HYLINAE, Rafinesque, 1815

FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, Bulletin of the American Museum of Natural History 2005 (294), pp. 1-240 : 75-76

HYLINAE RAFINESQUE, 1815

SYNONYMS: See sections for tribes.

DIAGNOSIS: This subfamily is diagnosed by 32 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. Possible morphological synapomorphies are the tendo superficialis digiti V (manus) with an additional tendon that arises ventrally from m. palmaris longus (da Silva, 1998, as cited by Duellman, 2001).

COMMENTS: The 2 n 5 24 chromosomes may be another putative synapomorphy of this clade, but it will be necessary to better understand its distribution in the more basal members of the different tribes.

COPHOMANTINI HOFFMANN, 1878

Cophomantina Hoffmann, 1878. Type genus: Cophomantis Peters, 1870 .

DIAGNOSIS: This tribe is diagnosed by 65 transformations in nuclear and mitochondrial

proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. Possible morphological synapomorphies include a ventral oral disc, and complete marginal papillae in larvae, (these character states subsequently transform in more inclusive groups within this clade).

COMMENTS: This tribe includes the genera Aplastodiscus , Bokermannohyla new genus, Hyloscirtus , Hypsiboas , and Myersiohyla new genus.

The increase in the number of labial tooth rows is likely another synapomorphy of Cophomantini , because all known larvae of Hyloscirtus and Myersiohyla new genus colectively have a minimum of 6/7 labial tooth rows. However, at this time the minimum number of labial tooth rows that is synapomorphic for Cophomantini is ambiguous, because the tadpole is still unknown in H. kanaima .

An enlarged prepollex is present in all species of Hyloscirtus and in most species of Myersiohyla , new genus. (Unlike species of the H. aromatica group, in H. kanaima , the prepollex is not enlarged.) This characteristic could be a synapomorphy of Cophomantini as an intermediate state leading to the enlarged prepollex with a projecting spine, as proposed by Duellman et al. (1997). In order to understand whether this character state is a synapomorphy of Cophomantini , further research is required, including (1) more osteological work to define the character states involved, and (2) additional studies on the phylogenetic relationships within Myersiohyla , new genus, to understand whether the character state present in the former H. kanaima could be interpreted as a reversal.

Burton (2004) suggested that the tendo superficialis hallucis tapering from an expand­ ed corner of the aponeurosis plantaris, with fibers of the m. transversus plantae distalis originating on distal tarsal 2–3 inserting on the lateral side of the tendon, provides evidence of monophyly of a group composed of the H. albomarginata , H. albopunctata , H. boans , H. geographica , and H. pulchella groups. The lack of information on the taxonomic distribution of this character state within several terminals of Cophomantini renders its optimization ambiguous in all our most parsimonious trees. While it is clear that it is a synapomorphy of some component of Cophomantini , at this point we do not know its level of inclusiveness. (Burton points out its presence in H. phyllognatha , the only member of the H. bogotensis group that he studied.) The same point holds for the presence of an accessory tendon of the m. lumbricalis longus digiti III, which Burton (2004) considered characteristic of those same species groups.

There are other character states that were observed in exemplars of this tribe whose taxonomic distribution needs to be assessed in its most basal taxa in order to know with more precision the limits of the clade or clades they diagnose. One of these is the point of insertion of the tendon of the m. extensor brevis medius digiti IV that Faivovich (2002) found to insert in the medial proximal margin of phalanx 2 in the exemplars of this tribe that he studied ( Aplastodiscus perviridis , Hyla albopunctata , H. faber , and H. raniceps ). In other hyloids this tendon is known to insert in the anterior medial margin of metacarpal IV (Burton, 1996, 1998b; Faivovich, 2002). Subsequently, this character state was observed in other species available for studies ( H. albomarginata , H. andina , H. circumdata , H. clepsydra , H. geographica , H. granosa , H. multifasciata , and H. polytaenia ; Faivovich, personal obs.).

There are at least two other character states whose taxonomic distribution within this tribe deserve further scrutiny. The first of these is the presence in the dorsal surface of the larval oral cavity of an anteromedial loop of the prenarial wall into the prenarial arena. Wassersug (1980) described and reported it in Hyla rufitela, Spirandeli Cruz (1991) in Aplastodiscus perviridis , H. faber , H. lundii , and H. prasina , and D’Heursel and de Sá (1999) in H. geographica and H. semilineata . The second character state is the presence of one (most frequently) or more (a few species of the H. bogotensis group; Mi­ jares­Urrutia, 1992b) fleshy projections of variable shape (triangular, round, or elliptic; sometimes called papillae) in the inner margin of the nostrils of the larvae, which various authors (Kenny, 1969; Peixoto, 1981; Peixoto and Cruz, 1983; Lavilla, 1984; Mijares­Urrutia, 1992b; Wild, 1992; Ayarzagüena and Señaris, ‘‘1993’’ [1994]; de Sa´, 1995, 1996; Duellman et al., 1997; Faivovich, 2002; Gomes and Peixoto, 2002; Faivovich, personal obs.) noticed in several species: Aplastodiscus perviridis , Hyla albofrenata , H. albomarginata , H. albopunctata , H. albosignata , H. alemani , H. andina , H. aromatica , H. charazani , H. balzani , H. carvalhoi , H. circumdata , H. faber , H. fasciata , H. granosa , H. inparquesi , H. jahni , H. leucopygia , H. multifasciata , H. palaestes , H. platydactyla , H. punctata , H. raniceps , H. sibleszi , and an undescribed species of the H. aromatica group. Although not explicitly mentioned in the descriptions, this character state seems evident in illustrations of other larvae: H. goiana , H. joaquini , H. marginata , H. polytaenia , and H. pulchella (Eterovick et al., 2002; Gallardo, 1964; Garcia et al., 2001b, 2003).