[Herpetology • 2019] Bothrops sonene • A New Species of Bothrops (Serpentes: Viperidae: Crotalinae) from Pampas del Heath, southeastern Peru, with Comments on the Systematics of the Bothrops neuwiedi species group

Bothrops sonene 

Carrasco, Grazziotin, Farfán, Koch, Ochoa, Scrocchi, Leynaud & Chaparro, 2019.

 DOI:  10.11646/zootaxa.4565.3.1

Abstract

We describe a new species of pitviper of the genus Bothrops from the Peruvian Pampas del Heath, in the Bahuaja-Sonene National Park. Pampas del Heath is an area of seasonally flooded savannas and a northwestern extension of the Gran Chaco Boliviano-Paraguayo. The new species is easily distinguished from its congeners by the exclusive combination of dorsal color pattern of body consisting of small C-shaped blotches, postocular stripe originating posteriorly to the eye, covering posterior supralabials, dorsum of the head with paired markings arranged symmetrically, venter cream heavily speckled with brown, prelacunal scale discrete in contact with second supralabial, three to five prefoveals, subfoveal single usually present, postfoveals absent to two, canthals two, seven intersupraoculars, one or two suboculars, two or three postoculars, seven or eight supralabials, nine to eleven infralabials, 26–27 interrictals, 23–25 middorsal scales, 172 ventrals in the female and 169–173 in males, 45 subcaudals in the female and 50 in males. We performed separate and combined phylogenetic analyses based on morphology and five mitochondrial genes and recovered the new species as a member of the Bothrops neuwiedi species group. All lineages of this clade inhabit the South American dry diagonal. This novel species of pitviper increases the known diversity of the genus Bothrops and adds to the number of described taxa from the unique and scarcely known ecosystem of Pampas del Heath.

Keywords: Reptilia, Bothrops mattogrossensis, species delimitation, molecular phylogeny, morphological characters, total evidence

Bothrops sonene

Paola A. Carrasco, Felipe G. Grazziotin, Roy Santa Cruz Farfán, Claudia Koch, José Antonio Ochoa, Gustavo J. Scrocchi, Gerardo C. Leynaud and Juan C. Chaparro. 2019. A New Species of Bothrops (Serpentes: Viperidae: Crotalinae) from Pampas del Heath, southeastern Peru, with Comments on the Systematics of the Bothrops neuwiedi species group. Zootaxa. 4565(3); 301–344. DOI:  10.11646/zootaxa.4565.3.1

[Herpetology • 2019] Atractaspis branchi • A New Stiletto Snake (Lamprophiidae, Atractaspidinae) from Liberia and Guinea, West Africa

Atractaspis branchi 

Rödel, Kucharzewski, Mahlow, Chirio, Pauwels, Carlino, Sambolah & Glos, 2019

Branch’s Stiletto Snake  ||  DOI: 10.3897/zse.95.31488

Abstract

We describe a new stiletto snake, Atractaspis, from western Liberia and southeastern Guinea. The new species shares with morphologically similar western African Atractaspis species, A. reticulata and A. corpulenta, the fusion of the 2nd infralabial with the inframaxillary. From A. corpulenta the new species differs by a more slender body (276–288 ventrals and 19 or 20 dorsal scale rows versus 178–208 ventrals with 23–29 dorsal scale rows), a divided anal plate and divided subcaudal scales (both non-divided in A. corpulenta). The new species differs from most A. reticulata by having 19 or 20 dorsal scale rows at midbody (versus 21–23, rarely 19), and a lower ventral count (276–288 versus 304–370). The new species thus has a relatively longer tail: snout-vent-length / tail-length in the female holotype (15.7) and paratype (21.5) versus a mean of 23.6 in seven female A. reticulata. The new Atractaspis likely is endemic to the western part of the Upper Guinea forest zone and thus adds to the uniqueness of this diverse and threatened biogeographic region.

Key Words: Biodiversity hotspot, biogeography, rainforest, Reptilia, Squamata, species delimitation, Upper Guinea forest

Figure 1. Life coloration of the Atractaspis branchi sp. n. holotype (ZMB 88529). 

Figure 3. Holotype of Atractaspis branchi sp. n. (ZMB 88529) 1 head scalation in dorsal (a), lateral (b), and ventral (c) views 2 ct-scan of skull in dorsal (a), lateral (b), and ventral (c) views; lower jaw virtually removed; green: pterygoid, yellow: palatine, orange: vomer. Scale bar: 1 mm.

Atractaspis branchi sp. n.

Diagnosis: External morphology, skull anatomy and molecular data (see below) clearly supports the position within the genus Atractaspis. The new species can be only mistaken morphologically with species from Laurent’s (1950) section ‘D’, his reticulata-group. In particular it differs from all other species of the genus, except A. reticulata and A. corpulenta (including the West African A. c. leucura), by the fusion of the 2nd infralabial with the inframaxillary. From A. corpulenta it differs by a much higher ventral count (276–288 vs 178–208), lower number of dorsal scale rows at midbody (19 vs 23–29), divided anal plate and subcaudals, and the absence of a white colored tail tip (present in A. c. leucura); from A. reticulata it can be distinguished by a lower ventral count (276–288 vs 304–370), and 19 (the paratype has mostly 19 scale rows, but 20 at midbody) dorsal scales rows at midbody (19 scale rows present in the A. reticulata holotype, other vouchers having 21–23 rows) (Table 1). The new species further differs from A. corpulenta by a more slender body and from A. reticulata by a longer tail compared to body length.

Figure 6. Type locality of Atractaspis branchi sp. n. in north-western Liberia. The holotype specimen was found at night. It was moving along the steep slope on the left bank of the small creek.

 Figure 7. Localities of Atractaspis branchi sp. n. and A. reticulata ssp.
Records are based on museum specimens, literature and database (GBIF) records; large closed symbols represent the type localities of the different taxa, stars: A. branchi sp. n., circles: A. reticulata records without reference to subspecies; triangles: A. r. reticulata; quadrats: A. r. heterochilus; diamonds: A. r. brieni; country borders indicted as white lines; background of map: major biomes based on Olson et al. (2001).

Natural history: We found the holotype at night. It was slowly moving along the steep slope of the bank of a small rocky creek in primary lowland evergreen rainforest (Fig. 6). When handled, the snake first tried to hide its head below body loops; the head was bend down at an almost right angle and with fangs partly visible outside of the mouth. In this head position, the snake repeatedly tried to strike. Either it tried to move slowly away from the human observers or it abruptly coiled and uncoiled, often jumping distances equaling almost its entire body length, similar to wolf snakes of the genus Lycophidion (Rödel et al. 1995; Greene 1997). The two snakes from south-eastern Guinea were collected in plantations of banana, manioc and coffee, which were planted under the few remaining high trees of the former forest. No other data on biology and ecology of the new species are known.

Distribution: So far the new species is known from the type locality and two additional sites in south-eastern Guinea. These latter two sites are about 27 km apart (Fig. 7).

Etymology: We name this new snake to honor our recently deceased friend and colleague, William Roy “Bill” Branch, for his outstanding contributions to African herpetology. MOR and OSGP are particularly pleased to name the species in memory of Bill. We remember our outstanding field trips with him, unforgettable discussions with a large portion of special humor, and his friendship. The dedication of this species of stiletto snake to Bill is particularly appropriate. After Bill turned from cancer research to herpetology (see “William R. Branch” in Li Vigni 2013), the subject of his first herpetological research, on the serotaxonomy and hemipeneal morphology of stiletto snakes, was presented in two contributions at a symposium of herpetology and ichthyology in Kruger National Park in 1975 (Branch 1975a, b). As the vernacular name, we suggest Branch’s Stiletto Snake.

 Mark-Oliver Rödel, Christoph Kucharzewski, Kristin Mahlow, Laurent Chirio, Olivier Pauwels, Piero Carlino, Gordon Sambolah and Julian Glos. 2019. A New Stiletto Snake (Lamprophiidae, Atractaspidinae, Atractaspis) from Liberia and Guinea, West Africa. Zoosystematics and Evolution. 95(1): 107-123.  DOI: 10.3897/zse.95.31488

[Entomology • 2018] Revision of the Genus Megacraspedus Zeller, 1839 (Lepidoptera, Gelechiidae), A Challenging Taxonomic Tightrope of Species Delimitation

[73-74]  Megacraspedus gallicus sp. n., [75-76] M. ribbeella (Caradja, 1920), [77-78] M. libycus sp. n. 

Huemer & Karsholt, 2018

  DOI: 10.3897/zookeys.800.26292 

Abstract

The taxonomy of the Palearctic genus Megacraspedus Zeller, 1839 (Lepidoptera, Gelechiidae) is revised, based on external morphology, genitalia and DNA barcodes. An integrative taxonomic approach supports the existence of 85 species which are arranged in 24 species groups (disputed taxa from other faunal regions are discussed). Morphology of all species is described and figured in detail. For 35 species both sexes are described; for 46 species only the male sex is reported, in one species the male is unknown, whereas in three species the female adult and/or genitalia morphology could not be analysed due to lack of material.

DNA barcode sequences of the COI barcode fragment with > 500 bp were obtained from 264 specimens representing 62 species or about three-quarters of the species. Species delimitation is particularly difficult in a few widely distributed species with high and allegedly intraspecific DNA barcode divergence of nearly 14%, and with up to 23 BINs in a single species. Deep intraspecific or geographical splits in DNA barcode are frequently not supported by morphology, thus indicating a complex phylogeographic history or other unresolved molecular problems.

The following 44 new species (22 of them from Europe) are described: Megacraspedus bengtssoni sp. n. (Spain), M. junnilaineni sp. n. (Turkey), M. similellus sp. n. (Bulgaria, Romania, Turkey), M. golestanicus sp. n. (Iran), M. tokari sp. n. (Croatia), M. neli sp. n. (France, Italy), M. faunierensis sp. n. (Italy), M. gredosensis sp. n. (Spain), M. bidentatus sp. n. (Spain), M. fuscus sp. n. (Spain), M. trineae sp. n. (Portugal, Spain), M. skoui sp. n. (Spain), M. spinophallus sp. n. (Spain), M. occidentellus sp. n. (Portugal), M. granadensis sp. n. (Spain), M. heckfordi sp. n. (Spain), M. tenuiuncus sp. n. (France, Spain), M. devorator sp. n. (Bulgaria, Romania), M. brachypteris sp. n. (Albania, Greece, Macedonia, Montenegro), M. barcodiellus sp. n. (Macedonia), M. sumpichi sp. n. (Spain), M. tabelli sp. n. (Morocco), M. gallicus sp. n. (France, Spain), M. libycus sp. n. (Libya, Morocco), M. latiuncus sp. n. (Kazahkstan), M. kazakhstanicus sp. n. (Kazahkstan), M. knudlarseni sp. n. (Spain), M. tenuignathos sp. n. (Morocco), M. glaberipalpus sp. n. (Morocco), M. nupponeni sp. n. (Russia), M. pototskii sp. n. (Kyrgyzstan), M. feminensis sp. n. (Kazakhstan), M. kirgizicus sp. n. (Afghanistan, Kazakhstan, Kyrgyzstan), M. ibericus sp. n. (Portugal, Spain), M. steineri sp. n. (Morocco), M. gibeauxi sp. n. (Algeria, Tunisia), M. multipunctellus sp. n. (Turkey), M. teriolensis sp. n. (Croatia, Greece, Italy, Slovenia), M. korabicus sp. n. (Macedonia), M. skulei sp. n. (Spain), M. longivalvellus sp. n. (Morocco), M. peslieri sp. n. (France, Spain), M. pacificus sp. n. (Afghanistan), and M. armatophallus sp. n. (Afghanistan). Nevadia Caradja, 1920, syn. n. (homonym), Cauloecista Dumont, 1928, syn. n., Reichardtiella Filipjev, 1931, syn. n., and Vadenia Caradja, 1933, syn. n. are treated as junior synonyms of Megacraspedus. Furthermore the following species are synonymised: M. subdolellus Staudinger, 1859, syn. n., M. tutti Walsingham, 1897, syn. n., and M. grossisquammellus Chrétien, 1925, syn. n. of M. lanceolellus (Zeller, 1850); M. culminicola Le Cerf, 1932, syn. n. of M. homochroa Le Cerf, 1932; M. separatellus (Fischer von Röslerstamm, 1843), syn. n. and M. incertellus Rebel, 1930, syn. n. of M. dolosellus (Zeller, 1839); M. mareotidellus Turati, 1924, syn. n. of M. numidellus (Chrétien, 1915); M. litovalvellus Junnilainen, 2010, syn. n. of M. imparellus (Fischer von Röslerstamm, 1843); M. kaszabianus Povolný, 1982, syn. n. of M. leuca (Filipjev, 1929); M. chretienella (Dumont, 1928), syn. n., M. halfella (Dumont, 1928), syn. n., and M. arnaldi (Turati & Krüger, 1936), syn. n. of M. violacellum (Chrétien, 1915); M. escalerellus Schmidt, 1941, syn. n. of M. squalida Meyrick, 1926. Megacraspedus ribbeella (Caradja, 1920), comb. n., M. numidellus (Chrétien, 1915), comb. n., M. albella (Amsel, 1935), comb. n., M. violacellum (Chrétien, 1915), comb. n., and M. grisea (Filipjev, 1931), comb. n. are newly combined in Megacraspedus.

Keywords: Brachyptery, DNA barcoding, Gelechiidae, Lepidoptera, Megacraspedus, new species, Palearctic, taxonomy

Figures 25–30. Megacraspedus adults in dorsal view.
25 M. similellus sp. n. – Holotype, male (RCJJ) 26 M. similellus sp. n. – Paratype, female (RCJJ) 27 M. golestanicus sp. n. – Paratype, male (TLMF) 28 M. tokari sp. n. – Holotype, male, Croatia (RCZT) 29 M. dolosellus (Zeller, 1839) – male, Slovenia (RCZT) 30 M. dolosellus (Zeller, 1839) – female (form dolosellus), Slovenia (RCZT). 

Figures 37–42. Megacraspedus adults in dorsal view.
37 M. dolosellus (Zeller, 1839) – male (form incertellus), Bulgaria (RCZT) 38 M. dolosellus (Zeller, 1839) – male, Greece (ZMUC) 39 M. neli sp. n. – Holotype, male, France (TLMF) 40 M. faunierensis sp. n. – Paratype, male, Italy (TLMF) 41 M. faunierensis sp. n. – Paratype, female, Italy (TLMF) 42 M. gredosensis sp. n. – Paratype, male, Spain (TLMF).

 Figures 73–78. Megacraspedus adults in dorsal view.
 73 Megacraspedus gallicus sp. n. – Holotype, male, Spain (TLMF) 74 M. gallicus sp. n. – Paratype, female, France (TLMF) 75 M. ribbeella (Caradja, 1920) – male, Spain (ZMUC) 76 M. ribbeella (Caradja, 1920) – female, Spain (NHMW) 77 M. libycus sp. n. – Holotype, male, Libya (ZMUC) 78 M. libycus sp. n. – Paratype, female, Morocco (ZSM).

Peter Huemer and Ole Karsholt. 2018. Revision of the Genus Megacraspedus Zeller, 1839, A Challenging Taxonomic Tightrope of Species Delimitation (Lepidoptera, Gelechiidae). ZooKeys. 800: 1-278.  DOI: 10.3897/zookeys.800.26292

      

[Mammalogy • 2018] Talpa martinorum • News from the Balkan refugium: Thrace has An Endemic Mole Species (Mammalia: Talpidae)

 Talpa martinorum

Kryštufek, Nedyalkov, Astrin & Hutterer, 2018

 Bonn zoological Bulletin. 67(1)

Abstract

 We utilized 1084 bp sequences of the cytochrome b gene to assess the taxonomic status of small blind moles from eastern Thrace in Bulgaria and European Turkey. So far, these moles were classified either as Talpa caeca or as T. levantis. Our study showed them to be genetically closer to T. europaea, T. aquitania, and T. occidentalis, albeit not being part of any of these species. We describe them as a new species, Talpa martinorum. n. sp. The new species differs from T. europaea, another mole occupying Thrace, by having a sealed palpebral fissure and a 1st upper molar with no parastyle, and by being smaller. The contemporary distribution range of T. martinorum n. sp. is small and restricted to the Black Sea coast between Burgas (Bulgaria) and Istanbul (Turkey). The species name is an eponym to the married couple Vladimir and Evgeniya Martino, two early students of Balkan mammals.

Key words. Balkans, cytochrome b, cryptic species, species delimitation, Talpa martinorum n. sp. N

Fig. 3. Head (a) in lateral view, tip of nose (b) in dorsal (left) and ventral (right) views, and ventral side of tail in  Talpa martinorum n. sp. Note that the palpebral fissure is covered by transparent skin (a). Museum vouchers PMS 25631 (a), ZFMK 2017.1152 (b) and ZFMK 2017.1151 (c). Not to scale.

Talpa martinorum n. sp. 

Diagnosis. A member of the subgenus Talpa. Medium-sized species with palpebral fissure sealed by a transparent skin (Fig. 3a). First upper molar (M1 ) lacks parastyle (Fig. 6a); the mesostyle is indistinctly bifurcate (Fig. 6a). Pairwise interspecific p-distances (> 9%) are within the range observed between other species of moles (e.g., within Talpa, interspecific distances average ca. 12%).

Etymology. Talpa martinorum n. sp. is an eponym to Vladimir Emmanuilovich Martino (Владимир Эммануилович Мартино, 1888–1961) and Evgeniya Veniaminovna Martino (Евгения Вениаминовнa Мартино, 1894–1979) née Stepanova (Степановa), ethnic Russians who in 1920 escaped the October Revolution by emigrating to the Kingdom of Serbs, Croats and Slovenes (Kingdom of Yugoslavia since 1929). In politically insecure and frequently violent Eastern Europe of the 20th century, the Martinos were refugees for more than three decades. In 1949 they moved to Bulgaria and in 1955 returned to Russia (at that time still Soviet Union). Inspired by G. S. Miller’s (1912) “Catalogue of the Mammals of Western Europe” they initiated mammal research in south-eastern Europe and named, among others, two Balkan endemics: Dinaromys bogdanovi (V. Martino & E. Martino, 1922) (Martino & Martino 1922: 413) and Talpa stankovici V. Martino & E. Martino, 1931 (Martino & Martino 1931: 53) (Fig. 8). While Vladimir had a degree in Biology from the University in Novorossiysk (1913), Evgeniya had no formal academic education. Despite this, she attained competency in mammalogy and successfully collaborated with her spouse, both in the field and cabinet. Although they published several joint papers (as V. and E. Martino), the contribution by Evgeniya is mainly ignored and she is nearly anonymous today (cf. Beolens et al. 2009). At least nine subspecific names for mammals with the epithet martinoi (see Appendix 3) were proposed by mammalogists between 1935 and 1971, and all are eponyms to Vladimir Martino. With the name martinorum n. sp. we stress the equal share by Evgeniya in the tandem “V. et E. Martino” and correct the injustice done to her contribution in the past decades. Along with Dorothea Bate (1878–1951) and Gabriele Neuhäuser (1911–1998) Evgeniya was one of the early women who studied the taxonomy of Palaearctic mammals already between the two great wars. In addition to their publications, a renowned legacy of Vladimir and Evgeniya Martino is their meticulously prepared and carefully labelled mammal collection, deposited primarily in the Natural History Museum London and the Zoological Institute and Museum of the Russian Academy of Sciences in St. Petersburg. For biographies of Vladimir Martino, see Mezentsev (1961), Paspalev (1962), Pusanov (1962), Zimmermann (1962), Gus’kov (1965), Taranenko (1999), and Boreiko (2001).

Boris Kryštufek, Nedko Nedyalkov, Jonas J. Astrin and Rainer Hutterer. 2018. News from the Balkan refugium: Thrace has An Endemic Mole Species (Mammalia: Talpidae).  Bonn zoological Bulletin. 67(1); 41–57. 

ZoologicalBulletin.de/BzB_Volumes/Volume_67_1/041_057_BzB67_1_krystufek_et_al.pdf

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[Mammalogy • 2018] Multi-locus Phylogeny of the Tribe Tragelaphini (Mammalia, Bovidae) and Species Delimitation in Bushbuck: Evidence for Chromosomal Speciation Mediated by Interspecific Hybridization

in Hassanin, Houck, Tshikung,et al., 2018. 

 DOI: 10.1016/j.ympev.2018.08.006  

Highlights

• Two species of bushbuck: Tragelaphus scriptus in NW Africa and T. sylvaticus in SE Africa.

• The two species have 2n = 57M/58F and 2n = 33M/34F chromosomes, respectively.

• T. scriptus is related to T. angasii with mtDNA, and to T. sylvaticus with nuDNA.

 • Mitochondrial introgressive hybridization in the common ancestor of T. scriptus.

• Evidence for chromosomal speciation after an event of interspecific hybridization.

Abstract

The bushbuck is the most widespread bovid species in Africa. Previous mitochondrial studies have revealed a polyphyletic pattern suggesting the possible existence of two distinct species.

To assess this issue, we have sequenced 16 nuclear genes and one mitochondrial fragment (cytochrome b gene + control region) for most species of the tribe Tragelaphini, including seven bushbuck individuals belonging to the two divergent mtDNA haplogroups, Scriptus and Sylvaticus. Our phylogenetic analyses show that the Scriptus lineage is a sister-group of Sylvaticus in the nuclear tree, whereas it is related to Tragelaphus angasii in the mitochondrial tree. This mito-nuclear discordance indicates that the mitochondrial genome of Scriptus was acquired by introgression after one or several past events of hybridization between bushbuck and an extinct species closely related to T. angasii. The division into two bushbuck species is supported by the analyses of nuclear markers and by the karyotype here described for T. scriptus (2n= 57M/58F), which is strikingly distinct from the one previously found for T. sylvaticus (2n= 33M/34F). Molecular dating estimates suggest that the two species separated during the Early Pleistocene after an event of interspecific hybridization, which may have mediated massive chromosomal rearrangements in the common ancestor of T. scriptus.

Keywords: Spiral-horned antelopes, species complex, introgressive hybridization, chromosomes, cytogenetics

Figure 3. Bayesian divergence times (in million years ago, Mya) estimated using the nuclear concatenation of 16 genes (A) or the mitochondrial fragment (B). Divergence times were estimated with BEAST 2.4.7 (see main text for details). Taxa other than Tragelaphini were removed from the figures. Bold values at the nodes are mean ages. Grey bars and values between brackets represent the 95% Highest Posterior Density (HPD) interval.

 Alexandre Hassanin, Marlys L. Houck, Didier Tshikung, Blaise Kadjo, Heidi Davis and Anne Ropiquet. 2018. Multi-locus Phylogeny of the Tribe Tragelaphini (Mammalia, Bovidae) and Species Delimitation in Bushbuck: Evidence for Chromosomal Speciation Mediated by Interspecific Hybridization. Molecular Phylogenetics and Evolution. In Press.  DOI: 10.1016/j.ympev.2018.08.006 

[Herpetology • 2018] Computational Molecular Species Delimitation and Taxonomic Revision of the Gecko Genus Ebenavia Boettger, 1878

Ebenavia safari

Hawlitschek, Scherz, Ruthensteiner, Crottini & Glaw, 2018

 DOI: 10.1007/s00114-018-1574-9 

 MarkScherz.com

Abstract

Cryptic species have been detected in many groups of organisms and must be assumed to make up a significant portion of global biodiversity. We study geckos of the Ebenavia inunguis complex from Madagascar and surrounding islands and use species delimitation algorithms (GMYC, BOLD, BPP), COI barcode divergence, diagnostic codon indels in the nuclear marker PRLR, diagnostic categorical morphological characters, and significant differences in continuous morphological characters for its taxonomic revision. BPP yielded ≥ 10 operational taxonomic units, whereas GMYC (≥ 27) and BOLD (26) suggested substantial oversplitting. In consequnce, we resurrect Ebenavia boettgeri Boulenger 1885 and describe Ebenavia tuelinae sp. nov., Ebenavia safari sp. nov., and Ebenavia robusta sp. nov., increasing the number of recognised species in Ebenavia from two to six. Further lineages of Ebenavia retrieved by BPP may warrant species or subspecies status, but further taxonomic conclusions are postponed until more data become available. Finally, we present an identification key to the genus Ebenavia, provide an updated distribution map, and discuss the diagnostic values of computational species delimitation as well as morphological and molecular diagnostic characters.

Keywords: BOLD, Operational Taxonomic Unit, Madagascar clawless gecko, Integrative taxonomy, Taxonomic inflation, Species complex 

Genus Ebenavia Boettger, 1878 

Type species: Ebenavia inunguis Boettger, 1878

Diagnosis and description: A comprehensive diagnosis and description of the genus Ebenavia are provided in Nussbaum and Raxworthy (1998). Our data on the genus leads us to agree with the findings of that study, except that we were not able to confirm the presence of claws on the pes of any of the females we examined, as reported by these authors, in agreement with Boettger (1878) and many subsequent authors. We did not examine any specimens of E. maintimainty. 

Content: Ebenavia inunguis Boettger, 1878; E. boettgeri Boulenger, 1885; E. maintimainty Nussbaum & Raxworthy, 1998; and three new species described below. 

Distribution: Madagascar except most of the central and western areas; all major island areas of the Comoros Archipelago; Pemba Island (Tanzania); Mauritius.

Ebenavia inunguis Boettger, 1878 

Ebenavia boettgeri Boulenger, 1885, bona species 

Ebenavia tuelinae sp. nov. 

Etymology: A matronym dedicated to the first author’s partner Tülin (alternative spelling Tuelin) for her ceaseless support of this and other works and for her excellent spotting abilities in the field.

Ebenavia safari, a newly described species from northern Madagascar, which also occurs on the Tanzanian island of Pemba

 DOI: 10.1007/s00114-018-1574-9  

MarkScherz.com 

Ebenavia safari sp. nov.

Etymology: ‘Safari’ means ‘voyage’ in the Kiswahili and Comoran (Shimaoré) languages spoken across the range of this species outside Madagascar. The name was chosen because this species dispersed over surprisingly long distances across the open ocean. It is treated as an unlatinised, invariable noun in apposition.

Ebenavia robusta sp. nov. 

Etymology: The specific name is the feminine form of the Latin adjective ‘robustus’, meaning ‘robust’ or ‘sturdy’. It was chosen because this species is the largest and most sturdily built member of this genus of small geckos.

Oliver Hawlitschek, Mark D. Scherz, Bernhard Ruthensteiner, Angelica Crottini and Frank Glaw. 2018. Computational Molecular Species Delimitation and Taxonomic Revision of the Gecko Genus Ebenavia Boettger, 1878. The Science of Nature. 105:49. DOI: 10.1007/s00114-018-1574-9 

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[Ichthyology • 2018] Spectrolebias gracilis • A New Miniature Cryptic Species of the Seasonal Killifish Genus Spectrolebias (Cyprinodontiformes, Aplocheilidae) from the Tocantins River basin, central Brazil

Spectrolebias gracilis  Costa & Amorim, 2018 

DOI: 10.3897/zse.94.28085 

Abstract

The miniature seasonal killifish Spectrolebias costae, first described for the middle Araguaia River basin, has been also recorded from two areas in the middle Tocantins River basin, from where male specimens exhibit some differences in their colour pattern. Analyses directed to species delineation (GMYC and bPTP), using a fragment of the mitochondrial gene COI, strongly support two species, S. costae from the Araguaia River basin and a new species from the Tocantins River basin. Spectrolebias gracilis sp. n. is described on the basis of specimens collected from two localities separated by about 530 km, Canabrava River floodplains near Alvorada do Tocantins and Tocantins River floodplains near Palmeirante. Field inventories were unsuccessful in finding additional populations in the region, which is attributed to the high environmental degradation, including several large dams that have permanently inundated typical killifish habitats. Spectrolebias gracilis is member of a clade also including S. costae, S. inaequipinnatus, and S. semiocellatus, diagnosed by having the dorsal and anal fins in males with iridescent dots restricted to their basal portion, caudal fin in males hyaline, and caudal-fin base with two pairs of neuromasts. Within this clade, a single miniaturisation event is supported for the most recent common ancestor of the subclade comprising S. costae and S. gracilis, which differ from other congeners by reaching only about 20 mm standard length as maximum adult size.

Key Words: Amazon, Biodiversity conservation, Integrative taxonomy, Miniaturization, Molecular taxonomy, Species delimitation

Taxonomic accounts

Spectrolebias gracilis sp. n.

Diagnosis: Spectrolebias gracilis is member of a clade endemic to the Araguaia-Tocantins River System, also including S. costae, S. semiocellatus Costa & Nielsen, 1997 and S. inaequipinnatus Costa & Brasil, 2008, and morphologically diagnosed by: dorsal and anal fins in males with iridescent dots restricted to the basal portion of fins (vs. scattered over the whole fin), caudal fin in males hyaline (vs. variably coloured, usually dark red or grey), caudal-fin base with two pairs of neuromasts (vs. one). Spectrolebias gracilis is similar to S. costae and distinguished from S. semiocellatus and S. inaequipinnatus by having dorsal fin rounded in males (vs. pointed), dark brown to black pigmentation on the flank in males (vs. light brownish grey), and a subdistal bright blue stripe on the dorsal and anal fins in males (vs. subdistal bright blue absent). Spectrolebias gracilis differs from S. costae by the iridescent light blue colour pattern in males, comprising the presence of 10–12 small blue spots irregularly arranged on opercle, surrounded by diffuse blue iridescence (Fig. 4; vs. 6–8 small blue spots, usually arranged in three vertical series, contrasting with dark brown colour ground, Fig. 3) and one or two series of dots irregularly arranged on the basal portion of the dorsal fin (Fig. 4; vs. blue dots arranged in single longitudinal row close to fin base, Fig. 3).

....

Figure 4. Spectrolebias gracilis sp. n., UFRJ 6440, holotype, male, 19.2 mm SL; Canabrava floodplains. 

Figure 5. Spectrolebias gracilis sp. n., UFRJ 6441, paratype, female, 17.8 mm SL; Canabrava floodplains.

Etymology: From the Latin gracilis, meaning thin, referring to the thin body of the small-sized new species.

Distribution and habitat: Spectrolebias gracilis is known from temporary pools of two localities of the middle Tocantins River basin, central Brazil (Fig. 6). In both localities pools were shallow, about 80 cm in deeper places, and densely occupied by aquatic vegetation.

Figure 3. Specrolebias costae, UFRJ 3549, male, 18.8 mm SL; das Mortes River floodplains.

Wilson J. E. M. Costa and Pedro F. Amorim. 2018. A New Miniature Cryptic Species of the Seasonal Killifish Genus Spectrolebias from the Tocantins River basin, central Brazil (Cyprinodontiformes, Aplocheilidae).  Zoosystematics and Evolution. 94(2): 359-368.  DOI: 10.3897/zse.94.28085

[Botany • 2018] Argyreia gyrobracteata • Species Delimitation of Some Argyreia (Convolvulaceae) Using Phenetic Analyses: Insights from Leaf Anatomical Data Reveal A New Species from northeastern Thailand

Argyreia gyrobracteata Traiperm & Chitchak

in Chitchak, Traiperm, Staples, et al., 2018. 

   DOI:  10.1139/cjb-2017-0108 

ABSTRACT
Argyreia Lour. is one of the most taxonomically complex genera of the morning glory family (Convolvulaceae). The number of named species is now 135, and new species are regularly being described. There are several species complexes that are morphologically similar and difficult to tell apart. Therefore, the aim of this study is to explore the species identification criteria for Argyreia, especially new sources for taxonomically informative characters. Ten accessions representing three morphologically similar Argyreia operational taxonomic units (OTUs) were collected and their anatomical characters were investigated using the leaf peeling technique and paraffin sectioning method. Anatomical character states were analyzed using two phenetic analysis methods: clustering analysis (CA) and principal component analysis (PCA). Three distinct clusters were clearly separated in both PCA and CA at the internal similarity coefficient of 0.48 with a high R-value of 0.89757. Nineteen effectively distinguishable character states were derived from the high loadings of the first two components. In conclusion, two of the separated groups were matched with known species, and the third separated group is here delineated as a new species. Therefore, a new species, Argyreia gyrobracteata Traiperm & Chitchak, is described and illustrated together with ecological data and a preliminary conservation assessment.

Keywords: cluster analysis, cryptic species, morphometrics, principal component analysis, species delimitation

Fig. 7. Argyreia gyrobracteata Traiperm & Chitchak sp. nov.:
 (A) flower in front view; (B) flower in side view; (C) interaction with an insect visitor, oriental carpenter bee (Xylocopa nasalis); (D) plant habit

(all photos taken by P. Rattanakrajang from live plants vouchered as P. Rattanakrajang et al. 104).

Argyreia gyrobracteata Traiperm & Chitchak, sp. nov. 

TYPE: Thailand. Ubon Ratchathani, Sirindhorn district, ..., in the edge of dipterocarp forest, August 2016, P. Rattanakrajang, N. Chitchak & P. Traiperm 110 (holotype BKF!; isotypes K!, QBG!)

DIAGNOSIS: The new species is similar to A. mekongensis in having a white campanulate corolla, but differs from that species by the linear-oblong to narrowly lanceolate bract shape (versus lanceolate or oblong-lanceolate), the curly or twisted bract orientation (versus falcate), the larger sepals, and the multicellular, uniseriate, villous trichomes restricted to a small, dense, triangular patch on the adaxial side of the filaments, above the insertion point of the filaments on the corolla tube (versus dispersed in a band 3–5 mm wide surrounding the free filament, above the insertion point on the corolla tube).
....

DISTRIBUTION: Known so far from discrete populations in two different districts within Ubon Ratchathani province, Thailand. One population is close to the border of Thailand–Laos and possibly A. gyrobracteata occurs across the border in Laos.

ETYMOLOGY: The specific epithet refers to the curly/twisted bracts of this species, which have not been observed in any other known species of Argyreia. 

 Natthaphong Chitchak, Paweena Traiperm, G. Staples, Pantamith Rattanakrajang and Pirada Sumanona. 2018. Species Delimitation of Some Argyreia (Convolvulaceae) Using Phenetic Analyses: Insights from Leaf Anatomical Data Reveal A New Species. Botany. 96(4); 217-233.  DOI:  10.1139/cjb-2017-0108 

Researchgate.net/publication/324267590_Species_delimitation_of_some_Argyreia_Convolvulaceae_using_phenetic_analyses_insights_from_leaf_anatomical_data_reveal_a_new_species

[Ornithology • 2018] Molecular Phylogenetics and Species Limits in A Cryptically Coloured Radiation of Australo-Papuan Passerine Birds (Pachycephalidae: Colluricincla)

in Marki, Fjeldså, Irestedt & Jønsson, 2018.

  DOI: 10.1016/j.ympev.2018.02.029 

Highlights: 

• A time-calibrated molecular phylogeny of all shrikethrushes (Colluricincla).

• C. megarhyncha consists of seven unrecognized species.

 • A new taxonomy for C. megarhyncha is proposed.

 • C. megarhyncha melanorhyncha belongs in the genus Pachycephala.

Abstract

Detailed knowledge of species limits is an essential component of the study of biodiversity. Although accurate species delimitation usually requires detailed knowledge of both genetic and phenotypic variation, such variation may be limited or unavailable for some groups. In this study, we reconstruct a molecular phylogeny for all currently recognized species and subspecies of Australasian shrikethrushes (Colluricincla), including the first sequences of the poorly known C. tenebrosa. Using a novel method for species delimitation, the multi-rate Poisson Tree Process (mPTP), in concordance with the phylogenetic data, we estimate species limits in this genetically diverse, but phenotypically subtly differentiated complex of birds. In line with previous studies, we find that one species, the little shrikethrush (C. megarhyncha) is characterized by deep divergences among populations. Delimitation results suggest that these clades represent distinct species and we consequently propose a new classification. Furthermore, our findings suggest that C. megarhyncha melanorhyncha of Biak Island does not belong in this genus, but is nested within the whistlers (Pachycephala) as sister to P. phaionota. This study represents a useful example of species delimitation when phenotypic variation is limited or poorly defined.

Keywords: Passerine birds, Corvides, Australia, New Guinea, Cryptic species, Species delimitation

Fig. 2. A time-calibrated maximum clade credibility tree of Colluricincla shrikethrushes derived from the divergence estimation of the one mitochondrial gene and three nuclear introns in BEAST. Posterior probabilities are shown for major nodes. Error bars show the 95% highest posterior density intervals for the divergence time estimates. Vertical bars indicate the species identified by the mPTP-approach. Species names reflect the new taxonomy proposed in this study. Illustrations are watercolours by Jon Fjeldså and show all eleven delimited species. For species that exhibit significant within-species morphological variation, multiple illustrations are shown.

Fig. 1. Sampling localities for the 129 sequences included in this study. Colours and taxon names refer to the eleven species delimited in this study. 

Conclusion: 

In this study, we present a densely sampled molecular phylogeny for the Australasian shrikethrushes. Our results suggest that species diversity within this complex is underestimated, and we consequently propose a revised classification. Nonetheless, we view our proposed taxonomy as preliminary and hope that this study may stimulate further study of species limits in this group. In particular, we believe that increased study of behaviour, contact zone dynamics and vocalizations coupled with the analysis of genome-wide data are likely to be promising in this respect.

 Petter Z. Marki, Jon Fjeldså, Martin Irestedt and Knud A. Jønsson. 2018. Molecular Phylogenetics and Species Limits in A Cryptically Coloured Radiation of Australo-Papuan Passerine Birds (Pachycephalidae: Colluricincla). Molecular Phylogenetics and Evolution. 124; 100-105. DOI: 10.1016/j.ympev.2018.02.029 

[Herpetology • 2018] A Pan-Amazonian Species Delimitation: High Species Diversity within the Genus Amazophrynella (Anura: Bufonidae)

 (A–B) Amazophrynella minuta; (C–D) A. teko sp. nov.; (E–F) A. siona sp. nov.; (G–H) A. xinguensis sp. nov.;

(O) A. matses; (Q) A. javierbustamantei; (S) A. vote; (U) A. moisesii sp. nov.

Rojas, Fouquet, Ron, Hernández-Ruz, Melo-Sampaio, et al​., 2018. 

 DOI: 10.7717/peerj.4941

photos by Rommel R. Rojas, Antoine Fouquet, Santiago R. Ron, Emil Hernándes-Ruz, Juan Carlos Chapparro,  Robson W. Ávila & Paulo R. Melo-Sampaio.

Abstract

Amphibians are probably the most vulnerable group to climate change and climate-change associate diseases. This ongoing biodiversity crisis makes it thus imperative to improve the taxonomy of anurans in biodiverse but understudied areas such as Amazonia. In this study, we applied robust integrative taxonomic methods combining genetic (mitochondrial 16S, 12S and COI genes), morphological and environmental data to delimit species of the genus Amazophrynella (Anura: Bufonidae) sampled from throughout their pan-Amazonian distribution. Our study confirms the hypothesis that the species diversity of the genus is grossly underestimated. Our analyses suggest the existence of eighteen linages of which seven are nominal species, three Deep Conspecific Lineages, one Unconfirmed Candidate Species, three Uncategorized Lineages, and four Confirmed Candidate Species and described herein. We also propose a phylogenetic hypothesis for the genus and discuss its implications for historical biogeography of this Amazonian group.

Amazophrynella teko sp. nov.

 Amazonella sp. Guianas (Fouquet et al., 2012a: 829, French Guiana [in part])

Amazophrynella sp. Guianas (Fouquet et al., 2012b: 68, French Guiana [in part])

Amazophrynella sp. Guianas (Rojas et al., 2015: 85, French Guiana [in part])

Amazophrynella sp1. (Fouquet et al., 2015: 365, French Guiana [in part])

Amazophrynella sp. aff. manaos (Rojas et al., 2016: 49, French Guiana [in part])

Diagnosis. An Amazophrynella with (1) SVL12.9–15.8 mm in males, 17.9–21.5 mm in females; (2) snout acute in lateral view; upper jaw, in lateral view, protruding beyond lower jaw; (3) texture of dorsal skin granular; (4) cranial crest, vocal slits and nuptial pads absent; (5) dorsum covered by abundant rounded granules; (6) abundance of granules on tympanic area, on edges of upper arms and on dorsal surface of arms; (7) ventral skin highly granular; (8) fingers slender, basally webbed; (9) finger III relatively short (HAL/SVL 0.2–0.22 mm, n = 30); (10) finger I shorter than finger II; (11) palmar tubercle protruding and elliptical; (12) hind limbs relatively short (TAL/SVL 0.48–0.49, n = 30); (13) toes slender, basally webbed; in life: (14) venter cream; small blotches on venter.

Distribution and natural history. Amazophrynella teko sp. nov. have been recorded from the district of Saint Laurent du Marioni, Saint Georges and Camopi, French Guiana, the state of Amapá, Brazil and in the southern region of Suriname (A Fouquet, pers. obs., 2017). It occurs at elevations ranging from 70 m a.s.l. to 350 m a.s.l. The species is diurnal and crepuscular but is also active at night during peak breeding period, which normally occurs at the beginning of the rainy season (January–February). This species shows a conspicuous sexual dimorphism, with males being much smaller than females. The conservation status of this species remains unknown. The habitat destruction and pollution must affect their populations; however, due to its abundance we believe that this species probably needs not be classified above Least Concern category.

Etymology. The specific epithet is a noun in apposition and refers to the name of the Teko Amerindians who occupy the southern half of French Guiana; the area occupied by the Teko tribe also encompasses the type locality.

Amazophrynella siona sp. nov.

 Atelopus minutus: (Duellman & Lynch, 1969: 238, Sarayacu [Ecuador])

Dendrophryniscus minutus (Duellman, 1978: 120, Santa Cecilia [Ecuador])

Dendrophryniscus minutus (Duellman & Mendelson III 1995: 336, vicinities of San Jacilllo and Teniente Lopez [Peru])

Amazonela cf. minutus “western Amazonia” (Fouquet et al., 2012a: 829, “western Amazonia”, Ecuador [in part])

Amazophrynella cf. minutus “western Amazonia” (Fouquet et al., 2012a: 68, “western Amazonia”, Ecuador [in part])

Amazophrynella aff. minuta “western Amazonia” (Rojas et al., 2015: 84, “western Amazonia”, Ecuador [in part])

Amazophrynella aff. minuta (Rojas et al., 2016: 49, “western Amazonia”, Ecuador [in part])

Diagnosis. An Amazophrynella with (1) SVL 11.5–14.7 mm in males, 16.1–20.0 mm in females; (2) snout acute in lateral view; upper jaw, in lateral view, protruding beyond lower jaw; (3) texture of dorsal skin finely granular; (4) cranial crests, vocal slits and nuptial pads absent; (5) small granules from the outer edge of the mouth to upper arm; (6) ventral skin granular; (7) tiny granules on ventral surfaces; (8) fingers slender, basally webbed; (9) finger III relative short (HAL/SVL 0.20–0.21, n = 62); (10) finger I shorter than finger II; (11) palmar tubercle rounded; (12) hind limbs relatively large (TAL/SVL 0.5–0.52, n = 62); (13) toes lacking lateral fingers; in life: (14) venter reddish brown; yellow blotches on venter.

Distribution and natural history. Amazophrynella siona sp. nov. have been recorded from Ecuador, in Provinces of Orellana, Sucumbíos and Pastaza and Peru in the Province Andoas, northern Loreto Department. It occurs at elevations ranging from 200–900 m a.s.l. The species is found in the leaf litter of primary and secondary forest, terra firme or flooded forest, and swamps. It is active during the day; at night individuals rest on leaves, usually less than 50 cm above ground. It breeds throughout the year (Duellman, 1978). This species shows conspicuous sexual dimorphism, with males being much smaller than females. The amplexus is axillar. Eggs are pigmented; males call from amidst leaf litter. Duellman & Lynch (1969) reported that this species deposited its eggs in gelatinous strands 245–285 mm long, with 245–291 eggs. It can be abundant at some sites (e.g., Cuyabeno reserve; SR Ron, pers. obs., 2018) Given its large distribution range (>20,000 km2) which also includes vast protected areas and locally abundant populations, we suggest assignment this species to the Least Concern category.

Etymology. The specific epithet is a noun in apposition and refers to the Siona, a western Tucanoan indigenous group that inhabits the Colombian and Ecuadorian Amazon. The Siona inhabit the Cuyabeno Lakes region, an area where Amazophrynella siona sp. nov. is be abundant. While working in his undergraduate thesis in the early 1990s, SRR lived with the Siona at Cuyabeno. The Siona chief, Victoriano Criollo, had an encyclopedic knowledge of the natural history of the Amazonian forest, superior in extent and detail to that of experienced biologists. His death, a few years ago, represents one of many instances of irreplaceable loss of traditional knowledge triggered by cultural change among Amazonian Amerindians.

  (C–D) Amazophrynella teko sp. nov. photo by Antoine Fouquet; 
(E–F) A. siona sp. nov. photo by Santiago R. Ron;
(G–H) A. xinguensis sp. nov. photo by Emil Hernándes-Ruz; 
(U–V) A. moisesii sp. nov. photo by Paulo R. Melo-Sampaio.

Amazophrynella xinguensis sp. nov.

Diagnosis. An Amazophrynella with (1) SVL 17.0–20.0 mm in males, 22.4–26.3 mm in females; (2) snout pointed in lateral view; (3) upper jaw, in lateral view, protruding beyond lower jaw; 4) tympanums, vocal sac, parotid gland and cranial crest not evident; (5) texture of dorsal skin highly granular; (6) abundance of small tubercles on dorsum, on upper arm and on arms; (7) texture of ventral skin granular; (8) fingers I and II basally webbed; (9) finger III relative short (HAL/SVL = 0.20–0.22, n = 18); (10) thumb larger and robust; (11) finger I larger or equal than finger II, FI = 2.1 vs. FII = 2.1 in adult males, n = 5 and FI = 2.8 mm, vs. FII = 2.9 mm, in adult females, n = 13; (12) palmar tubercle ovoid; (13) toes slender, basally webbed; in life: (14) venter greyish; black dots on venter.

Distribution and natural history. Amazophrynella xinguensis sp. nov. have been recorded from State of Pará, Brazil, at three localities: PDS Virola Jatoba, municipality of Anapú, Fazenda Paraiso, municipality of Senador José Porfirio (right bank of Xingu River) and Ramal dos Cocos, municipality of Altamira (left bank of Xingu River), all of them in area of influence of the Belo Monte dam. It occurs in elevations of 86–106 m a.s.l. This species is found amidst leaf litter. The amplexus is axillar (Fig. 18C). Reproduction occurs in the rainy season in tiny puddles. Males were found hidden in the leaf litter. Tadpoles and advertisement call are unknown. The conservation status of this species remains unknown, but the recent construction of the Belo Monte hydroelectric complex on the Xingu River represents a threat to the population status of this species.

Etymology. The specific epithet refers to geographic distribution of the species within the lower Xingu River basin, Brazil.

Amazophrynella moisesii sp. nov.

Dendrophryniscus minutus (Bernarde et al. 2011: 120 plate 2, Fig. d)

Amazophrynella minuta (Bernarde et al. 2013: 224, 227 plate 7 Fig. c; Miranda et al. 2015: 96)

Diagnosis. An Amazophrynella with (1) SVL 12.2–15.8 mm in males, 16.4–20.9 mm in females; (2) snout acuminate in lateral view, upper jaw, in lateral view, protruding beyond lower jaw; (3) snout length protuberant, large for the genus (SL/HL = 0.48–0.5); (4) cranial crest, vocal slits and nuptial pads absent; (5) small tubercles on upper arms and posterior area of tympanums; (6) texture of dorsal skin tuberculate; (7) texture of ventral skin highly granular (8) finger III relative large (HAL/SVL 0.23–0.25, n = 28); (9) fingers slender, basally webbed; (10) finger I shorter than finger II; (11) palmar tubercle elliptic; (12) hind limbs relatively large (TAL/SVL 0.51–0.53, n = 28); (13) toes slender basally webbed; in life: (14) venter pale yellow; small irregular dots on venter.

Distribution and natural history. Amazophrynella moisesii sp. nov. have been recorded from Brasil. State of Acre: municipalities of Cruzeiro do Sul, Mâncio Lima, Porto Walter and Tarauacá; State of Amazonas: municipality of Envira. Peru: Department of Huanuco, Panguana, Rio Llullapichis. Due to its abundance and presence in conservation units of Brazil (Floresta Estadual do Gregório, Reserva Extrativista do Alto Juruá and Parque Nacional da Serra do Divisor) we recommend the IUCN Least Concern category.

Etymology. The specific epithet refers to Dr. Moisés Barbosa de Souza, a Brazilian biologist, professor and friend at the Universidade Federal do Acre (UFAC), to whom we dedicate this species in recognition of his contributions to herpetological research and amphibian conservation in the state of Acre, Brazil.

Figure 25: Confirmed candidate species (CCS) of Amazophrynella. (A–B) Amazophrynella minuta photo by Rommel R. Rojas; (C–D) A. teko sp. nov. photo by Antoine Fouquet; (E–F) A. siona sp. nov. photo by Santiago R. Ron; (G–H) A. xinguensis sp. nov. photo by Emil Hernándes-Ruz; (I–J) A. bokermanni photo by Marcelo Gordo; (K–L) A. manaos photo by Rommel R. Rojas. (M–N) A. amazonicola photo by Rommel R. Rojas. (O–P) A. matses photo by Rommel R. Rojas; (Q–R) A. javierbustamantei photo by Juan Carlos Chapparro; (S–T) A. vote photo by Robson W. Ávila; (U–V) A. moisesii sp. nov. photo by Paulo R. Melo-Sampaio.

Rommel R. Rojas, Antoine Fouquet, Santiago R. Ron, José Hernández-Ruz, Paulo R. Melo-Sampaio, Juan C. Chaparro, Richard C. Vogt, Vinicius Tadeu de Carvalho, Leandra Cardoso Pinheiro, Robson W. Avila, Izeni Pires Farias, Marcelo Gordo and Tomas Hrbek​. 2018. A Pan-Amazonian Species Delimitation: High Species Diversity within the Genus Amazophrynella (Anura: Bufonidae). PeerJ. 6:e4941  DOI: 10.7717/peerj.4941