Morphology of the synlophe and genital cone of Parostertagia heterospiculum (Trichostrongylidae) with comments on the subfamilial placement of the genus

The genus Parostertagia is referred to the Cooperiinae with reference to a suite of shared characters of the synlophe and genital cone postulated as derived homologies for the subfamily. The genus has traditionally been relegated to the Graphidiinae or Ostertagiinae, but unique structural attributes of the synlophe and genital cone indicate close affinities with the Cooperiinae. The synlophe of Parostertagia heterospiculum is characterised by a relatively low number of ridges in the cervical zone (16–20 at the excretory pore), a frontal axis of orientation, sequential increase in ridges posteriorly, lateral addition of ridges, minuscule lateral-most ridges and hypertrophy of specific ridges in the lateral field of females. The genital cone with a tripartite “0” papilla, a pair of dorsal cloacal papillae (a character potentially homologous with lateral protuberances in some cooperiines, but reported for the first time in this study), and paired “7” papillae supporting a small accessory bursal membrane appear unique among the trichostrongylids. Additionally, Parostertagia was found to possess minuscule cervical papillae and a rudimentary (or reduced?) cephalic and cervical expansion. Inclusion of Parostertagia within the Cooperiinae is based on shared characters with this subfamily (postulated synapomorphies) rather than differences that may be utilized to separate this genus from members of other subfamilies of the Trichostrongylidae.     

An exploration of diversity among the Ostertagiinae (Nematoda: Trichostrongyloidea) in ungulates from Sub-Saharan Africa with a proposal for a new genus

Abomasal nematodes (Ostertagiine: Trichostrongyloidea) representing a previously unrecognized genus and species are reported in African buffalo (Syncerus caffer caffer) from Kenya, Uganda, and South Africa. Africanastrongylus buceros gen. nov. et sp. nov. is characterized by a symmetrical tapering synlophe in the cervical region and a maximum of 60 ridges in males and females. Bursal structure is 2-2-1, with subequal Rays 4/5, massive Rays 8, and Rays 9/10, and a massive dorsal lobe that is reduced in length, laterally and dorsally inflated, and positioned ventral to externodorsal rays. Spicules are tripartite, and the gubernaculum is broadly alate in the anterior. A proconus is present. Among ostertagiines with a 2-2-1 bursa (Cervicaprastrongylus, Hyostrongylus, Mazamastrongylus, Sarwaria, Spiculopteragia, and Teladorsagia) specimens of Africanastrongylus are differentiated from respective genera based on the structure of the cervical synlophe, patterns of dorsal, externodorsal, lateral, and ventral rays, and configuration of the genital cone, gubernaculum, and spicules. Among 13 genera of the Ostertagiinae in the global fauna, 3 are entirely limited in distribution to Africa, including Africanastrongylus, Longistrongylus, and Pseudomarshallagia. Another 5 genera including Cervicaprastrongylus, Hyostrongylus, Marshallagia, Ostertagia, and Teladorsagia are represented as mosaics, with diversity centered in Eurasia or the Holarctic. Genera not represented in the African fauna include Camelostrongylus among Caprinae and some Antelopinae from Eurasia, Mazamastrongylus and Spiculopteragia in Cervidae from the Holarctic and Eurasia, respectively, Orloffia in Cervidae and Bovidae from the Holarctic, and Sarwaria among Tragulidae and Bovinae in southern Asia. The diverse nature of the ostertagiine fauna, with a disproportionate number of endemic genera relative to other regions of the northern hemisphere, may reflect the timing of episodic expansion events for artiodactyls into Africa from Eurasia during the Tertiary and Quaternary.     

Polymorphism in the Ostertagiinae Lopez-Neyra, 1947 and comments on the systematics of these nematodes

A list of 19 polymorphic species of the generaOstertagia, Orloffia, Teladorsagia, Marshallagia andSpiculopteragia is presented. The morphs of the polymorphic species in question have identical females and show similarities in some male features, e.g. the structure of the oesophagus, the synlophe and the shape of the rays of the copulatory bursa. However, they differ by characters which are recognised as generic features, e.g. the structure of spicules, gulbernaculum and genital cone. The presence or lack of minor morphs is a generic feature in the Ostertagiinae. Each polymorphic genus has a defined type of minor morph.     

Synlophe patterns of the Haemonchinae of ruminants (Nematoda: Trichostrongyloidea)

The pattern of longitudinal ridges (synlophe) on the external cuticular surface of trichostrongylid nematodes has been shown to be of value for distinguishing species and determining relationships among higher taxa. In the process of studying Mecistocirrus digitatus, the large stomach worm of bovids of Asia that has been imported and established in the Americas, we observed remarkably similar synlophe patterns to those described for 3 species of Haemonchus and to those we examined in a species of Ashworthius. In all 3 genera, the synlophe is absent from the posterior part of the body. Only in Haemonchus does the synlophe extend beyond midbody. In both M. digitatus and Ashworthius sidemi, the synlophe extends posteriorly only about 1/4 of body length. In all 3 genera, the synlophe consists of about 30 ridges in the region of the esophagus with variation among species in specific areas, including additional pairs of subventral and subdorsal ridges and different lengths of sublateral ridges. This information is useful for identifying species and determining relationships among these large stomach worm parasites of cattle, sheep, goats, and farmed and wild cervids.     

Onchocerca eberhardi n. sp. (Nematoda: Filarioidea) from sika deer in Japan; relationships between species parasitic in cervids and bovids in the Holarctic region

Onchocerca eberhardi n. sp. from the sika deer, Cervus nippon, in Japan is described. Adult worms lived in the carpal ligament; infection reached high levels (up to 25 female and 16 male worms in a single carpal limb). Skin dwelling microfilariae were mainly found in the ears. Prevalence of infection was 81% at the type locality, Mt. Sobo, in Kyushu. The new material was compared to the 31 species of Onchocerca presently known. Onchocerca eberhardi n. sp. females were characterized by a long slender anterior end and a thin esophagus < or =1 mm long with no or only a slight glandular region. The vulva was located near the level of the mid-esophagus and the cuticle had transverse external ridges and internal striae (two striae between adjoining ridges). The most similar species were O. stilesi (re-examined), O. lienalis, and to a lesser extent O. gutturosa, all from bovids (cattle). Two main lineages of Onchocerca are recognized in cervids with either primitive or with derived characteristics (as exemplified by the new species). The species in both lineages are not restricted to cervids but are also found in bovids in the Holarctic region, suggesting that the species diversified in the two host groups simultaneously, when these host groups lived in the some geographic area.     

Abomasal parasites in wild sympatric cervids, red deer, Cervus elaphus and fallow deer, Dama dama, from three localities across central and western Spain: relationship to host density and park management

A survey of abomasal parasites in cervids from Central Spain was conducted at 3 sites, Quintos de Mora (Toledo), Malué?ez de Arriba (Cáceres), and La Herguijuela (Cáceres). Commonly occurring helminths belonged to 3 polymorphic species of the Ostertagiinae: Spiculopteragia asymmetricalS. quadrispiculata, Ostertagia leptospicularis/O. kolchida, and O. drozdzi/O. ryjikovi. Trichostrongylus axei was found in very few cases. Ostertagia drozdzi/O. ryjikovi and the minor male morphotype, S. quadrispiculata, are reported for the first time in red deer from Spain. The 3 ostertagiine species are also reported for the first time in fallow deer from Spain. These 3 species of Ostertagiinae are primarily parasites of cervids, and nematode species characteristic of domestic ruminants were not present. Prevalence of infection by gastrointestinal parasites in cervids was high, ranging from 97.5 to 100%, across the 3 areas sampled. Mean intensity of infection and abundance showed a positive relationship to the population density of red deer. Helminth burdens were higher in fallow deer than in the sympatric red deer and may reflect the gregarious social structure and different foraging patterns of fallow deer.     

Pseudostertagia bullosa (Nematoda: Trichostrongyloidea) in artiodactyl hosts from North America: redescription and comments on systematics

A relationship for Pseudostertagia bullosa within the trichostrongyloids has been enigmatic or unresolved. Studies of the synlophe in males and females of P. bullosa revealed a tapering system anterior to the deirids and a pattern of parallel ridges extending to near the caudal extremity in both lateral and median fields. Structurally, the synlophe differs considerably from that seen among the Cooperiinae and exhibits homoplasy with respect to ridge systems among some Ostertagiinae. Other structural characters due to symplesiomorphy, homoplasy or because they represent autapomorphies do not serve to reveal the putative relationships for P. bullosa with other trichostrongyloids. Although somewhat equivocal, the 2-2-1 pattern of the bursa and position of rays 2 and 3 suggest an association with the Cooperinae, as postulated by Durette-Desset and others. Pseudostertagia bullosa appears to be a species that has survived in the pronghorn, Antilocapra americana, a relictual pecoran artiodactyl that occurs in xeric regions of western North America; pronghorn are the sole remnant of the late Tertiary radiation for Antilocapridae across North America. Pseudostertagia bullosa may occur in mixed infections with a number of ostertagiines in the abomasa of mule deer (Odocoileus hemionus) and domestic sheep (Ovis aries) in regions of sympatry for pronghorn and these artiodactyl hosts.     

A comparison of cuticular ridge patterns and other morphological characters of Mazamastrongylus odocoilei and Mazamastrongylus pursglovei (Nematoda: Trichostrongyloidea) from white-tailed deer Odocoileus virginianus

The synlophes of Mazamastrongylus odocoilei (Dikmans, 1931) and M. pursglovei (Davidson & Prestwood, 1979) from Odocoileus virginianus were described to provide characteristics for their differentiation from related species in domestic and wild ruminants. The synlophes of M. odocoilei and M. pursglovei were identical. Both had single continuous dorsal, ventral and lateral ridges. Other ridges in the lateral fields in the anterior third of the body angled posteriorly ending adjacent to the single lateral ridge. The ridge system in Mazamastrongylus spp. appears to be unique in lacking the continuous subventral and subdorsal ridges present in all previously studied medium stomach worms from domestic ruminants. Only two characters, spicule length and structure, are known to be useful for differentiating males of M. odocoilei and M. pursglovei, and females cannot be differentiated. Evidence that M. odocoilei and M. pursglovei are separate species include: (1) the two species are frequently found together in the same individual host, but each is also found in the absence of the other; (2) no overlap in spicule size or intergrades in shape have been found although numerous populations have been sampled; and (3) the two species have overlapping, but different, geographical ranges in eastern North America. No differences in the synlophe were found between these two species. The importance of this finding is that the absence of differences in the synlophe is not an unequivocal indicator that nematodes are conspecific.     

Redescriptions of Haemonchus mitchelli and Haemonchus okapiae (Nematoda: Trichostrongyloidea) and description of a unique synlophe for the haemonchinae

In the course of a revision of Haemonchus Cobb, 1898 (Nematoda), commonly referred to as large stomach worms, significant new morphological information was discovered that allows the recognition of 2 species believed for more than 50 yr to be synonymous. Both species, Haemonchus mitchelli Le Roux, 1929, from the eland Taurotragus oryx and other African ruminants and H. okapiae van den Berghe, 1937, from the okapi Okapia johnstoni, have a synlophe of 42 ridges, but the synlophe of H. mitchelli is longer than that of H. okapiae. The distal tip of the left spicule of H. mitchelli bears a barb that is about twice as long as the short barb and half as long as the long barb on the right spicule. In contrast, the barb on the left spicule of H. okapiae is similar in size to the short barb and about 25% as long as the long barb of the right spicule. The dorsal ray of H. mitchelli is bifurcated distally for 25-39% (32%) of its length and its stem is expanded proximally, but the dorsal ray of H. okapiae is bifurcated 37-50% (42%) and its stem is of uniform thickness.     

Two new species ofOdilia (Nematoda: Heligmonellidae) from Australian rodents,with comments onO. bainae Beveridge & Durette-Desset, 1992

Odilia tasmaniensis n. sp.,O. praeputialis n. sp. andO. bainae Beveridge & Durette-Desset, 1992 from the small intestine of Australian murids are described and illustrated. The two new species are distinguished from the other eight species in the genus, namelyO. mackerrasae (Mawson, 1961),O. brachybursa (Mawson, 1961),O. emanuelae (Mawson, 1961),O. melomyos (Mawson, 1961),O. polyrhabdote (Mawson, 1961),O. uromyos (Mawson, 1961),O. mawsonae (Durette-Desset, 1969) andO. bainae Beveridge & Durette-Desset, 1992.O. tasmaniensis n. sp. fromRattus lutreolus is characterised by the longitudinal cuticular ridges being continuous, the presence of 18 ridges in cross-section in the middle region of the body, the joined distal ends of the spicules forming a curved bluntly rounded tip, a short genital cone with a single ventral papilla and a pair of laterally curving dorsal raylets and the posterior end of the female tapering sharply.O. praeputialis n. sp. fromZyzomys woodwardi is characterised by continuous longitudinal cuticular ridges, the presence of 22–35 ridges in cross-section in the middle of the body, the sharply pointed joined distal tips of the spicules, a complex genital cone with a flat membraneous proconus, a ventral papilla projecting from an extension of the body wall, a pair of short straight dorsal raylets and the presence of a praepuce on the posterior end of the female.O. bainae is characterised by the longitudinal cuticular ridges being continuous, the presence of 17–22 ridges in cross-section in the middle region of the body, the joined distal ends of the spicules surrounded in an oval transparent cap, a long genital cone with a single ventral papilla and a pair of laterally curving dorsal raylets, and the absence of a praepuce on the posterior end of the female.Rattus lutreolus and the pseudomyine rodentsPseudomys higginsi andMastacomys fuscus are new host records for this species.     

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.     

Two new species of the Trichostrongylidae (Nematoda: Trichostrongyloidea), parasitic in Gorilla gorilla beringei in Uganda

Two new species of trichostrongylid parasites of a mountain gorilla in Uganda are described. Paralibyostrongylus kalinae n. sp. is characterised by a short dorsal ray, a dorsal oesophageal tooth and two median longitudinal ridges in the vulvar region. The species is more closely related to leporid parasites than to P. hebrenicutus, another species known from the gorilla. A “transfuge” phenomenon, i.e. transfer to a new host without speciation is suggested. Hyostrongylus kigeziensis n. sp. has several specific morphological characteristics and in particular a synlophe with many strong ridges, rays 4 and 5 divergent at the apex, and the dorsal ray with two branches. Hyostrongylus appears to be an Ethiopian genus which has radiated in many archaic mammals. With the technical collaboration of Mrs. N. Caillaud and Mr R. Tchesprakoff. With the technical collaboration of Mrs. N. Caillaud and Mr R. Tchesprakoff.     

A new species of Oswaldocruzia (Molineidae: Nematoda) in Chaunus marinus (Amphibian: Bufonidae) (Linneaus, 1758) from Brazil

Oswaldocruzia belenensis n. sp. (Strongylida: Molineidae) from the small intestine of Chaunus marinus (L.) from Belém, Pará State, Brazil is described and illustrated by light and scanning electron microscopy. Oswaldocruzia belenensis n. sp. is a neotropical species of this genus, harboring caudal bursa Type II, spicules divided in 3 branches, i.e., a blade, shoe, and fork. The blade is divided in 4 points, of which at least 2 are bifurcated. Cervical alae are absent; there is a simple cephalic vesicle and synlophe with low ridges perpendicular to the body without chitinous supports. The most closely related species are O. bonsi and O. lescurei. Oswaldocruzia belenensis n. sp. differs from O. lescurei and O. bonsi by the number and location of cephalic papillae, rays 2-3 and 5-6 running parallel and slightly separated, ray 6 not overlapping ray 8, and body structure morphometry. Oswaldocruzia belenensis n sp. also differs from O. lescurei by the discontinuity of the longitudinal ridges, the number of subdivisions of the blade, and the absence of extra processes at the bifurcation level of the fork of the spicules. The new species differs from O. bonsi by male and female body dimensions, the symmetry of the caudal bursa, dimension and subdivisions of the spicules without extra processes of the fork, 2 extra processes at the distal division of the blade, and location of ray 7 at the anterior margin of the cloacal aperture. Oswaldocruzia belenensis n. sp. represents the 82nd species assigned to the genus.     

Description of a new species of Heligmonina Baylis, 1928 (Nematoda: Heligmonellidae) a parasite of Mastomys natalensis (Rodentia: Muridae) from Swaziland and new data on the synlophe of Heligmonina chabaudi (Desset, 1966)

A new species of heligmosomoid nematode belonging to the subfamily Nippostrongylinae Durette-Desset, 1970 is described: Heligmonina wakelini n. sp., a parasite from the small intestine of the commensal rodent Mastomys natalensis (Smith, 1834) from Swaziland. It differs from the most closely related species H. boomkeri Durette-Desset & Digiani, 2005 by the number of the cuticular ridges in the female synlophe (10 vs 12), the width of the left ala, larger than the body diameter in the male, and the inclination of the axis of orientation of the ridges in both sexes (53 degrees vs 70 degrees). New morphological data (head and synlophe) on Heligmonina chabaudi (Desset, 1964), also a parasite of Mastomys natalensis in the Republic of Congo, are provided in order to compare with the new species.     

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 degrees 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 degrees from the sagittal axis, and a caudal bursa with the right lobe enlarged vertically.     

DNA barcoding and morphology reveal three cryptic species of Anania (Lepidoptera: Crambidae: Pyraustinae) in North America,all distinct from their European counterpart

Anania coronata (Hufnagel), a Holarctic species of pyraustine crambid moth, has long been treated as having two geographically separated subspecies – the nominotypical Anania coronata in the Palaearctic Region and Anania coronata tertialis (Guenée) in the Nearctic Region. Maximum likelihood and Bayesian inference analysis of mitochondrial DNA barcodes both recover four well‐supported, reciprocally monophyletic groups within Anania coronata. Qualitative and quantitative analyses of genital structures reveal diagnostic differences that correspond to the four barcode lineages. On the basis of both molecular and morphological evidence, we conclude that Anania coronata is actually a complex of four species. Anania coronata (Hufnagel) is restricted to Europe, whereas three species occur in North America: Anania tertialis (Guenée), Anania plectilis (Grote & Robinson) and Anania tennesseensis sp.n. Yang.     

CONTRIBUTION OF POLLEN MORPHOLOGY TO SYSTEMATICS OF COLLOMIA (POLEMONIACEAE)

Pollen morphology of 14 species of Collomia (Polemoniaceae) was examined by light microscopy, and by both scanning and transmission electron microscopy. Four distinct pollen types were observed which are based principally upon 1) shape, number and distribution of apertures, and 2) surface sculpturing: Type 1—zonocolporate with striate ridges; Type 2—zonocolporate with striato-reticulate ridges; Type 3—pantoporate with radiate ridges; Type 4—pantoporate with irregularly reticulate ridges. Evaluation of pollen morphology reveals considerable discrepancy with respect to presently accepted sectional classification. Collomia grandiflora of sect. Collomia has a pollen type similar to that of members of sect. Collomiastrum and is now interpreted as representing an independent evolutionary line derived from the latter section. Collomia diversifolia of sect. Courtoisia has a pollen morphology similar to that of sect. Collomia. whereas C. heterophylla of the same section possesses pollen unique within the genus. This last pollen type shows close similarity to the pollen of members of Polemonium, Gilia, Leptodactylon, and Ipomopsis. Pollen of C. tinctoria and C. tracyi of sect. Collomia are anomalous within Polemoniaceae. No significant difference in exine stratification was discernible among the four pollen types.