quite different from previous observations in other orders of Adenophorea. Two sets of intracuticular canals were observed as well as a peculiar type of junction between cuticle and somatic muscles. These results confirm previous studies on the anatomy of trichodorids and support the withdrawal of the family Trichodoridae from Dorylaimida. Siddiqi's (1983) classification of the trichodorids under the order Triplonchida should be considered with attention.
Nematodes are increasingly being used in environmental studies. One of the potential parameters to measure the impact of disturbances
and to monitor changes in structure and functioning of the below-ground ecosystem is the nematode Maturity Index; an index
based on the proportion of colonizers (r-strategists s.l.) and persisters (K-strategists s.l.) in samples. In this paper the
original allocation of nematode taxa on the colonizer-persister scale, and the tolerance and sensitivity of colonizers and
persisters are discussed from an evolutionary viewpoint. The phenomenon that neither relative egg size nor body length is
an unequivocal character to scale nematodes suggests that the main selection for life history traits occurred independently
in the major evolutionary branches.
Cardiac fibrillation (spontaneous, asynchronous contractions of cardiac muscle fibres) is the leading cause of death in the industrialized world, yet it is not clear how it occurs. It has been debated whether or not fibrillation is a random phenomenon. There is some determinism during fibrillation, perhaps resulting from rotating waves of electrical activity. Here we present a new algorithm that markedly reduces the amount of data required to depict the complex spatiotemporal patterns of fibrillation. We use a potentiometric dye and video imaging to record the dynamics of transmembrane potentials at many sites during fibrillation. Transmembrane signals at each site exhibit a strong periodic component centred near 8 Hz. This periodicity is seen as an attractor in two-dimensional-phase space and each site can be represented by its phase around the attractor. Spatial phase maps at each instant reveal the 'sources' of fibrillation in the form of topological defects, or phase singularities, at a few sites. Using our method of identifying phase singularities, we can elucidate the mechanisms for the formation and termination of these singularities, and represent an episode of fibrillation by locating singularities. Our results indicate an unprecedented amount of temporal and spatial organization during cardiac fibrillation.
Nematodes are important: parasitic nematodes threaten the health of plants, animals and humans on a global scale; interstitial nematodes pervade sediment and soil ecosystems in overwhelming numbers; and Caenorhabditis elegans is a favourite experimental model system. A lack of clearly homologous characters and the absence of an informative fossil record have prevented us from deriving a consistent evolutionary framework for the phylum. Here we present a phylogenetic analysis, using 53 small subunit ribosomal DNA sequences from a wide range of nematodes. With this analysis, we can compare animal-parasitic, plant-parasitic and free-living taxa using a common measurement. Our results indicate that convergent morphological evolution may be extensive and that present higher-level classification of the Nematoda will need revision. We identify five major clades within the phylum, all of which include parasitic species. We suggest that animal parasitism arose independently at least four times, and plant parasitism three times. We clarify the relationship of C. elegans to major parasitic groups; this will allow more effective exploitation of our genetic and biological knowledge of this model species.
L'ultrastructure de la cuticule est décrite chez #Trichodorus primitivus$, #T. similis$, #T. hooperi$, #T. viruliferus$, #Paratrichodorus pachydermus$ et #P. nanus$. La cuticule présente une structure largement semblable à celle décrite récemment chez quatre espèces d'Afrique occidentale mais, selon les espèces, une différence dans l'épaisseur des couches 4 et 6 est observée. La structure de la région des spicules est nettement différente chez #P. pachydermus$ et chez #P. primitivus$, confirmant ainsi la valeur diagnostique du caractère présence ou absence d'ailes caudales pour différencier les genres didelphes. (Résumé d'auteur)
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