Checklist of helminth parasites of Chiroptera of North America north of Mexico.

A parasite-host, host-parasite and distribution-based checklist of helminths found in bats (Chiroptera) of North America north of Mexico is presented. The parasite-host checklist includes a total of 93 species (including records without a species identification) of helminth parasites reported in the literature from 30 species of bats. These include 54 trematodes, 11 cestodes, and 28 nematodes. Each helminth species is listed under its most current accepted name, with all known synonyms, distribution by state/province, and references for each geographic location. Lists of helminths reported from individual species of bats as well as states of the United States and provinces/territories of Canada are also provided. The following new combinations are proposed: Paralecithodendrium alaskensis (Neiland, 1962) n. comb. for Prosthodendrium alaskensis Neiland, 1962; Paralecthodendrium longiforme (Bhalerao, 1926) n. comb. for Lecithodendrium longiforme Bhalerao, 1926; and Paralecithodendrium singularium (Byrd & Macy, 1942) n. comb. for Prosthodendrium singularium Byrd & Macy, 1942. The state of knowledge of helminths of bats in North America is briefly discussed.

A reconstruction of parasite burden reveals one century of climate-associated parasite decline.

Long-term data allow ecologists to assess trajectories of population abundance. Without this context, it is impossible to know whether a taxon is thriving or declining to extinction. For parasites of wildlife, there are few long-term data-a gap that creates an impediment to managing parasite biodiversity and infectious threats in a changing world. We produced a century-scale time series of metazoan parasite abundance and used it to test whether parasitism is changing in Puget Sound, United States, and, if so, why. We performed parasitological dissection of fluid-preserved specimens held in natural history collections for eight fish species collected between 1880 and 2019. We found that parasite taxa using three or more obligately required host species-a group that comprised 52% of the parasite taxa we detected-declined in abundance at a rate of 10.9% per decade, whereas no change in abundance was detected for parasites using one or two obligately required host species. We tested several potential mechanisms for the decline in 3+-host parasites and found that parasite abundance was negatively correlated with sea surface temperature, diminishing at a rate of 38% for every 1 °C increase. Although the temperature effect was strong, it did not explain all variability in parasite burden, suggesting that other factors may also have contributed to the long-term declines we observed. These data document one century of climate-associated parasite decline in Puget Sound-a massive loss of biodiversity, undetected until now.

Molecular phylogeny supports invalidation of Didelphodiplostomum and Pharyngostomoides (Digenea: Diplostomidae) and reveals a Tylodelphys from mammals.

Alaria, Didelphodiplostomum and Pharyngostomoides are among genera of diplostomid digeneans known to parasitize mammalian definitive hosts. Despite numerous recent molecular phylogenetic studies of diplostomids, limited DNA sequence data is available from diplostomids parasitic in mammals. Herein, we provide the first 28S rDNA and cox1 mtDNA sequences from morphologically identified, adult specimens of Didelphodiplostomum and Pharyngostomoides. Newly generated 28S sequences were used to infer the phylogenetic interrelationships of these two genera among other major lineages of diplostomoideans. The phylogeny based on 28S and a review of morphology clearly suggests that Pharyngostomoides should be considered a junior synonym of Alaria, while Didelphodiplostomum should be considered a junior synonym of Tylodelphys. Pharyngostomoides procyonis (type species), Pharyngostomoides adenocephala and Pharyngostomoides dasyuri were transferred into Alaria as Alaria procyonis comb. nov., Alaria adenocephala comb. nov. and Alaria dasyuri comb. nov.; Didelphodiplostomum variabile (type species) and Didelphodiplostomum nunezae were transferred into Tylodelphys as Tylodelphys variabilis comb. nov. and Tylodelphys nunezae comb. nov. In addition, Alaria ovalis comb. nov. (formerly included in Pharyngostomoides) was restored and transferred into Alaria based on a morphological study of well-fixed, adult specimens and the comparison of cox1 DNA sequences among Alaria spp. The diplostomid genus Parallelorchis was restored based on review of morphology.

Parasitic nematodes of the genus Syphacia Seurat, 1916 infecting Cricetidae in the British Isles: the enigmatic status of Syphacia nigeriana.

Oxyurid nematodes (Syphacia spp.) from bank (Myodes glareolus) and field/common (Microtus spp.) voles, from disparate geographical sites in the British Isles, were examined morphologically and genetically. The genetic signatures of 118 new isolates are provided, based primarily on the rDNA internal transcribed spacers (ITS1-5.8S-ITS2) region and for representative isolates also on the small subunit 18S rDNA region and cytochrome c oxidase subunit 1 (cox-1) gene locus. Genetic data on worms recovered from Microtus spp. from the European mainland and from other rodent genera from the Palaearctic, North America and West Africa are also included. We test historical hypotheses indicating that S. nigeriana is a generalist species, infecting a range of different rodent genera. Our results establish that S. nigeriana is a parasite of both bank and field voles in the British Isles. An identical genotype was also recorded from Hubert's multimammate mouse (Mastomys huberti) from Senegal, but Mastomys spp. from West Africa were additionally parasitized by a related, although genetically distinct Syphacia species. We found no evidence for S. petrusewiczi in voles from the British Isles but isolates from Russia and North America were genetically distinct and formed their own separate deep branch in maximum likelihood molecular phylogenetic trees.

The gastrointestinal nematodes of plains and Grevy's zebras: Phylogenetic relationships and host specificity.

Equids are chronically infected with parasitic strongyle nematodes. There is a rich literature on horse strongyles, but they are difficult to identify morphologically and genetic studies on strongyles infecting other equid species are few, hampering studies of host specificity. We sequenced expelled worms from two sympatric zebra species in central Kenya to expand the strongyle phylogeny and used DNA metabarcoding on faecal samples to genetically characterize zebra nemabiomes for the first time. We generated sequences for several species new to public genetic reference databases, all of which are typical strongyles in wild zebras (i.e., the three species of Cylindropharynx and Cyathostomum montgomeryi), and identified their closest relatives. We also discovered an apparent fungus infecting a quarter of the expelled Crossocephalus viviparus worms, a hyperabundant nematode species in the family Atractidae, hinting at the possibility that zebra host-parasite dynamics may involve a zebra-fungus mutualism. The two zebra species had similar nemabiomes; we found a complete overlap in the list of nematode species they carry and very similar prevalence (i.e., proportion of hosts infected) for the different nematode species. Our study suggests limited host-specificity in zebra strongyles and high potential for transmission between the plains zebra and the endangered Grevy's zebra.

Eimeria longirostris (Apicomplexa: Eimeriidae) from Vagrant Shrew, Sorex vagrans (Mammalia: Eulipotyphla: Soricidae), from Montana, USA.

PURPOSE: Nothing is known about the coccidian parasites of vagrant shrews, Sorex vagrans Baird, 1868. Here we report, for the first time, the occurrence of Eimeria longirostris Hertel and Duszynski, 1987 from faecal contents of S. vagrans from Montana, USA. METHODS: Faecal samples, collected in July and August 2020 from six pitfall-trapped vagrant shrews as well as faeces from two masked shrews, Sorex cinereus Kerr, 1792, and one American pygmy shrew, Sorex hoyi Baird, 1857 from Missoula County, Montana, USA, were examined for coccidian parasites. Samples were placed in individual vials containing aqueous potassium dichromate. They were examined for coccidia after flotation in Sheather's sugar solution, measured, and photographed. RESULTS: Three (50%) S. vagrans and one (50%) S. cinereus were found to be passing oocysts of Eimeria longirostris Hertel and Duszynski, 1987; the single S. hoyi was negative. Oocysts from S. vagrans were subspheroidal and measured (average L × W) 16.1 × 14.4 µm with an L/W ratio of 1.1. One (typically) to two polar granules was present but a micropyle and oocyst residuum were absent. Sporocysts were ovoidal and measured 9.6 × 6.2 µm with an L/W ratio of 1.6. A Stieda body was present but subStieda and paraStieda bodies were absent. The sporocyst residuum was composed of various sized granules typically scattered between and across the sporozoites but sometimes formed a loose aggregate or compact mass. CONCLUSION: We document a new host and new geographic record for E. longirostris from S. vagrans and report the coccidian from S. cinereus for the third time but the first report from specimens from Montana. This coccidian has now been reported from at least 12 species of shrews within the genus Sorex in 14 US states and two provinces in Canada.

Parasite spillover: indirect effects of invasive Burmese pythons.

Identification of the origin of parasites of nonindigenous species (NIS) can be complex. NIS may introduce parasites from their native range and acquire parasites from within their invaded range. Determination of whether parasites are non-native or native can be complicated when parasite genera occur within both the NIS' native range and its introduced range. We explored potential for spillover and spillback of lung parasites infecting Burmese pythons (Python bivittatus) in their invasive range (Florida). We collected 498 indigenous snakes of 26 species and 805 Burmese pythons during 2004-2016 and examined them for lung parasites. We used morphology to identify three genera of pentastome parasites, Raillietiella, a cosmopolitan form, and Porocephalus and Kiricephalus, both New World forms. We sequenced these parasites at one mitochondrial and one nuclear locus and showed that each genus is represented by a single species, R. orientalis, P. crotali, and K. coarctatus. Pythons are host to R. orientalis and P. crotali, but not K. coarctatus; native snakes are host to all three species. Sequence data show that pythons introduced R. orientalis to North America, where this parasite now infects native snakes. Additionally, our data suggest that pythons are competent hosts to P. crotali, a widespread parasite native to North and South America that was previously hypothesized to infect only viperid snakes. Our results indicate invasive Burmese pythons have affected parasite-host dynamics of native snakes in ways that are consistent with parasite spillover and demonstrate the potential for indirect effects during invasions. Additionally, we show that pythons have acquired a parasite native to their introduced range, which is the initial condition necessary for parasite spillback.

A review of Gongylonema spp. (Nematoda: Gongylonematidae) in North American rodents with description of a new species from the cotton rat, Sigmodon hispidus (Mammalia: Cricetidae).

Gongylonema archboldi n. sp. (Nematoda: Gongylonematidae) is described from tunnels in the gastric mucosa of the stomach of the cotton rat (Sigmodon hispidus) from Highlands County, Florida, U.S.A. Measurements are also given for specimens from cotton mice (Peromyscus gossypinus), oldfield mice (Peromyscus polionotus), Florida mice (Podomys floridanus), and golden mice (Ochrotomys nuttalli) from the same locality. Additional specimens were collected from the cotton rat and the rice rat (Oryzomys palustris) from Berry Island, San Patricio County, Texas. The new species is differentiated from congeners by a combination of the following characters: length of the left spicule, length and shape of the gubernaculum, distribution of cuticular bosses, length of esophagus, and distance of the vulva from the posterior end. The status of the genus Gongylonema in North American rodents is reviewed.

Helminth communities of herons (Aves: Ardeidae) in southern Italy.

The helminth communities of nine species of herons from southern Italy were studied and compared. Of 24 taxa found including seven digeneans, seven nematodes, six cestodes and four acanthocephalans, only five taxa were found in more than one heron species, and five of the 21 taxa that could be identified to species level were classified as 'heron specialists'. The total number of helminth species per heron species ranged from 1 in Botaurus stellaris to 9 in Ixobrychus minutus with infection levels generally low. A statistical comparison was carried out for herons with a sample size >5. At the infracommunity level, only I. minutus clearly differed from other heron species. Diversity parameters of heminth infracommunities did not significantly differ among heron species. Species richness ranged from just 0.3 to 2.3 helminth taxa per individual host, and the Brillouin index, from 0 to 0.3. Total helminth abundance did not exceed 40 worms per host except in a single case of Ardeola ralloides. Infracommunities clearly were dominated by single helminth species. The present study confirms a depauperate helminth community in herons from southern Italy. Comparison with data from Spain and the Czech Republic showed strong quantitative similarities with values obtained in the present study. Results also suggest that the composition of local helminth communities are strongly variable depending on geographical location as is demonstrated by comparison with data from other European areas. However, whether herons in Europe naturally host depauperate helminth communities or these communities are depauperate because of other factors is unknown.

New host, geographic records, and histopathologic studies of Angiostrongylus spp (Nematoda: Angiostrongylidae) in rodents from Argentina with updated summary of records from rodent hosts and host specificity assessment.

To date, 21 species of the genus Angiostrongylus (Nematoda: Angiostrongylidae) have been reported around the world, 15 of which are parasites of rodents. In this study, new host, geographic records, and histopathologic studies of Angiostrongylus spp in sigmodontine rodents from Argentina, with an updated summary of records from rodent hosts and host specificity assessment, are provided. Records of Angiostrongylus costaricensis from Akodon montensis and Angiostrongylus morerai from six new hosts and geographical localities in Argentina are reported. The gross and histopathologic changes in the lungs of the host species due to angiostrongylosis are described. Published records of the genus Angiostrongylus from rodents and patterns of host specificity are presented. Individual Angiostrongylus species parasitise between one-19 different host species. The most frequent values of the specificity index (STD) were between 1-5.97. The elevated number of host species (n = 7) of A. morerai with a STD = 1.86 is a reflection of multiple systematic studies of parasites from sigmodontine rodents in the area of Cuenca del Plata, Argentina, showing that an increase in sampling effort can result in new findings. The combination of low host specificity and a wide geographic distribution of Angiostrongylus spp indicates a troubling epidemiological scenario although, as yet, no human cases have been reported.

New host, geographic records, and histopathologic studies ofAngiostrongylus spp (Nematoda: Angiostrongylidae) in rodents from Argentina with updated summary of records from rodent hosts and host specificity assessment

To date, 21 species of the genus Angiostrongylus (Nematoda: Angiostrongylidae) have been reported around the world, 15 of which are parasites of rodents. In this study, new host, geographic records, and histopathologic studies of Angiostrongylus spp in sigmodontine rodents from Argentina, with an updated summary of records from rodent hosts and host specificity assessment, are provided. Records of Angiostrongylus costaricensis from Akodon montensis andAngiostrongylus morerai from six new hosts and geographical localities in Argentina are reported. The gross and histopathologic changes in the lungs of the host species due to angiostrongylosis are described. Published records of the genus Angiostrongylus from rodents and patterns of host specificity are presented. Individual Angiostrongylusspecies parasitise between one-19 different host species. The most frequent values of the specificity index (STD) were between 1-5.97. The elevated number of host species (n = 7) of A. morerai with a STD = 1.86 is a reflection of multiple systematic studies of parasites from sigmodontine rodents in the area of Cuenca del Plata, Argentina, showing that an increase in sampling effort can result in new findings. The combination of low host specificity and a wide geographic distribution of Angiostrongylus spp indicates a troubling epidemiological scenario although, as yet, no human cases have been reported.

Endohelminths in Bird Hosts from Northern California and an Analysis of the Role of Life History Traits on Parasite Richness.

The life history characteristics of hosts often influence patterns of parasite infection either by affecting the likelihood of parasite exposure or the probability of infection after exposure. In birds, migratory behavior has been suggested to affect both the composition and abundance of parasites within a host, although whether migratory birds have more or fewer parasites is unclear. To help address these knowledge gaps, we collaborated with airports, animal rescue/rehabilitation centers, and hunter check stations in the San Francisco Bay Area of California to collect 57 raptors, egrets, herons, ducks, and other waterfowl for parasitological analysis. After dissections of the gastrointestinal tract of each host, we identified 64 taxa of parasites: 5 acanthocephalans, 24 nematodes, 8 cestodes, and 27 trematodes. We then used a generalized linear mixed model to determine how life history traits influenced parasite richness among bird hosts, while controlling for host phylogeny. Parasite richness was greater in birds that were migratory with larger clutch sizes and lower in birds that were herbivorous. The effects of clutch size and diet are consistent with previous studies and have been linked to immune function and parasite exposure, respectively, whereas the effect of migration supports the hypothesis of "migratory exposure" rather than that of "migratory escape."

A new genus and species of Heligmonellidae (Nematoda: Trichostrongylina) parasitic in Delomys dorsalis (Rodentia: Sigmodontinae) from Misiones, Argentina.

Alippistrongylus bicaudatus gen. et sp. n. (Nematoda: Heligmonellidae) is described from the striped Atlantic forest rat, Delomys dorsalis (Hensel) (Rodentia: Sigmodontinae), from the province of Misiones in Argentina. The new genus and species is characterised by a synlophe of 21 unequal ridges in both sexes without a gradient in size, with two ridges weakly sclerotised and oriented perpendicularly in the dorsal left quadrant; males with a highly dissymmetrical bursa with a hypertrophied right lobe, and females with a dorsal conical appendage just posterior to the vulva, conferring a two-tailed appearance to the female worms.

A megafauna's microfauna: gastrointestinal parasites of New Zealand's extinct moa (Aves: Dinornithiformes).

WE PERFORM THE FIRST MULTIDISCIPLINARY STUDY OF PARASITES FROM AN EXTINCT MEGAFAUNAL CLADE USING COPROLITES FROM THE NEW ZEALAND MOA (AVES: Dinornithiformes). Ancient DNA and microscopic analyses of 84 coprolites deposited by four moa species (South Island giant moa, Dinornis robustus; little bush moa, Anomalopteryx didiformis; heavy-footed moa, Pachyornis elephantopus; and upland moa, Megalapteryx didinus) reveal an array of gastrointestinal parasites including coccidians (Cryptosporidium and members of the suborder Eimeriorina), nematodes (Heterakoidea, Trichostrongylidae, Trichinellidae) and a trematode (Echinostomida). Parasite eggs were most prevalent and diverse in coprolites from lowland sites, where multiple sympatric moa species occurred and host density was therefore probably higher. Morphological and phylogenetic evidence supports a possible vicariant Gondwanan origin for some of the moa parasites. The discovery of apparently host-specific parasite taxa suggests paleoparasitological studies of megafauna coprolites may provide useful case-studies of coextinction.

Outbreak of Philophthalmus gralli in four greater rheas (Rhea americana).

Using slit-lamp biomicroscopy, conjunctival biopsy, and morphological identification, a flock of four Greater rheas (Rhea americana) in Arizona were diagnosed with conjunctivitis secondary to Philophthalmus gralli (P. gralli) infection. Aquatic snails from the exhibit's water source were identified as Melanoides tuberculatus, a known vector for P. gralli. Comparison of partial sequences of DNA regions from P. gralli adults removed from the rheas and metacercariae from the aquatic snails demonstrated a 100% match, confirming the source of infection. The flock was divided into two treatment groups: the most severely affected rheas received both manual removal of trematodes and praziquantel 1% ointment OU q12 h and the least severely affected rheas were only given praziquantel 1% ointment OU q12 h. The rheas were permanently relocated away from the infected water source and aquatic snails. Initial resolution was seen at 17 weeks in the most severely affected rhea, which had 675 adult P. gralli removed and topical praziquantel. The two rheas that only received topical praziquantel showed resolution within 3 and 15 weeks. Current recommendations for treating P. gralli include: manual removal of trematodes, topical praziquantel 1% ointment, and relocation away from infected water sources and aquatic snails.

Morphological and molecular differentiation of Staphylocystis clydesengeri n. sp. (Cestoda, Hymenolepididae) from the vagrant shrew, Sorex vagrans (Soricomorpha, Soricidae), in North America.

Staphylocystis clydesengeri n. sp. is described from shrews Sorex vagrans in Montana and Washington, United States. It differs from the only previously known North American representative of the genus, S. schilleri, in having more numerous (37-42 vs. 22-30) and larger (39-44 microm vs. 27-30 microm) rostellar hooks. The two species also differ in several other important characters such as relative length of the cirrus pouch, position of gonads and shape of mature proglottides. Morphological differentiation of the new species from all previously known Palearctic species of Staphylocystis from Sorex is also provided. Differentiation from Staphylocystis parasitic in crocidurine shrews is not provided due to the high level of specificity among shrew hymenolepidids to the host genera and much greater levels of sequence divergence between Staphylocystis from the two groups of shrews. Molecular differentiation based on 2,800 base pair long sequences of nuclear ribosomal RNA (complete ITS region and partial 28S region), 663 base pair long sequences of mitochondrial nad1 gene and 542 base pair long sequences of mitochondrial ribosomal 16S gene strongly support the status of Staphylocystis clydesengeri n. sp. Relative utility of the DNA fragments used in this study for reliable differentiation among closely related species of mammalian hymenolepidids is discussed. Nuclear ribosomal RNA region appears to be too conserved for this purpose. Use of at least one mitochondrial gene in addition to nuclear ribosomal RNA or without it, is recommended. Vampirolepis novosibirskiensis Sawada & Kobayashi, 1994 is transferred to Staphylocystis as a junior synonym of S. furcara (Stieda, 1862). Rodentolepis gnoskei Greiman & Tkach, 2012 is transferred to Pararodentolepis Makarikov and Gulyaev, 2009 as a new combination Pararodentolepis gnoskei (Greiman & Tkach, 2012) n. comb.

Helminth collection and identification from wildlife.

Wild animals are commonly parasitized by a wide range of helminths. The four major types of helminths are "roundworms" (nematodes), "thorny-headed worms" (acanthocephalans), "flukes" (trematodes), and "tapeworms" (cestodes). The optimum method for collecting helminths is to examine a host that has been dead less than 4-6 hr since most helminths will still be alive. A thorough necropsy should be conducted and all major organs examined. Organs are washed over a 106 µm sieve under running water and contents examined under a stereo microscope. All helminths are counted and a representative number are fixed (either in 70% ethanol, 10% buffered formalin, or alcohol-formalin-acetic acid). For species identification, helminths are either cleared in lactophenol (nematodes and small acanthocephalans) or stained (trematodes, cestodes, and large acanthocephalans) using Harris' hematoxylin or Semichon's carmine. Helminths are keyed to species by examining different structures (e.g. male spicules in nematodes or the rostellum in cestodes). The protocols outlined here can be applied to any vertebrate animal. They require some expertise on recognizing the different organs and being able to differentiate helminths from other tissue debris or gut contents. Collection, preservation, and staining are straightforward techniques that require minimal equipment and reagents. Taxonomic identification, especially to species, can be very time consuming and might require the submission of specimens to an expert or DNA analysis.