Gymnophiona

Class: Amphibia > Order: Gymnophiona
224 species

Apoda Oppel, 1811, Ordn. Fam. Gatt. Rept.: 76; Oppel, 1811, Ann. Mus. Hist. Nat. Paris, 16: 409. Unavailable family-group name composed solely of caecilians.

Apoda Fischer von Waldheim, 1813, Zoognosia Tab. Synopt. Ill., Ed. 3, 1: 57. Unranked taxon (above family-group) for snakes, amphisbaenians, and caecilians.

Gymnophia Rafinesque, 1814, Specchio Sci., 2, 2: 104. Order for caecilians.

Gymnodermia Rafinesque, 1815, Analyse Nat.: 78. Heterogenous taxon. Unavailable family-group name for caecilians and amphisbaenians.

Pseudophydiens Blainville, 1816, Bull. Soc. Philomath., Paris, Ser. 3, 3: 111. Order (non-Latinized) for caecilians. Placed in synonymy of Apoda Merrem, 1820, by Gray, 1825, Ann. Philos., London, Ser. 2, 10: 217.

Apoda Merrem, 1820, Tent. Syst. Amph.: 163, 166. Order containing solely caecilians. Presumably a reranking of Apoda Oppel, 1811.

Amblyopes Goldfuss, 1820, Handb. Zool., 2: 137. Unvailable family-group name for caecilians, amphisbaenians, and typhlopid snakes.

PseudophidiiGray, 1825, Ann. Philos., London, Ser. 2, 10: 217. Latinization of Pseudophydiens Blainville, 1816.

Batrachophides Latreille, 1825, Fam. Nat. Regne Animal: 102. "Section" (above family-group) to include solely caecilians.

Gymnophides Latreille, 1825, Fam. Nat. Regne Animal: 103. Unavailable family-group name for all caecilians.

Nuda Fitzinger, 1826, Neue Class. Rept.: 35, 63. "Tribus" containing only Coecilioidea.

Georynchii Wagler, 1828, Isis von Oken, 21: 859. Heterogenous order coined to include caecilians, burrowing lizards, and some snakes.

Caeciliae Wagler, 1830, Nat. Syst. Amph.: 131. Order for caecilians.

Gymnophidia Müller, 1831, Isis von Oken, 24: 711. Order for caecilians. Presumably a reranking of Gymnophides Latreille, 1825.

Coeciliae Müller, 1831, Isis von Oken, 24: 711. Order for caecilians. Alternative name for Gymnophidia.

Batrachophides Bonaparte, 1831, Giorn. Arcad. Sci. Lett. Arti, Roma, 49: 75. Family-group name (and therefore unavailable) to contain caecilians.

Gymnophiona Müller, 1832, Z. Physiol., 4: 198. Apparent replacement name for Gymnophidia Müller, 1831, preoccupied for caecilians, although the 1831 name was not mentioned in the 1832 work. Considered to be an emendation of Gymnophia Rafinesque, 1814, by Dubois, 2004, Alytes, 22: 1-14; this was rejected by Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 356, because neither the name nor the publication of Rafinesque, 1814, were mentioned by Müller, 1832.

Coeciliae Müller, 1832, Isis von Oken, 25: 504. Order for caecilians. Given as a synonym of Gymnophiona.

Nuda Bonaparte, 1832, Saggio Dist. Metod. Animal. Vert.: 9. "Sezione" (over the level of Order) containing solely the Order Batrachophidii (caecilians).

Batrachophidii Bonaparte, 1832, Saggio Dist. Metod. Animal. Vert.: 9, 24; Bonaparte, 1840, Mem. Accad. Sci. Torino, Ser. 2, 2: 388, 394; Bonaparte, 1840, Nuovi Ann. Sci. Nat., Bologna, 4: 100; Bonaparte, 1850, Conspect. Syst. Herpetol. Amph.: 1. Order containing solely caecilians.

Pseudo-ophidiaBlainville, 1835, Nouv. Ann. Mus. Hist. Nat., Paris, 4: 282. Latinization of Pseudophydiens Blainville, 1816. Order for caecilians.

Batracophidii Bonaparte, 1838, Iconograph. Fauna Ital., 2 (Fasc. 22): unnumberd page (2nd page of Hyla viridis account). Section (above family-group) to contain caecilians. Incorrect subsequent spelling.

Batrachophidii Bonaparte, 1838, Nuovi Ann. Sci. Nat., Bologna, 1: 392. Order for caecilians and amphisbaenians.

Abranchia Hogg, 1838, Ann. Nat. Hist., London, 1: 152. Explicit Order, containing solely family Caecilidae [sic].

GymnophionaTschudi, 1838, Classif. Batr.: 25. Order for caecilians, attributed to Müller.

Caeciliae Tschudi, 1838, Classif. Batr.: 26. Order for caecilians.

Coeciliae Tschudi, 1838, Classif. Batr.: 55. Order for caecilians. Alternative spelling of Caeciliae.

Apoda Hogg, 1839, Mag. Nat. Hist., N.S., 3: 271. Order for caecilians.

Batrachopidii — Bonaparte, 1839, Mem. Soc. Sci. Nat. Neuchâtel, 2: 16. Incorrect subsequent spelling. 

Aphanobranchiata Leuckart, 1840, Froriep’s Neue Notizen, 13: 20. Unavailable family-group name proposed to contain caecilians.

Péromèles Duméril and Bibron, 1841, Erp. Gen., 6: plate opposite page 53. Suborder for caecilians.

Ophiosomes Duméril and Bibron, 1841, Erp. Gen., 6: plate opposite page 53. "Group" (between suborder and family) for caecilians. Anura Fitzinger, 1843, Syst. Rept.: 34. Name coined as a "Section" (above family, below Order) including branchiosaurus (temnospondyli) and caecilians.

Celatibranchia Hogg, 1841, Ann. Mag. Nat. Hist., Ser. 1, 7: 361. A "tribe" (in this context a taxon over the level of family-group) of Caducibranchia to contain only Caeciliadae.

Peromela Bonaparte, 1850, Conspect. Syst. Herpetol. Amph.: 1. Subclass to contain caecilians (Batrachophidii) and Batrachosaurus (Batrachosaurii).

Pseudophidia Gray, 1850, Cat. Spec. Amph. Coll. Brit. Mus., Batr. Grad.: 6. Order for caecilians.

Ophiomorpha Van der Hoeven, 1855, Handb. Dierkd., Ed. 2, 2: 460. Order for caecilians.

Peromela Van der Hoeven, 1855, Handb. Dierkd., Ed. 2, 2: 460. Alternative name for Ophiomorpha.

Ophiosoma Lichtenstein and Martens, 1856, Nomencl. Rept. Amph. Mus. Zool. Berol.: 35. Unranked taxon (between order and family, in context, although possibly a family), for caecilians.

Ophiosoma Jan, 1857, Cenni Mus. Civ. Milano: 52. Latinization of Ophiosomes Duméril and Bibron, 1841.

Anguinea Leunis, 1860, Synops. Drei Naturr., Zool., Ed. 2: 342. Unavailable family name for caecilians (parallel with "families" Caudata and Ecaudata).

Batrachopidii Duméril, 1863, Mem. Soc. Imp. Sci. Nat. Cherbourg, 9: 303. Order containing caecilians. Likely an incorrect subsequent spelling of Batrachophidii.

Celatibranchia Duméril, 1863, Mem. Soc. Imp. Sci. Nat. Cherbourg, 9: 300. "Tribu" (above the level of family in context), including only caecilians.

Peromela Duméril, 1863, Mem. Soc. Imp. Sci. Nat. Cherbourg, 9: 303. Subclass for caecilians and a fossil group, the Batrachosaurii.

Ophiomorpha Huxley, 1863, Q. J. Geol. Soc. London, 19: 68; Owen, 1866, Anat. Vert., 1: 15. Suborder of Batrachia for caecilians.

Phractamphibia Haeckel, 1866, Gen. Morphol. Organ., 2: cxxx. Subclass including caecilians (Order Peromela) and various fossil groups.

Peromela Haeckel, 1866, Gen. Morphol. Organ., 2: cxxxi. Order composed of caecilians.

Gymnophidia Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 20: 209. Order of caecilians, content not stated. Presumably a secondary use of Gymnophidia Müller, 1831.

Adélobatraciens Gouriet, 1868, Rev. Mag. Zool., Paris, Ser. 2, 20: 206. Heterogenous series to include lungfish and caecilians.

Ophiomorpha Fatio, 1872, Fauna Vert. Suisse, 3: 230. Order for caecilians.

Caeciloidea Sarasin and Sarasin, 1890, Ergebn. Naturwiss. Frosch. Ceylon 1884–1886, 2(4): 245. Suborder for caecilians and Amphiuma.

Apoda Goodrich, 1930, Stud. Struct. Develop. Vert.: xxi. Subclass and Order including caecilians.

Batrachophidia Miranda-Ribeiro, 1937, O Campo, 8: 66. Presumably an alternative spelling for Batrachophidii Bonaparte, 1832.

Apoda Tamarunov, 1964, in Orlov (ed.), Osnovy Paleontologii, 12: 163. Order for caecilians.

Epicriidei Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 152. Infraorder for their Epicriidae (Ichthyophiidae) and Uraeotyphlidae.

Caecilioidei Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 159. Suborder to contain their infraorders Siphonopidei and Caeciliidei.

Caeciliidei Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 167. Infraorder containing their Caeciliidae, Oscaeciliidae, Potamotyphlidae, and Typhlonectidae.

Rhinatrematidei Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 152. Infraorder for Rhinatrematidae and Scolecomorphidae.

Rhinatrematoidei Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 152. Suborder containing their infraorders Rhinatrematidei and Epicriidei.

Siphonopidei Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 162. Infraorder containing their families Geotrypeteidae and Siphonopidae.

Stegokrotaphia Cannatella and Hillis, 1993, Herpetol. Monogr., 7: 2. Unranked taxon, above the family-group, coined as the ancestor of all caecilians, excluding Rhinatrematidae, and all of the descendants of the nearest ancestral species.

Neocaecilia Wilkinson and Nussbaum, 2006, In Exbrayat (ed.), Reprod. Biol. Phylog. Gymnophiona: 44. Unranked taxon composed of non-rhinatrematid caecilians, a synonym of Stegokrotaphia.

Diatriata Wilkinson and Nussbaum, 2006, In Exbrayat (ed.), Reprod. Biol. Phylog. Gymnophiona: 46. An unranked taxon composing a monotypic family, Uraeotyphlidae (Uraeotyphlus) and the paraphyletic residue "Ichthyophiidae".

Teresomata Wilkinson and Nussbaum, 2006, In Exbrayat (ed.), Reprod. Biol. Phylog. Gymnophiona: 46. An unranked taxon composed of Typhlonectidae + Scolecomorphidae + "Caeciliidae" (paraphyletic residue). Restricted to caecilians excluding Ichthyophiidae, and Rhinatrematidae by San Mauro, Gower, Müller, Loader, Zardoya, Nussbaum, and Wilkinson, 2014, Mol. Phylogenet. Evol., 73: 182. 

Plesiophiona Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 114. Suborder for Rhinatrematidae. 

Pseudophiona — Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 267, suborder in Gymnophiona, the sister taxon of Plesiophiona, and the name a reformulation of spelling Pseudophydiens of Blainville, 1816.

Common Names

Caecilians (Cochran, 1961, Living Amph. World: 5).

Limbless Amphibians (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 23).

Distribution

Pantropical, except for Oceania and Australia.

Comment

The most comprehensive revision of caecilians was by Taylor, 1968, Caecilians of the World, although that work is now very out-dated. Carroll and Currie, 1972, Zool. J. Linn. Soc., 57: 229–247, suggested a lepospondyl origin of this group; see Nussbaum, 1983, J. Zool., London, 199: 545–554, for more discussion and disagreement with this view. Evans and Sigogneau-Russell, 2001, Palaeontology, 44: 259–273, discussed the fossil evidence for the origin of the group and their relationships, and suggested that the name Gymnophiona be used for the fossil antecedents as well as the crown-group (living taxa) and that Apoda be restricted to use for the living crown-group. The nomenclature of this group name was discussed by Dubois, 1984, Alytes, 3: 111–116, and Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 1–370 (the latter who coined the name Parabatrachia for the fossil and living caecilians, rejected the use of the name Apoda, and restricted the name Gymnophiona to the living crown-group taxa. Wake, 1985, Zoomorphology, 105: 277–295, reported on comparative morphology and evolution of eyes in Gymnophiona. Laurent, 1986, in Grassé and Delsol (eds.), Traite de Zool., 14: 595–608, and Lescure, Renous, and Gasc, 1986, Mem. Soc. Zool. France, 43: 145–177, erected several new families and genera, largely based on geographic reasoning. These were rejected by Nussbaum and Wilkinson, 1989, Herpetol. Monogr., 3: 1–42. Nussbaum, 1991, in Green and Sessions (eds.), Amph. Cytogenet. Evol.: 33–61, reported on karyological evidence as it bears on the evolutionary history of caecilians. Nussbaum, 1979, Occas. Pap. Mus. Zool. Univ. Michigan, 687, and Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 1–370 (115), discussed phylogeny within Gymnophiona and redelimited Caeciliidae (to include Scolecomorphidae and Typhlonectidae as subfamiles) and Ichthyophiidae (to include Uraeotyphlidae as a synonym). Roelants, Gower, Wilkinson, Loader, Biju, Guillaume, Moriau, and Bossuyt, 2007, Proc. Natl. Acad. Sci. USA, 104: 887–892, on the basis of less inclusive sampling, a moderately different molecular dataset, and a different analytical method, recovered largely congruent results. Liu, Wang, and Zhou, 2004, Zool. Res., Kunming, 25: 185–190, discussed the phylogenetic placement of Gymnophiona on the basis of tRNA study. Hass, Nussbaum, and Maxson, 1993, Herpetol. Monogr., 7: 56–63, discussed the immunological evidence for relationships among the families, and Hedges, Nussbaum, and Maxson, 1993, Herpetol. Monogr., 7: 64–76, discussed the phylogenetic evidence from mitochondrial DNA sequences and San Mauro, Gower, Oommen, Wilkinson, and Zardoya, 2004, Mol. Phylogenet. Evol., 33: 413–427, reported on phylogenetics based on a complete mitochondrial genoma and nuclear RAG1 genes. The preponderance of evidence suggests that caecilians are the sister taxon of Batrachia (salamanders + frogs); see Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 1–370, for a general review of the literature and evidence. See also Wake, 1993, Herpetol. Monogr., 7: 42–55, for discussion of other lines of phylogenetic evidence. Wilkinson, 1997, Biol. Rev. Cambridge Philos. Soc., 72: 423–470, discussed the congruence of neuroanatomical data with traditional morphological characters. Lynch, 2000 "1999", Rev. Acad. Colomb. Cienc. Exact. Fis. Nat., 23: 317–337, provided a key to the species in Colombia and briefly discussed their systematics and distribution. Wilkinson, Sheps, Oommen, and Cohen, 2002, Mol. Phylogenet. Evol., 23: 401–407, discussed previous biogeographic hypotheses and systematics of caecilians. Lescure, 1986, Mem. Soc. Zool. France, 43: 11–19, reviewed the history of classification of this group. Wilkinson, Loader, Gower, Sheps, and Cohen, 2003, Afr. J. Herpetol., 52: 83–92, provided a mtDNA tree of caecilians: Rhinatrematidae ((Ichthyophiidae + Uraeotyphlidae) ("Caeciliidae" + Typhlonectidae + Scolecomorphidae))), in other words, largely corroborating earlier work but providing evidence that Caeciliidae is paraphyletic with respect to both Typhlonectidae and the Scolecomorphidae. Wake, Parra-Olea, and Sheen, 2005, in Donnelly et al. (eds.), Ecol. Evol. Tropics: 48–64, discussed phylogeny of Central and South American caecilians and presented additional evidence for the paraphyly of Caeciliidae to Typhlonectidae. Wilkinson and Nussbaum, 2006, In Exbrayat (ed.), Reprod. Biol. Phylog. Gymnophiona: 39–78, provided a detailed discussion of the evidence for phylogeny within Gymnophiona. Wollenberg and Measey, 2009, J. Evol. Biol., 22: 1046–1056, provided additional perspective on the molecular evolution within Gymnophiona (although the use of midpoint rooting results in a novel placement of Scolecomorphinae—DRF). Zhang and Wake, 2009, Mol. Phylogenet. Evol., 53: 479–491, provided a phylogenetic analysis of caecilians, further corroborating the paraphyly of the "Caeciliidae" with respect to nominal Typhlonectidae, but found Scolecomorphinae to be the sister taxon of all other caeciliids, unlike Wilkinson, Loader, Gower, Sheps, and Cohen, 2003, Afr. J. Herpetol., 52: 83–92, who found Scolecomorphinae within Caeciliidae. Taxon sampling and data sets were similar but differed in some respects, so at this point in the time the reason(s) for the different results remain problematic. Carroll, 2009, The Rise of Amph., reviewed the paleontology of caecilians. San Mauro, 2010, Mol. Phylogenet. Evol., 56: 554–561, provided a multilocus analysis of the age and relationships of major amphibian groups. Wilkinson, San Mauro, Sherratt, and Gower, 2011, Zootaxa, 2874: 41–64, discussed the history of family-group taxonomy, and provided a revised, monophyletic taxonomy of families. Maciel and Hoogmoed, 2011, Zootaxa, 2984: 1–53, provided detail accounts, spot maps, and an identification key to the species of Brazilian Amazonia. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583 (see comment in Amphibia record) provided as part of a larger analysis a tree of Gymnophiona: Rhinatrematidae + (Ichthyophiidae + (Scolecomorphidae + (Herpelidae + ((Typhlonectidae + Caeciliidae) + (Indotyphlidae + (Dermophiidae + Siphonopidae)))))) but could not confirm that Gymnophiona is the sister taxon of Anura + Caudata. Blackburn and Wake, 2011, In Zhang (ed.), Zootaxa, 3148: 39–55, briefly reviewed the taxonomic history of this taxon. Chen, Wang, Liu, Xie, and Jiang, 2011, Curr. Zool., Chengdu, 57: 785–805, provided a phylogenetic trees based on 11 protein-coding mtDNA genes. San Mauro, Gower, Cotton, Zardoya, Wilkinson, and Massingham, 2012, Syst. Biol., 61: 661–674, confirmed the familial relationships posited by Pyron and Wiens (2011) and discussed aspects of phylogenetic analysis of the group. Maddin, Russell, and Anderson, 2012, Zool. J. Linn. Soc., 166: 160–201, discussed the evolutionary morphology of the gymnophionan braincase and provided a cladogram of combined morphological and molecular evidence for the group. Cole, Townsend, Reynolds, MacCulloch, and Lathrop, 2013, Proc. Biol. Soc. Washington, 125: 317–578, provided identification keys and accounts for the species in Guyana. Vitt and Caldwell, 2014, Herpetology, 4th Ed., provided a general summary of taxonomy and life history. Köhler, 2011, Amph. Cent. Am.: 29–35, provided keys, maps, brief accounts, and maps for the species of Central America. Sherratt, Gower, Klingenberg, and Wilkinson, 2014, Evol. Biol., 41: 528–545, reported on caecilian phylogenetics and the evolution of skull shape. San Mauro, Gower, Müller, Loader, Zardoya, Nussbaum, and Wilkinson, 2014, Mol. Phylogenet. Evol., 73: 177–189, provided a mitogenomic phylogeny of the group that confirmed their earlier results and discussed life-history evolution. Venu, 2014, Internatl. J. Adv. Res., 2: 78–83, reported on karyology of Indian species. Fei and Ye, 2016, Amph. China, 1: i–xv, 1–1040, reviewed the caecilians of China. Bardua, Wilkinson, Gower, Sherratt, and Goswami, 2019, BMC Evol. Biol., 19(30): 1–23, reported on morphological evolution and modularity in the caecilian skull. Torres-Sanchez, Gower, Alvarez-Ponce, Creevey, Wilkinson, and San Mauro, 2019, BMC Genomics, 20 (354): 1–13, reported on genomic evolution, largely those genes correlated with burrowing ability. Torres-Sanchez, Gower, Alvarez-Ponce, Creevey, Wilkinson, and San Mauro, 2019, BMC Genomics, 20 (354): 1–13, reported on the genomic underpinnings of the gymnophionan radiation. Santos, Laurin, and Zaher, 2020, Biol. J. Linn. Soc., 131: 737–755, reviewed the fossil record of caecilians. Zhang, Miao, Hu, Sun, Ding, Ji, Guo, Yan, Wang, Kan, and Nie, 2021, BMC Ecol. Evol., 21 (19): 1–14, examined mitochondrial rearrangement in amphibians in a phylogenetic context. Hime, Lemmon, Lemmon, Prendini, Brown, Thomson, Kratovil, Noonan, Pyron, Peloso, Kortyna, Keogh, Donnellan, Mueller, Raxworthy, Kunte, Das, Gaitonde, Green, Labisko, Che, and Weisrock, 2021, Syst. Biol., 70: 49–66, provided a sequence-dense tree of familial relationships and a time-tree. Dubois, Ohler, and Pyron, 2021, Megataxa, 5: 1–738, discussed the phylogenetics and nomenclature of the group. Zug, 2022, Smithson. Contrib. Zool., 653: 5, provided a key to the species of Myanmar. Barcelos and Santos, 2023, Paleodivers. Palaeoenvironm., 103: 341–405, reviewed the fossil localities and the literature of the fossil history of this group in South America. Lowie, De Kegel, Wilkinson, Measey, O'Reilly, Kley, Gaucher, Adriaens, and Herrel, 2023, J. Anat., 242: 3312–326, reported on comparative anatomy of head muscles in a phylogenetic context in order to evaluate differences due to ecology and phylogeny. Serra Silva, 2024, J. Syst. Palaeontol., 22(1: 2321620): 1–21, discussed the analytical issues and pitfalls involved in understanding the phylogenetics of fossil and living Lissamphibia. Marjanović, Maddin, Olori, and Laurin, 2024, Fossil Record, 27: 55–94, discussed Chinlestegophis (Triassic fossil) rejecting its relevance to understanding caecilian evolution. 

Contained taxa (224 sp.):

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