Search

Rivera-Correa, Mauricio, Baldo, Diego, Candioti, Florencia Vera, Orrico, Victor Goyannes Dill, Blackburn, David C., Castroviejo-Fisher, Santiago, Chan, Kin Onn, Gambale, Priscilla, Gower, David J., Quah, Evan S.H., Rowley, Jodi J. L., Twomey, Evan, Vences, Miguel (2021): Amphibians in Zootaxa: 20 salamanders, and caecilians years documenting the global diversity of frogs. Zootaxa 4979 (1): 57-69, DOI: https://doi.org/10.11646/zootaxa.4979.1.9

Pseudohemisus longimanus Angel, 1930 Name-bearing type: Lectotype by present designation, MNHN 1929.237, from “Ambongo” according to the original description. The original syntype series included a second item, MNHN 1929.236, with probably non-conspecific tadpoles (see below). The type locality of this nomen, Ambongo, is located in north-western Madagascar, not far from the karstic massif of the Tsingy de Bemaraha, according to Blommers-Schl̂sser & Blanc (1991). The nomen was originally based on two syntypes. One of these, MNHN 1929.237, is a specimen in a moderately poor state of preservation that is probably an adult and is clearly assignable to the genus Scaphiophryne. The second number, MNHN 1929.236, refers to a series of about eight tadpoles in poor state of preservation, conserved in the same jar as MNHN 1929.237. Most of these tadpoles, upon superficial examination, show distinct blackish jaw sheaths and some are large; consequently, by these characters and general appearance clearly do not correspond to tadpoles of the subgenus Pseudohemisus in Scaphiophryne (Grosjean et al. 2007). For this reason, we hereby designate MNHN 1929.237 as lectotype of Pseudohemisus longimanus Angel, 1930. This stabilizes this nomen, in the event that future work on the S. calcarata complex reveals the existence of further species in addition to S. calcarata and S. obscura, for which this nomen may then be available. Morphology. For measurements of the lectotype, see Table 1. The tarsal tubercle and enlarged inner metatarsal tubercle are clearly visible (Fig. 9). Subarticular tubercles are difficult to recognize, but are not fully flattened (as remarked in the original description). On the left side of the body, a dark stripe runs from the eye backwards for a few millimeters, reminiscent of the dorsolateral fold typical for the S. calcarata complex. Osteological examination revealed close resemblance to the other Pseudohemisus specimens examined here (Fig. 10, Supplementary Figs S2–S 3). Assignment. This taxon was considered to be a synonym of S. calcarata by Guibé (1978) and Blommers-Schl̂sser & Blanc (1991). However, it is important to mention that due to the somewhat elongated head with a distinct neck, indistinct subarticular tubercles, and very large inner metatarsal tubercle, the lectotype specimen is also reminiscent of S. brevis, and some S. brevis also show a thin dorsolateral line or dorsolateral ridge in the anterior part of the body (as recognizable in the longimanus lectotype). According to Raselimanana (2008), S. brevis does occur in the Tsingy de Bemaraha area (along with populations of the S. calcarata complex). Based on its locality, it therefore cannot be ruled out that longimanus is a junior synonym of S. brevis. However, considering the probable presence of a dorsolateral fold, we here follow Guibé (1978) and Blommers-Schl̂sser & Blanc (1991) in assigning the nomen to the S. calcarata complex. Micro-CT scans of S. brevis, which were not produced for this study, might help to clarify this further. No recently collected material and no DNA sequences of the S. calcarata complex are available from the Tsingy de Bemaraha area, which is located geographically between the known distribution areas of the western and the north-western lineages of S. obscura. While the data are insufficient to assign the nomen to either of these deep conspecific lineages, based on geography we assign Pseudohemisus longimanus Angel, 1930 as junior synonym to Scaphiophryne obscura., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on page 409, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Angel, F. (1930) Description de deux especes nouvelles de batraciens de Madagascar, appartenant au genre Pseudohemisus. Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 2, 2, 70 - 74.","Grosjean, S., Glos, J., Teschke, M., Glaw, F. & Vences, M. (2007) Comparative larval morphology of Madagascan toadlets of the genus Scaphiophryne: phylogenetic and taxonomic inferences. Zoological Journal of the Linnean Society, 151, 555 - 576. https: // doi. org / 10.1111 / j. 1096 - 3642.2007.00329. x","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","Raselimanana, A. P. (2008) Herpetofaune des forets seches malgaches. Malagasy Nature, 1, 46 - 75."]}

Calophrynus calcaratus Mocquard, 1895 Name-bearing type: Lectotype by present designation, MNHN 1895.294, from “ Madagascar ” according to the original description. Originally, there were four syntype specimens under the number 95.294 (= MNHN 1895.294), of which three were re-catalogued as MNHN 1991.2862, MNHN 1991.2863 and MNHN 1991.2864 (now paralectotypes). Historically, this is the second described species assignable to the subgenus Pseudohemisus, and unfortunately only has the imprecise type locality “ Madagascar ”. The original syntypes are in an extremely poor state of preservation and probably juveniles. Due to the lack of clear morphological distinctions between the lineages within the S. calcarata complex, it is not possible to unambiguously define to which of these the name calcarata should apply on the basis of morphology or biogeography. The largest of the original syntypes is the one that originally bore the number “95.294”. SVL of the four specimens is as follows: MNHN 1895.294: 13.7 mm (Table 1); MNHN 1991.2862: 11.8 mm; MNHN 1991.2863: 9.8 mm; MNHN 1991.2864: 8.7 mm. We here designate the largest specimen, MNHN 1895.294, as lectotype to stabilize this nomen because a few more taxonomically relevant morphological characters are visible in this specimen than in the others. Morphology. Despite its small size and extremely poor state of preservation, the lectotype can easily be recognized as belonging to Scaphiophryne by the presence of an enlarged inner metatarsal tubercle and a distinct tarsal tubercle (Fig. 9). No traces of vomerine or maxillary teeth are recognizable, but the anterior region of the head is largely destroyed, so this remains inconclusive. The subarticular tubercle on the first finger is slightly longer and higher than those on the other fingers. There are no enlarged terminal disks on fingers or toes. Determining osteological characters of this specimen proved exceptionally difficult because its skeleton is largely destroyed and fragmented, especially the skull, which is highly disarticulated and has several elements missing altogether (Fig. 10). Nevertheless, there is some osteological support for its identification as a scaphiophrynine microhylid, such as the T-shaped squamosal, diplasiocoelous vertebral column, terminal phalanges of fingers and toes robust and knobbed and not T-shaped, and pectoral girdle without ossified sternum and omosternum but with ossified clavicles (Supplementary Figs S2–S 3). Assignment. Due to the lack of precise information on the geographical provenance of these specimens, assigning them to a particular lineage of the subgenus Pseudohemisus is not straightforward. First of all, we considered whether the types of Calophrynus calcaratus could be conspecific with the lineage usually named Scaphiophryne brevis. Blommers-Schl̂sser & Blanc (1991) noted that the subarticular tubercles in S. brevis are flat whereas in S. calcarata they are protruding. In the calcaratus lectotype they are rather protruding, as already stated in the original description. Furthermore, and more importantly, on the left side of the lectotype the boundary between the dorsal and lateral skin is folded in a way that may correspond to the dorsolateral fold typical for those specimens usually assigned to S. calcarata, but not for most of those assigned to S. brevis. We therefore conclude that the lectotype of Calophrynus calcaratus belongs to one of the evolutionary lineages herein subsumed in the S. calcarata complex (Fig. 1). To decide to which of these lineages the lectotype belongs, we can only apply a combination of arguments of historical plausibility and nomenclatural stability. We have already assigned the older nomen obscurus to the lineage occurring in north-western and western Madagascar. The range of the second lineage occurring in southern Madagascar includes several well-sampled sites such as Tolagnaro and Toliara. The original type series of Calophrynus calcaratus was collected by Grandidier who is known to have collected in the area (Betsileo) where the southern lineage is known to occur (e.g., Ambalavao, Anja, Isalo). In addition, the publication by Mocquard (1895), which includes the description of Calophrynus calcaratus, also features descriptions of several other species originating from Betsileo. Therefore, we propose to apply the name Scaphiophryne calcarata (Mocquard, 1895) to the southern lineage in the S. calcarata complex, i.e., to the form called Scaphiophryne calcarata B in Glaw & Vences (2007) and Scaphiophryne sp. 1 in Vieites et al. (2009). Besides being plausible based on what is known of Grandidier’s collections, this taxonomic proposal also has the advantage of largely preserving the prevalent use of this name., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on page 408, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Mocquard, F. (1895) Sur les reptiles recueillis Madascar de 1867 a 1885 par M. Grandidier. Bulletin de la Societe Philomathique de Paris, Series 8, 7, 93 - 111.","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. K ˆ ln, Vences & Glaw, 496 pp.","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the United States of America, 106, 8267 - 8272. https: // doi. org / 10.1073 / pnas. 0810821106"]}

Pseudohemisus granulosus Guibé, 1952 Name-bearing type: Holotype MNHN 1953.238, from “Grotte d’Andranoboka, baie Mahajamba, Nord Majunga” according to the original description (Guibé 1952). Morphology. The holotype of this nomen is an adult in a good state of preservation and clearly conforms to the definition of the species in the S. calcarata complex. The inner metatarsal tubercle is enlarged, a tarsal tubercle is present (Fig. 9), a dorsolateral ridge runs along most of body, the dorsal skin is covered densely with very small tubercles. Measurements are given in Table 1. Osteologically, the specimen is consistent with the other Pseudohemisus specimens examined here (Fig. 10, Supplementary Figs S2–S 3). Assignment. Based on geographical provenance of the holotype and morphology (dorsolateral ridge along most of the body) this nomen can be confidently assigned to the northern lineage of S. obscura. We therefore consider Pseudohemisus granulosus Guibe, 1952 to be a junior synonym of Scaphiophryne obscura., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on page 411, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Guibe, J. (1952) Recherches sur les batraciens de Madagascar. Memoires de l'Institut Scientifique de Madagascar. Serie A, Biologie Animale, 7, 109 - 116."]}

Pseudohemisus longimanus var. melanopleura Angel, 1934 Name-bearing type: Lectotype by present designation, MNHN 1933.149 from “Antsingy, prov. Menabe, 300 m ” according to the original description. The original type series consisted of two syntypes. The second of these, MNHN 1933.148 has a new (red) label that erroneously reads “1933.0149”, but its identity can be clearly assessed based on its paper label. We here designate the larger of the two syntypes, MNHN 1933.149, as lectotype, to allow unambiguous definition of this nomen in case future studies reveal the existence of more than two species in the S. calcarata complex. Morphology. The two original syntypes of this nomen (Frost 2020) are in a rather good state of preservation and are clearly assignable to Scaphiophryne based on the tarsal tubercle and enlarged inner metatarsal tubercle (Fig. 9). They bear clear diagnostic characters of the S. calcarata complex, particularly, a dorsolateral ridge running along the entire body. Osteological examination of the lectotype revealed close resemblance to the other Pseudohemisus specimens examined here (Fig. 10, Supplementary Figs S2–S 3). Assignment. As with Pseudohemisus longimanus, the type locality of this nomen is in an area between the known ranges of the western and the north-western lineage of S. obscura. While it is therefore not possible to assign the nomen to either of these intraspecific lineages, based on its provenance we propose considering Pseudohemisus longimanus var. melanopleura Angel, 1934 a junior synonym of Scaphiophryne obscura., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on pages 410-411, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Angel, F. (1934) Description d'un Viperide nouveau du Congo Belge et de deux batraciens de Madagascar. Bulletin de la Societe Zoologique de France, 59, 169 - 172.","Frost, D. R. (2020) Amphibian Species of the World: an Online Reference. Version 6.1. Electronic Database accessible at American Museum of Natural History, New York. Available from: https: // amphibiansoftheworld. amnh. org / index. php (accessed 7 June 2020) https: // doi. org / 10.5531 / db. vz. 0001"]}

Hemisus obscurus Grandidier, 1872 Name-bearing type: Lectotype by present designation, MNHN 1895.280, from “Côte N. O. de Madagascar ” according to the original description. Originally, there were four syntypes, all catalogued under MNHN 1895.280; one of these specimens today has an attached label with the number 1895.280 (here designated as lectotype, see below), and two other original syntypes (now paralectotypes) probably correspond to 280A and 280B (see below). The fourth specimen (according to the MNHN catalogue used for a skeleton preparation) was not examined as it could not be retrieved in the MNHN collection. The type locality “Côte N. O. de Madagascar ” means “Côte Nord Ouest”, i.e., the north-west coast. Hemisus obscurus, the type species of the subgenus Pseudohemisus, is the earliest available name for a Malagasy microhylid species from western Madagascar and therefore must be a valid taxon for reasons of priority. However, probably due to the short description and bad state of preservation of the types, the species has been considered as dubious by Blommers-Schl̂sser & Blanc (1991) and not further discussed in subsequent publications. In fact, Mocquard (1895) already stated that the types were in very poor condition when he erected Pseudohemisus as new genus name for this taxon. Upon examination in 2020, the three syntypes of Hemisus obscurus were in an extremely poor state of preservation. The largest of the specimens (SVL 23.7 mm; see Table 1) bears the label 1895.280; two additional labels 1895.280A and 280B are included in the same jar and probably refer to the other two specimens, which bear no labels and which measure 19.9 mm and 16.1 mm in SVL, respectively. Because these specimens are smaller, probably not adults, bear no labels, and are in an even worse state of preservation, the largest specimen with the label 1895.280 is here designated as lectotype to stabilize this nomen. Morphology. Only a few measurements could be taken due to the bad state of preservation; these are included in Table 1. The enlarged inner metatarsal tubercle typical for Scaphiophryne, and the size and shape of subarticular tubercles, is not recognizable in the lectotype, probably because most of the skin around the feet is destroyed (Fig. 9). However, it is just possible recognize a tarsal tubercle in the lectotype, which is typical for Scaphiophryne, and the enlarged inner metatarsal tubercle is still faintly visible in one of the paralectotypes. Furthermore, the triangular shape of the head, becoming distinctly narrower towards the snout, which is rather truncate in dorsal and ventral views, and the absence of vomerine and maxillary teeth, indicate that the lectotype and the two paralectotypes indeed are Scaphiophryne. The head of the lectotype bears what might be the remains of a light vertebral line as is typical for most specimens of the S. calcarata complex (e.g., Glaw & Vences 2007). The designated lectotype also agrees osteologically with traits of the subgenus Pseudohemisus based on the deepened septomaxilla, T-shaped squamosal, ossified and anteriorly extended sphenethmoid, maxillary and vomerine teeth absent, non-divided vomer (divided in Cophylinae), sacral diapophysis flattened and broadened (as diagnostic for microhylids), vertebral column diplasiocoelous, terminal phalanges of fingers and toes knobbed and not T-shaped as in species in subgenus Scaphiophryne with expanded finger and toe tips, and pectoral girdle without ossified sternum and omosternum but with ossified clavicles (Fig. 10, Supplementary Figs S2–S 3). Assignment. The type locality of Hemisus obscurus is, rather imprecisely, the north-west coast of Madagascar. In this region, no frogs of the lineage assigned to S. brevis (see below) occur, and therefore it is clear that the name obscurus should be assigned to a lineage of the S. calcarata complex. Hence, we apply the name Scaphiophryne obscura (Grandidier, 1872) to the north-western/western lineage of the S. calcarata complex, i.e., to the form called Scaphiophryne calcarata A in Glaw & Vences (2007) and Scaphiophryne sp. 2 in Vieites et al. (2009)., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on pages 405-406, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Grandidier, A. (1872) Description de quelques reptiles nouveaux decouverts a Madagascar en 1870. Annales des Sciences Naturelles. Zoologie et Paleontologie, Paris, Serie 5, 15, 6 - 11.","Mocquard, F. (1895) Sur les reptiles recueillis Madascar de 1867 a 1885 par M. Grandidier. Bulletin de la Societe Philomathique de Paris, Series 8, 7, 93 - 111.","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. K ˆ ln, Vences & Glaw, 496 pp.","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the United States of America, 106, 8267 - 8272. https: // doi. org / 10.1073 / pnas. 0810821106"]}

Pseudohemisus verrucosus Angel, 1930 Name-bearing type: Lectotype by present designation, MNHN 1929.233, from “Lavenombato, bords de l’Onilahy” according to the original description. The original syntype series contained five syntypes; MNHN 1929.231 – 232 and 234–235 are to be considered as paralectotypes. This nomen has been considered a dubious species by Blommers-Schl̂sser & Blanc (1991) and is considered to be a nomen inquirendum in Frost (2020). The original type series of five syntypes was collected at a locality in south-western Madagascar, where S. brevis is common and where the S. calcarata complex has only recently been documented (D’Cruze et al. 2009). Four of the specimens (MNHN 1929.231–234) are juveniles, in a satisfactory state of preservation, and clearly belong to Scaphiophryne based on the presence of an enlarged inner metatarsal tubercle and presence of a tarsal tubercle. The fifth number (MNHN 1929.235) refers to a series of five tadpoles, which probably do not belong to Scaphiophryne. Because the types of verrucosus are not adults, it is difficult to ascertain their identity. We here designate the specimen MNHN 1929.233 (with the most strongly expressed tubercular state of the dorsum) as lectotype of Pseudohemisus verrucosus Angel, 1930 in an effort to stabilize this nomen given the heterogeneity of the type series. Morphology. The most distinctive character of MNHN 1929.231 – 234 is its strongly tubercular dorsum. The large tubercles are distinct, especially in the lectotype MNHN 1929.233 but less so in the paralectotype MNHN 1929.231. However, in all four specimens these tubercles are recognizable, and they even form a pair of distinct, parallel dorsal ridges above the forelimb insertion. A dorsolateral ridge is also recognizable towards the front of the flank. The tips of the fingers and toes are not enlarged; the subarticular tubercle of the first toe does not appear to be distinctly enlarged or prominent (Fig. 9). Osteologically, the lectotype appears to be a juvenile, because several components of its skeleton are not fully ossified. However, it does have all the hallmarks of Pseudohemisus identified in the other specimens examined here, although the type of vertebral articulation is not ascertainable due to the damage to the specimen (Fig. 10, Supplementary Figs S2–S 3). Assignment. As far as known, distinct dorsal tubercles and ridges are not known from Scaphiophryne calcarata sensu lato, whereas specimens of S. brevis, especially in life, can have a tubercular dorsum (see photos in Glaw & Vences 2007). In addition, a photograph taken by N’ D’Cruze at Sept Lacs, not far from the Onilahy/ Toliara region (reproduced in Glaw & Vences 2007: page 448) shows a specimen assigned to S. brevis with several of the characters of the verrucosus types, i.e., rather distinct dorsolateral ridge in the anterior part of body, and dorsum with tubercles forming short dorsal ridges. D’Cruze et al. (2009) collected voucher specimens of S. brevis and S. “ calcarata ”. Examining these specimens and carrying out additional collections in the region will be necessary to understand the variability of juvenile S. brevis and to understand whether an additional, previously unrecognized species of Scaphiophryne with tubercular dorsum may occur in this region to which the nomen verrucosus may be assignable. Considering the general variability observed within Scaphiophryne species in characters regarding shape of subarticular tubercles and dorsal skin texture (e.g., tubercular vs. smooth in specimens of S. madagascariensis from different localities), we propose to consider the nomen Pseudohemisus verrucosus Angel, 1930 as a junior synonym of Scaphiophryne brevis, which appears to be the most plausible hypothesis in light of the current state of knowledge., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on page 410, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Angel, F. (1930) Description de deux especes nouvelles de batraciens de Madagascar, appartenant au genre Pseudohemisus. Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 2, 2, 70 - 74.","Frost, D. R. (2020) Amphibian Species of the World: an Online Reference. Version 6.1. Electronic Database accessible at American Museum of Natural History, New York. Available from: https: // amphibiansoftheworld. amnh. org / index. php (accessed 7 June 2020) https: // doi. org / 10.5531 / db. vz. 0001","D'Cruze, N., Olsonn, A., Henson, D., Kumar, S. & Emmett, D. (2009) Amphibians and reptiles of the Onilahy River valley. Herpetological Conservation and Biology, 4, 62 - 79.","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. K ˆ ln, Vences & Glaw, 496 pp."]}

Calophrynus brevis Boulenger, 1896 Name-bearing type: Holotype BMNH 1947.2.7.42, from “S.W. Madagascar ” according to the original description. The holotype of this nomen is in good state of preservation, allowing the examination of most or all characters relevant for taxonomic assignment. Although it is smaller than most other adult specimens typically assigned to S. brevis and therefore may be a subadult, its size is similar to one small-sized adult male collected by us (ZSM 577/2000; Table 1). The general body shape and absence of dorsolateral folds or a sharp dorsolateral color border (i.e., a clearly delimited boundary between the colors of the dorsum and flanks) differentiate the specimen from those of the S. calcarata complex, and confirm that Scaphiophryne brevis (Boulenger, 1896) has been correctly applied by previous researchers (e.g., Blommers-Schl̂sser & Blanc 1991; Glaw & Vences 2007; Vieites et al. 2009). D’Cruze et al. (2009) reported this species from the Onilahy River valley. Scaphiophryne brevis seems to exhibit at least two pronounced color morphs, one with a diffuse dorsolateral color border and typically a light midvertebral stripe (first row of Fig. 3), and one that is more marbled (second row of Fig. 3). A potential third color morph with green elements is figured in Glaw & Vences 2007 (page 448), and discussed in the context of Pseudohemisus verrucosus below. These color morphs are not related to sex., Published as part of Scherz, Mark D., Schmidt, Luca, Crottini, Angelica, Miralles, Aurélien, Rakotoarison, Andolalao, Raselimanana, Achille P., Köhler, Jörn, Glaw, Frank & Vences, Miguel, 2021, Into the Chamber of Horrors: A proposal for the resolution of nomenclatural chaos in the Scaphiophryne calcarata complex (Anura: Microhylidae), with a new species-level phylogenetic hypothesis for Scaphiophryninae, pp. 392-420 in Zootaxa 4938 (1) on page 409, DOI: 10.11646/zootaxa.4938.4.2, http://zenodo.org/record/4574869, {"references":["Boulenger, G. A. (1896) Descriptions of new batrachians in the British Museum. Annals and Magazine of Natural History, Series 6, 17, 401 - 406. https: // doi. org / 10.1080 / 00222939608680389","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. K ˆ ln, Vences & Glaw, 496 pp.","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the United States of America, 106, 8267 - 8272. https: // doi. org / 10.1073 / pnas. 0810821106","D'Cruze, N., Olsonn, A., Henson, D., Kumar, S. & Emmett, D. (2009) Amphibians and reptiles of the Onilahy River valley. Herpetological Conservation and Biology, 4, 62 - 79."]}

Galápagos marine iguanas are primarily associated with the marine environment and show special nutritional adaptations. They are the only lizards worldwide that forage on marine macroalgae. Until now, consumed algae have been identified by direct observations during their feeding activities and microscopic identification in faeces samples. In this study, we use a novel DNA metabarcoding approach to identify consumed algal species from the faeces of marine iguanas. We developed primers for the ribulose-bisphosphate carboxylase (rbcL) gene and applied a metabarcoding approach to 25 individual faeces samples collected in four representative sites of two subspecies (Amblyrhynchus cristatus mertensi and A. c. godzilla), found in the San Cristóbal Island. We detected 18 consistently occurring macroalgal operational taxonomic units (OTUs). Most of the OTUs were assigned to Rhodophyta (red algae) and only one OTU to Chlorophyta (green algae). Despite the number of consumed algal species did not differ between two subspecies (OTU richness; P = 0.383), diet overlap level between A. c. mertensi and A. c. godzilla was low (Schoener index = 0.345), suggesting that both subspecies consumed different algal species in their natural environment. Further studies are needed to understand whether the difference of consumed algae reflects disparities in the abundance of algal species between sites, or whether iguanas of the two genetically differentiated subspecies prefer distinct algal species.

Mantellid frogs of the Madagascar-endemic Gephyromantis plicifer complex consist of three nominal species (G. luteus, G. plicifer and G. sculpturatus) as well as several genetically divergent lineages (candidate species), but uncertainties surround the identity of the name-bearing types of all three established nomina. We applied laboratory techniques to isolate archival DNA from the three old, liquid-preserved, name-bearing type specimens and conducted target-enriched DNA sequencing of a fragment of the mitochondrial 16S rRNA gene as basis for the revision of their taxonomy. Along with 16S sequences of 161 fresh samples, sequences of a fragment of the nuclear-encoded Rag-1 gene for 97 of these samples, a multi-gene data set of selected samples, and a comprehensive analysis of available advertisement call recordings, the new data suggest a novel taxonomic hypothesis: (1) The name G. plicifer applies to a highland clade of populations from the southern Central East, previously referred to as Gephyromantis sp. Ca21, with G. sculpturatus as a junior synonym. (2) The name G. luteus applies to a widespread clade distributed along most of Madagascar's eastern rainforest band, and more specifically, is assigned to a subclade occurring mostly in coastal regions of the northern Central East and North East. (3) Large-sized specimens from Ranomafana (southern Central East) and Andohahela (South East) previously considered to belong to G. plicifer in fact represent a hitherto undescribed species, which we here formally name as G. pedronoi sp. nov. The three species are distinguished by very high genetic distances (over 8 % in the 16S gene), an absence of haplotype sharing in Rag-1, as well as morphological and bioacoustic characteristics. Moreover, several infra-specific clades with 16S divergences > 4 % may represent distinct species, especially within the widespread G. luteus, but cannot be unambiguously delimited by available data. We suggest dedicated sampling at contact zones and more extensive bioacoustic study of genotyped call vouchers to clarify their taxonomic status.

Uroplatus fivehy sp. nov. LSID: urn:lsid:zoobank.org:act: AC24C7D3-1AB5-451F-9DE2-935DE7174FDD Figs 3, 4, 6���8 Remarks. This new species corresponds to the lineage previously named Uroplatus sp. A by Raxworthy et al. (2008), Uroplatus sp. 4 by Ratsoavina et al. (2011), and Uroplatus ebenaui [Ca4] by Ratsoavina et al. (2012, 2013, 2015, 2017, 2019a,b). Holotype. ZSM 1721 /2012 (field number FGZC 3590), adult male, collected on 26 November 2012 by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina and A. Razafimanantsoa on the Sorata Massif, Sava Region, northern Madagascar, in the vicinity of the campsite, near 13.6805��S, 49.4451��E, 1417 m a.s.l. Paratypes. A total of 12 specimens: ZSM 1722 /2012 (field number FGZC 3591), adult female, with same collection data as the holotype; UADBA-R 70851 (FGZC 3694), adult male, collected by same collectors as holotype on 29 November 2012 on the Sorata Massif at 13.6868��S, 49.4412��E, 1272 m a.s.l.; UADBA-R 70850 (FGZC 3715), adult male, collected by same collectors as holotype on 30 November 2012 on the Sorata Massif near a creek at 13.6829��S, 49.4403��E, 1325 a.s.l.; ZSM 634 /2009 (ZCMV 11308), adult male, and UADBA-R 70966 (ZCMV 11309), adult female, collected between 22���25 June 2009 by F. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison, D. R. Vieites and M. Vences near Angozongahy campsite on the western side of the Makira plateau in the Northern Central East of Madagascar, near 15.437��S, 49.1167��E, 1009 m a.s.l.; UADBA 29428 (APR 05955) and UADBA 29427 (APR 05921), two adult males collected on 22 November 2004 and UADBA 29426 (APR 05859), subadult, collected on 19 November 2004 by A.P. Raselimanana at Anjiabolo site (ca. 16.2800��S, 48.8017��E; respectively at 1100 m and 1000 m a.s.l) in Marotandrano Special Reserve, Sofia Region, Northern Central East Madagascar; UADBA 29430 (APR 05984), UADBA 29431 (APR 05985), two males and UADBA 29429 (APR 05983), adult female, collected on 25 November 2004; and UADBA 29432 (APR 06118), adult male, collected on 29 November 2004 by A.P. Raselimanana at Riamalandy site (16.2850��S, 48.8150��E, between 825���875 m a.s.l.) in Marotandrano Special Reserve. Etymology. The species epithet fivehy is a noun in apposition to the genus name, meaning ���paddle��� in local Malagasy dialect, and referring to the paddle-like shape of its tail. Diagnosis. Uroplatus fivehy sp. nov. is assigned to the Uroplatus ebenaui species group based on its relatively small size, leaf-like, laterally-compressed body shape, short tail, and triangular head with supraocular spines. The following unique combination of characters characterizes the new species: (1) short (TAL/SVL 0.21���0.41) and slender (TAW/SVL 0.06���0.13) tail, (2) oral mucosa without black pigment, and (3) relatively smooth (not spiny) appearance in males. Additionally, it is distinguished by high genetic pairwise-distances> 10% in DNA sequences of the mitochondrial 16S gene from all other nominal species of Uroplatus. By its overall leaf-like appearance, the species can easily be distinguished from all members of the U. fimbriatus species group, from U. lineatus, and from the phenetic U. alluaudi species group. Within the U. ebenaui species group, U. fivehy sp. nov. can be distinguished from U. finaritra and U. malama by smaller body size (SVL 60.8���72.4 mm versus 77.5���95.3 mm); from U. malama, U. phantasticus, U. finiavana and U. finaritra by shorter tail (TAL/SVL 0.21���0.41 versus 0.48���0.76); from U. fangorn, U. ebenaui, U. fetsy, and U. kelirambo by its unpigmented oral mucosa (pale pinkish all over versus black or deep red pigment present); from U. kelirambo by more expanded tail edges (versus strongly spear-shaped; TAW/SVL 0.06���0.13 versus 0.04���0.05 in U. kelirambo). U. fivehy sp. nov. is morphologically most similar to U. fiera and to U. fotsivava, which however concordantly differ in mitochondrial and nuclear DNA sequences and are not its immediate relatives based on molecular phylogenetic analysis; it differs from U. fotsivava by a generally longer and wider tail, although the values do overlap (TAL/SVL 0.21���0.41 versus 0.15���0.32; TAW/SVL 0.06���0.13 versus 0.05���0.10), and from U. fiera possibly in a slightly larger ratio TAL/TAW (0.22���0.37 versus 0.19���0.21). Description of the holotype. Adult male in good condition with an intact tail and everted hemipenes. SVL 63.1 mm, tail length 25.9 mm, maximum tail width 5.7 mm, for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis recognizable and almost straight; snout sloping strongly and continuously downward anteriorly; snout weakly depressed, short; eyes large (eye diameter 4.6 mm), not bulging above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small but distinct (horizontal diameter 0.6 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in lateral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and practically without spines on forelimbs and hindlimbs, except for a small flap on the knee; forelimb reaches beyond tip of snout when adpressed forward and does not reach the groin when adpressed backwards along body (forelimb length/axilla���groin distance 27.7/ 29.7 mm = 93%), hindlimb reaches beyond axilla when adpressed forward along body (hindlimb length/axilla���groin distance 37.4/ 29.7 mm = 126%); original tail length 41% of snout���vent length, membranous borders of the denticulated tail rather asymmetrical, broadening on either side and then narrowing again toward the tip), completely absent from the slightly spatulate tail-tip. Nares separated from each other by eight small granular scales, from the first supralabial by one scale, and from the rostral scale by two scales; the first supralabial taller than the others; rostral entire, much wider than tall; mental scale small, not differentiated from infralabial scales; 21/20 (right/left) supralabials and 19/21 infralabials (as defined in methods above); scales at the periphery of the chin (toward the infralabial scales) slightly larger than central chin scales; the dorsal scales of head, neck, body, limbs and tail small, granular, juxtaposed and largely of homogeneous size interspersed with very few larger and partly raised tubercles; the numerous curved transverse lines on the head and dorsal side of the body are not marked by significantly enlarged scales. Very few spines on head and hindlimbs, a dermal flap on each knee, bearing no spine; no flaps or spines on elbow and forelimb; a prominent pointed flap on the posterior portion of each upper eyelid (supraocular spine); posterior border of eye fringed. Axillary pits not recognizable. Scales of the ventral abdomen distinctly larger than dorsal body scales and arranged more homogeneously. Hemipenes everted, bearing two lobes. Calyx with protuberance bearing honeycomb-like structures, especially on the asulcal side. Area of sulcus spermaticus is smooth. Each lobe with a dense field of pointed papillae at its apex. Coloration of the holotype. After eight years of preservation in 70% ethanol the color pattern remains similar to that of the living animal (Figs 6H, 8) but its vividness and intensity have slightly faded. All dorsal surfaces are mottled beige and blackish in base color. The tail has a beige pattern on its dorsal pygal section, bordered by blackish color. The dorsum has rather distinctly defined posteroventrally oriented lighter and darker beige chevrons. The dorsal surfaces of the head are dark brown with a series of six distinct transverse narrow beige lines, running either straight or converging anteriorly or posteriorly. No dark band between the eyes, but a dark V-shaped area on the nape is well distinguishable. Additional dark flecks are present on the mid-dorsum, and several smaller asymmetrical spots on other parts of the body, including legs and nape. One light cream marking is present posteriorly below each eye, descending to the supralabials. The supralabials and the infralabials are generally dark gray in color. The venter, including the ventral limbs except for the lower hindlimbs, is slightly lighter beige than the dorsum, and less mottled. The pectoral region bears two elongated, poorly delimited whitish spots. The postpygal section of the tail is of similar color dorsally and ventrally. The oral mucosa is unpigmented and whitish in preservative. The iris of the eye is gray, and the vertical pupil is blackish. Variation. As described for U. fangorn above, there is a substantial color variation (Figs 6���7), and sexual dimorphism in the edges of the tail. The color of the eye varies from beige/gray to dark brown and reddish-brown. Dorsal color can be uniform beige to reddish-brown with or without white lichen-like markings. Two sharply delimited vertebral black patches on the anterior and posterior part of the dorsum are present in several individuals. Relative tail length and tail width are characterized by substantial variability among individuals (Fig. 4). Here, we have included in the paratype series and in the morphological comparisons a series of individuals from Marotandrano Special Reserve that we attribute to this species. However, it must be stressed that the species identity has been confirmed by DNA sequences only for one of these (UADBA 29428 = APR5955). In the unlikely case that some of these individuals will turn out to belong to another species of Uroplatus, the morphometric variation of U. fivehy may be narrower than reported here. ......continued on the next page Distribution, natural history and conservation status. According to the material collected and examined by us, U. fivehy is known from (1) the type locality on the Sorata Massif, (2) the western slope of the Makira Reserve, and (3) Marotandrano Special Reserve. In addition, mitochondrial DNA sequences published by Ratsoavina et al. (2012) suggest its occurrence in (4) the Anjanaharibe-Sud Massif, and mitochondrial DNA sequences from Raxworthy et al. (2008) furthermore suggest the occurrence of the species in (5) Marojejy, (6) Ankitsika and (7) Betaolana. The confirmed elevational range from our own samples is between 1009 and 1417 m a.s.l., and the record from Ankitsika would extend this range downwards to 830 m a.s.l., characterizing the species as rather wide-ranging from the Northern Central East to the North East of Madagascar from mid-elevations to highlands (see also Ratsoavina et al. 2013). All specimens were found in intact rainforest, at night, perched on branches of trees. At Marotandrano, an adult male was found hidden in a Pandanus leaf axil during the day in closed canopy humid forest, and two adult individuals (male and female) were found at night close to each other in the same tree on 25 November 2004, suggesting it was probably the reproduction period for this species. Individuals were often observed active on small branches, leaves, and lianas at 0.5���3 m above the forest floor. All individuals were collected on slope or ridge forest. In Marotandrano, the species appeared to be quite common in forests associated with vine-like bamboo and shrub vegetation. A minimum convex polygon of the known distribution of this species covers an area of ca 15,100 km 2, qualifying the species for a status of Vulnerable under the IUCN Red List criterion B (IUCN 2012). This species is known from several protected areas (Marotandrano Special Reserve, Makira Reserve, COMATSA Nord, Marojejy National Park, Anjanaharibe-Sud National Park) but cumulatively just seven locations (B subcriterion a). These locations, and especially the unprotected areas between them, are experiencing substantial ongoing declines in the extent and quality of habitat (B subcriterion b(iii)). Therefore, we propose to list this species as Vulnerable under IUCN criterion B1ab(iii)., Published as part of Ratsoavina, Fanomezana M., Glaw, Frank, Raselimanana, Achille P., Rakotoarison, Andolalao, Vieites, David R., Hawlitschek, Oliver, Vences, Miguel & Scherz, Mark D., 2020, Towards completion of the species inventory of small-sized leaf-tailed geckos: two new species of Uroplatus from northern Madagascar, pp. 251-271 in Zootaxa 4895 (2) on pages 261-268, DOI: 10.11646/zootaxa.4895.2.5, http://zenodo.org/record/4322511, {"references":["Raxworthy, C. J., Pearson, R. G., Zimkus, B. M., Reddy, S., Deo, A. J., Nussbaum, R. A. & Ingram, C. M. (2008) Continental speciation in the tropics: contrasting biogeographic patterns of divergence in the Uroplatus leaf-tailed gecko radiation of Madagascar. Journal of Zoology, 275, 423 - 440. https: // doi. org / 10.1111 / j. 1469 - 7998.2008.00460. x","Ratsoavina, F. M., Louis, E. E. Jr., Crottini, A., Randrianiaina, R. D., Glaw, F. & Vences, M. (2011) A new leaf tailed gecko species from northern Madagascar with a preliminary assessment of molecular and morphological variability in the Uroplatus ebenaui group. Zootaxa, 3022 (1), 39 - 57. https: // doi. org / 10.11646 / zootaxa. 3022.1.3","Ratsoavina, F. M., Vences, M. & Louis, E. E. Jr. (2012) Phylogeny and phylogeography of the Malagasy leaf-tailed geckos in the Uroplatus ebenaui group. African Journal of Herpetology, 61, 143 - 158. https: // doi. org / 10.1080 / 21564574.2012.729761","Ratsoavina, F. M., Raminosoa, N. R., Louis, E. E. Jr., Raselimanana, A. P., Glaw, F. & Vences, M. (2013) An overview of Madagascar's leaf tailed geckos (genus Uroplatus): species boundaries, candidate species and review of geographical distribu- tion based on molecular data. Salamandra, 49, 115 - 148.","Ratsoavina, F. M., Ranjanaharisoa, F. A., Glaw, F., Raselimanana, A. P., Miralles, A. & Vences, M. (2015) A new leaf-tailed gecko of the Uroplatus ebenaui group (Squamata: Gekkonidae) from Madagascar's central eastern rainforests. Zootaxa, 4006 (1), 143 - 160. https: // doi. org / 10.11646 / zootaxa. 4006.1.7","Ratsoavina, F. M., Gehring, P. S., Scherz, M. D., Vieites, D. R., Glaw, F. & Vences, M. (2017) Two new species of leaf-tailed geckos (Uroplatus) from the Tsaratanana mountain massif in northern Madagascar. Zootaxa, 4347 (3), 446 - 464. https: // doi. org / 10.11646 / zootaxa. 4347.3.2","Ratsoavina, F. M., Raselimanana, A. P., Scherz, M. D., Rakotoarison, A., Razafindraibe, J. H., Glaw, F. & Vences, M. (2019 a) Finaritra! A new leaf-tailed gecko (Uroplatus) species from Marojejy National Park in north-eastern Madagascar. Zootaxa, 4545 (4), 563 - 577. https: // doi. org / 10.11646 / zootaxa. 4545.4.7","IUCN. (2012) IUCN Red List Categories and Criteria: Version 3.1. IUCN, Gland and Cambridge."]}