Invasion, establishment, and range expansion of two parasitic nematodes in the Canadian Arctic.

Climate warming is occurring at an unprecedented rate in the Arctic and is having profound effects on host-parasite interactions, including range expansion. Recently, two species of protostrongylid nematodes have emerged for the first time in muskoxen and caribou on Victoria Island in the western Canadian Arctic Archipelago. Umingmakstrongylus pallikuukensis, the muskox lungworm, was detected for the first time in 2008 in muskoxen at a community hunt on the southwest corner of the island and by 2012, it was found several hundred kilometers east in commercially harvested muskoxen near the town of Ikaluktutiak. In 2010, Varestrongylus sp., a recently discovered lungworm of caribou and muskoxen was found in muskoxen near Ikaluktutiak and has been found annually in this area since then. Whereas invasion of the island by U. pallikuukensis appears to have been mediated by stochastic movement of muskoxen from the mainland to the southwest corner of the island, Varestrongylus has likely been introduced at several times and locations by the seasonal migration of caribou between the island and the mainland. A newly permissive climate, now suitable for completion of the parasite life cycles in a single summer, likely facilitated the initial establishment and now drives range expansion for both parasites.

Hamulonema gen. nov. for Teladorsagia hamata and Ostertagia kenyensis in the ostertagiine fauna (Nematoda: Trichostrongyloidea) from African ungulates.

Hamulonema gen. nov. is proposed for Teladorsagia hamata and Ostertagia kenyensis in the ostertagiine nematode fauna found in artiodactyl hosts from Africa. Monomorphic species representing this genus are characterized by a bilaterally symmetrical and parallel synlophe in males and females, a 2-2-1 bursal formula, an accessory bursal membrane that is strongly cuticularized and reduced, a strongly reduced dorsal lobe and ray, and robust spicules with a simple, weakly pointed, ventral process, and curved, hooklike dorsal process. Species referred to Hamulonema nov. gen. are immediately distinguished from those of Camelostrongylus, Longistrongylus, Marshallagia, Orloffia, Ostertagia, and Pseudomarshallagia in which the bursal formula is 2-1-2 in males. Hamulonema nov. gen. is distinguished from those genera having a 2-2-1 bursa, including Africanastrongylus, Cervicaprastrongylus, Hyostrongylus, Mazamastrongylus, Sarwaria, Spiculopteragia, and Teladorsagia by the structure of the synlophe, bursa, genital cone, "0" and "7" papillae, dorsal lobe, and spicules in specific instances. In the global fauna, 4 of 14 ostertagiine genera are endemic to Africa. African genera may represent morphologically divergent and discrete or historically isolated lineages reflecting a pattern of geographic and host colonization as a driver for diversification since the Miocene.

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.

Emended description and redetermination of Sarwaria caballeroi n. comb. (nematoda: Ostertagiinae) with details of the synlophe and esophageal characters.

Among genera of the ostertagiine nematodes, structural attributes of Spiculopteragia caballeroi are consistent with criteria that diagnose the genus Sarwaria. Specifically, the following characters are compatible with referral to this genus: (1) species characterized by monomorphic males; (2) tapering lateral synlophe in the cervical zone; (3) minuscule, thornlike cervical papillae; (4) length of Ray "4" < Ray "5"; (5) relatively thick or robust Ray "4"; (6) a substantially reduced dorsal lobe and dorsal ray that are disposed or curved ventrally relative to Rays "8"; and (7) a broad, laterally inflated dorsal lobe. Consequently, we propose Sarwaria caballeroi n. comb., and we provide further validation for the genus Sarwaria. Additionally, we propose that the ostertagiines are represented by 12 valid genera, including those characterized by a bursal formula of 2-2-1 (Cervicaprastrongylus, Hyostrongylus, Mazamastrongylus, Spiculopteragia, and Teladorsagia in addition to Sarwaria), and those in which the lateral rays describe a 2-1-2 pattern (Camelostrongylus, Longistrongylus, Marshallagia, Orloffia, Ostertagia, and Pseudomarshallagia).

Parelaphostrongylus odocoilei in Columbian black-tailed deer from Oregon.

Documenting the occurrence of Parelaphostrongylus odocoilei has historically relied on the morphological examination of adult worms collected from the skeletal muscle of definitive hosts, including deer. Recent advances in the knowledge of protostrongylid genetic sequences now permit larvae to be identified. Dorsal-spined larvae (DSLs) collected in 2003-2004 from the lung and feces of six Columbian black-tailed deer (Odocoileus hemionus columbianus) from Oregon were characterized genetically. The sequences from unknown DSLs were compared to those from morphologically validated adults and larvae of P. odocoilei at both the second internal transcribed spacer (ITS-2) of ribosomal DNA and the mitochondrial cytochrome oxidase II gene. We provide the first unequivocal identification of P. odocoilei in Columbian black-tailed deer from Oregon. The broader geographic distribution, prevalence, and pathology of P. odocoilei are not known in populations of Oregon deer.

Ashworthius Patriciapilittae n. sp. (Trichostrongyloidea: Haemonchinae), an abomasal nematode in Odocoileus virginianus from Costa Rica, and a new record for species of the genus in the Western Hemisphere.

A species of Ashworthius is reported for the first time in the Western Hemisphere, and A. patriciapilittae n. sp. is described on the basis of specimens in white-tailed deer Odocoileus virginianus from Costa Rica. Among 8 known species, A. patriciapilittae is morphologically similar to A. tuyenquangi in red muntjac Muntjacas muntjak from northern Vietnam. The synlophe in A. patriciapilittae is composed of 26 ridges in the cervical zone and is continuous to the caudal extremity in males and females. Males are characterized by a complex dorsal ray and narrow trifurcate spicules (351-356 microm long) lacking an "eyelet." with dissimilar ventral and dorsal processes; the gubernaculum is 45-48% of the spicule length. Females have a prominent linguiform flap at the vulva and large eggs (108-142 microm long). The presence of A. patriciapilittae in Costa Rica is examined in the context of competing hypotheses for cospeciation or contemporary host-switching in cervids: either A. patriciapilittae is a component of an endemic Central and South American fauna that has diversified through coevolution of Ashworthius and cervid hosts or it has been introduced. Among haemonchines in the Western Hemisphere, specimens of A. patriciapilittae may be confused with 3 species of Haemonchus, including H. contortus, H. placei, and H. similis, that occur in both domestic and wild ruminants. Discovery of A. patriciapilittae emphasizes the continued need for survey and inventory to define the structure and distribution of parasite faunas in wild and domestic ruminants from the Nearctic and Neotropical regions.

Discovery and description of the "davtiani" morphotype for Teladorsagia boreoarcticus (Trichostrongyloidea: Ostertagiinae) abomasal parasites in muskoxen, Ovibos moschatus, and caribou, Rangifer tarandus, from the North American Arctic: implications for parasite faunal diversity.

Collections to explore helminth diversity among free-ranging ungulates in the North American Arctic revealed the occurrence of a third male, or " davtiani ," morphotype for Teladorsagia boreoarcticus . Designated as T. boreoarcticus forma (f.) minor B, the males occurred with T. boreoarcticus f. major and T. borearcticus f. minor A in endemic populations of muskoxen (Ovibos moschatus wardi) and barrenground caribou (Rangifer tarandus groenlandicus) on Victoria Island, Nunavut, Canada, and in muskoxen and Peary caribou ( Rangifer tarandus pearyi ) on Banks Island, Northwest Territories, Canada. These specimens differ from conspecific morphotypes in the structure of the genital cone and Sjöberg's organ. Relative to T. boreoarcticus f. minor A, specimens of T. boreoarcticus f. minor B are consistently smaller, and mean dimensions for the bursa and spicules do not overlap. The robust spicules are similar in form, particularly in the relative length of the dorsal and ventral processes, but mean total length is substantially less in specimens of T. boreoarcticus f. minor B. Differences that distinguish the minor morphotypes of T. boreoarcticus parallel those demonstrated for the T. trifurcata and T. davtiani morphotypes in association with T. circumcincta sensu stricto. New host and geographic records include the 3 morphotypes of T. boreoarcticus in muskoxen and Peary caribou from Banks Island and in barrenground caribou from Victoria Island. Recognition of the ubiquitous nature of cryptic species emphasizes the need to effectively develop and use our collections-based resources and museum archives to build a robust understanding of the biosphere. Field inventory should include provisions for integrative approaches that preserve specimens suitable for comparative morphology, multi-faceted molecular investigations, and population genetics.

Discovery and description of a new trichostrongyloid species (Nematoda: Ostertagiinae), abomasal parasites in mountain goat, Oreamnos americanus, from the Western Cordillera of North America.

Marshallagia lichtenfelsi sp. n. is a dimorphic ostertagiine nematode occurring in the abomasum of mountain goats, Oreamnos americanus, from the Western Cordillera of North America. Major and minor morphotype males and females are characterized and distinguished relative to the morphologically similar Marshallagia marshalli / Marshallagia occidentalis from North America and Marshallagia dentispicularis, along with other congeners, from the Palearctic region. The configuration of the convoluted and irregular synlophe in the cervical region of males and females of M. lichtenfelsi is apparently unique, contrasting with a continuous and parallel system of ridges among those species of Marshallagia, including M. marshalli/M. occidentalis, which have been evaluated. Specimens of M. lichtenfelsi are further defined by the rectangular form of the accessory bursal membrane (width > length) in the major morphotype and by the trapezoidal Sjöberg's organ in the minor morphotype, in addition to specific attributes of the spicules and spicule tips. We regard 12 species, including the proposed new taxon, to be valid. Primary diagnostic characters are reviewed for Marshallagia and a framework is presented for standardization of future descriptions incorporating the synlophe in males and females and the structure of the spicules and genital cone in major and minor morphotype males. The center of diversity for species of Marshallagia is the mountain-steppe region of central Eurasia where 11 species (including the Holarctic M. marshalli) are recognized in association with Caprini, Rupicaprini, and Antelopinae; only 2 species occur in the Nearctic. In this assemblage, M. lichtenfelsi is endemic to North America and limited in host distribution to mountain goats. An intricate history for refugial isolation and population fragmentation demonstrated for mountain goats and wild sheep indicates the potential for considerable cryptic diversity for Marshallagia and other nematodes. Shifting patterns of contact and sympatry among assemblages of ungulates during the Pleistocene are consistent with geographic and host colonization as a process involved in diversification of these parasites.

A new species of trichostrongyloid in African buffalo (Syncerus caffer) (Artiodactyla: Bovinae) from Uganda.

Africanastrongylus giganticus n. sp. is described based on large ostertagiine nematodes occurring in the abomasum of African buffalo, Syncerus caffer, from Uganda; this represents the second species recognized in the genus. Specimens of A. giganticus are characterized by large size (15-19 mm in total length), a strongly tapering synlophe in the cervical region, and a great number of ridges at all levels of the body (maximum 72 attained in the third quarter); numbers of ridges exceed that reported among any known genera and species of the Ostertagiinae. We refer A. giganticus to this genus based on a strongly tapering lateral synlophe, relatively large numbers of ridges at all levels of the body, miniscule cervical papillae, poorly demarcated divisions of the ovejector, absence of vulval cuticular inflations, and the presence of slightly protruding lips at the vulva. It is distinguished from its congener, Africanastrongylus buceros, in total length, maximum number of ridges (68-72 vs. 53, respectively), structure and disposition of the synlophe, presence of strongly spiraled ovarian tracks, and eggs that are distributed in 3 or more rows in the uterus. A superficial resemblance to Longistrongylus meyeri, the only other large ostertagiine in the African fauna, is evident; these species, however, are distinct based on the synlophe and other characters. Recognition of a second species of Africanastrongylus represented by nematodes of large size suggests that prior reports of L. meyeri in Syncerus caffer may be attributable to A. giganticus.

Synlophe structure in Pseudomarshallagia elongata (Nematoda: Trichostrongyloidea), abomasal parasites among Ethiopian ungulates, with consideration of other morphological attributes and differentiation within the Ostertagiinae.

The independence of Pseudomarshallagia and its placement among the medium stomach worms of ungulates, Ostertagiinae, is confirmed based on comparative morphological studies of the synlophe and genital attributes among male and female specimens. An emended description of Pseudomarshallagia elongata is presented based on a series of specimens in sheep from northern Ethiopia. Pseudomarshallagia elongata is retained among the 15 genera of the Ostertagiinae based on presence of a prominent esophageal-intestinal valve, paired "0" papillae, a modified accessory bursal membrane containing the paired "7" papillae, and configuration of the copulatory bursa. The structure of the synlophe in males and females is also typical and within the range of variation demonstrated for Type II and Type A cervical patterns among other ostertagiines. We emphasize the importance of continued survey and inventory of parasite faunas to establish the limits of diversity on local, regional, and global scales. Consequently we strongly encourage routine deposition of voucher specimens from all survey activities designed to explore faunal diversity. Although this is particularly important in regions that remain poorly known, such as Africa, the principle certainly applies more broadly.

Robustostrongylus aferensis gen. nov. et sp. nov. (Nematoda: Trichostrongyloidea) in kob (Kobus kob) and hartebeest (Alcelaphus buselaphus jacksoni) (Artiodactyla) from sub-Saharan Africa, with further ruminations on the Ostertagiinae.

Abomasal nematodes (Ostertagiinae: Trichostrongyloidea), representing a previously unrecognized genus and species, were discovered in kob (Kobus kob) and kongoni (hartebeest) (Alcelaphus buselaphus jacksoni) from Uganda during surveys of ungulate parasites in the 1960s. Robustostrongylus aferensis gen. nov. et sp. nov. is characterized by a ventriculus-like, bilobed valve at the junction of the esophagus and intestine, a synlophe with unusually robust ridges, cervical papillae and excretory pore situated posterior to the mid-length of the esophagus, a unique body form and large diameter in males and females, a relatively anterior position for the vulva, and strongly convoluted and spiraled ovarian tracks in females. Bursal structure is 2-1-2, with subequal Rays 2/3, strongly reduced and robust Rays 8, and relatively narrow Rays 9/10 contained within a reduced, laterally inflated dorsal lobe. Spicules are filamentous and tripartite; the gubernaculum is cryptic, alate, and heart-shaped in the anterior. Robustostrongylus aferensis, with narrow filamentous spicules that trifurcate distally near 80%, paired arcuate "0" papillae that terminate in bulbous expansions, and a reduced dorsal lobe and ray most closely resembles species of Longistrongylus. A suite of unique characters, consistent in males and females, however, unequivocally distinguishes specimens of R. aferensis from all ostertagiines with either a 2-1-2 or 2-2-1 bursal pattern. Among 15 genera of the Ostertagiinae in the global fauna, 5 are entirely limited in distribution to Africa, including Africanastrongylus, Hamulonema, Longistrongylus, Pseudomarshallagia, and Robustostrongylus gen. nov.; species among 5 additional genera, including Cervicaprastrongylus, Hyostrongylus, Marshallagia, Ostertagia, and Teladorsagia, also occur in Africa, but they are represented as mosaics, with diversity centered in Eurasia or the Holarctic.

Synlophe structure for species of Longistrongylus (Nematoda: Trichostrongyloidea), abomasal parasites among ungulates from sub-Saharan Africa, with comparisons to the global ostertagiine fauna.

The synlophe, or system of longitudinal cuticular ridges characteristic of some trichostrongyloid nematodes, is examined in detail for 6 of 8 species in Longistrongylus (Ostertagiinae) that occur in ungulates across sub-Saharan Africa. Among the species of Longistrongylus examined, 5 are characterized by a tapering pattern laterally in the cervical zone (anterior to the esophageal-intestinal junction), which is largely consistent among multiple male and female specimens; in contrast, for Longistrongylus meyeri the lateral pattern is parallel. The synlophe is bilaterally symmetrical, with ridges extending from the base of the cephalic expansion to near the caudal extremity in males and females. Ridges are acutely pointed, with perpendicular orientation and absence of gradient as viewed in transverse section. Species-specific patterns in conjunction with the numbers of ridges may serve to augment an array of diagnostic characters for species of Longistrongylus and contribute to increasingly accurate identification of female specimens. Among 5 of 6 species examined in the current study, the numbers of ridges in males was equal to or exceeded that observed in females, a pattern seen only in Africanastrongylus among the 15 genera of the Ostertagiinae. The differential numbers of ridges in males and females may represent another character among the suite of attributes that in part diagnose the genus Longistrongylus .