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Cophylinae
	Anodonthyla boulengerii
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Amphibia
Order:	Anura
Family:	Microhylidae
Subfamily:	Cophylinae; Cope, 1889
Type genus
	Cophyla; Boettger, 1880
Genera
	See text

Cophylinae is a subfamily of microhylid frogs endemic to Madagascar. It has over 100 species in eight genera.^[1]^[2] Members of this subfamily range from minute (< 10 mm adult body size) to fairly large (> 100 mm adult body size), and they are highly ecologically diverse. DNA barcode research has revealed a significant taxonomic gap in this subfamily, and an estimated 70+ candidate species were identified.^[3] Many of these have subsequently been described, as well as numerous new discoveries (e.g. 26 species of Stumpffia described in 2017).^[4]

Genera

As of December 2019, the following genera are recognised in the subfamily Cophylinae:^[1]^[2]^[5]

Anilany Scherz, Vences, Rakotoarison, Andreone, Köhler, Glaw & Crottini, 2016 (monotypic)
Anodonthyla Müller, 1892 (12 species)
Cophyla Boettger, 1880 (21 species)
Madecassophryne Guibé, 1974 (monotypic)
Mini Scherz, Hutter, Rakotoarison, Riemann, Rödel, Ndriantsoa, Glos, Roberts, Crottini, Vences & Glaw, 2019 (3 species)
Plethodontohyla Boulenger, 1882 (11 species)
Rhombophryne Boettger, 1880 (9 species)
Stumpffia Boettger, 1881 (41 species)

Biology

Cophylines are characterized by a derived mode of larval development: whereas most microhylids have a specialized filter-feeding tadpole, cophylines have non-feeding tadpoles that develop either in tree holes, terrestrial foam nests, or terrestrial jelly nests.^[6] Most cophylines have very simple advertisement calls, consisting of single melodious notes that are repeated after regular intervals and for long periods of time, usually lasting several minutes. Correlated to the reproductive mode of the various cophyline lineages is their arboreal versus terrestrial or fossorial ecology, and apparently, multiple evolutionary shifts between arboreal and terrestrial habits have occurred in this subfamily.^[7]

Origins and Evolution

There is little doubt that the Cophylinae originated on Madagascar, as they are restricted to the island. Their affinities with other subfamilies of the diverse Microhylidae have been a matter of some debate,^[8]^[9]^[10]^[11]^[12] and only recently has a tentative consensus emerged that they are most closely related to the Scaphiophryninae, another Madagascar-endemic subfamily.^[10]^[12] Thus, two subfamily units of Microhylidae are endemic to and probably originated on the island of Madagascar. What is also clear is that the third Madagascar-endemic subfamily, Dyscophinae, is not closely related to these two subfamilies, so microhylids colonised Madagascar at least twice. It is not however clear which subfamily is most closely related to Cophylinae+Scaphiophryninae, in part because the topology of deep nodes of the microhylid evolutionary tree is not satisfactorily resolved and remains unstable.^[11]

The subfamily Cophylinae has its centre of diversity in the rainforests of northern Madagascar,^[13]^[14] and today only few species are known from more arid areas in the drier west of the island.^[15] As its sister subfamily Scaphiophryninae also has most of its centre of diversity in this area,^[13] it is probable that these subfamilies originated in northern Madagascar and radiated thence outward across the island. It remains to be seen if the description of the 70+ undescribed candidate species of Cophylinae^[3] will influence this pattern once described and incorporated into spatial analyses.

The Cophylinae are highly ecologically labile over evolutionary time: in several instances, phylogenetic relationships based on DNA sequence data clearly indicate repeated transitions from e.g. terrestrial to arboreal habits and back.^[16] This has resulted in extensive homoplasy in ecologically-related morphological traits among the frogs, which led also to taxonomic confusion when only sparse morphological data were available;^[9]^[14]^[16] a problem that has been mostly but not fully rectified by the increased availability of DNA sequence data.^[2] Among the repeated ecomorphs that have evolved is miniaturised body size. In 2019, five new species of miniaturised frogs were described, including the genus Mini, and members of the genera Rhombophryne and Anodonthyla.^[17] The relationships of these frogs within the subfamily indicate that at least four different groups within this subfamily have independently evolved to be among the smallest frogs in the world.^[17]

References

^ ^a ^b Frost, Darrel R. (2016). "Cophylinae Cope, 1889". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 10 May 2016.
^ ^a ^b ^c Scherz, Mark D.; Vences, Miguel; Rakotoarison, Andolalao; Andreone, Franco; Köhler, Jörn; Glaw, Frank; Crottini, Angelica (2016). "Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae)". Molecular Phylogenetics and Evolution. 100: 372–381. Bibcode:2016MolPE.100..372S. doi:10.1016/j.ympev.2016.04.019. PMID 27085671.
^ ^a ^b Vieites, David R.; Wollenberg, Katharina C.; Andreone, Franco; Köhler, Jörn; Glaw, Frank; Vences, Miguel (2009-05-19). "Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory". Proceedings of the National Academy of Sciences. 106 (20): 8267–8272. doi:10.1073/pnas.0810821106. ISSN 0027-8424. PMC 2688882. PMID 19416818.
^ Rakotoarison, A.; Scherz, M.D.; Glaw, F.; Köhler, J; Andreone, F.; Franzen, M.; Glos, J.; Hawlitschek, O.; Jono, T.; Mori, A.; Ndriantsoa, S.H.; Raminosoa Rasoamampionona, N.; Riemann, J.C.; Rödel, M.-O.; Rosa, G.M.; Vieites, D.R.; Crottini, A.; Vences, M. (2017). "Describing the smaller majority: Integrative fast-track taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar". Vertebrate Zoology. 67 (3): 271–398. doi:10.3897/vz.67.e31595.
^ "AmphibiaWeb". Retrieved 2019-03-28.
^ Blommers-Schlösser RMA (1975). "Observations on the larval development of some Malagasy frogs, with notes on their ecology and biology (Anura: Dyscophinae, Scaphiophryninae and Cophylinae)". Beaufortia. 24: 7–26.
^ Andreone F; Vences M; Vieites DR; Glaw F; Meyer M (2005). "Recurrent ecological adaptations revealed through a molecular analysis of the secretive cophyline frogs of Madagascar" (PDF). Molecular Phylogenetics and Evolution. 34 (2): 315–322. Bibcode:2005MolPE..34..315A. doi:10.1016/j.ympev.2004.10.013. PMID 15619444.
^ de Sá, Rafael O.; Streicher, Jeffrey W.; Sekonyela, Relebohile; Forlani, Mauricio C.; Loader, Simon P.; Greenbaum, Eli; Richards, Stephen; Haddad, Célio F. B. (2012-12-10). "Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera". BMC Evolutionary Biology. 12 (1): 241. Bibcode:2012BMCEE..12..241D. doi:10.1186/1471-2148-12-241. ISSN 1471-2148. PMC 3561245. PMID 23228209.
^ ^a ^b Parker, H. W. (1934). Monograph of the frogs of the family Microhylidae. Trustees of the British Museum, London, UK.
^ ^a ^b van der Meijden, Arie; Vences, Miguel; Hoegg, Simone; Boistel, Renaud; Channing, Alan; Meyer, Axel (2007-09-01). "Nuclear gene phylogeny of narrow-mouthed toads (Family: Microhylidae) and a discussion of competing hypotheses concerning their biogeographical origins". Molecular Phylogenetics and Evolution. 44 (3): 1017–1030. Bibcode:2007MolPE..44.1017V. doi:10.1016/j.ympev.2007.02.008. PMID 17369057.
^ ^a ^b Peloso, Pedro L.V.; Frost, Darrel R.; Richards, Stephen J.; Rodrigues, Miguel T.; Donnellan, Stephen; Matsui, Masafumi; Raxworthy, Cristopher J.; Biju, S.d.; Lemmon, Emily Moriarty (2016-04-01). "The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in narrow-mouthed frogs (Anura, Microhylidae)". Cladistics. 32 (2): 113–140. doi:10.1111/cla.12118. ISSN 1096-0031. PMID 34732021. S2CID 84925667.
^ ^a ^b Feng, Yan-Jie; Blackburn, David C.; Liang, Dan; Hillis, David M.; Wake, David B.; Cannatella, David C.; Zhang, Peng (2017-07-03). "Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary". Proceedings of the National Academy of Sciences. 114 (29): E5864–E5870. Bibcode:2017PNAS..114E5864F. doi:10.1073/pnas.1704632114. ISSN 0027-8424. PMC 5530686. PMID 28673970.
^ ^a ^b Brown, Jason L.; Sillero, Neftali; Glaw, Frank; Bora, Parfait; Vieites, David R.; Vences, Miguel (2016-01-06). "Spatial Biodiversity Patterns of Madagascar's Amphibians and Reptiles". PLOS ONE. 11 (1): e0144076. Bibcode:2016PLoSO..1144076B. doi:10.1371/journal.pone.0144076. ISSN 1932-6203. PMC 4703303. PMID 26735688.
^ ^a ^b Wollenberg, Katharina C.; Vieites, David R.; Van Der Meijden, Arie; Glaw, Frank; Cannatella, David C.; Vences, Miguel (2008-08-01). "Patterns of Endemism and Species Richness in Malagasy Cophyline Frogs Support a Key Role of Mountainous Areas for Speciation". Evolution. 62 (8): 1890–1907. doi:10.1111/j.1558-5646.2008.00420.x. ISSN 1558-5646. PMID 18485110. S2CID 205781693.
^ Andreone, Franco; Randrianirina, Jasmin E. (2008). "An unexpected Rhombophryne record at Tsingy de Bemaraha confirms the presence of cophyline frogs in western Madagascar". Zootaxa. 1812: 46–48. doi:10.11646/zootaxa.1812.1.2.
^ ^a ^b Andreone, Franco; Vences, Miguel; Vieites, David R.; Glaw, Frank; Meyer, Axel (2005-02-01). "Recurrent ecological adaptations revealed through a molecular analysis of the secretive cophyline frogs of Madagascar" (PDF). Molecular Phylogenetics and Evolution. 34 (2): 315–322. Bibcode:2005MolPE..34..315A. doi:10.1016/j.ympev.2004.10.013. PMID 15619444.
^ ^a ^b Scherz, Mark D.; Hutter, Carl R.; Rakotoarison, Andolalao; Riemann, Jana C.; Rödel, Mark-Oliver; Ndriantsoa, Serge H.; Glos, Julian; Hyde Roberts, Sam; Crottini, Angelica (2019-03-27). Crowther, Mathew S. (ed.). "Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera". PLOS ONE. 14 (3): e0213314. Bibcode:2019PLoSO..1413314S. doi:10.1371/journal.pone.0213314. ISSN 1932-6203. PMC 6436692. PMID 30917162.