The genus Oswaldocruzia represents a taxonomically diverse group of nematodes with global distribution. Although Oswaldocruzia species are widespread and exhibit a remarkably wide host range in some species, their genetic diversity and biogeographic patterns remain poorly understood. This study investigated the genetic variability and distribution of Oswaldocruzia spp. in nine anuran species from the genera Bufo, Bufotes, Pelophylax, and Rana across Central Europe and the Balkans. Two species were identified: Oswaldocruzia filiformis and O. ukrainae, each exhibiting a different range of host associations. Phylogenetic analyses based on mitochondrial COI sequences revealed significant haplotype diversity in the generalist O. filiformis, with low geographic and host-associated genetic structuring. In contrast, O. ukrainae, which is closely associated with Bufotes viridis, exhibited only one genetic variant across all samples, highlighting its restricted genetic diversity. The findings emphasize contrasting genetic diversities among nematode parasites exhibiting different levels of host-specificity and expand the known distribution of O. filiformis into new regions of the Balkans. In addition, they highlight the need for additional studies on the ecological and evolutionary factors that influence the genetic diversity of parasites in amphibians.

TITLE: Aperçu de la diversité génétique et de la répartition des espèces des nématodes Oswaldocruzia (Trichostrongylida : Molineidae) en Europe : absence apparente de structuration géographique et populationnelle chez les amphibiens. ABSTRACT: Le genre Oswaldocruzia représente un groupe de nématodes taxonomiquement diversifié, réparti à l’échelle mondiale. Bien que les espèces d’Oswaldocruzia soient répandues et présentent une gamme d’hôtes remarquablement large chez certaines espèces, leur diversité génétique et leurs profils biogéographiques restent mal compris. Cette étude a examiné la variabilité génétique et la répartition des espèces d’Oswaldocruzia chez neuf espèces d’anoures des genres Bufo, Bufotes, Pelophylax et Rana en Europe centrale et dans les Balkans. Deux espèces ont été identifiées : Oswaldocruzia filiformis et O. ukrainae, chacune présentant une gamme différente d’associations d’hôtes. Les analyses phylogénétiques basées sur les séquences COI mitochondriales ont révélé une diversité d’haplotypes significative chez l’espèce généraliste O. filiformis, avec une faible structuration génétique géographique et associée à l’hôte. En revanche, O. ukrainae, étroitement associé à Bufotes viridis, ne présentait qu’une seule variante génétique dans tous les échantillons, soulignant sa diversité génétique limitée. Ces résultats mettent en évidence les diversités génétiques contrastées parmi les nématodes parasites présentant différents niveaux de spécificité d’hôte et étendent la répartition connue d’O. filiformis à de nouvelles régions des Balkans. De plus, ils soulignent la nécessité de mener des études complémentaires sur les facteurs écologiques et évolutifs qui influencent la diversité génétique des parasites des amphibiens.

Department of Animal Physiology Faculty of Science Pavol Jozef Šafárik University in Košice Šrobárova 2 040 01 Košice Slovakia

Department of Zoology Faculty of Natural Sciences Comenius University in Bratislava Ilkovičova 6 842 15 Bratislava Slovakia

Department of Zoology Faculty of Natural Sciences Comenius University in Bratislava Ilkovičova 6 842 15 Bratislava Slovakia Department of Botany and Zoology Faculty of Science Masaryk University Kotlářská 2 611 37 Brno Czechia Unit for Environmental Sciences and Management North West University Potchefstroom 2520 South Africa

Department of Zoology Faculty of Science Pavol Jozef Šafárik University in Košice Šrobárova 2 040 01 Košice Slovakia

Institute of Parasitology Slovak Academy of Sciences 04001 Košice Slovakia

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Alali F, Hussein M, Jawad M. 2024. Molecular survey on the parasitic nematode Contracaecum spp. in fish (Planiliza abu) destined for human consumption. Veterinarski Glasnik, 78, 142–154.

Andrew RL, Ostevik KL, Ebert DP, Rieseberg LH. 2012. Adaptation with gene flow across the landscape in a dune sunflower. Molecular Ecology, 21, 2078–2091. PubMed

Archie EA, Ezenwa VO. 2011. Population genetic structure and history of a generalist parasite infecting multiple sympatric host species. International Journal for Parasitology, 41, 89–98. PubMed

Attir B, Mammeri A, Kechroud N, Mazouz K, Chabira D, Chenchouni H. 2024. A survey of nematodes in the European hake (Merluccius merluccius) intended for human consumption. Journal of Parasitic Diseases, 48, 347–357. PubMed PMC

Baker MR. 1981. On three Oswaldocruzia spp. (Trichostrongyloidea: Molineidae) in amphibians from Africa. Canadian Journal of Zoology, 59, 246–251.

Barton DP. 1994. A checklist of helminth parasites of Australian amphibia. Records of the South Australian Museum, 27, 13–30.

Belaich MN, Buldain D, Ghiringhelli PD, Hyman B, Micieli MV, Achinelly MF. 2015. Nucleotide sequence differentiation of Argentine isolates of the mosquito parasitic nematode Strelkovimermis spiculatus (Nematoda: Mermithidae). Journal of Vector Ecology, 40, 415–418. PubMed

Birbele E, Di Marzio A, Grauda D, Vimercati G, Deksne G. 2024. Genetic diversity of European tree frogs (Hyla arborea group): a systematic review. European Journal of Ecology, 10(1), 1–16.

Blouin MS, Yowell CA, Courtney CH, Dame JB. Host movement and the genetic structure of populations of parasitic nematodes. Genetics, 141, 1007–1014. PubMed PMC

Borkovcová M, Kopřiva J. 2005. Parasitic helminths of reptiles (Reptilia) in South Moravia (Czech Republic). Parasitology Research, 95, 77–78. PubMed

Bowles J, Blair D, McManus DP. 1992. Genetic variants within the genus Echinococcus identified by mitochondrial DNA sequencing. Molecular and Biochemical Parasitology, 54(2), 165–173. PubMed

Buchvarov GK. 1977. Catalogue des helminthes des Amphibies en Bulgarie. Plovdiv, Bulgaria: University of Plovdiv “P. Hilendarski”; (in Bulgarian: ).

Bursey CR, Goldberg SR. 2011. New species of Oswaldocruzia (Nematoda: Molineidae) and other helminths in Bolitoglossa subpalmata (Caudata: Plethodontidae) from Costa Rica. Journal of Parasitology, 97, 286–292. PubMed

Bursey CR, Goldberg SR, Vitt LJ. 2006. New species of Oswaldocruzia (Nematoda: Molineidae) in Ameiva festiva (Squamata: Teiidae) from Nicaragua. Journal of Parasitology, 92, 350–352. PubMed

Bush AO, Lafferty KD, Lotz JM, Shostak AW. 1997. Parasitology meets ecology on its own terms: Margolis et al. Revisited. Journal of Parasitology, 83, 575. PubMed

Chikhlyaev I, Ruchin A. 2014. The helminth fauna study of European common brown frog (Rana temporaria Linnaeus, 1758) in the Volga basin. Acta Parasitologica, 59, 459–471. PubMed

Chikhlyaev IV, Ruchin AB, Fayzulin AI. 2016. The helminth fauna study of European common toad in the Volga Basin. Nature Environment and Pollution Technology, 15, 1103–1109.

Chikhlyaev IV, Ruchin AB, Kirillov AA. 2020. Ecological analysis of the helminth fauna in Bufo bufo (Amphibia: Anura) from various habitats. Nature Conservation Research, 5, 1–10.

Cole R, Viney M. 2018. The population genetics of parasitic nematodes of wild animals. Parasites & Vectors, 11, 590. PubMed PMC

Crispo E, Bentzen P, Reznick DN, Kinnison MT, Hendry AP. The relative influence of natural selection and geography on gene flow in guppies. Molecular Ecology, 15, 49–62. PubMed

Dufresnes C. 2019. Amphibians of Europe, North Africa & The Middle East, A photographic guide. London, UK: Bloomsbury Publishing Plc.

Durette-Desset MC, Alves Dos Anjos L, Vrcibradic D. 2006. Three new species of the genus Oswaldocruzia Travassos, 1917 (Nematoda, Trichostrongylina, Molineoidea) parasites of Enyalius spp. (Iguanidae) from Brazil. Parasite, 13, 115–125. PubMed

Durette-Desset MC, Batcharov G, Ben Slimane B, Chabaud A. 1993. Some Oswaldocruzia (Nematoda Trichostrongyloidae) parasites of Amphibia in Bulgaria. Redescription of Oswaldocruzia bialata (Molin, 1860). Helminthologia, 30, 99–104.

Düşen S, Oğuz M. 2010. Metazoan endoparasites of three species of anurans collected from the Middle Black Sea Region of Turkey. Helminthologia, 47, 226–232.

Düşen S, Öz M. 2013. Helminth fauna of the Eurasian marsh frog, Pelophylax ridibundus (Pallas, 1771) (Anura: Ranidae), collected from Denizli Province, Inner-West Anatolia Region, Turkey. Helminthologia, 50, 57–66.

Falk BG, Perkins SL. 2013. Host specificity shapes population structure of pinworm parasites in Caribbean reptiles. Molecular Ecology, 22, 4576–4590. PubMed

Foll M, Gaggiotti O. 2006. Identifying the environmental factors that determine the genetic structure of populations. Genetics, 174, 875–891. PubMed PMC

Gómez A, Serra M, Carvalho GR, Lunt DH. 2002. Speciation in ancient cryptic species complexes: Evidence from the molecular phylogeny of Brachionus plicatilis (Rotifera). Evolution, 56, 1431–1444. PubMed

Griffin CT. 1988. The effect of constant and changing temperatures on the development of the eggs and larvae of Oswaldocruzia filiformis (Nematoda: Trichostrongyloidea). Journal of Helminthology, 62, 281–292.

Griffin CT. 1989. Oswaldocruzia filiformis (Nematoda: Trichostrongyloidea) in frogs (Rana temporaria) from three locations in Ireland. Journal of Helminthology, 63, 53–62.

Guerrero R. 2013. Two new species of Oswaldocruzia (Nematoda: Trichostrongylina: Molineoidea) parasites of the cane toad Rhinella marina (Amphibia: Anura) from Peru. Acta Parasitologica, 58, 30–36. PubMed

Günther R. 1990. Die Wasserfrösche Europas. Wittenberg Lutherstadt, Germany: A. Ziemsen Verlag; (German: ).

Gustafson KD, Newman RA, Rhen T, Tkach VV. 2018. Spatial and genetic structure of directly‐transmitted parasites reflects the distribution of their specific amphibian hosts. Population Ecology, 60, 261–273.

Hanna AA, Joy JE. 2003. Oswaldocruzia pipiens (Nematoda: Trichostrongyloidea) from the eastern American toad, Bufo americanus americanus (Salienta: Bufonidae), in West Virginia, U.S.A. Comparative Parasitology, 70, 92–96.

Hendrikx WML. 1983. Observations on the routes of infection of Oswaldocruzia filiformis (Nematoda: Trichostrongylidae) in amphibia. Zeitschrift für Parasitenkunde, 69, 119–126. PubMed

Herczeg D, Vörös J, Végvári Z, Kuzmin Y, Brooks DR. 2016. Helminth parasites of the Pelophylax esculentus Complex (Anura: Ranidae) in Hortobágy National Park (Hungary). Comparative Parasitology, 83, 36–48.

Hoffmann A, Plötner J, Pruvost NBM, Christiansen DG, Röthlisberger S, Choleva L, Mikulíček P, Cogălniceanu D, Sas‐Kovács I, Shabanov D, Morozov‐Leonov S, Reyer H. 2015. Genetic diversity and distribution patterns of diploid and polyploid hybrid water frog populations (Pelophylax esculentus complex) across Europe. Molecular Ecology, 24, 4371–4391. PubMed

Huyse T, Poulin R, Théron A. 2005. Speciation in parasites: a population genetics approach. Trends in Parasitology, 21, 469–475. PubMed

Jablonski D, Jandzik D, Mikulíček P, Džukić G, Ljubisavljević K, Tzankov N, Jelić D, Thanou E, Moravec J, Gvoždík V. 2016. Contrasting evolutionary histories of the legless lizards slow worms (Anguis) shaped by the topography of the Balkan Peninsula. BMC Evolutionary Biology, 16, 99. PubMed PMC

Jablonski D, Najbar B, Grochowalska R, Gvoždík V, Strzała T. 2017. Phylogeography and postglacial colonization of Central Europe by Anguis fragilis and Anguis colchica . Amphibia-Reptilia, 38, 562–569.

Jorge F, Roca V, Perera A, Harris DJ, Carretero MA, et al. 2011. A phylogenetic assessment of the colonisation patterns in Spauligodon atlanticus Astasio-Arbiza et al., 1987 (Nematoda: Oxyurida: Pharyngodonidae), a parasite of lizards of the genus Gallotia Boulenger: no simple answers. Systematic Parasitology, 80, 53–66. PubMed

Karakaş M. 2015. Helminth parasites of Bufo viridis, Rana ridibunda and Hyla arborea collected from the different regions of Turkey. Manas Journal of Agriculture Veterinary and Life Sciences, 5(1), 1–6.

Katoh K. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066. PubMed PMC

Kirillova NY, Kirillov AA. 2020. Morphological variability of Oswaldocruzia filiformis (Nematoda: Molineidae) in reptiles inhabiting the protected areas of the Republic of Mordovia (Russia). IOP Conference Series: Earth and Environmental Science, 607, 012007.

Kirillova NY, Kirillov AA, Chikhlyaev IV. 2021. Morphological variability of Oswaldocruzia filiformis (Nematoda: Molineidae) in amphibians from European Russia. IOP Conference Series: Earth and Environmental Science, 818.

Kirillova NY, Kirillov AA, Shchenkov SV, Chikhlyaev IV. 2020. Oswaldocruzia filiformis sensu lato (Nematoda: Molineidae) from amphibians and reptiles in European Russia: Morphological and molecular data. Nature Conservation Research, 5, 41–56.

Kirillova NY, Kirillov AA, Shchenkov SV, Chikhlyaev IV. 2023. Oswaldocruzia ukrainae (Nematoda: Molineidae) – A parasite of European green toad Bufotes viridis: morphological and molecular data. Biology, 12, 772. PubMed PMC

Koprivnikar J, Marcogliese DJ, Rohr JR, Orlofske SA, Raffel TR, Johnson PTJ. 2012. Macroparasite infections of amphibians: What can they tell us? EcoHealth, 9, 342–360. PubMed

Kudlai O, Rakauskas V, Baker NJ, Pantoja C, Lisitsyna O, Binkienė R. 2024. Helminth parasites of invasive freshwater fish in Lithuania. Animals, 14, 3293. PubMed PMC

Kuzmin Y, Dmytrieva I, Marushchak O, Morozov-Leonov S, Oskyrko O, Nekrasova O. 2020. Helminth species and infracommunities in frogs Pelophylax ridibundus and P. esculentus (Amphibia: Ranidae) in northern Ukraine. Acta Parasitologica, 65, 341–353. PubMed

Larrat YM, de Vasconcelos Melo FT, Furo Gomes TF, Wilkens Y, Nascimento dos Santos J. 2018. Oswaldocruzia lanfrediae n. sp. (Strongylida: Molineidae), a parasite of Leptodactylus paraensis Heyer (Anura: Leptodactylidae) in Brazil. Systematic Parasitology, 95, 871–879. PubMed

Lee C-R, Mitchell-Olds T. 2011. Quantifying effects of environmental and geographical factors on patterns of genetic differentiation. Molecular Ecology, 20, 4631–4642. PubMed PMC

Leigh JW, Bryant D. 2015. Popart: full‐feature software for haplotype network construction. Methods in Ecology and Evolution, 6, 1110–1116.

Li WX, Wang GT, Nie P. 2008. Genetic variation of fish parasite populations in historically connected habitats: Undetected habitat fragmentation effect on populations of the nematode Procamallanus fulvidraconis in the catfish Pelteobagrus fulvidraco . Journal of Parasitology, 94, 643–647. PubMed

Librado P, Rozas J. 2009. DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452. PubMed

Mashaii N, Balouch M, Mobedi I. 2008. A report about helminth parasites of some amphibians (Anura: Ranidae, Bufonidae) from the north and northeast of Iran. Journal of Science (University of Tehran), 33, 9–13.

Mazé-Guilmo E, Blanchet S, Rey O, Canto N, Loot G. 2016. Local adaptation drives thermal tolerance among parasite populations: a common garden experiment. Proceedings of the Royal Society B: Biological Sciences, 283, 20160587. PubMed PMC

Mendoza-Roldan JA, Perles L, Filippi E, Szafranski N, Montinaro G, Carbonara M, Scalera R, de Abreu Teles PP, Walochnik J, Otranto D. 2024. Parasites and microorganisms associated with the snakes collected for the “Festa dei serpari” in Cocullo, Italy. PLoS Neglected Tropical Diseases, 18, e0011973. PubMed PMC

Mikulíček P, Mešková M, Cyprich M, Jablonski D, Papežík P, Hamidi D, Pekşen ÇA, Vörös J, Herczeg D, Benovics M. 2021. Weak population‐genetic structure of a widely distributed nematode parasite of frogs in the western Palearctic. Journal of Zoological Systematics and Evolutionary Research, 59, 1689–1702.

Nadler SA. 1995. Microevolution and the genetic structure of parasite populations. Journal of Parasitology, 81, 395. PubMed

Nieberding C, Libois R, Douady CJ, Morand S, Michaux JR. 2005. Phylogeography of a nematode (Heligmosomoides polygyrus) in the western Palearctic region: persistence of northern cryptic populations during ice ages? Molecular Ecology, 14, 765–779. PubMed

Nosil P, Egan SP, Funk DJ. 2008. Heterogeneous genomic differentiation between walking-stick ecotypes: “Isolation by adaptation” and multiple roles for divergent selection. Evolution, 62, 316–336. PubMed

Nosil P, Vines TH, Funk DJ. 2005. Reproductive isolation caused by natural selection against immigrants from divergent habitats. Evolution, 59, 705–719. PubMed

Novokhatskaya O. 2008. On the occurrence of the nematode Oswaldocruzia filiformis (Strongylida: Molineidae) in Karelia. Parazitologiia, 204–209 (in Russian). PubMed

Oksanen J, Simpson GL, Blanchet F, Kindt R, Legendre P, Minchin PR, O’Hara RB, Solymos P, Stevens MHH, Szoecs E, Wagner H, Barbour M, Bedward M, Bolker B, Borcard D, Carvalho G, Chirico M, Caceres De M, Durand S, Evangelista HBA, Fitz John R, Friendly M, Furneaux B, Hannigan G, Hill MO, Lahti L, McGlinn D, Ouellette M-H, Cunha ER, Smith T, Stier A, Ter Braak CJF, Weedon J. 2024. vegan: Community Ecology Package. R package version 2.7-0. Available at https://CRAN.R-project.org/package=vegan.

Okulewicz A. 1976. Oswaldocruzia filiformis (Goeze, 1782) Travassos 1917 – Nowy pasożyt jaszczurki zwinki (Lacerta agilis L.) Wiadomości Parazytologiczne, 22, 297–301. PubMed

Papežík P, Kubala M, Jablonski D, Doležálková-Kaštánková M, Choleva L, Benovics M, Mikulíček P. 2021. Morphological differentiation of endemic water frogs (Ranidae: Pelophylax) from the southwestern Balkans. Salamandra, 57, 105–123.

Papežík P, Mikulíček P, Benovics M, Balogová M, Choleva L, Doležálková-Kaštánková M, Lymberakis P, Mizsei E, Papežíková S, Poulakakis N, Saçdanaku E, Szabolcs M, Šanda R, Uhrin M, Vukić J, Jablonski D. 2023. Comparative mitochondrial phylogeography of water frogs (Ranidae: Pelophylax spp.) from the southwestern Balkans. Vertebrate Zoology, 73, 525–544.

Paradis E, Schliep K. 2019. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics, 35, 526–528. PubMed

Patterson JEH, Ruckstuhl KE. 2013. Parasite infection and host group size: A meta-analytical review. Parasitology, 140, 803–813. PubMed PMC

Plötner J. 2005. Die westpaläarktischen Wasserfrösche. Bielefeld, DE: Laurenti Verlag.

Plötner J, Uzzell T, Beerli P, Spolsky C, Ohst T, Litvinchuk SN, Guex G-D, Reyer H-U, Hotz H. 2008. Widespread unidirectional transfer of mitochondrial DNA: a case in western Palaearctic water frogs. Journal of Evolutionary Biology, 21, 668–681. PubMed PMC

Poulin R. 2005. Relative infection levels and taxonomic distances among the host species used by a parasite: Insights into parasite specialization. Parasitology, 130, 109–115. PubMed

R Core Team. 2022. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. https://www.R-Project.Org/.

Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904. PubMed PMC

Räsänen K, Hendry AP. 2008. Disentangling interactions between adaptive divergence and gene flow when ecology drives diversification. Ecology Letters, 11, 624–636. PubMed

Reiczigel J, Marozzi M, Fábián I, Rózsa L. 2019. Biostatistics for parasitologists – A primer to quantitative parasitology. Trends in Parasitology, 35, 277–281. PubMed

Ringbauer H, Kolesnikov A, Field DL, Barton NH. 2018. Estimating barriers to gene flow from distorted isolation-by-distance patterns. Genetics, 208, 1231–1245. PubMed PMC

Ronquist F, Teslenko M, Mark Pvander, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542. PubMed PMC

Rousset F. 1997. Genetic differentiation and estimation of gene flow from F-statistics under isolation by distance. Genetics, 145, 1219–1228. PubMed PMC

Rózsa L, Reiczigel J, Majoros G. 2000. Quantifying parasites in samples of hosts. Journal of Parasitology, 86, 228–232. PubMed

Ruiz-González MX, Bryden J, Moret Y, Reber-Funk C, Schmid-Hempel P, Brown MJF. 2012. Dynamic transmission, host quality, and population structure in a multihost parasite of bumblebees. Evolution, 66, 3053–3066. PubMed

Rujkorakarn P, Suvannachart P, Patamatamkul S, Thanchomnang T, Pramual P, Saijuntha W, Maleewong W, Uni S. 2024. Morphologic and molecular identification of human ocular infection caused by Pelecitus Nematodes, Thailand. Emerging Infectious Diseases, 30, 1763–1769. PubMed PMC

Sakka H, Henttonen H, Baraket G, Amel S-H, Michaux J. 2015. Phylogeography analysis and molecular evolution patterns of the nematode parasite Heligmosomum mixtum based on mitochondrial DNA sequences. Acta Parasitologica, 60, 85–98. PubMed

Sanchis V, Roig JM, Carretero MA, Roca V, Llorente GA. 2000. Host-parasite relationships of Zootoca vivipara (Sauria: Lacertidae) in the Pyrenees (North Spain). Folia Parasitologica, 47, 118–122. PubMed

Santos JN, Giese EG, Maldonado A, Lanfredi RM. 2008. A new species of Oswaldocruzia (Molineidae: Nematoda) in Chaunus marinus (Amphibian: Bufonidae) (Linneaus, 1758) from Brazil. Journal of Parasitology, 94, 264–268. PubMed

Sattmann VH. 1984. Über die Helminthenfauna von Triturus alpestris Laurenti 1768 und Rana temporaria L. aus Almtümpeln in Oberösterreich (Amphibia, Plathelminthes und Nemathelminthes). Annalen des Naturhistorischen Museums in Wien, 87, 193–196.

Shaw CL, Bilich R, Duffy MA. 2024. A common multi-host parasite shows genetic structuring at the host species and population levels. Parasitology, 151, 557–566. PubMed PMC

Shaw CL, Bilich R, O’Brien B, Cáceres CE, Hall SR, James TY, Duffy MA. 2021. Genotypic variation in an ecologically important parasite is associated with host species, lake and spore size. Parasitology, 148, 1303–1312. PubMed PMC

Shimalov VV, Shimalov VT. 2000. Helminth fauna of snakes (Reptilia, Serpentes) in Belorussian Polesye. Parasitology Research, 86, 340–341. PubMed

Shimalov VV, Shimalov VT, Shimalov AV. 2000. Helminth fauna of lizards (Reptilia, Sauria) in the southern part of Belarus. Parasitology Research, 86, 343. PubMed

Shimalov VV, Shimalov VT, Shimalov AV. 2001. Helminth fauna of newts in Belorussian Polesie. Parasitology Research, 87, 356. PubMed

Sillero N, Campos J, Bonardi A, Corti C, Creemers R, Crochet PA, Isailović JC, Denoël M, Ficetola GF, Gonçalves J, Kuzmin S, Lymberakis P, Pous P De, Rodríguez A, Sindaco R, Speybroeck J, Toxopeus B, Vieites DR, Vences M. 2014. Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia Reptilia, 35, 1–31.

Sinsch U, Heneberg P, Těšínský M, Balczun C, Scheid P. 2018. Helminth endoparasites of the smooth newt Lissotriton vulgaris: linking morphological identification and molecular data. Journal of Helminthology, 93, 332–341. PubMed

Slatkin M. 1993. Isolation by distance in equilibrium and non-equilibrium populations. Evolution, 47, 264–279. PubMed

Slimane BB, Durette-Desset M-C. 1996. Four new species of Oswaldocruzia (Nematoda: Trichostrongylina, Molineoidea) parasitizing amphibians and lizards from Ecuador. Memórias do Instituto Oswaldo Cruz, 91, 317–328.

Slimane BB, Durette-Desset M-C, Chabaud AG. 1993. Oswaldocruzia (Trichostrongyloidea) parasites d’amphibiens des collections du Muséum de Paris. Annales de Parasitologie Humaine et Comparée, 68, 88–100.

Stamatakis A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690. PubMed

Stamatakis A. 2014. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313. PubMed PMC

Svitin R. 2017. Two new species of Oswaldocruzia (Nematoda, Molineidae) parasitising lizards in Ukraine. Zootaxa, 4263, 358–368. PubMed

Thorpe RS, Surget-Groba Y, Johansson H. 2008. The relative importance of ecology and geographic isolation for speciation in anoles. Philosophical Transactions of the Royal Society B: Biological Sciences, 363, 3071–3081. PubMed PMC

Varella K, Vilela RD, Gentile R, dos Santos Cardoso T, da Costa-Neto SF, Júnior AM. 2022. Population genetic structure and phenotypic diversity of Aspidodera raillieti (Nematoda: Heterakoidea), a parasite of Didelphini marsupials in Brazil’s South and Southeast Atlantic Forest. Parasites & Vectors, 15, 203. PubMed PMC

Vieira FM, Pereira FB, Ribeiro LB, Oliveira JB, Silva DCN, Muniz-Pereira LC, Felix-Nascimento G. 2023. Oswaldocruzia franciscoensis n. sp. (Nematoda: Molineidae) in Leptodactylus macrosternum Miranda-Ribeiro, 1926 (Anura: Leptodactylidae) from Caatinga morphoclimatic domain, Brazil: morphological and molecular characterisation. Journal of Helminthology, 97, e104. PubMed

Vojtková L. 1961. K poznání Helmintofauny žab v okolí Komárna. Biologia, 1, 25–30.

Vojtková L. 1963. Zur Kenntnis der Helminthenfauna der Schwanzlurchen (Urodela) der Tschechoslowakei. Institut Für Zoologie Der Landwirtschaftlichen Hochschule in Brünn, 27(1), 20–30.

Vojtková L. 1976. Nematoda (Hlístice) obojživelníku ČSSR. Folia, Facultatis Scientiarum Naturaluim Universitatis Purkynianae Brunensis. Biologia, 55, 5–77.

Vojtková L, Moravec F, Křivanec K. 1972. Příspěvek k poznáni larválních stadií hlístic (Nematoda) z obojživelníků v ČSSR. Facultatis Scientiarum Naturalium Universitatis Purkynianae Brunensis. Biologia, 36, 79–92.

Wang IJ, Bradburd GS. 2014. Isolation by environment. Molecular Ecology, 23, 5649–5662. PubMed

Wang IJ, Summers K. 2010. Genetic structure is correlated with phenotypic divergence rather than geographic isolation in the highly polymorphic strawberry poison-dart frog. Molecular Ecology, 19, 447–458. PubMed

Waser LE, Schweizer M, Schmidt BR, Hertwig ST. 2015. Phylogeography of the common toad (Bufo bufo, Lissamphibia: Anura) in Switzerland. Amphibia Reptilia, 36, 425–436.

Weinstein SB, Lafferty KD. 2015. How do humans affect wildlife nematodes? Trends in Parasitology, 31, 222–227. PubMed

Wickham H. 2016. ggplot2: Elegant graphics for data analysis. New York: Springer-Verlag. ISBN 978-3-319-24277-4, https://ggplot2.tidyverse.org.

Willkens Y, Furtado AP, dos Santos JN, Melo FT de V. 2021. Do host habitat use and cospeciation matter in the evolution of Oswaldocruzia (Nematoda, Molineidae) from neotropical amphibians? Journal of Helminthology, 95, e33. PubMed

Willkens Y, Maldonado A, dos Santos JN, Maschio GF, Vasconcelos Melo FT de V. 2016. Redescription of Oswaldocruzia chambrieri (Strongylida: Molineidae) from Rhinella margaritifera (Anura: Bufonidae) in Caxiuanã National Forest, Brazil. Acta Parasitologica, 61, 567–575. PubMed

Wright S. 1943. Isolation by distance. Genetics, 28, 114–138. PubMed PMC

Yildirimhan HS. 1996. Reseaches on parasitic helminths of Bufo viridis Laurenti, 1768 in Turkey. Turkish Journal of Zoology, 23, 177–195.

Yildirimhan HS. 2008. An investigation of the helminth fauna of Triturus vittatus (Jenyns, 1835) and Triturus karelinii (Strauch, 1870). Türkiye Parazitoloji Dergisi, 32, 158–160. PubMed

Yildirimhan HS, Bursey CR, Altunel FN. 2011. Helminth parasites of the Balkan green lizard, Lacerta trilineata Bedriaga 1886, from Bursa, Turkey. Turkish Journal of Zoology, 35(4), 519–535.

Yildirimhan HS, Bursey CR, Goldberg SR. 2006. Helminth parasites of the Taurus Frog, Rana holtzi, and the Uludag Frog, Rana macrocnemis, with remarks on the helminth community of Turkish anurans. Comparative Parasitology, 73, 237–248.

Yildirimhan HS, Bursey CR, Goldberg SR. 2009. Helminth parasites of the Caucasian parsley frog, Pelodytes caucasicus, from Turkey. Comparative Parasitology, 76, 247–257.

Zulfikar Z, Umar S, Freasyi T, Tafsin M, Sidabukke S. 2024. Environmental factors influence the infestation of gastrointestinal nematodes in cattle in Aceh Province. Jurnal Agripet, 24, 29–35.