An annotated checklist of the genus Pterygodermatites Wedl, 1861 (Nematoda: Rictulariidae), with notes on hosts and geographical distribution.

Species of rictulariid nematodes belonging to the genus Pterygodermatites Wedl, 1861 are parasites of mammals with a wide geographical distribution. The taxonomy of this speciose genus is complex and based mainly on morphological traits of the cephalic structures and cuticular processes. Currently, the known species are allocated to five subgenera [P. (Paucipectines), P. (Neopaucipectines), P. (Multipectines), P. (Pterygodermatites) and P. (Mesopectines)], based on the classification proposed in 1969 by Quentin. Studies on life cycles, taxonomy, biogeography and association of these parasites with mammals peaked at the end of the 1960s. Although the momentum was lost, new species and geographic records have been added in the last 50 years. Therefore, an updated checklist of species of Pterygodermatites is presented here, based on a literature survey. Information on classification, geographical distribution and host range obtained for 68 species is presented, including a distribution map. Moreover, the main differential morphological traits related to each species are tabulated. More than 140 species from 10 different mammalian orders (Carnivora, Chiroptera, Didelphimorphia, Erinaceomorpha, Macroscelidae, Microbiotheria, Paucituberculata, Primates, Rodentia and Xenarthra) have been reported as hosts of Pterygodermatites spp. The taxonomic information here compiled can contribute to future studies on the biogeography, in addition to evolutionary and ecological aspects of this diverse but poorly known genus of parasitic nematodes of mammals.

DIPLOTRIAENA OBTUSA (NEMATODA: DIPLOTRIAENIDAE) FROM BARN SWALLOWS (HIRUNDO RUSTICA) AND CLIFF SWALLOWS (PETROCHELIDON PYRRHONOTA) COLLECTED DURING MORTALITY EVENTS IN THE UPPER MIDWEST, USA.

Several mortality events involving barn swallows (Hirundo rustica) and cliff swallows (Petrochelidon pyrrhonota) were reported in the Upper Midwestern states in 2017 and 2018. Barn swallow mortality followed unseasonal cold snaps, with the primary cause of death being emaciation with concurrent air sac nematodiasis. Lesions in cliff swallows were consistent with blunt force trauma from suspected car impacts. Examination of air sac nematodes from both bird species revealed morphological characters consistent with Diplotriaena obtusa. Sequence analysis of the partial 18S rRNA gene indicated the samples clustered with other species in the genus Diplotriaena. These nematodes provide a link between morphological specimens and DNA sequence data for D. obtusa.

Deep in the systematics of Camallanidae (Nematoda): using integrative taxonomy to better understand the phylogeny and consistency of diagnostic traits.

Due to conflicts between classic and molecular systematics of Camallanidae, different data types were used for the first time, to better understand the evolutionary history and taxa consistency within this family. Genetic [18S and 28S rDNA; cytochrome c oxidase subunit I (COI) mtDNA], morphological and life history traits were used to infer phylogenies using Bayesian inference, reconstructed from separated and concatenated datasets. The consistency of tree and morphological traits was evaluated using the consistency index. Characters were mapped on the trees and the phylogenetic informativeness of genetic markers was estimated. Phylogenetic informativeness of 18S provided better resolution for outer nodes, COI for inners and 28S had an intermediate profile. New sequences for two camallanid species were obtained. Phylogenies of genetic and concatenated data largely agreed, showing more divergence in the COI dataset, due to its higher mutation rate vs stable morphology for diagnosing higher taxa. No genus sustained monophyly. The lack of autapomorphy and phylogenetic proximity supported the partition of Batrachocamallanus as synonym of Procamallanus and Spirocamallanus, which should not be considered as subgenera. Although traits of buccal capsule, male tail, habitat, host and biogeographic were highly consistent, intrinsic patterns varied according to different taxa assemblages. Morphological systematics of Camallanidae, based on buccal capsule, is artificial for certain taxa.

Description and Molecular Differentiation of a New Skrjabinoptera (Nematode: Physalopteridae) from Eutropis macularia (Sauria: Scincidae) in North-Central Vietnam.

Skrjabinoptera vietnamensis n. sp. is described from specimens recovered from the stomach of Eutropis macularia in north-central Vietnam. The new species is characterized by the medium-sized male worms (6.7-8.7 mm in length and 154-182 µm in width) relative to known members of the genus, 2 pointed spicules of unequal length (87-112 µm and 56-72 µm in length), and 10 pairs of caudal papillae. Female worms are larger than male worms (10.7-18.4 mm in length and 264-411 µm in width), with the vulva situated in the anterior part, and embryonated, elliptical eggs, 35-46 µm long by 20-24 µm wide. Skrjabinoptera vietnamensis n. sp. represents the ninth species assigned to the genus and the first species recorded from the Oriental region. Partial sequences of the 18S ribosomal RNA gene (rDNA), and cytochrome c oxidase subunit 1 (COI) are provided for the new species. The molecular phylogenetic position of the genus Skrjabinoptera is briefly discussed.

First Documentation of Nematode Dispharynx sp. in Scaled Quail.

Quail populations in the United States have been declining for several decades, and the role that parasites may be playing in this decline is not well understood. The goal of this study was to document novel parasites that inhabited the scaled quail, Callipepla squamata, of the Trans-Pecos ecoregion of Texas. To do this, quail were collected by hunter-harvest, night-netting, and funnel-trapping and were necropsied in the laboratory to determine the parasites they hosted. After analyzing 386 birds, we identified Dispharynx sp. in one of the samples. This specimen is the first to be officially documented in scaled quail.

Description and Molecular Differentiation of a New Falcaustra (Nematode: Kathlaniidae) from the Indochinese Water Dragon, Physignathus cocincinus (Squamata: Agamidae) in North-central Vietnam.

Falcaustra vietnamensis n. sp. is described from the small intestine of Physignathus cocincinus from north-central Vietnam. The new species is characterized by the large male worms (20.2-28.8 mm in length and 557-724 µm in width) relative to known members of the genus, 2 sharply pointed alate spicules of equal length (1,128-1,256 µm in length), gubernaculum including 2 separate pieces, 1 ventral with a pointed distal end and 1 dorsal with a blunt distal end (164-192 µm and 155-172 µm in length, respectively), and 12 pairs of caudal papillae. Female worms are larger than male worms (24.2-34.1 mm in length and 532-735 µm in width), with the vulva situated in the posterior half of body, and elliptical eggs, 60-70 µm long by 42-47 µm wide. Falcaustra vietnamensis n. sp. represents the 38th species assigned to the genus and the third species recorded from a lizard host in the Oriental biogeographical region. Partial sequences of the 18S ribosomal RNA gene (rDNA), internal transcribed spacer regions (ITS), and cytochrome c oxidase subunit 1 (COI) are provided for the new species. The molecular phylogenetic position of the genus Falcaustra is briefly discussed.

Redescription of Serratospiculum seurati Bain & Mawson, 1981 (Nematoda; Diplotriaenidae) from Peregrine falcon (Falco peregrinus Tunstall, 1771).

Air sac nematodes from birds are known for more than 200 years now and Filaria attenuata was the first described species from falcons, owl and corvid birds. The superficial description and the loss of the original material made F. attenuata a species inquirenda. Seurat (1915) redescribed the species with material from lanner falcon and pallid harrier from Algeria and based on this description Bain and Mawson, Rec S Aust Mus 18:265-28, (1981) created a new species, Serratospiculum seurati, by adding some, slightly divergent, measurements. The current paper is based on light and scanning electron microscopy of five male and 10 female S. seurati specimens from a Peregrine falcon that acquired the infection in Pakistan. The length of the slender male and female nematodes varied between 42-70 and 165-221 mm, respectively, spicules of unequal shape and length measured 292-325 and 638-785 µm. S. seurati was also found in Saker, Barbary and crossbreed falcons.

Cucullanus carettae Baylis, 1923, in a loggerhead sea turtle (Caretta caretta) from the Adriatic sea: first detection and molecular characterization.

Cucullanus carettae Baylis, 1923 (Nematoda: Cucullanidae) is found worldwide in loggerhead turtles (Caretta caretta). Regarding the Mediterranean, C. carettae has been identified in the Tyrrhenian and the Ionian Sea and a unique description of a Cucullanus sp. specimen in loggerheads from the Adriatic Sea has been reported in the literature so far. In the framework of a bio-monitoring project of the Abruzzo and Molise coasts, a parasitological survey was performed on stranded and by-caught sea turtles, at the Istituto Zooprofilattico of Abruzzo and Molise "G. Caporale." During necropsy, the gastrointestinal system of 72 stranded loggerhead turtles was analyzed for the presence of endoparasites and fecal samples were collected for coprological examination. Adult C. carettae (n = 123) was found in the upper intestine of one loggerhead turtle, associated with chronic lymphoplasmocytic enteritis. Additionally, five stool samples (6.9%) were positive for Cucullanus sp. eggs. Molecular characterization of adult nematodes was carried out to study phylogenetic relationships among the Cucullanus species. To our knowledge, this is the first morphological and molecular identification of C. carettae in loggerhead turtles from the Adriatic Sea. Additional studies on the distribution of this parasite in the Mediterranean are encouraged.

Development of conventional multiplex PCR method for discrimination between Dispharynx nasuta and Cheilospirura hamulosa (Nematoda: Acuariidae) parasitizing poultry.

The poultry infections caused by Dispharynx nasuta and Cheilospirura hamulosa nematodes are difficult to be diagnosed by fecal examination because of their egg similarity. In this study, we analyzed DNA sequences of nuclear ribosomal 18S-ITS1-5.8S-ITS2-28S region of D. nasuta and C. hamulosa and developed conventional multiplex PCR method using species-specific primers for discriminating between the two species. The method amplified 455-bp and 319-bp fragments specific to D. nasuta and C. hamulosa, respectively, and did not produce them against the other chicken nematode species, Ascaridia galli, Oxyspirura mansoni, Heterakis gallinarum, Heterakis beramporia, and Heterakis indica, suggesting that the multiplex PCR is sensitive and available for species diagnosis.

Molecular Phylogeny of Thelastomatoidea (Nematoda) with the Description of a New Genus and Two New Species of Hystrignathidae Associated with Bess Beetles (Coleoptera: Passalidae) from Oaxaca, Mexico.

Bess beetles (Passalidae) display important roles in forestall ecosystems, particularly in energy extraction from dead wood. These organisms maintain complex biological interactions with their gut symbiotic communities, including bacteria, protists, and metazoans. Very little is known about symbionts since most of the species of Passalidae haven't been studied from a parasitological point of view. Here we describe a new genus and 2 new species of nematodes of the family Hystrignathidae associated with 2 beetle species of the tribe Proculini collected in the State of Oaxaca, Mexico. Tuhmai garciaprietoi n. gen., n. sp., found in Vindex agnoscendus is characterized by the presence of an unarmed cervical cuticle, a subcylindrical procorpus and a conspicuous isthmus, a monodelphic-prodelphic reproductive system, and a short subulate tail. Urbanonema osorioi n. sp., found in Verres hageni mainly differs from other species of Urbanonema by the number and disposition of cervical spines, as well as by a subulate tail. For each new taxon, we describe the external and internal morphology, and we generated molecular data (nuclear ribosomal DNA) to place the new taxa in a phylogenetic context.

An integrative approach assesses the intraspecific variations of Procamallanus (Spirocamallanus) inopinatus, a common parasite in Neotropical freshwater fishes, and the phylogenetic patterns of Camallanidae.

Integrative taxonomy was used to evaluate two component populations of Procamallanus (Spirocamallanus) inopinatus in Brazil and the phylogeny Camallanidae. Parasite populations were collected in the characiform Anostomoides passionis from River Xingu (Amazon basin) and Megaleporinus elongatus from River Miranda (Paraguay basin). Morphology was analysed using light and scanning electron microscopy (SEM). Genetic characterization was based on partial sequences of the 18S and 28S rDNA, and COI mtDNA. Phylogenies were based on 18S and COI due to data availability. Generalized Mixed Yule Coalescent (GMYC), Poisson Tree Process (PTP) and *BEAST were used for species delimitation and validation. SEM revealed for the first time the presence of minute denticles and pore-like structures surrounding the oral opening, phasmids in females and confirmed other important morphological aspects. Statistical comparison between the two-component populations indicated morphometric variations, especially among males. The different component population of P. (S.) inopinatus showed variable morphometry, but uniform morphology and were validated as conspecific by the GMYC, PTP and *BEAST. Some camallanid sequences in GenBank have incorrect taxonomic labelling. Host, environment and geographic aspects seem to be related to some lineages within Camallanidae; however, their real phylogenetic meanings are still unclear.

Surveillance for Heterakis spp. in Game Birds and Cage-Free, Floor-Raised Poultry in Pennsylvania.

Histomoniasis is a significant disease of gallinaceous birds caused by Histomonas meleagridis. Transmission of this parasite is dependent on use of the cecal nematode Heterakis gallinarum. To define the host range of this nematode, cecal contents from 399 game birds and poultry, representing eight species, were examined for Heterakis spp. The majority of these species (five of eight) were infected with Heterakis nematodes. Heterakis gallinarum was detected in free-ranging wild turkeys (Meleagridis gallopovo), captive-raised ring-necked pheasants (Phasianus colchicus), chukars (Alectoris chukar), and domestic chickens (Gallus gallus domesticus), whereas H. isolonche was found in ruffed grouse (Bonasa umbellus). No Heterakis species were identified in the domestic turkey (Meleagridis gallopovo), American woodcock (Scolopax minor), and dabbling duck (Anas spp.) samples. Genetic characterization indicated that nematodes identified as H. gallinarum were present in two distinct clades. One clade of H. gallinarum sequenced from this study grouped with chicken-derived sequences from other countries. The other group of sequences consisted of a sister clade to a group of parasites morphologically identified as H. isolonche. Currently it is unknown if this group represents a genetic variant of H. gallinarum, a variant of H. isolonche, or a novel species. These results indicate Heterakis infection varies among poultry and game bird species but is common among select gallinaceous species in Pennsylvania.

Nota de investigación- Vigilancia de Heterakis spp. en aves de caza y aves comerciales criadas en piso sin jaulas en Pennsylvania. La histomoniasis es una enfermedad importante de las aves gallináceas causada por Histomonas meleagridis. La transmisión de este parásito depende de la interacción con el nematodo cecal Heterakis gallinarum. Para definir el rango de hospedadores de este nematodo, se examinaron los contenidos cecales de 399 aves de caza y aves domésticas, que representaron a ocho especies, para detectar Heterakis spp. La mayoría de estas especies (cinco de ocho) estaban infectadas con nematodos Heterakis. Heterakis gallinarum se detectó en pavos silvestres (Meleagridis gallopavo), faisanes comunes criados en cautiverio (Phasianus colchicus), perdiz chukar (Alectoris chukar) y pollos domésticos (Gallus gallus domesticus), mientras que H. isolonche se encontró en el grévol engolado (Bonasa umbellus). No se identificaron especies de Heterakis en las muestras de pavo doméstico (Meleagridis gallopavo), chocha americana (Scolopax minor) y pato chapuceadores (Anas spp.). La caracterización genética indicó que los nematodos identificados como H. gallinarum estaban presentes en dos clados distintos. Un clado de H. gallinarum secuenciado de este estudio agrupado con secuencias derivadas de pollos de otros países. El otro grupo de secuencias consistió en un clado hermano de un grupo de parásitos identificados morfológicamente como H. isolonche. Actualmente se desconoce si este grupo representa una variante genética de H. gallinarum, una variante de H. isolonche o una especie nueva. Estos resultados indican que la infección por Heterakis varía entre las aves domésticas y las especies de aves de caza, pero es común entre las especies de gallináceas seleccionadas en Pennsylvania.

Vertical Transmission of a Nematode from Female Lizards to the Brains of Their Offspring.

Parasites have evolved a diversity of lifestyles that exploit the biology of their hosts. Some nematodes that parasitize mammals pass via the placenta or milk from one host to another. Similar cases of vertical transmission have never been reported in avian and nonavian reptiles, suggesting that egg laying may constrain the means of parasite transmission. However, here we report the first incidence of transovarial transmission of a previously undescribed nematode in an egg-laying amniote, the common wall lizard (Podarcis muralis). Nematodes enter the developing brain from the female ovary early in embryonic development. Infected lizard embryos develop normally and hatch with nematodes residing in their braincase. We present a morphological and molecular phylogenetic characterization of the nematode and suggest that particular features of lizard biology that are absent from birds and turtles facilitated the evolutionary origin of this novel life history.

Morphology and Molecular Characterization of Parabronema smithii (Cobbold, 1882) (Nematoda: Habronematidae) from Wild Asian Elephant (Elephas maximus maximus) of Sri Lanka.

PURPOSE: The aim of the present study was to carry out a detailed study of morphological features and to determine the phylogenetic position of Parabronema smithii (Cobbold, 1882) found in wild elephants in Sri Lanka. METHODS: Adult worms were collected from stomach ulcers at postmortem examination of wild elephants in the Udawalawe National Park, Sri Lanka. The detailed morphology of P. smithii was studied using light microscopy and, for the first time, scanning electron microscopy. Fifteen morphological characteristics were investigated. The phylogenetic analysis was conducted using the second internal transcribed spacer region (ITS2), and portions of the large subunit ribosomal DNA (28S) and cytochrome c oxidase subunit 1 (cox1). Furthermore, the present study provides a comparison of morphology and morphometrics of Parabronema species that occur in different hosts. CONCLUSION: Parabronema smithii isolated from wild elephants exhibited the key morphological features. Phylogenetic analysis of selected genes revealed that P. smithii is closely associated with P. skrjabini and Habronema spp. Findings of the present study enhance our understanding of the biology and taxonomy of P. smithii in wild elephant in Sri Lanka and will contribute to future phylogeographic studies.

A New Species of Triumphalisnema (Nematoda: Coronostomatoidea) Parasite of Proculejus hirtus (Coleoptera: Passalidae) from Mexico, and the Position of the Family in the Oxyuridomorpha.

A new species of the nematode Triumphalisnema Kloss, 1962 (Oxyuridomorpha), is described from the wood beetle Proculejus hirtus Truqui from the mountain mesophilic forest in Hidalgo State, Mexico. Triumphalisnema zuuei n. sp. is distinguished from the other 4 congeners species by the presence of an expanded cervical ring, well-developed lateral alae, an obtuse cauda with a short and bifurcated caudal appendage, series of cuticular folds at ventral and dorsal body surface from excretory pore level to anal region, and ellipsoidal eggs ornamented with numerous small mushroom-like structures over the surface. The phylogenetic position of the new species is inferred based on a Maximum Likelihood and Bayesian Inference analysis of partial sequences of 18S SSU rRNA. The phylogenetic analysis showed that Triumphalisnema zuuei n. sp., the only representative of the Traklosiidae in our study, is closely related to Coynema poeyi and species of Longior, Hystrignathus, and Lepidonema, all of them members of Hystrignathidae. These relationships are supported by high support values. The present study increases to 5 the number of species assigned to Triumphalisnema, all of them parasites of Passalidae. Additionally, a taxonomic key to the species of the genus is provided.

Redescription of Dichelyne ( Cucullanellus) Dichelyneformis (Nematoda: Cucullanidae) and Description of a New Congeneric Species in Eleginops maclovinus (Pisces: Eleginopsidae) Collected in the Bay of Corral, Chile.

Dichelyne ( Cucullanellus) dichelyneformis ( Szidat, 1950 ) Petter, 1974 , was collected in the intestine of the rock cod, Eleginops maclovinus, in the Bay of Corral, Chile, and redescribed based on light and scanning electron microscopy. The correct arrangement of caudal papillae and post deirids, the shape of gubernaculum, and the morphology and number of denticles surrounding the oral opening are described for the first time. These characteristics along with the excretory pore position, size of spicules, and presence of a nodular thickening near the distal end of the spicules differentiate D. ( C.) dichelyneformis from the other congeners reported in South America and in the Antarctic. The new congeneric species found in E. maclovinus differs from D. ( C.) dichelyneformis based on the larger males, larger and more numerous denticles with a smooth surface, weakly developed sclerotized plates in esophastome, excretory pore and deirids posterior to nerve ring, shorter gubernaculum and spicules with no thickening on their distal region, lower spicules/body length ratio (10.4-15.1%), and a papilla-like phasmids closer to pairs 8 and 10.

Cosmocerca bengalensis sp. nov. (Nematoda: Cosmocercidae) in Hoplobatrachus tigerinus (Daudin, 1803) (Amphibia, Anura, Dicroglossidae) from West Bengal, India.

Cosmocerca bengalensis sp. nov. (Ascaridida, Cosmocercidae) recovered from the rectum of an Indian bullfrog, Hoplobatrachus tigerinus (Daudin, 1803), collected from Hetampur town in Birbhum district of West Bengal, India, is described and illustrated. This species is similar to C. acanthurum, C. banyulensis, C. cruzi, C. japonica, C. kalesari, C. microhylae, C. novaeguineae, C. ornata, C. paraguayensis, C. parva, C. podicipinus and C. travassosi in having 5 pairs of plectanes supporting preanal papillae but differs from these species by general morphometry, absence of somatic papillae in females, absence of gubernaculum and having only one pair of adanal papillae in males and one pair postanal papillae in females except C. microhylae. Present parasites differ from C. microhylae by absence of gubernaculum and general morphometry. Cosmocerca bengalensis sp. nov. represents the thirtheith species assigned to the genus, seventh from Oriental region and fifth species from India.

A new species of Orientatractis (Nematoda: Atractidae) from the tortoise Indotestudo elongata (Blyth) in China.

A new species of nematode, Orientatractis longicaudata n. sp. is described from the intestine of Indotestudo elongata (Blyth) (Testudinidae) from Zoo of Tianjin, Tianjin, China. The new species can be easily distinguished from its congers by having longer tail, by the length of gubernaculum and left spicule, and by the numbers of caudal papillae. This is first species of Orientatractis Petter, 1966 reported in China. A key to the species of Orientatractis is provided.

Redescription of Dichelyne (Cucullanellus) jialaris (Nematoda: Cucullanidae), a parasite of seabreams (Perciformes: Sparidae) in East Asia.

An insufficiently known nematode species Dichelyne (Cucullanellus) jialaris Luo, Guo, Fang et Huang, 2004 (Cucullanidae), originally described from the red seabream Pagrus major (Temminck et Schlegel) (Sparidae) in the Taiwan Strait, off China, is redescribed based on light and scanning electron microscopical (SEM) examinations of specimens collected from the intestine of three species of sparid fishes, Acanthopagrus schlegelii (Bleeker), Evynnis tumifrons (Temminck et Schlegel) and P. major, in Hiroshima Bay, Japan. The present redescription provides exact data on the cephalic structure, character of cephalic papillae and amphids, location of the excretory pore, deirids and phasmids, and the number and distribution of caudal papillae in the male; the presence of the dorsal postdeirid in this species is reported for the first time. The SEM study revealed that the anterior cloacal lip has a median lobe-like elevation bearing two minute papillae, whereas the posterior cloacal lip is somewhat depressed, with a median longitudinal rod-like elevation probably serving to direct the movement of spicules. The size of eggs is reported for the first time. Dichelyne (C.) jialaris differs from the morphologically similar D. (C.) pleuronectidis (Yamaguti, 1935) mainly in the shape of the oesophagus. The present finding of D. (C.) jialaris represents the first record of this nematode from fishes in Japanese waters and A. schlegelii and E. tumifrons are its new host records.

New Species of Falcaustra (Nematoda: Kathlaniidae) in Batagur trivittata (Testudines: Geoemydidae) from Myanmar.

Falcaustra tintlwini sp. nov. (Ascaridida, Kathlaniidae) from the large intestine of Batagur trivittata (Testudines, Geoemydidae) is described and illustrated. Falcaustra tintlwini represents the 20th Oriental species assigned to the genus and is distinguished from other Oriental species by the distribution pattern of the caudal papillae (6 precloacal, 12 postcloacal, and 1 median), length of spicules (0.43-0.50 mm) and absence of a pseudosucker.