Boana claresignata, AND BOK. CLEPSYDRA AS

BOK. CLARESIGNATA AND BOK. CLEPSYDRA AS BOANA

Our phylogenetic analyses recover Bok. claresignata deeply nested in Boana , as the sister taxon of the Boana pulchella group, with 85% jackknife support ( Fig. 1; Supporting Information, Appendix S5). Although we included only Bok. claresignata in our analysis, the monophyly of this species and Bok. clepsydra , in what has been called the Bok.claresignata group, is supported by phenotypic evidence. For this reason, we consider that the recovered position of Bok. claresignata can be extended to Bok. clepsydra .

Our result conflicts with the tentative taxonomic placement of the former Hyla claresignata group in Bokermannohyla by Faivovich et al. (2005), but it does not imply more phenotypic character conflict than did its placement in Bokermannohyla . Although Boana is diagnosable from the other genera of Cophomantini (combination of prepollical spine, projected or not, with expanded sacral diapophyses), no phenotypic synapomorphies are known for this genus.

An expanded sacral diapophysis occurs in Aplastodiscus , Boana and Hyloscirtus , as opposed to a round or unexpanded diapophysis in Bokermannohyla , Myersiohyla and, homoplastically, in a few species deeply nested in Hyloscirtus (unknown in Nesorohyla and in the Hyloscirtus jahni group; Kizirian et al., 2003; Coloma et al., 2012; A.C.C. Lourenço, M. Rivera-Correa, J. Faivovich and P.D.P. Pinheiro, pers. obs.). The origin of the expanded sacral diapophyses optimizes ambiguously in Cophomantini , being equally parsimonious to interpret it as a synapomorphy of the common ancestor of Aplastodiscus , Boana , Bokermannohyla and Hyloscirtus , with a reversal in Bokermannohyla , or two independent origins in Hyloscirtus and the common ancestor of Aplastodiscus and Boana . The expanded diapophysis occurs also in Bok. claresignata (MNRJ 24028) and Bok. clepsydra (MZUSP 112612).

Besides the molecular evidence, there is one character state that supports the position of the former Hyla claresignata group as the sister taxon of the Boana pulchella group. This is the absence of the slip of the m. depressor mandibulae that originates at the level of the dorsal fascia of the m. levator scapulae (Supporting Information, Appendix S2: Fig. S2; see also Pinheiro et al., 2018: fig. 2). J. Faivovich and P.C.A. Garcia (in Faivovich et al., 2005) identified this character state as a synapomorphy of the Boana pulchella group. Subsequent observations indicate that in Cophomantini it is homoplastic only with the clade including Boana atlantica (Caramaschi & Velosa, 1996) , Boana cinerascens (Spix, 1824) and Boana punctata (Schneider, 1799) (P.D.P. Pinheiro, pers. obs.)

Pinheiro et al. (2018) stated that the absence of the anterolateral process of the hyoid plate is known so far only in the Boana pulchella group. An anterolateral process is also absent in Bok. clepsydra (MZUSP 112612) but present in Bok. claresignata (MNRJ 24028). The taxonomic distribution of this character state requires more study; it is still unknown in a number of species of the Boana pulchella group [ Boana aguilari ( Lehr, Faivovich & Jungfer, 2010) , Boana balzani (Boulenger, 1898) , Boana cambui ( Pinheiro, Pezzuti, Leite, Garcia, Haddad & Faivovich, 2016) , Boana melanopleura (Boulenger, 1912) and Boana palaestes ( Duellman, De la Riva & Wild, 1997) ], and the anterolateral process is present in Boana freicanecae (Carnaval & Peixoto, 2004) (P.D.P. Pinheiro, pers. obs.); therefore, its polarity is still not clear.

The hyoid plates of Bok. claresignata and Bok. clepsydra lack the posterolateral processes. The available evidence indicates that this process is absent in some species of the Boana pulchella group ( Pinheiro et al., 2018: fig 3D) and present at least in some species of the Boana albopunctata and Boana faber groups ( Pinheiro et al., 2018: fig. 3A–C). Although it could be another putative synapomorphy supporting the monophyly of the former Hyla claresignata group and the Boana pulchella group, its taxonomic distribution still requires more study.