identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
AC6D87D2FFFE157EFF05FAC67F27FDD4.text	AC6D87D2FFFE157EFF05FAC67F27FDD4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhinella	<div><p>SYSTEMATICS OF RHINELLA</p> <p>For decades, all South American true toads were part of the formerly large and poorly defined genus Bufo, which included a heterogeneous group of toads distributed throughout Africa, America, and Eurasia (e.g., Blair, 1972; Graybeal, 1997). Frost et al. (2006) partitioned this polyphyletic genus into monophyletic units mostly on the basis of the results of their phylogenetic analysis but also on the results of previous studies (e.g., Graybeal, 1997; Pauly et al., 2004). Frost et al. (2006) resurrected Rhinella for the species of the former Bufo margaritifer Group, which they recovered as distantly related to the other species of South American true toads included in their analysis, including Chaunus and Rhaebo (both also resurrected by Frost et al., 2006). Frost et al. (2006) noted that Bufo margaritifer was nested within Chaunus in a previous phylogenetic study (Pauly et al., 2004), a finding that was subsequently supported by Pramuk (2006) and Chaparro et al. (2007). Therefore, Rhinella was later redefined to include the species of Chaunus and Rhamphophryne as well (Chaparro et al., 2007).</p> <p>The species groups of the former Bufo now referred to Rhinella were all recognized primarily on the basis of osteological characters and external morphology that were interpreted without quantitative phylogenetic analyses (Tihen, 1962; Cei, 1972a; R.F. Martin, 1972a, 1972b; Duellman and Schulte, 1992), including the R. crucifer, R. granulosa, R. margaritifera, R. marina, R. spinulosa, and R. veraguensis Groups. Pramuk (2006) studied the phylogenetic relationships of these toads on the basis of a combined analysis of morphological (mostly osteological) and molecular evidence. She rejected the monophyly of some of these species groups (e.g., the R. veraguensis Group is polyphyletic with respect to R. ocellata, the R. margaritifera Group, and Rhamphophryne), but did not modify their composition or diagnosis.</p> <p>The subsequent increase in the knowledge of relations within Rhinella was limited to the addition of available sequences of some species in extensive phylogenetic analyses of Bufonidae or Anura (e.g., van Bocxlaer et al., 2010; Pyron and Wiens, 2011; Pyron, 2014; Jetz and Pyron, 2018). Figure 1 summarizes the main results of the more inclusive analyses of Rhinella.</p> <p>For well over a decade, the systematics of Rhinella as a whole has languished, although several efforts focusing on the relationships and taxonomy of parts of the genus have been undertaken. These include phylogenetic analyses of presumptively monophyletic species groups (i.e., the R. crucifer, R. granulosa, and R. marina Groups; Maciel et al., 2006, 2010; Thomé et al., 2010, 2012; Vallinoto et al., 2010; Pereyra et al., 2016a) or fractions of the diversity of certain groups (i.e., the R. festae and R. margaritifera Groups; Fouquet et al., 2007a; Moravec et al., 2014; Santos et al., 2015; Cusi et al., 2017; Avila et al., 2018). Most recent studies on Rhinella aimed primarily to resolve species-level taxonomic problems (e.g., Fouquet et al., 2007a; Narvaes and Rodrigues, 2009; Jansen et al., 2011; Grant and Bolívar-G., 2014; Moravec et al., 2014; Cusi et al., 2017). Consequently, more than a decade after Pramuk’s (2006) revision, species groups remain poorly defined, several species cannot be assigned to any of them, and few additional phenotypic synapomorphies have been proposed for Rhinella or its internal clades (Hoogmoed, 1986; 1990; La Marca and Mijares- Urrutia, 1996; Pramuk, 2006; Chaparro et al., 2007; Padial et al., 2009; Blotto et al., 2014; Grant and Bolívar-G., 2014; Pereyra et al., 2016a).</p> <p>Natural hybridization is common in several groups of Bufonidae, including many species of Rhinella (Blair, 1972; Feder, 1979; Haddad et al., 1990; Masta et al., 2002; Azevedo et al., 2003; Green and Parent, 2003; Yamazaki et al., 2008; Fontenot et al., 2011; Guerra et al., 2011), and mitochondrial and nuclear introgression have been corroborated in some of these clades (e.g. Green and Parent, 2003; Yamazaki et al., 2008; Fontenot et al., 2011; Dufresnes et al., 2019). Pereyra et al. (2016a) demonstrated the occurrence of hybridization events in the R. granulosa Group and unidirectional mitochondrial introgression of R. dorbignyi into R. bernardoi. A similar situation might exist between R. marina and R. diptycha, although the evidence is not conclusive (Sequeira et al., 2011; Vallinoto et al., 2017). The impact of these phenomena on the inference of phylogenetic relationships (Hennig, 1966; McDade, 1992; Posada and Crandall, 2002) could be mitigated, at least partially, if detected. A detailed evaluation of the discordance between mitochondrial and nuclear genomes together with a critical taxonomic evaluation provide an effective way to detect hybridization/introgression (Pereyra et al., 2016a).</p> <p>In this paper, we present a densely sampled phylogenetic analysis of Rhinella, including 83 of its 92 species, using molecular (four mitochondrial and five nuclear genes) and phenotypic characters (90 characters from multiple character systems). The goals of this study are to (1) perform a stringent test of the monophyly of Rhinella as well as similar tests on all its species groups, (2) identify phenotypic synapomorphies to diagnose the species groups of Rhinella, and (3) to evaluate the taxonomic status of several taxa.</p> </div>	https://treatment.plazi.org/id/AC6D87D2FFFE157EFF05FAC67F27FDD4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pereyra, Martín O.;Blotto, Boris L.;Baldo, Diego;Chaparro, Juan C.;Ron, Santiago R.;Elias-Costa, Agustín J.;Iglesias, Patricia P.;Venegas, Pablo J.;C. Thomé, Maria Tereza;Ospina-Sarria, Jhon Jairo;Maciel, Natan M.;Rada, Marco;Kolenc, Francisco;Borteiro, Claudio;Rivera-Correa, Mauricio;Rojas-Runjaic, Fernando J. M.;Moravec, Jiří;De La Riva, Ignacio;Wheeler, Ward C.;Castroviejo-Fisher, Santiago;Grant, Taran;Haddad, Célio F. B.;Faivovich, Julián	Pereyra, Martín O., Blotto, Boris L., Baldo, Diego, Chaparro, Juan C., Ron, Santiago R., Elias-Costa, Agustín J., Iglesias, Patricia P., Venegas, Pablo J., C. Thomé, Maria Tereza, Ospina-Sarria, Jhon Jairo, Maciel, Natan M., Rada, Marco, Kolenc, Francisco, Borteiro, Claudio, Rivera-Correa, Mauricio, Rojas-Runjaic, Fernando J. M., Moravec, Jiří, De La Riva, Ignacio, Wheeler, Ward C., Castroviejo-Fisher, Santiago, Grant, Taran, Haddad, Célio F. B., Faivovich, Julián (2021): Evolution In The Genus Rhinella: A Total Evidence Phylogenetic Analysis Of Neotropical True Toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History 2021 (447): 1-157, DOI: 10.1206/0003-0090.447.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-447/issue-1/0003-0090.447.1.1/Evolution-in-the-Genus-Rhinella--A-Total-Evidence-Phylogenetic/10.1206/0003-0090.447.1.1.full
AC6D87D2FFDC155CFF9FF9D57A00FC59.text	AC6D87D2FFDC155CFF9FF9D57A00FC59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhinella AND ITS	<div><p>RHINELLA AND ITS INTERNAL RELATIONSHIPS</p> <p>In the parsimony total evidence analysis, Rhinella was recovered as monophyletic (after trans- ferring R. ceratophrys to Rhaebo) and well supported (JGC and JAF = 98%). The monophyly of Rhinella was previously recovered by several phylogenetic studies that used fewer taxa (e.g., Pauly et al., 2004: fig. 4; Pramuk, 2006; Pyron and Wiens, 2011; Pyron, 2014: suppl. information “amph_shl.tre”; Portik and Papenfuss, 2015; Jetz and Pyron, 2018: suppl. information “amph_ shl_new.tre”). In contrast to all previous studies, we found that Rhinella is composed of two major, well-supported clades (figs. 11–14; see below). Our results support the R. crucifer, R. granulosa, and R. marina Groups as monophyletic. Otherwise, the R. spinulosa Group is recovered paraphyletic due to the nested position of R. gallardoi (a species unassigned to any group). The R. margaritifera Group is polyphyletic due to the position of the former R. ceratophrys nested in Rhaebo. The R. veraguensis Group is polyphyletic due to the position of several taxa (i.e., R. arborescandens, R. chavin, R. lilyrodriguezae, R. manu, R. multiverrucosa, R. nesiotes, R. tacana, and R. yanachaga) more closely related to the R. margaritifera Group, and with the monophyletic R. acrolopha Group nested within them. The ML analysis of the molecular + phenotypic datasets supported most of these results (supplementary data 4.2–4.5), and we only discuss the relevant differences between analyses. Below, we provide a revised account and comments for Rhinella and its main clades and species groups on the basis of these results.</p> </div>	https://treatment.plazi.org/id/AC6D87D2FFDC155CFF9FF9D57A00FC59	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pereyra, Martín O.;Blotto, Boris L.;Baldo, Diego;Chaparro, Juan C.;Ron, Santiago R.;Elias-Costa, Agustín J.;Iglesias, Patricia P.;Venegas, Pablo J.;C. Thomé, Maria Tereza;Ospina-Sarria, Jhon Jairo;Maciel, Natan M.;Rada, Marco;Kolenc, Francisco;Borteiro, Claudio;Rivera-Correa, Mauricio;Rojas-Runjaic, Fernando J. M.;Moravec, Jiří;De La Riva, Ignacio;Wheeler, Ward C.;Castroviejo-Fisher, Santiago;Grant, Taran;Haddad, Célio F. B.;Faivovich, Julián	Pereyra, Martín O., Blotto, Boris L., Baldo, Diego, Chaparro, Juan C., Ron, Santiago R., Elias-Costa, Agustín J., Iglesias, Patricia P., Venegas, Pablo J., C. Thomé, Maria Tereza, Ospina-Sarria, Jhon Jairo, Maciel, Natan M., Rada, Marco, Kolenc, Francisco, Borteiro, Claudio, Rivera-Correa, Mauricio, Rojas-Runjaic, Fernando J. M., Moravec, Jiří, De La Riva, Ignacio, Wheeler, Ward C., Castroviejo-Fisher, Santiago, Grant, Taran, Haddad, Célio F. B., Faivovich, Julián (2021): Evolution In The Genus Rhinella: A Total Evidence Phylogenetic Analysis Of Neotropical True Toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History 2021 (447): 1-157, DOI: 10.1206/0003-0090.447.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-447/issue-1/0003-0090.447.1.1/Evolution-in-the-Genus-Rhinella--A-Total-Evidence-Phylogenetic/10.1206/0003-0090.447.1.1.full
AC6D87D2FFD31558FF89FC287FAAF91C.text	AC6D87D2FFD31558FF89FC287FAAF91C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhinella	<div><p>RHINELLA</p> <p>DIAGNOSIS: The long third pair of external gills (char. 79.2) optimizes as the only phenotypic synapomorphy of Rhinella in all the MPTs, which reverts to short third pair of external gills, the plesiomorphic bufonid condition, in the R. granulosa Group. An unequivocal diagnosis of this genus is obscured by the large phenotypic variation within Rhinella that overlaps with the diversity of many of the related bufonid genera. Nevertheless, this genus can be diagnosed from most of the related bufonids by the combination of the following phenotypic characters: (1) nasals and frontoparietal heavily ornamented with pits, striations, and rugosities (char. 9.2); (2) presence of a row of dorsolateral tubercles on skin (char. 57.1); and (3) nucleolar organizer regions (NORs) located on interstitial position of the short arms of the chromosome pair 7 (char. 89.4).</p> <p>SISTER CLADE: The well-supported clade composed of Anaxyrus + Incilius (JGC = 96%, JAF = 97%).</p> <p>DISTRIBUTION: Mostly Neotropical, ranging from the southern United States to southern South America. Rhinella marina is a highly invasive species introduced in many countries and islands outside its native distribution (e.g., Antilles, Australia, Hawaii, Philippines, Taiwan, etc.; see Frost, 2020; IUCN, 2020).</p> <p>COMMENTS: The exclusion of the former Bufo ceratophrys renders Rhinella monophyletic. The two large clades of Rhinella were not recovered in previous phylogenetic analyses (e.g., Pramuk, 2006: fig. 4; Chaparro et al., 2007; Pramuk et al., 2008; van Bocxlaer et al., 2010: suppl. information S1; Pyron and Wiens, 2011; Pyron, 2014: supp. information “amph_shl.tre”; Portik and Papenfuss, 2015; Pereyra et al., 2016a; Jetz and Pyron, 2018: suppl. information “amph_shl_new. tre”). Vera Candioti et al. (2016) proposed the long third pair of external gills as a putative synapomorphy of Rhinella in the context of a review of embryonic morphology of Bufonidae. Our TE analysis supports this character state as synapomorphy of the genus, although the embryonic morphology of many genera of Bufonidae and species of the R. margaritifera Clade (see below) is unknown. This synapomorphy of Rhinella reverts to the plesiomorphic state (short third pair of external gills) in the R. granulosa Group.</p> <p>As a result of our TE analysis (also see ML result), we define two major clades, the Rhinella marina Clade and the R. margaritifera Clade, composed of eight species groups within Rhinella. The R. marina Clade includes (1) the R. arunco Group (new species group); (2) the R. crucifer Group; (3) the R. granulosa Group; (4) the R. marina Group; and (5) the R. spinulosa Group as redefined here. The second clade, the R. margaritifera Clade, is composed of (1) R. sternosignata, a species unassigned to any group; (2) the R. festae Group as redefined here; (3) the R. margaritifera Group as redefined here; and (4) the R. veraguensis Group as redefined here. Below, we provide diagnoses, content, and comments on the distribution and systematics of each of the newly defined major clades and all species groups of Rhinella. The clades and species group are presented in the order described above and correspond to the sequence in which they appear in the TE tree (figs. 10–14) from base to tip and top to bottom.</p> <p>THE RHINELLA MARINA CLADE (figs. 11, 12)</p> <p>DIAGNOSIS: This clade is moderately supported (JGC = 88%, JAF = 92%) and diagnosed by a phenotypic synapomorphy: larval otic capsule with poorly developed processus anterolateralis with a rounded aspect (char. 76.1), with one instance of homoplasy in Sclerophrys regularis.</p> <p>SISTER CLADE: The Rhinella margaritifera Clade (figs. 13, 14).</p> <p>CONTENTS: The Rhinella marina Clade is composed of the R. crucifer, R. granulosa, and R. marina Groups, the R. spinulosa Group as redefined here, and the R. arunco Group, a new group defined here (see below). Moreover, we found a divergent mitochondrial lineage introgressed into R. horribilis (hereafter referred to as GIM [ghost introgressed mitochondrion], see below and discussion) that does not seem to belong to any recognized extant species of Rhinella and was recovered as sister clade of the R. marina + R. crucifer Groups (see fig. 12), although with poor support (see below).</p> <p>DISTRIBUTION: The species of this clade naturally occur in all main biogeographic regions of the Neotropics.</p> <p>COMMENTS: The Rhinella marina Clade is composed of two subclades. One is poorly supported (JGC = 68%, JAF = 82%) and includes the R. arunco + R. spinulosa Groups (fig. 11). It is diagnosed by four phenotypic synapomorphies: (1) the supraorbital flange on frontoparietal does not extend laterally beyond the lateral margin of the sphenethmoid (char. 17.0, with instances of homoplasy in R. quechua and some outgroups); (2) the m. extensor digitorum on the metacarpophalangeal joint of digiti III (char. 44.1, with instances of homoplasy in Anaxyrus woodhousii [polymorphic], Rhinella hoogmoedi, R. jimi, and R. rumbolli); (3) parotoid gland round to ovoid, mostly symmetrical (char. 56.2, with instances of homoplasy in R. bergi and several species of the R. margaritifera Clade); and (4) vocal sac absent in adult males (char. 58.0, with instances of homoplasy within Rhinella and outgroups). The other subclade is well supported (JGC and JAF = 99%) and includes the R. crucifer, R. granulosa, and R. marina Groups, and the GIM (figs. 11, 12). Three phenotypic synapomorphies are recovered for this subclade: (1) occurrence of a well-developed supraorbital crest (char. 1.2, with instances of homoplasy in several bufonids); (2) occipital artery pathway completely covered with bone (char. 10.2, with instances of homoplasy in bufonids); and (3) general pattern of coloration of caudal musculature of larvae uniformly dark except an unpigmented longitudinal stripe along the inferior edge (char. 69.1, with instances of homoplasy in R. quechua, R. veraguensis, and some outgroups).</p> <p>Previous phylogenetic studies including less complete sampling of Rhinella (Pramuk, 2006; Pramuk et al., 2008; van Bocxlaer et al., 2010; Pyron and Wiens, 2011; Pyron, 2014; Portik and Papenfuss, 2015; Pereyra et al., 2016a; Jetz and Pyron, 2018) never found a sister relation between the clade composed of the R. arunco + R. spinulosa Groups and the clade composed of the R. granulosa + (R. crucifer + R. marina) Groups. Instead, these studies found the R. arunco and R. spinulosa Groups as: (1) the sister clade of the species of the R. margaritifera Clade as defined here (Pramuk, 2006), (2) as sister clade of the remaining species of Rhinella (Pramuk et al., 2008; Pereyra et al., 2016a), or (3) as successive sister clades of the remaining species of Rhinella (van Bocxlaer et al., 2010; Pyron and Wiens, 2011; Pyron, 2014; Portik and Papenfuss, 2015; Jetz and Pyron, 2018). In contrast to all these analyses, our ML analysis recovers the R. arunco Group as the sister clade of the remain- der of the R. marina Clade, whereas the R. spinulosa Group is the sister taxon of the clade composed of R. granulosa + (R. crucifer + R. marina) Groups. This last clade has always been recovered as monophyletic in previous phylogenetic analyses (Pramuk, 2006; van Bocxlaer et al., 2010; Pyron and Wiens, 2011; Pyron, 2014; Portik and Papenfuss, 2015; Pereyra et al., 2016a; Jetz and Pyron, 2018).</p> <p>THE RHINELLA ARUNCO GROUP</p> <p>DIAGNOSIS: Two phenotypic synapomorphies diagnose this strongly supported (JGC and JAF = 100%) species group: (1) jaw articulation opposite to the fenestra ovalis (char. 16.1, with instances of homoplasy in some species of the Rhinella granulosa Group, the R. margaritifera Clade, and in Nannophryne variegata); and (2) anterior edge of sacral diapophyses perpendicular to the midline axis of the vertebral column (char. 30.1, with instances of homoplasy in R. crucifer, R. quechua, R. rubescens, R. spinulosa, and R. vellardi). The presence of an insertion of the m. extensor digitorum longus on metatarsophalangeal joint of digit V (char. 40.1) and the presence of an insertion of the m. extensor digitorum on metacarpophalangeal joint of digiti V (char. 45.1) could represent two additional synapomorphies of this group or an internal clade. Moreover, species of the R. arunco Group can be distinguished from members of the other species groups of Rhinella by the following combination of character states: (1) preorbital crest weak (char. 0.1), (2) occipital artery pathway uncovered with bone (char. 10.0), (3) frontoparietal that does not extend laterally beyond the lateral margin of the sphenethmoid (char. 17.0), (4) medial ramus of the pterygoid fused with the anterolateral margin of the parasphenoid (char. 21.1), (5) m. extensor digitorum longus with an insertion on the metatarsophalangeal joint of the digit IV (char. 39.1), (6) m. extensor digitorum with an insertion on the metacarpophalangeal joint of digiti III (char. 44.1), (7) inguinal fat bodies present (char. 51.1), (8) row of dorsolateral tubercles absent (char. 57.0), (9) vocal sac absent in adult males (char. 58.0), and (10) eggs biserially disposed in strings (char. 87.1).</p> <p>SISTER CLADE: The Rhinella spinulosa Group.</p> <p>CONTENTS (3 SPECIES): Rhinella arunco (Molina, 1782), R. atacamensis (Cei, 1962), and R. rubropunctata (Guichenot, 1848).</p> <p>DISTRIBUTION: Species of the Rhinella arunco Group are distributed in Argentina and Chile: Rhinella arunco and R. atacamensis in the Atac- ama Desert region, R. rubropunctata in the Austral Temperate Forest region (Cei, 1962, 1980; Correa et al., 2013). See map 1 (available at https://doi.org/10.5531/sd.sp.46) for type localities and sampled localities.</p> <p>COMMENTS: Pramuk (2006) found the Rhinella spinulosa Group (sensu Duellman and Schulte, 1992) as nonmonophyletic and excluded the species now placed in Nannophryne (i.e., N. apolobambica, N. cophotis, N. corynetes, and N. variegata; see Frost et al., 2006; Frost, 2020). The remaining species constituted a well-supported clade in her combined (morphological and molecular) analysis, being the sister taxon to all the remaining species of Rhinella. A subsequent molecular phylogeny (Pramuk et al., 2008) considering a similar taxon sampling and mitochondrial genes, but several different nuclear genes with respect to Pramuk (2006; cxcr4 and rag1 -a vs pomc and rag1 -a), recovered this redelimited group as monophyletic with poor support. Previous and subsequent analyses with slightly increased taxon and gene sampling, however, found this group as paraphyletic with respect to all remaining species groups of Rhinella (Frost et al., 2006; van Bocxlaer et al., 2010; Pyron and Wiens, 2011; Pyron, 2014; Portik and Papenfuss, 2015; Jetz and Pyron, 2018), or as the (poorly supported) sister taxon of all other species of Rhinella (Pereyra et al., 2016a). In our TE analysis, the former R. spinulosa Group (including R. gallardoi, see below) was recovered as monophyletic but poorly supported (JGC = 68%, JAF = 82%). Moreover, the individual monophyly of its sister subclades is strongly supported (both with JGC and JAF = 100%) and can be diagnosed by phenotypic synapomorphies (see Diagnosis of both groups). Our ML analysis found the former R. spinulosa Group paraphyletic with respect to the remaining species groups of the R. marina Clade (supplementary data 4.2). Based on these observations, we restrict the R. spinulosa Group to the strongly supported clade containing most species of the former R. spinulosa Group (and including R. gallardoi), and exclude the extra- Andean species R. arunco, R. atacamensis, and R. rubropunctata that constitute another well-supported clade, herein recognized as the R. arunco Group. The southernmost distributed species R. arunco and R. rubropunctata are recovered as sister taxa, although with poor support (JGC = 25, JAF &lt;50%). The three species of this group show a high genetic differentiation in comparison to other species groups of the R. marina Clade (see tables 3–6). Natural hybridization between R. arunco and R. atacamensis was reported by Correa et al. (2012, 2013), but they did not find mitochondrial and nuclear introgression outside a narrow hybrid zone.</p> <p>THE RHINELLA SPINULOSA GROUP</p> <p>DIAGNOSIS: The following character states optimize as phenotypic synapomorphies of this strongly supported group (JGC and JAF = 100%) in our TE analysis: (1) pretympanic crest absent or indistinguible (char. 2.0, with instances of homoplasy in Rhinella arunco, R. castaneotica, R. festae, and some outgroups); (2) nasal and frontoparietal bones articulating only laterally (char. 8.1, homoplastic in R. quechua, R. rubropunctata, R. veraguensis, R. yanachaga, Rhinella sp. 14, and some outgroups); (3) lightly exostosed dermal roofing bones (char. 9.1, homoplastic in the R. festae Group, in several species of the R. marina Group, and outgroups); and (4) slightly enlarged otic ramus of squamosal, overlapping with the dorsal surface of the crista parotica (char. 11.1). In addition, species of the R. spinulosa Group can be distinguished from members of the other species groups of Rhinella by the following combination of character states: (1) occipital artery pathway not covered by bone (char. 10.0), (2) frontoparietal that does not extend laterally beyond the lateral margin of the sphenethmoid (char. 17.0), (3) medial ramus of the pterygoid fused with the anterolateral margin of the parasphenoid (char. 21.1), (4) m. extensor digitorum longus with an insertion on metatarsophalangeal joint of digiti IV (char. 39.1), (5) m. extensor digitorum with an insertion on the metacarpophalangeal joint of digiti III (char. 44.1), (6) inguinal fat bodies present (char. 51.1), (7) mul- tiserial configuration of eggs in the jelly string (char. 87.2), (8) tarsal fold present (char. 65.1), and (9) adhesive gland divided after fusion of the operculum in embryo (char. 82.2).</p> <p>SISTER CLADE: The Rhinella arunco Group.</p> <p>CONTENTS (9 SPECIES): Rhinella achalensis (Cei, 1972b), R. altiperuviana (Gallardo, 1961) new status, R. amabilis (Pramuk and Kadivar, 2003), R. gallardoi (Carrizo, 1992), R. limensis (Werner, 1901), R. papillosa (Philippi, 1902), new status, R. spinulosa (Wiegmann, 1834) [including R. arequipensis (Vellard, 1959), new synonymy, see below], R. trifolium (Tschudi, 1845) new status, and R. vellardi (Leviton and Duellman, 1978).</p> <p>DISTRIBUTION: This species group is mostly distributed in arid regions along the Andes of Argentina, Bolivia, Ecuador, Chile, and Peru, except Rhinella gallardoi that inhabits the humid subandean forest of Argentina (Vellard, 1959; Córdova, 1999; Pramuk and Kadivar, 2003; Lavilla and Cei, 2001). Rhinella achalensis and R. limensis are the only species of this group with an extra-Andean distribution in the Sierras Pampeanas Centrales in the Pampas region of Argentina and Atacama Desert of Peru respectively (Vellard, 1959; Cei, 1972b). See map 2 (available at https://doi.org/10.5531/sd.sp.46) for type localities and sampled localities.</p> <p>COMMENTS: The Rhinella spinulosa Group as redelimited here is composed of some taxa with controversial taxonomies that are discussed in the context of our results. The widespread, polytypic, and poorly defined species R. spinulosa is recovered as nonmonophyletic, with R. achalensis, R. gallardoi, and R. arequipensis nested within it (fig. 11). Based on our results and considering that “ Peru ” is the type locality of R. spinulosa (and putatively confined to southern Peru, see Vellard, 1959), we restrict the species R. spinulosa s.s. to the well-supported lineage (JGC = 95%, JAF = 97%), composed of the populations from southern Peru and northern Bolivia. The lineage containing these populations of R. spinulosa also includes the sampled specimen of R. arequipensis from Departamento Arequipa, Peru. Rhinella arequipensis was originally described as a subspecies of R. spinulosa based only on differences in coloration and density of granular formations in the dorsal tegument (Vellard, 1959). Morrison (1992, 1994), Córdova (1999), and Aguilar and Gamarra (2004) did not find morphological, osteological, karyological, or larval differences that could discriminate between R. spinulosa and R. arequipensis. According to these observations and our results, we consider Bufo spinulosus arequipensis Vellard, 1959, a junior synonym of Rhinella spinulosa (Wiegmann, 1834). Thus, the species R. spinulosa is restricted to the populations distributed mainly along the Andean Puna of Peru and adjacent Bolivia.</p> <p>Populations of Rhinella spinulosa that had been considered as R. s. trifolium were recovered as a distinct and strongly supported lineage (JGC and JAF = 98%) sister to a poorly supported clade (JGC &lt;25%, JAF &lt;50%) containing R. spinulosa s.s. and several other species of the group (see below). There are several morphological differences between R. s. trifolium and R. spinulosa s.s. Vellard (1959) pointed out the disposition of the dorsal glands (longitudinal rows in R. s. trifolium and a uniform distribution in R. s. spinulosa) and the occurrence of a middorsal vertebral line in R. s. trifolium, as the main distinguishing characters. Morrison (1992, 1994), Sinsch (1986), Haas (2002), and Pramuk and Kadivar (2003) considered R. spinulosa s.s. and R. s. trifolium (and also R. s. flavolineata) as variations of a single species (see below), although all but Haas failed to provide detailed justification. The morphological comparisons were some superficial and a detailed reevaluation of the specimens and comparisons with topotypes is needed. Córdova (1999) and Aguilar and Gamarra (2004) did not find karyological or larval differences between R. s. spinulosa and R. s. trifolium; however, these character systems are conserved in related species of Rhinella (see Tolledo and Toledo, 2010; Kolenc et al., 2013; Blotto et al., 2014). The UPDs between the specimens of R. s. trifolium and R. spinulosa s.s. are relatively high for this species group (1.11%–1.30%, see table 4). Consequently, the differences in adult morphology proposed by Vellard (1959) and their genetic divergence support the recognition of Rhinella trifolium (Tschudi, 1845) as a distinct species.</p> <p>Some populations currently assigned to Rhinella spinulosa s.l. from Jujuy (Argentina) and La Paz (Bolivia) were recovered as another distinct and strongly supported lineage (JGC and JAF = 100%; see fig. 4) with a low UPD between them (0.18%). In the intermediate area of Puna between these localities (~ 800 km) lays the type locality of R. s. altiperuviana (Challapata, Oruro, Bolivia). Gallardo (1961) described this subspecies from two adult females; the characters used to differentiate it from R. spinulosa s.l. (i.e., tubercles structure, head shape, tarsal fringe development) show considerable variation, at least, in the studied female specimens from northwestern Argentina. Thus, we tentatively assign these populations to R. s. altiperuviana. In addition to the phylogenetic position, these specimens differ in UPDs (see table 4) and adult and larval external morphology (B.L.B., D.B., M.O.P., personal obs.) from other species of the group. For these reasons, these populations should be considered as a distinctive species, R. altiperuviana (Gallardo, 1961) from the Andean Puna of Argentina and Bolivia. A detailed taxonomic revision is beyond the scope of this work but will be discussed in a subsequent contribution (B.L.B. and M.O.P., in prep.).</p> <p>Populations of Rhinella spinulosa that had been considered as R. s. papillosa are recovered as a strongly supported lineage (JGC and JAF = 100%), sister taxon of R. achalensis. Both taxa differ in UPDs (1.10 to 1.47%, see table 4), and are morphologically differentiable from R. spinulosa s.s. (B.L.B. and M.O.P., in prep.). Thus, we consider R. papillosa (Philippi, 1902), a valid species from the austral Andes of Argentina and Chile.</p> <p>Rhinella gallardoi is deeply nested within the R. spinulosa Group. In the original description, Carrizo (1992) highlighted the “broad skull” of this species over the general morphological similarity with the species of the R. spinulosa Group and assigned it to the “ Bufo veraguensis-typho-</p> </div>	https://treatment.plazi.org/id/AC6D87D2FFD31558FF89FC287FAAF91C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pereyra, Martín O.;Blotto, Boris L.;Baldo, Diego;Chaparro, Juan C.;Ron, Santiago R.;Elias-Costa, Agustín J.;Iglesias, Patricia P.;Venegas, Pablo J.;C. Thomé, Maria Tereza;Ospina-Sarria, Jhon Jairo;Maciel, Natan M.;Rada, Marco;Kolenc, Francisco;Borteiro, Claudio;Rivera-Correa, Mauricio;Rojas-Runjaic, Fernando J. M.;Moravec, Jiří;De La Riva, Ignacio;Wheeler, Ward C.;Castroviejo-Fisher, Santiago;Grant, Taran;Haddad, Célio F. B.;Faivovich, Julián	Pereyra, Martín O., Blotto, Boris L., Baldo, Diego, Chaparro, Juan C., Ron, Santiago R., Elias-Costa, Agustín J., Iglesias, Patricia P., Venegas, Pablo J., C. Thomé, Maria Tereza, Ospina-Sarria, Jhon Jairo, Maciel, Natan M., Rada, Marco, Kolenc, Francisco, Borteiro, Claudio, Rivera-Correa, Mauricio, Rojas-Runjaic, Fernando J. M., Moravec, Jiří, De La Riva, Ignacio, Wheeler, Ward C., Castroviejo-Fisher, Santiago, Grant, Taran, Haddad, Célio F. B., Faivovich, Julián (2021): Evolution In The Genus Rhinella: A Total Evidence Phylogenetic Analysis Of Neotropical True Toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History 2021 (447): 1-157, DOI: 10.1206/0003-0090.447.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-447/issue-1/0003-0090.447.1.1/Evolution-in-the-Genus-Rhinella--A-Total-Evidence-Phylogenetic/10.1206/0003-0090.447.1.1.full
AC6D87D2FFC0154FFF89FCE67FAAF91F.text	AC6D87D2FFC0154FFF89FCE67FAAF91F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhinella sternosignata (Gunther 1858)	<div><p>RHINELLA STERNOSIGNATA</p> <p>DIAGNOSIS: Rhinella sternosignata (Günther, 1858b) was recovered as the sister taxon of all other species of the R. margaritifera Clade, with poor support (JGC = 49%, JAC = 71%). Phenotypic autapomorphies are: (1) acuminate anterior margins of nasals (char. 6.1), with instances of homoplasy in Incilius coniferus and the R. margaritifera Group; (2) articulation of the zygomatic ramus of the squamosal with the maxilla (char. 15.1), with instances of homoplasy in the R. granulosa Group and Peltophryne lemur; (3) articulation between the medial ramus of the pterygoid and parasphenoid alae with a jagged suture (char. 22.1) with instances of homoplasy in R. achalensis, R. atacamensis, some species of the R. festae and R. margaritifera Groups, and in the R. marina Group; (4) parotoid gland round to ovoid mostly symmetrical (char. 56.2); (5) large size of adult males with respect to adult females (char. 66.1), with instances of homoplasy in R. yanachaga, and in several species of the R. marina Clade; and (6) unpigmented eggs (char. 88.0), with instances of homoplasy in Ansonia longidigita, Rhinella justinianoi, R. stanlaii, and in the R. festae Group.</p> <p>DISTRIBUTION: This species inhabits montane forests of the Cordillera de la Costa and the Andean Cordillera de Mérida of Venezuela (La Marca and Mijares-Urrutia, 1996; Barrio- Amorós et al., 2019). See map 5 (available at https://doi.org/10.5531/sd.sp.46) for type and sampled localities.</p> <p>SISTER CLADE: The clade composed of the Rhinella festae, R. margaritifera, and R. veraguensis Groups.</p> <p>COMMENTS: This species was tentatively associated with the Rhinella margaritifera (Cei, 1972a; Hoogmoed, 1990; Duellman and Schulte, 1992) or R. granulosa Groups (Gallardo, 1962). Pereyra et al. (2016a) rejected the inclusion of this species in any of these groups, but they could not determine its relationships rigorously due to the poor sampling of Rhinella. This species was wrongly reported for many localities outside the Cordillera de la Costa montane forests region in Venezuela as discussed by La Marca and Mijares-Urrutia (1996). Vélez-Rodríguez (1999) recorded this species in error for Colombia (see Vélez-Rodríguez, 2004b, 2005). Additionally, there are a large number of recent reports of R. sternosignata for Colombia (Acosta-Galvis et al., 2006; Romero et al., 2008; Acosta-Galvis, 2012a, 2012b). Analyzed specimens tentatively assigned to this species from the eastern slope of the Cordillera Oriental in Colombia (MAR 1314, Boyacá and MAR 1955, Caquetá) were unrelated to the specimen of R. sternosignata from Venezuela in the phylogenetic analyses, and instead, they represent an undescribed species along with other specimens of the R. margaritifera Group from Loreto, Peru, and Miranda, Venezuela (Rhinella sp. 13, see below). These results, and the absence of comprehensive comparative studies considering topotypical material of R. sternosignata, indicate that there is no evidence to consider its occurrence in Colombia.</p> <p>THE RHINELLA VERAGUENSIS GROUP DIAGNOSIS: No phenotypic synapomorphies were recovered for this strongly supported group (JGC and JAF = 100%). This is mainly due to the lack of detailed information for one of its two con-</p> </div>	https://treatment.plazi.org/id/AC6D87D2FFC0154FFF89FCE67FAAF91F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pereyra, Martín O.;Blotto, Boris L.;Baldo, Diego;Chaparro, Juan C.;Ron, Santiago R.;Elias-Costa, Agustín J.;Iglesias, Patricia P.;Venegas, Pablo J.;C. Thomé, Maria Tereza;Ospina-Sarria, Jhon Jairo;Maciel, Natan M.;Rada, Marco;Kolenc, Francisco;Borteiro, Claudio;Rivera-Correa, Mauricio;Rojas-Runjaic, Fernando J. M.;Moravec, Jiří;De La Riva, Ignacio;Wheeler, Ward C.;Castroviejo-Fisher, Santiago;Grant, Taran;Haddad, Célio F. B.;Faivovich, Julián	Pereyra, Martín O., Blotto, Boris L., Baldo, Diego, Chaparro, Juan C., Ron, Santiago R., Elias-Costa, Agustín J., Iglesias, Patricia P., Venegas, Pablo J., C. Thomé, Maria Tereza, Ospina-Sarria, Jhon Jairo, Maciel, Natan M., Rada, Marco, Kolenc, Francisco, Borteiro, Claudio, Rivera-Correa, Mauricio, Rojas-Runjaic, Fernando J. M., Moravec, Jiří, De La Riva, Ignacio, Wheeler, Ward C., Castroviejo-Fisher, Santiago, Grant, Taran, Haddad, Célio F. B., Faivovich, Julián (2021): Evolution In The Genus Rhinella: A Total Evidence Phylogenetic Analysis Of Neotropical True Toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History 2021 (447): 1-157, DOI: 10.1206/0003-0090.447.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-447/issue-1/0003-0090.447.1.1/Evolution-in-the-Genus-Rhinella--A-Total-Evidence-Phylogenetic/10.1206/0003-0090.447.1.1.full
