Hostname: page-component-6bf8c574d5-t27h7 Total loading time: 0 Render date: 2025-02-22T22:02:31.704Z Has data issue: false hasContentIssue false
  • English
  • Français

First molecular characterisation of a species of Pharyngodon Diesing, 1861 (Nematoda: Pharyngodonidae), and description of Pharyngodon ameivulum n. sp. in a lizard from Caatinga in Brazil

Published online by Cambridge University Press:  14 February 2025

A.C.S. Ferreira Open the ORCID record for A.C.S. Ferreira [Opens in a new window] ,
F.M. Vieira Open the ORCID record for F.M. Vieira [Opens in a new window] ,
M.C. Diniz Open the ORCID record for M.C. Diniz [Opens in a new window] ,
D.C.N. Silva ,
L.B. Ribeiro Open the ORCID record for L.B. Ribeiro [Opens in a new window]  and
L.C. Muniz-Pereira Open the ORCID record for L.C. Muniz-Pereira [Opens in a new window]
A.C.S. Ferreira
Affiliation:
Laboratório de Helmintos Parasitos de Vertebrados, Instituto Oswaldo Cruz, FIOCRUZ. Av. Brasil 4365, Rio de Janeiro, CEP 21040-900, Brazil Programa de Pós-graduação em Biodiversidade e Saúde (PPGBS), Instituto Oswaldo Cruz, FIOCRUZ. Av. Brasil 4365, Rio de Janeiro, CEP 21040-900, Brazil
F.M. Vieira*
Affiliation:
Universidade Federal do Vale do São Francisco (UNIVASF), Rodovia BR-407, KM 12 Lote 543 S/n Projeto de Irrigação Nilo Coelho, Petrolina, 56300-000, Brazil
M.C. Diniz
Affiliation:
Universidade Federal do Vale do São Francisco (UNIVASF), Rodovia BR-407, KM 12 Lote 543 S/n Projeto de Irrigação Nilo Coelho, Petrolina, 56300-000, Brazil
D.C.N. Silva
Affiliation:
Universidade Federal do Vale do São Francisco (UNIVASF), Rodovia BR-407, KM 12 Lote 543 S/n Projeto de Irrigação Nilo Coelho, Petrolina, 56300-000, Brazil
L.B. Ribeiro
Affiliation:
Universidade Federal do Vale do São Francisco (UNIVASF), Rodovia BR-407, KM 12 Lote 543 S/n Projeto de Irrigação Nilo Coelho, Petrolina, 56300-000, Brazil
L.C. Muniz-Pereira
Affiliation:
Laboratório de Helmintos Parasitos de Vertebrados, Instituto Oswaldo Cruz, FIOCRUZ. Av. Brasil 4365, Rio de Janeiro, CEP 21040-900, Brazil Programa de Pós-graduação em Biodiversidade e Saúde (PPGBS), Instituto Oswaldo Cruz, FIOCRUZ. Av. Brasil 4365, Rio de Janeiro, CEP 21040-900, Brazil
*
Corresponding author: F.M. Vieira; Email: [email protected]
Get access Rights & Permissions [Opens in a new window]

Abstract

This study describes a new species of Pharyngodon Diesing, 1961 (Nematoda: Pharyngodonidae) in teiid lizards Ameivula ocellifera (Spix, 1895) (Squamata: Teiidae) from a Caatinga morphoclimatic domain in the state of Pernambuco, Brazil. Pharyngodon ameivulum n. sp., like 11 other species of the genus, features males without spicules, females with truncated eggs, and no tail spines. However, the new species is distinguished by a unique set of morphological characteristics, such as males possessing three pairs of caudal papillae, with the first pair precloacal, the second adcloacal, and the third postcloacal (arrangement 1:1:1), the second pair (adcloacal) of papillae having a bifurcated distal end, and females with lateral body alae. Molecular analysis of the 18S rDNA, 28S rDNA, and 18S + 28S concatenated sequences genes reveals that P. ameivulum n. sp. clusters with representatives of Pharyngodonidae from the genera Skrjabinodon Inglis, 1968 and Spauligodon Skrjabin, Schikhobalova & Lagodovska, 1960, forming a basal clade to the clade composed of Spauligodon spp. and Skrjabinodon trimorphi Ainsworth, 1990. These are the first phylogenetic assays to include a species of Pharyngodon.

Keywords

Biodiversitynematodesphylogenytaxonomy
Type
Research Paper
Information
Journal of Helminthology , Volume 99 , 2025 , e27
Copyright
© The Author(s), 2025. Published by Cambridge University Press

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Abdel-Ghaffar, F, Varjabedian, KG, Fol, M, Talal, N, Abdel-Gaber, R and Al Quraishy, S (2020a) Morphological redescription and phylogenetic assessment of Spauligodon aspiculus (Oxyuroidea: Pharyngodonidae) infecting the white-spotted gecko, Tarentola annularis (Squamata: Gekkonidae). Microscopy Research and Technique 83, 901–919. doi: 10.1002/jemt.23484.Google ScholarPubMed
Abdel-Ghaffar, F, Varjabedian, KG, Al Quraishy, S, Abdel-Gaber, R, Fol, M and Talal, N (2020b) Morphological description and phylogenetic assessment of 28S rRNA for Thelandros chalcidiae sp. nov. from Chalcides ocellatus. Molecular Biology Reports 47, 3705–3718. doi: 10.1007/s11033-020-05412-8.CrossRefGoogle ScholarPubMed
Abdel-Ghaffar, F, Varjabedian, KG, Fol, M, Talal, N, Al Quraishy, S and Abdel-Gaber, R (2022) Pharyngodon mamillatus (Nematoda: Pharyngodonidae) in the ocellated skink Chalcides ocellatus (Squamata: Scincidae) from Egypt: Light and scanning electron microscopic studies. Microscopy Research and Technique 85, 2579–2586. doi: 10.1002/jemt.24112.CrossRefGoogle ScholarPubMed
Adamson, ML (1984) Descriptions of Pharyngodon asterostoma n. sp. and P. tiliquae Baylis, 1930 (Nematoda: Oxyuroidea) from Egernia cunninghami (Scincidae) in Australia. Systematic Parasitology 6, 39–46. doi: 10.1007/BF00752615.CrossRefGoogle Scholar
Ajala, OO (2015) The occurrence of Pharyngodon mamillatus (Linstow, 1897) (Oxyuroidea, Pharyngodonidae) in a tropical wall gecko. International Journal of Applied and Natural Sciences 4, 47–50.Google Scholar
Alho, CJR and Moura, VP (1970) Oxyurata de lagartos do Planalto Central. Ocorrência de Pharyngodon cesarpintoi Pereira, 135 (Nematoda, Subuluroidea). Atas da Sociedade Biológica do Rio de Janeiro 13, 67–69.Google Scholar
Amer, SO and Bursey, CR (2008) On the oxyurid nematode, Pharyngodon mamillatus in the skink, Novoeumeces schneideri (Lacertilia: Scincidae) from Egypt. Comparative Parasitology 75, 333–338. doi: 10.1654/4338.1.CrossRefGoogle Scholar
Anderson, RC, Chabaud, AG and Willmott, S (2009) Keys to the Nematode Parasites of Vertebrates . Archival Volume. Wallingford, UK: CABI International.CrossRefGoogle Scholar
Araújo Filho, JA, Teixeira, AAM, Teles, DA, Rocha, SM, Almeida, WO, Mesquita, DO and Lacerda, ACF (2020) Using lizards to evaluate the influence of average abundance on the variance of endoparasites in semiarid areas: Dispersion and assemblage structure. Journal of Helminthology 94, e121. doi: 10.1017/S0022149X19001147.CrossRefGoogle ScholarPubMed
Ashour, AA, Koura, EA, El-Alfy, AM and Abdel-Aal, Z (1992) On the morphology of the oxiurid nematode Pharyngodon mamillatus (Linstow, 1897) from Chalcides ocellatus from Egypt. Journal of Egyptian Society of Parasitology 22, 801–806.Google ScholarPubMed
Ávila, RW and Silva, RJ (2010) Checklist of helminths from lizards and amphisbaenians (Reptilia, Squamata) of South America. Journal of Venomous Animals and Toxins Including Tropical Diseases 16, 543–572. doi: 10.1590/S1678-91992010000400005.CrossRefGoogle Scholar
Ávila, RW, Souza, FL and Silva, RJ (2010) Helminths from seven species of lizards (Reptilia: Squamata) at the Cerrado of Mato Grosso do Sul State, Brazil. Comparative Parasitology 77, 67–71. doi: 10.1654/4414.1.Google Scholar
Baker, MR (1987) Synopsis of the Nematoda parasitic in amphibians and reptiles. Memorial University of Newfoundland Occasional Papers in Biology 11, 1–325.Google Scholar
Baylis, HA (1923) Report on a collection of parasitic nematodes, mainly from Egypt. Part II. Oxyuridae. Parasitology 15, 14–23. doi: 10.1017/S0031182000014463.CrossRefGoogle Scholar
Baylis, HA (1930) Some Heterakidae and Oxyuridae from Queensland. Annals and Magazine of Natural History 10, 354–366.CrossRefGoogle Scholar
Binh, TT, Bursey, CR and Goldberg, SR (2007) Two new species of Pharyngodonidae (Nematoda, Oxyuroidea) in Gekko ulikovskii from Vietnam. Acta Parasitologica 52, 363–367. doi: 10.2478/s11686-007-0045-9.CrossRefGoogle Scholar
Brito, SV, Corso, G, Almeida, AM, Ferreira, FS, Almeida, WO, Anjos, LA, Mesquita, DO and Vasconcellos, A (2014a) Phylogeny and micro-habitats utilized by lizards determine the composition of their endoparasites in the semiarid Caatinga of Northeast Brazil. Parasitology Research 113, 3963–3972. doi: 10.1007/s00436-014-4061-z.CrossRefGoogle ScholarPubMed
Brito, SV, Ferreira, FS, Ribeiro, SC, Anjos, LA, Almeida, WO, Mesquita, DO and Vasconcellos, A (2014b) Spatial-temporal variation of parasites in Cnemidophorus ocellifer (Teiidae) and Tropidurus hispidus and Tropidurus semitaeniatus (Tropiduridae) from Caatinga areas in northeastern Brazil. Parasitology Research 113, 1163–1169. doi: 10.1007/s00436-014-3754-7.CrossRefGoogle ScholarPubMed
Bursey, CR and Goldberg, SR (1996) Pharyngodon lepidodactylus sp. n. (Nematoda: Pharyngodonidae) from the mourning gecko, Lepidodactylus lugubris (Lacertilia: Gekkonidae), from Hawaii. Journal of the Helminthological Society of Washington 63, 51–55.Google Scholar
Bursey, CR and Goldberg, SR (1999) Pharyngodon oceanicus sp. n. (Nematoda: Pharyngodonidae) from the Oceanic gecko, Gehyra oceanica (Sauria: Gekkonidae) of the Pacific Islands. Journal of the Helminthological Society of Washington 66, 37–40.Google Scholar
Bursey, CR and Goldberg, SR (2008) Gastrointestinal helminths of three species of Dicrodon (Squamata: Teiidae) from Peru. Herpetological Bulletin 106, 4–6.Google Scholar
Bursey, CR, Goldberg, SR and Kraus, F (2008) A new species of Pharyngodon (Nematoda, Pharyngodonidae) and other helminths in Cyrtodactylus louisiadensis (Sauria, Gekkonidae) from Papua New Guinea. Acta Parasitologica 53, 41–45. doi: 10.2478/s11686-008-0005-z.CrossRefGoogle Scholar
Bush, AO, Lafferty, KD, Lotz, JM and Shostak, AW (1997) Parasitology meets ecology on its own terms: Margolis et al. revisited*. Journal of Parasitology 83, 575–583. doi: 10.7939/R3J38KV04.CrossRefGoogle Scholar
Castillo, GN, Acosta, J, Gonzales–Rivas, C and Ramallo, G (2020) Checklist of nematode parasites of reptiles from Argentina. Annals of Parasitology 66, 425–432. doi: 10.17420/ap6604.282.Google ScholarPubMed
Castillo, GN, González–Rivas, CJ and Acosta, JC (2022) Primer registro de Pharyngodon travassosi (Nematoda, Pharyngodonidae) en Teius teyou (Squamata, Teiidae) en Argentina. Arxius de Miscellània Zoològica 20, 41–46. doi: 10.32800/amz.2022.20.0041.CrossRefGoogle Scholar
Conselho Nacional de Controle de Experimentação Animal (2018) ResoluçãoNormativa CONCEA nº. 37, de 15 de fevereiro de 2018. Baixa a Diretriz da Prática de Eutanásia do Conselho Nacional de Controle de Experimentação Animal—Concea. Diário Oficial da União 1, 5. https://www.gov.br/mcti/ptbr/acompanhe-o-mcti/concea/arquivos/pdf/legislacao/resolucao-normativa-no-37-de-15-de-fevereiro-de-2018.pdf/view.Google Scholar
De Sousa, A, Jorge, F, Carretero, MA, Harris, DJ, Roca, V and Perera, A (2018) The importance of integrative approaches in nematode taxonomy: The validity of Parapharyngodon and Thelandros as distinct genera. Journal of Helminthology 93, 616–628. doi: 10.1017/S0022149X1800069X.CrossRefGoogle ScholarPubMed
Er-Rguibi, O, Harris, DJ, Aglagane, A, Laghzaoui, EM, Kimdil, L, Abbad, A, and El Mouden, EH (2022). A phylogenetic assessment of nematodes (Oxyuroidea: Pharyngodonidae) infecting Moroccan lizards. Helminthologia 17, 275–283. doi: 10.2478/helm-2022-0022.CrossRefGoogle Scholar
Falk, BG and Perkins, SL (2013) Host specificity shapes population structure of pinworm parasites in Caribbean reptiles. Molecular Ecology 22, 4576–4590. doi: 10.1111/mec.12410.CrossRefGoogle ScholarPubMed
Fenner, AL, Smales, LR and Bull, CM (2008) Pharyngodon wandillahesis n. sp. (Nematoda: Pharyngodonidae) from the Endangered Pygmy Bluetongue Lizard Tiliqua adelaidensis Peters, 1863 (Sauria: Scincidae), South Australia, Australia. Comparative Parasitology 75, 69–75. doi: 10.1654/4316.1.CrossRefGoogle Scholar
Floyd, RM, Rogers, AD, Lambshead, PJD and Smith, CR (2005) Nematode-specific PCR primers for the 18S small subunit rRNA gene. Molecular Ecology Notes 5, 611–612. doi: 10.1111/j.1471-8286.2005.01009.x.CrossRefGoogle Scholar
Folmer, O, Black, M, Hoeh, W, Lutz, R and Vrijenhoek, R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3, 294–299.Google ScholarPubMed
Freitas, JFT and Ibáñez, N (1963) Fauna helmintológica del Perú – Notas sobre “Pharyngodonidae” Travassos, 1920 y descripción de Pharyngodon micrurus sp. n. Revista Universitaria 23–24, 3–10.Google Scholar
Guimarães, JF (1975) Contribuição ao estudo da fauna helmintológica do Estado da Bahia. Boletim do Instituto Biológico da Bahia 14, 44–47.Google Scholar
Hall, TA (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41, 95–98.Google Scholar
Harwood, PD (1932) The helminth parasitic in the Amphibia and Reptilia of Houston, Texas, and vicinity. Proceedings of the United States National Museum 81, 1–71. doi: 10.5479/si.00963801.81-2940.1.CrossRefGoogle Scholar
Hodda, M (2022) Phylum Nematoda: a classification, catalogue and index of valid genera, with a census of valid species. Zootaxa 5114, 1–289. doi: 10.11646/zootaxa.5114.1.1.CrossRefGoogle ScholarPubMed
Johnston, TH and Mawson, PM (1942) The Gallard Collection of parasitic nematodes in the Australian Museum. Records of the Australian Museum 21, 110–115.CrossRefGoogle Scholar
Jorge, F, Perera, A, Poulin, R, Roca, V and Carretero, MA (2018) Getting there and around: Host range oscillations during colonization of the Canary Islands by the parasitic nematode Spauligodon. Molecular Ecology 27, 533–549. doi: 10.1111/mec.14458.CrossRefGoogle ScholarPubMed
Jorge, F, Perera, A, Roca, V, Carretero, MA, Harris, DJ and Poulin, R (2014) Evolution of alternative male morphotypes in oxyurid nematodes: A case of convergence? Journal of Evolutionary Biology 27, 1631–1643. doi: 10.1111/jeb.12430.CrossRefGoogle ScholarPubMed
Jorge, F, Roca, V, Perera, A, Harris, DJ and Carretero, MA (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. doi: 10.1007/s11230-011-9311-1.CrossRefGoogle ScholarPubMed
Lopes, SG, Silva, LEM, Datas, EF and Almeida, WO (2007) Infecção por helmintos em três espécies de lagartos do nordeste brasileiro. Cadernos de Cultura e Ciência 1, 1–9.Google Scholar
Malysheva, SV (2016) Morphometrics and molecular analysis of Ozolaimus linstowi n. sp. (Oxyuroidea: Pharyngodonidae) from the green lizard Iguana iguana. Journal of Helminthology 90, 186–198. doi: 10.1017/S0022149X15000061.CrossRefGoogle Scholar
Mesquita, JMS, Oliveira, SS, Perez, R and Ávila, RW (2020) Helminths associated with Norops fuscoauratus (Squamata, Dactyloidae) in Highland marshes of the Brazilian semi-arid. Journal of Helminthology 94, e153. doi: 10.1017/S0022149X20000358.CrossRefGoogle Scholar
Mockett, S, Bell, T, Poulin, R and Jorge, F (2017) The diversity and evolution of nematodes (Pharyngodonidae) infecting New Zealand lizards. Parasitology 144, 680–691. doi: 10.1017/S0031182016002365.CrossRefGoogle ScholarPubMed
Moravec, F, Baruš, V and Ryšavý, B (1987) On parasitic nematodes of the families Heterakidae and Pharyngodonidae from reptiles in Egypt. Folia Parasitologica 34, 269–280.Google Scholar
Oliveira, EF, Gehara, M, São-Pedro, VA, Chen, X, Myers, EA, Burbrink, FT, Mesquita, DO, Garda, AA, Colli, GR, Rodrigues, MT, Arias, FJ, Zaher, H, Santos, RM and Costa, GC (2015) Speciation with gene flow in whiptail lizards from a Neotropical xeric biome. Molecular Ecology 24, 5957–5975. doi: 10.1111/mec.13433.CrossRefGoogle ScholarPubMed
Pereira, FB, Luque, JL and Tavares, LER (2017) Redescription of the nematode parasites of lizards: Strongyluris oscari Travassos, 1923 (Heterakidae) from Brazil and Pharyngodon mamillatus (Linstow, 1897) (Pharyngodonidae) from Egypt. Acta Parasitologica 62, 805–814. doi: 10.1515/ap-2017-0097.CrossRefGoogle ScholarPubMed
Pereira, FB, Luque, JL and Tavares, LER (2018) Integrative approach on Pharyngodonidae (Nematoda: Oxyuroidea) parasitic in reptiles: Relationship among its genera, importance of their diagnostic features, and new data on Parapharyngodon bainae. PLoS One 13, e0200494. doi: 10.1371/journal.pone.0200494.CrossRefGoogle ScholarPubMed
Pereira, C (1935) Os Oxyurata parasitos de Lacertilia do nordeste Brasileiro. Archivos do Instituto Biológico 6, 5–27.Google Scholar
Rabie, SAH, El-Din, MZ, El-Latif, A, Mohamed, NI and Al-Hissin, OFA (2014) Redescription of nematodes Pharyngodon mamillatus and Thelandros sp. from some reptiles of Qena, Egypt. International Journal of Science Research 3, 1368–1380.Google Scholar
Rambaudt, A, Drummond, AJ, Xie, D, Baele, G and Suchard, MA (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67, 901–904. doi: 10.1093/sysbio/syy032.CrossRefGoogle Scholar
Read, CP and Amrein, YU (1953) North American nematodes of the genus Pharyngodon Diesing (Oxyuridae). Journal of Parasitology 39, 365–370. doi: 10.2307/3274277.CrossRefGoogle ScholarPubMed
Rocha, CFD (1995) Nematode parasites of the Brazilian sand lizard, Liolaemus lutzae. Amphibia- Reptilia 16, 412–415. doi: 10.1163/156853895X00505.CrossRefGoogle Scholar
Ronquist, F and Huelsenbeck, J (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574. doi: 10.1093/bioinformatics/btg180.CrossRefGoogle ScholarPubMed
Silva, LAF, Manoel, PS, Uieda, VS, Ávila, RW and Silva, RJ (2019) Spatiotemporal variation in diet and its association with parasitic helminths in Ameivula pyrrhogularis (Squamata: Teiidae) from Northeast Brazil. Herpetological Conservation and Biology 14, 325–336.Google Scholar
Solera-Puertas, MA, Gonzalez-Santiago, PM, Carvajal-Gallardo, MM and Zapatero-Ramos, LM (1988) Description de Pharyngodon hierrensis n. sp. (Oxyuroidea, Pharyngodonidae) em Chalcides viridanus viridanus Steindachner, 1891 (Reptilia, Scincidae) em la isla do Hierro (Islas Canarias). Revista Iberica de Parasitologia 48, 387–392.Google Scholar
Solórzano-García, B, Falcón-Ordaz, J, Parra-Olea, G and Pérez-Ponce de León, G (2020) Batracholandros salamandrae (Oxyuroidea: Pharyngodonidae) in endemic salamanders (Amphibia: Plethodontidae) of the Trans-Mexican volcanic belt: Host range wide distribution or cryptic species complex? Journal of Parasitology 106, 633–643. doi: 10.1645/19-186.CrossRefGoogle ScholarPubMed
Specian, RD and Ubelaker, JE (1974) Two new species of Pharyngodon Diesing, 1861 (Nematoda: Oxyuridae) from lizards in west Texas. Proceedings of the Helminthological Society of Washington 41, 46–51.Google Scholar
SpeciesLink (2023) Species link network. Available at https://specieslink.net/search/ (accessed 17 July 2024).Google Scholar
Tamura, K, Stecher, G and Kumar, S (2021) MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution 38 3022–3027. doi: 10.1093/molbev/msab120.CrossRefGoogle ScholarPubMed
Teixeira, AAM, Riul, P, Brito, SV, Araujo-Filho, JA, Teles, DA, Almeida, WO and Mesquita, DO (2020) Ecological release in lizard endoparasites from the Atlantic Forest, northeast of the Neotropical Region. Parasitology 147, 491–500. doi: 10.1017/S0031182020000025.CrossRefGoogle ScholarPubMed
Vicente, JJ, Rodrigues, HO, Gomes, DC and Pinto, RM (1993) Nematoides do Brasil. Parte III: Nematoides de Répteis. Revista Brasileira de Zoologia 10, 19–168. doi: 10.1590/S0101-81751993000100003.CrossRefGoogle Scholar
Walker, MA, Bolek, MG, Zieman, EA, Langford, GJ, Reyda, FB and Jiménez, FA (2023) Molecular phylogenetic analysis of Gyrinicola batrachiensis (Nematoda: Oxyuroidea) lends support to the monophyly of the genus and the resurrection of Gyrinicolinae. Journal of Parasitology 109, 622–632. doi: 10.1645/22-108.CrossRefGoogle Scholar
Whiting, MF (2002) Mecoptera is paraphyletic: Multiple genes and phylogeny of Mecoptera and Siphonaptera. Zoologica Scripta 31, 93–104. doi: 10.1046/j.0300-3256.2001.00095.x.CrossRefGoogle Scholar
Xavier, MA, Anjos, LA, Gazêta, GS, Machado, IB, Storti-Melo, LM and Dias, EJR (2019) Influence of geographic distribution on parasite loads in teiid lizards (Squamata: Ameivula) from northeastern Brazil. Herpetologica 75, 315–322. doi: 10.1655/Herpetologica-D-19-00044.1.CrossRefGoogle Scholar
Xia, X (2017) DAMBE6: New tools for microbial genomics, phylogenetics and molecular evolution. Journal of Heredity 108, 431–437. doi: 10.1093/jhered/esx033.CrossRefGoogle ScholarPubMed