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3.3.1 Metachirus aritanai new species 3.3.1.1 Synonymy (Didelphys [Metachirus]) nudicaudata: Trouessart 1989:1236; part, not M. nudicaudatus (Geoffroy St. Hilaire 1803). [Philander] nudicaudatus: Pine 1973:391; part, not M. nudicaudatus (Geoffroy St. Hilaire 1803). M. nudicaudatus: Patton et al. 2000:61–64; not M. nudicaudatus (Geoffroy St. Hilaire 1803); first evidence that Metachirus from Pará represented a new taxon. M [etachirus]. n [udicaudatus]. nudicaudatus: Gardner and Dagosto 2007:37; part, not M. nudicaudatus (Geoffroy St. Hilaire 1803). Holotype: An adult male consisting of skin and skull (INPA 2831, original number CS 83), in addition to tissue sample preserved in ethanol, collected on 8 July 1999 by Maria Nazareth F. da Silva in the Tapirapé-Aquiri National Forest, Marabá, Pará, Brazil (05° 46′ 53.10″ S; 50° 31′ 48.60″ W; Figure 1). MC, male from the Carajás region; FC, females from the Carajás region; MN, male of M. nudicaudatus; FN, females of M. nudicaudatus; MM, males of M.myosuros; FM,females of M.myosuros; MCC,maximal cranial circumference;NAS,nasal bone length;RWJ,rostrum width between the jugal bones;RWF, rostrum width between the frontal bonés;NW, greater nasal width; CI, interorbital constriction; PC,postorbital constriction; WZ,width between zygomatic arches; GCW, greatest cranial width; MAX, length of the maxillary row of teeth; LUM, length of the upper molar series; CAN, width between the lateral margins of the upper canines; WPP, width of the palatal projection; WM, width between the external faces of the third molars; LLM, length of the lower molar series; JL, jaw length. Paratypes ( N = 19): Eight adult females (N = 8) consisting of skins, skulls and tissues preserved in ethanol; two adult males (N = 2) consisting of skins, skulls and tissues preserved in ethanol; and five sub-adult and young individuals (N = 9) consisting of specimens preserved in fluid, skull and tissues. Adult females: two specimens (PSA 06; PSA 200) were collected by Cleuton L. Miranda in the Tapirapé-Aquiri National Forest, municipality of Marabá, Pará, Brazil and will be cataloged in the mammal collection of MPEG. Three specimens were collected by Natalia C. Ardente, two (MN 78307; MN 79953) in the Carajás National Forest, municipality of Marabá, Pará, Brazil, and one (MN 75549) in the Tapirapé-Aquiri National Forest, municipality of Marabá, Pará, Brazil. The three remaining specimens (LBCE 14843, 14845, 14859) were collected by Fabiana Caramaschi, Cibele R. Bonvicino and Paulo Sérgio D’ Andrea in the Vila Santa Cruz, Araguatins municipality, State of Tocantins, Brazil, and will be cataloged in the mammal collection of MN. Adult male specimens: two specimens (MN 73864; MN 78309) were collected by Natalia C. Ardente in the Carajás National Forest, State of Pará, Brazil. Sub-adult and young specimens: nine specimens were collected by Marco Antonio R. Junior and Stepheson Abrantes; five specimens (MAR 890, 1165, 1292, 1407, 1409) were collected in the Caxiuanã National Forest, municipality of Portel, Pará, Brazil, and will be cataloged in the mammal collection of MPEG; Four specimens (CN 78, 83, 91, 240) were collected in Bom Futuro on the left margin of the middle Xingu River, Pará, Brazil, and will be cataloged in MPEG. Type locality: Tapirapé-Aquiri National Forest, municipality of Marabá, state of Pará, Brazil (05° 46′ 53.10″ S; 50° 31′ 48.60″ W; Figure 1). Etymology: The name M. aritanai is given in honor of the great leader Aritana Yawalapiti (1949–2020), of the Yawalapiti ethnicity, from the upper Xingu River region and in honour of all indigenous peoples of Brazil. The leader Aritana died in August 2020, a victim of covid-19, representing a great loss for the Xingu people, all indigenous peoples, and all Brazilians. Distribution: M. aritanai new species is restricted to the Xingu-Tocantins interfluve, the endemism center Xingu sensu Silva et al. (2005). As far as we know M. aritanai new species is endemic to this biogeographic region (Figure 1). The species is bounded to the north by the Amazon River, to the west by the Xingu River and to the east by the Tocantins River. The southern boundary cannot yet be clearly defined. According to the consensus topology and times of divergence presented in this study, a sister species of M. aritanai new species, M. nudicaudatus, occurs north of the Amazon River (Figures 2 and 3). Diagnosis: M. aritanai new species is characterized by the combination of the following characters: general colouration light brown with speckles of gold on the back; small amount of rust hair behind the ears; supra-ocular spots of cream colour; cheeks with the same colouration as supra-ocular spots; brown and gold-speckled face; absence of median stripe on the rostrum; the presence of a narrow, blackened dorsal line at the top of the head, usually extending to the nape of the neck; clear ventral surface, usually pure cream, wide interorbital region (average = 12.12 mm) and with incipient ridges; rounded and inflated cranial cavity, with large maxillopalatine (average = 24.68 mm) and fenestrae; small entoconids; absent or incipient temporal crest. Figures 5–7 show the mentioned characters, except for the morphology of the entoconid, which is similar to that of M. nudicaudatus and may be seen in Figure 15, page 46 in Voss et al. (2019). 3.4 Description 3.4.1 External morphology Combined length of adult head and body ca. 230 mm on average (ranging 243–275 mm); adult weight ca. 380 g. Midrostral fur cream coloured with apical portion dark brown (most of the cream-coloured hairs have black tips, but there are also completely blackened or cream hairs in lesser quantities, somewhat like the dorsal coat). Dark median rostral stripe absent or inconspicuous. Dark circumocular mask present (blackened), extending posteriorly to the base of the ear, continuous with the coronal fur. Narrow dark brown dorsal line starting at the coronal spot and extending to the end of the nape. Throat gland present in adult males. Small pure cream supraocular spots present, usually in the shape of an inverted triangle, varying in size. Pure cream-coloured cheeks, of the same colour as the supraocular spots. Completely brown ears. Presence of small amounts of rust hair behind the base of the ears. General colouration light brown speckled with gold. Dorsal colour pale brown, with grey-based hairs, cream stripe over most of its length and short blackish apex, giving the back a golden speckled appearance. Dorsal hairs usually do not exceed 10 mm in length. Dorsal guard hairs short, dark, and inconspicuous. Lateral portions of the body brown and lighter than the back. Ventral pelage usually pure cream, similar in colour to the supra-ocular spots and cheeks. Mammae 4–1–4 = 9, all abdominal inguinal. Pouch absent. Tail is light brown to dark brown, dorsal discolouring gradually towards the distal portion; ventral surface of the tail not pigmented. Scales without a consistent arrangement (both spiral and annular patterns coexisting). Tail longer than the length of the head and body, covered with 5–10 mm long hairs on all sides at the base. Hands and feet covered with cream-coloured hair, like the ventral coat; Manus mesaxonic (d III> d IV); Carpal tubercles absent in mature adult males. 3.4.2 Cranium-dental morphology Cranium wide and robust. Relatively elongated face. Nasal branch of the premaxilla developed. Rostral process of the premaxilla absent. Long nasal bone, widely enlarged in the region of the maxillary/frontal suture. Mandible developed, with elongated and deep infraorbital foramen at the height of the Pm2. Interorbital region-wide (average = 12.12 mm) with incipient ridges. Lacrimal bone extends in the rostrum to the anterior margin of the orbital cavity, surpassing the two exposed lacrimal foramina. Zygomatic arch well developed, composed by the jugal and squamosal. Postorbital process of the jugal absent. Postorbital process of the frontal absent. Palatine articulates ventrally with mandible and dorsally with frontal. Frontal quite developed extending from the posterior region of the nasal branch of the mandible to the posterior region of the cranium, where it articulates with the parietal. Supraorbital crest imperceptible. Absent or incipient temporal crest. When incipient, the crest most often extends from the frontal to the parietal, where it can be joined to the midline of the cranium and extend to the lambdoidal crest. Lamboidal crest not developed. Sagittal crests absent in the anterior part of the occiput. Parietal articulates anteriorly with the frontal, but they are separated by median sutures on the right and on the left. Parietal articulates posteriorly with the supra-occipital and mastoid. Squamosal in contact with the frontal on the side of the cranium. Parietal and alisphenoid not in contact. Petrosal not exposed laterally. Squamosal parietal fenestra absent. Parietal and mastoid normally in contact. Interparietal not in contact with squamosal. Palate long, wide and composed mostly by the mandible, which articulates with the palatal branch of the pre-mandible at the height of the well-developed incisor foramen. Mandible articulates anteriorly with the palate. Maxillopalatine large (average = 24.68 mm) and fenestrated. Mandible and alisphenoid not connected at the bottom of the orbital region, separated by the palatine. Foramen of transverse canal absent. Tympanic process of alisphenoid laterally compressed and non-globular, slightly pointed or rounded. Anterior member of ectotympanic suspended directly from the basicranium. Paraoccipital process broad, erect, and ventrally projected. Dorsal margin of the foramen magnum in contact with exoccipital. Well-developed mandible. Dental formula i5/4, c1/1, pm3/3, m4/4. First upper premolar smaller than the second, well developed and non-vestigial. Second and third upper premolars equivalent in height. Post-metacrist bigger than post-protocrist in molars. The lower third premolar, when deciduous, has three cusps. Entoconids small. 3.4.3 Morphological variation We did not observe differences in the external morphology of males and females within any of the considered age groups (young, sub-adult or adult). However, there were significant age-related morphological differences. Young and sub-adult individuals had a general greyish and darker colour than adult individuals. Yet, sub-adults also exhibit some diagnostic characters of M. aritanai new species, such as cream supra-ocular spots and cheeks of the same colour, absence of median stripe on the rostrum and cream-pure ventral surface (Table 5). 3.4.4 Comparisons M. aritanai new species differs from the nominal form, M. nudicaudatus (sensu Voss et al. 2019 and present study) and from M. myosuros from the Atlantic Forest (sensu present study) by their dorsal colouring (light brown speckled with gold in M. aritanai vs. brown speckled with black and light orange in M. nudicaudatus, vs. brown-orange to light brown in M. myosuros); ventral surface colouring (purecream in M. aritanai vs. yellow-cream in M. nudicaudatus, vs. yellowish hues in M. myosuros); colouring of the sides of the body (similar to the dorsum but slightly lighter in M. aritanai vs. orange-brown, visibly lighter than the dorsum with conspicuous blackish and orange hairs in M. nudicaudatus vs. yellow with a variety of light greyish and even golden hues of grey and golden in M. myosuros); colouring of the supra-ocular spots (cream in M. aritanai vs. cream and slightly orange, especially at the edges in M. nudicaudatus, vs. yellow to cream in M. myosuros); lighter hair behind the ears (yellowish and in small amounts in M. aritanai vs. rust and in large quantities in M. nudicaudatus, vs. yellowish hues in M. myosuros); hair colour on the tarsus (cream in M. aritanai vs. brownish-orange in M. nudicaudatus, vs. varying hues from blackish to brownish-grey with golden fur in M. myosuros); width and robustness of the skull (wider and more robust in M. aritanai vs. narrower and more delicate in both M. nudicaudatus and M. myosuros); nasal width in relation to the maxillary (much narrower than the pre-maxillary in the initial portion and similar width in the end portion with a rounder end near frontal bone in M. aritanai vs. much narrower than the pre-maxillary in the initial portion and similar width in the end portion with a tapered end near frontal in both M. nudicaudatus and M. myosuros); contour of the pre-maxillary (wider, short and pointed in M. aritanai vs. narrower, long and rounded in M. nudicaudatus, vs. narrower, long and pointed in M. myosuros); frontal edges (irregular in the posterior portion with the presence of a protuberance in the median region in M. aritanai vs. more regular and rounded limit, without the presence of a median protuberance in M. nudicaudatus, vs. usually developed and pointed in M. myosuros); average differences in mitochondrial sequences of the Cytb gene (between M. aritanai and M. nudicaudatus = 8.2%; between M. aritanai and M. myosuros = 14.4%; Table 5; Figures 5–7)., Published as part of Miranda, Cleuton Lima, Nunes, Mario da Silva, Fabrício Machado, Arielli, Farias, Izeni Pires, Menezes, Fernando Heberson, Ardente, Natalia Carneiro, Dos Santos-Filho, Manoel, Bredin, Yennie Katarina & da Silva, Maria Nazareth F., 2023, A new species of jupati, genus Metachirus Burmeister 1854 (Didelphimorphia, Didelphidae) for the Brazilian Amazon, pp. 172-189 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 87 (2) on pages 178-184, DOI: 10.1515/mammalia-2021-0176, http://zenodo.org/record/7837851, {"references":["Trouessart, E. - L. (1989). Catalogus mammalium tam viventium quam fossilium. Fasciculus V. Sirenia, Cetacea, Edentata, Marsupialia, Allotheria, Monotremata., 2. R. Friedlander & Sohn, Berolini, pp. 999 - 1264.","Pine, R. H. (1973). Anatomical and nomenclatural notes on opossums. Proc. Biol. Soc. Washington 86: 391 - 402.","Patton, J. L., da Silva, M. N. F., and Malcolm, J. R. (2000). Mammals of the Rio Jurua and the evolutionary and ecological diversification of Amazonia. Bull. Am. Mus. Nat. Hist. 244: 1 - 306.","Gardner, A. L. and Dagosto, M. (2007). Tribe Metachirini. In: Gardner, A. L. (Ed.). Mammals of South America. Volume 1: marsupials, xenarthrans, shrews, and bats. University of Chicago Press, Chicago, pp. 35 - 39.","Silva, J. M. C., Rylands, A. B., and da Fonseca, G. A. B. (2005). The fate of the Amazonian areas of rndemism. Conserv. Biol. 19: 689 - 694.","Voss, R. S., Fleck, D. W., and Jansa, S. A. (2019). Mammalian diversity and Matses ethnomammalogy in Amazonian Peru. Part 3. Marsupials (Didelphimorphia). Bull. Am. Mus. Nat. Hist. 432: 1 - 87."]}

3.3.1 Metachirus aritanai new species 3.3.1.1 Synonymy (Didelphys [Metachirus]) nudicaudata: Trouessart 1989:1236; part, not M. nudicaudatus (Geoffroy St. Hilaire 1803). [Philander] nudicaudatus: Pine 1973:391; part, not M. nudicaudatus (Geoffroy St. Hilaire 1803). M. nudicaudatus: Patton et al. 2000:61–64; not M. nudicaudatus (Geoffroy St. Hilaire 1803); first evidence that Metachirus from Pará represented a new taxon. M [etachirus]. n [udicaudatus]. nudicaudatus: Gardner and Dagosto 2007:37; part, not M. nudicaudatus (Geoffroy St. Hilaire 1803). Holotype: An adult male consisting of skin and skull (INPA 2831, original number CS 83), in addition to tissue sample preserved in ethanol, collected on 8 July 1999 by Maria Nazareth F. da Silva in the Tapirapé-Aquiri National Forest, Marabá, Pará, Brazil (05° 46′ 53.10″ S; 50° 31′ 48.60″ W; Figure 1). MC, male from the Carajás region; FC, females from the Carajás region; MN, male of M. nudicaudatus; FN, females of M. nudicaudatus; MM, males of M.myosuros; FM,females of M.myosuros; MCC,maximal cranial circumference;NAS,nasal bone length;RWJ,rostrum width between the jugal bones;RWF, rostrum width between the frontal bonés;NW, greater nasal width; CI, interorbital constriction; PC,postorbital constriction; WZ,width between zygomatic arches; GCW, greatest cranial width; MAX, length of the maxillary row of teeth; LUM, length of the upper molar series; CAN, width between the lateral margins of the upper canines; WPP, width of the palatal projection; WM, width between the external faces of the third molars; LLM, length of the lower molar series; JL, jaw length. Paratypes ( N = 19): Eight adult females (N = 8) consisting of skins, skulls and tissues preserved in ethanol; two adult males (N = 2) consisting of skins, skulls and tissues preserved in ethanol; and five sub-adult and young individuals (N = 9) consisting of specimens preserved in fluid, skull and tissues. Adult females: two specimens (PSA 06; PSA 200) were collected by Cleuton L. Miranda in the Tapirapé-Aquiri National Forest, municipality of Marabá, Pará, Brazil and will be cataloged in the mammal collection of MPEG. Three specimens were collected by Natalia C. Ardente, two (MN 78307; MN 79953) in the Carajás National Forest, municipality of Marabá, Pará, Brazil, and one (MN 75549) in the Tapirapé-Aquiri National Forest, municipality of Marabá, Pará, Brazil. The three remaining specimens (LBCE 14843, 14845, 14859) were collected by Fabiana Caramaschi, Cibele R. Bonvicino and Paulo Sérgio D’ Andrea in the Vila Santa Cruz, Araguatins municipality, State of Tocantins, Brazil, and will be cataloged in the mammal collection of MN. Adult male specimens: two specimens (MN 73864; MN 78309) were collected by Natalia C. Ardente in the Carajás National Forest, State of Pará, Brazil. Sub-adult and young specimens: nine specimens were collected by Marco Antonio R. Junior and Stepheson Abrantes; five specimens (MAR 890, 1165, 1292, 1407, 1409) were collected in the Caxiuanã National Forest, municipality of Portel, Pará, Brazil, and will be cataloged in the mammal collection of MPEG; Four specimens (CN 78, 83, 91, 240) were collected in Bom Futuro on the left margin of the middle Xingu River, Pará, Brazil, and will be cataloged in MPEG. Type locality: Tapirapé-Aquiri National Forest, municipality of Marabá, state of Pará, Brazil (05° 46′ 53.10″ S; 50° 31′ 48.60″ W; Figure 1). Etymology: The name M. aritanai is given in honor of the great leader Aritana Yawalapiti (1949–2020), of the Yawalapiti ethnicity, from the upper Xingu River region and in honour of all indigenous peoples of Brazil. The leader Aritana died in August 2020, a victim of covid-19, representing a great loss for the Xingu people, all indigenous peoples, and all Brazilians. Distribution: M. aritanai new species is restricted to the Xingu-Tocantins interfluve, the endemism center Xingu sensu Silva et al. (2005). As far as we know M. aritanai new species is endemic to this biogeographic region (Figure 1). The species is bounded to the north by the Amazon River, to the west by the Xingu River and to the east by the Tocantins River. The southern boundary cannot yet be clearly defined. According to the consensus topology and times of divergence presented in this study, a sister species of M. aritanai new species, M. nudicaudatus, occurs north of the Amazon River (Figures 2 and 3). Diagnosis: M. aritanai new species is characterized by the combination of the following characters: general colouration light brown with speckles of gold on the back; small amount of rust hair behind the ears; supra-ocular spots of cream colour; cheeks with the same colouration as supra-ocular spots; brown and gold-speckled face; absence of median stripe on the rostrum; the presence of a narrow, blackened dorsal line at the top of the head, usually extending to the nape of the neck; clear ventral surface, usually pure cream, wide interorbital region (average = 12.12 mm) and with incipient ridges; rounded and inflated cranial cavity, with large maxillopalatine (average = 24.68 mm) and fenestrae; small entoconids; absent or incipient temporal crest. Figures 5–7 show the mentioned characters, except for the morphology of the entoconid, which is similar to that of M. nudicaudatus and may be seen in Figure 15, page 46 in Voss et al. (2019). 3.4 Description 3.4.1 External morphology Combined length of adult head and body ca. 230 mm on average (ranging 243–275 mm); adult weight ca. 380 g. Midrostral fur cream coloured with apical portion dark brown (most of the cream-coloured hairs have black tips, but there are also completely blackened or cream hairs in lesser quantities, somewhat like the dorsal coat). Dark median rostral stripe absent or inconspicuous. Dark circumocular mask present (blackened), extending posteriorly to the base of the ear, continuous with the coronal fur. Narrow dark brown dorsal line starting at the coronal spot and extending to the end of the nape. Throat gland present in adult males. Small pure cream supraocular spots present, usually in the shape of an inverted triangle, varying in size. Pure cream-coloured cheeks, of the same colour as the supraocular spots. Completely brown ears. Presence of small amounts of rust hair behind the base of the ears. General colouration light brown speckled with gold. Dorsal colour pale brown, with grey-based hairs, cream stripe over most of its length and short blackish apex, giving the back a golden speckled appearance. Dorsal hairs usually do not exceed 10 mm in length. Dorsal guard hairs short, dark, and inconspicuous. Lateral portions of the body brown and lighter than the back. Ventral pelage usually pure cream, similar in colour to the supra-ocular spots and cheeks. Mammae 4–1–4 = 9, all abdominal inguinal. Pouch absent. Tail is light brown to dark brown, dorsal discolouring gradually towards the distal portion; ventral surface of the tail not pigmented. Scales without a consistent arrangement (both spiral and annular patterns coexisting). Tail longer than the length of the head and body, covered with 5–10 mm long hairs on all sides at the base. Hands and feet covered with cream-coloured hair, like the ventral coat; Manus mesaxonic (d III> d IV); Carpal tubercles absent in mature adult males. 3.4.2 Cranium-dental morphology Cranium wide and robust. Relatively elongated face. Nasal branch of the premaxilla developed. Rostral process of the premaxilla absent. Long nasal bone, widely enlarged in the region of the maxillary/frontal suture. Mandible developed, with elongated and deep infraorbital foramen at the height of the Pm2. Interorbital region-wide (average = 12.12 mm) with incipient ridges. Lacrimal bone extends in the rostrum to the anterior margin of the orbital cavity, surpassing the two exposed lacrimal foramina. Zygomatic arch well developed, composed by the jugal and squamosal. Postorbital process of the jugal absent. Postorbital process of the frontal absent. Palatine articulates ventrally with mandible and dorsally with frontal. Frontal quite developed extending from the posterior region of the nasal branch of the mandible to the posterior region of the cranium, where it articulates with the parietal. Supraorbital crest imperceptible. Absent or incipient temporal crest. When incipient, the crest most often extends from the frontal to the parietal, where it can be joined to the midline of the cranium and extend to the lambdoidal crest. Lamboidal crest not developed. Sagittal crests absent in the anterior part of the occiput. Parietal articulates anteriorly with the frontal, but they are separated by median sutures on the right and on the left. Parietal articulates posteriorly with the supra-occipital and mastoid. Squamosal in contact with the frontal on the side of the cranium. Parietal and alisphenoid not in contact. Petrosal not exposed laterally. Squamosal parietal fenestra absent. Parietal and mastoid normally in contact. Interparietal not in contact with squamosal. Palate long, wide and composed mostly by the mandible, which articulates with the palatal branch of the pre-mandible at the height of the well-developed incisor foramen. Mandible articulates anteriorly with the palate. Maxillopalatine large (average = 24.68 mm) and fenestrated. Mandible and alisphenoid not connected at the bottom of the orbital region, separated by the palatine. Foramen of transverse canal absent. Tympanic process of alisphenoid laterally compressed and non-globular, slightly pointed or rounded. Anterior member of ectotympanic suspended directly from the basicranium. Paraoccipital process broad, erect, and ventrally projected. Dorsal margin of the foramen magnum in contact with exoccipital. Well-developed mandible. Dental formula i5/4, c1/1, pm3/3, m4/4. First upper premolar smaller than the second, well developed and non-vestigial. Second and third upper premolars equivalent in height. Post-metacrist bigger than post-protocrist in molars. The lower third premolar, when deciduous, has three cusps. Entoconids small. 3.4.3 Morphological variation We did not observe differences in the external morphology of males and females within any of the considered age groups (young, sub-adult or adult). However, there were significant age-related morphological differences. Young and sub-adult individuals had a general greyish and darker colour than adult individuals. Yet, sub-adults also exhibit some diagnostic characters of M. aritanai new species, such as cream supra-ocular spots and cheeks of the same colour, absence of median stripe on the rostrum and cream-pure ventral surface (Table 5). 3.4.4 Comparisons M. aritanai new species differs from the nominal form, M. nudicaudatus (sensu Voss et al. 2019 and present study) and from M. myosuros from the Atlantic Forest (sensu present study) by their dorsal colouring (light brown speckled with gold in M. aritanai vs. brown speckled with black and light orange in M. nudicaudatus, vs. brown-orange to light brown in M. myosuros); ventral surface colouring (purecream in M. aritanai vs. yellow-cream in M. nudicaudatus, vs. yellowish hues in M. myosuros); colouring of the sides of the body (similar to the dorsum but slightly lighter in M. aritanai vs. orange-brown, visibly lighter than the dorsum with conspicuous blackish and orange hairs in M. nudicaudatus vs. yellow with a variety of light greyish and even golden hues of grey and golden in M. myosuros); colouring of the supra-ocular spots (cream in M. aritanai vs. cream and slightly orange, especially at the edges in M. nudicaudatus, vs. yellow to cream in M. myosuros); lighter hair behind the ears (yellowish and in small amounts in M. aritanai vs. rust and in large quantities in M. nudicaudatus, vs. yellowish hues in M. myosuros); hair colour on the tarsus (cream in M. aritanai vs. brownish-orange in M. nudicaudatus, vs. varying hues from blackish to brownish-grey with golden fur in M. myosuros); width and robustness of the skull (wider and more robust in M. aritanai vs. narrower and more delicate in both M. nudicaudatus and M. myosuros); nasal width in relation to the maxillary (much narrower than the pre-maxillary in the initial portion and similar width in the end portion with a rounder end near frontal bone in M. aritanai vs. much narrower than the pre-maxillary in the initial portion and similar width in the end portion with a tapered end near frontal in both M. nudicaudatus and M. myosuros); contour of the pre-maxillary (wider, short and pointed in M. aritanai vs. narrower, long and rounded in M. nudicaudatus, vs. narrower, long and pointed in M. myosuros); frontal edges (irregular in the posterior portion with the presence of a protuberance in the median region in M. aritanai vs. more regular and rounded limit, without the presence of a median protuberance in M. nudicaudatus, vs. usually developed and pointed in M. myosuros); average differences in mitochondrial sequences of the Cytb gene (between M. aritanai and M. nudicaudatus = 8.2%; between M. aritanai and M. myosuros = 14.4%; Table 5; Figures 5–7).

Rhinella angeli sp. nov. Synonyms Bufo sp. nov. (pinocho) (Rodrigues & Knel, 2003) Bufo sp. nov. (pinocho) (Gordo et al. 2006) Bufo sp. nov. (Pérez et al. 2006) Holotype. CRBIIAP -2126 adult male collected at Sacacaca, Yavari-Mirin basin (4° 32’ 3’’ S, 72° 30’ 17’’ W) Department of Loreto, Province of Ramon Castilla, District of Yavari, Peru by Pedro Perez Peña on 22 November 2011 at 9:00 am (Figure 2 A–E). Paratypes. MZUNAP-AN 551 adult female collected at Sierra del Divisor National Reserve (6° 21’ 59” S, 73° 27’ 43” W) Department of Loreto, Province of Requena, Perú by Jose Luis Ramirez. CRBIIAP-2127 adult male collected at Miricillo locality (4° 22’ 42” S, 73° 6’ 38” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña. CRBIIAP-2128 adult female collected at Panguana locality (4° 22’ 38” S, 73° 6’ 21” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña. CRBIIAP-2129 juvenile collected at Nueva Esperanza locality (4° 22’ 31” S, 73° 6’ 3” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña. CRBIIAP -2130 juvenile collected at Taguado locality (4° 22’ 12” S, 73° 5’ 42” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña (Figure 3 A-L). Diagnosis. Rhinella angeli sp. nov. is diagnosed by: (1) presence of a very long fleshy proboscis (20% of HL) on its snout that can hang flaccid or stand stiffly straight, (2) snout-vent length 36.2-37.5 mm in males and 40.1-48.6 mm in females, (3) cephalic crests poorly developed, (4) vertebral apophyses absent, (5) absence of bony knobs at the lower angle of jaws, (6) dorsolateral row of tubercles present, (7) texture of dorsum highly granular (Figure 2 A–H). Holotype measurements (mm). SVL 37.4; HL 11.8; HW 12.3; IND 2.3; END 3.4; ED 4.1; UEW 3.4; IOD 6.6; POCH 1.4; POCL 3.6; HTD 2.6; VTD 2.5; PGL 7.8; HAL 8.8; FAL 8.5; THL 16.5; TL 10.9; TAL 8.6; FL 13.2. Comparison with other species. Rhinella angeli sp. nov. differs from all other species belonging to the R. margaritifera species group by their pronounced fleshy proboscis in the nose (Table 3, Figure 4 A–H). From similar species (characteristics of other species in parentheses), two have been reported as having proboscis (less pronounced than the new species): R. dapsilis and R. proboscidea. The new species is additionally distinguished from R. dapsilis by snout-vent length smaller in males (36.2–37.5 mm) and females (40.1–48.6 mm) (vs. 50–59 mm in males and 63–94 mm in females) and poorly developed cephalic crests (well developed). From R. proboscidea the new species is promptly distinguished by the presence of pronounced fleshy proboscis (snout acute referred as a proboscis), by the granulose dorsal skin in males (smooth dorsal skin in males), and poorly developed crests (developed). Also, from its phylogenetically most closely related described species R. magnussoni the new species is distinguished by the absence of a dorsolateral line of tubercles that extends from behind the eye alongside the base of the legs (present), absence of bony knobs at angle of jaws (presence), presence of pronounced fleshy proboscis (absent). Finally, small-bodied toads from R. margaritifera group have been confused with R. castaneotica, from which the new species is also distinguished by the presence of external tympanum (not distinguished), the snout strongly acute in lateral view (protruding) and the presence of a long fleshy proboscis (absent). Description of holotype. CRBIIAP-2126, adult male. SVL 37.4 mm. Body robust. Head wider (12.3 mm) than longer (11.8 mm). Snout strongly acute in dorsal and lateral view, strongly acute with fleshy ridge extending to tip of snout in lateral view. Cephalic crests not developed, granulose with keratinized borders. Supraorbital crest not developed, granulose with a few coalescent granules. Supraorbital crest low, granulose, occasionally with a few coalescent granules. Preorbital and postorbital crest low. Infraorbital crest low, curved, granulose. Supratympanic crest low and short, straight, with granules of irregular size and form. Parietal crest absent. Canthus rostralis without crests, concave. Interorbital and inter nostril region flat, nostrils strongly protuberant directed dorsolaterally. Eye to nostril distance equal to eye diameter, larger than inter nostril distance. Upper eyelid width about 62% of eye diameter. Tympanum distinct, rounded; its horizontal diameter representing 88% of the vertical diameter and equal to inter nostril distance. Corner of mouth triangular. Bony knobs at the lower angle of jaws absent. Parotoid elongate. Vocal slits absent; choanae small, ovoid, lateral, widely separated; tongue free. Dorsolateral line of small-pointed granules extends from the posterior border of the parotoid gland to the groin. Vertebral apophyses absent. Skin on dorsum and limbs highly granular, with many small, rounded tubercles, irregularly distributed. Small tubercles covering lips, eyelids, interorbital and loreal region. Forearms slender, covered with small, scattered tubercles on the edges. Fingers free, with tips slightly enlarged; length of fingers 2Color in life (Figure 4 A–H). Doral coloration light brown. A light brown vertebral line from the interorbital region to the cloaca, surrounded by a dark brown spot. Black small spots distributed at midbody. Dark brown bands on arms and legs, and dark brown blotches on flanks and tympanic region. Venter cream with scattered dark brown blotches on chest. Loreal and tympanic region with dark brown blotches. Dark brown throat, arms and legs cream in ventral view. Iris gold, with black reticulations. Colour in preservative. Similar to that in life. In alcohol 70% the body coloration is slightly faded, mainly in dark brown blotches (Figure 3 A–L). Morphological variation. The type series includes 6 specimens (2 adult males, 2 adult females and 2 juveniles). The new species is phenotypically variable, although all of the specimens present the characteristic fleshy proboscis. Sexual dimorphism is present (Table 4). Patterning on the dorsum varies, with specimens presenting brown chevrons extending from the head to the vent. Dorsal coloration varies from dark to light brown (Figure 2–3). Specimens can present dark brown spots in the midbody. Dorsal tubercles present variable sizes, conical with keratinized apices in larger individuals. Dorsal skin texture does not vary between sexes, being covered with scattered conical tubercles in males and females. Some individuals have a thin clear brown line that extends from the rostrum to the cloaca along the central dorsum. Dorsolateral region may vary in color, from dark brown to light brown. Color of the gular and chest region varies from dark brown to creamy brown. Ventral coloration also varies from almost immaculate light brown to heavily pigmented with dark brown blotches. Tubercles on vent can vary and depend on individual size. Younger specimens present crests with highly scattered granules and reduced amount of keratin. Distribution and natural history. Rhinella angeli sp. nov. is known from the Loreto Department, Peru, western Amazonia (Figure 5) in the localities of Sacacaca, Miricillo, Panguana and Nueva Esperanza in the Yavari-Mirin river basin, as well as the localities of Taguado, Corrientes and Cauchuyo in the Tahuayo-Quebrada blanco river basin and the Sierra del Divisor Natural Reserve. We also reported specimens from the Matsés National Reserve, right bank of the Yaquerana River (reported by Gordo et al. 2006). Individuals were found in the primary forest, in upland (= terra firme) forests away from rivers, within root mass of bushes or in leaf litter. Specimens were visualized and collected in diurnal surveys. The species are found between 75 to 180 m a.s.l. When individuals were disturbed during surveys, they tried to camouflage in the litter by expanding their forelimbs and hindlimbs and becoming immobile (Figure 6 A–B). The color of the secretion of the parotoid glands is yellowish (Figure 6 C). Etymology. The specific epithet is honorific to Angel Ruiz Frias, professor of Biological Sciences at Universidad Nacional de la Amazonia Peruana-UNAP that left this world due to COVID 19. Vocalization. Unknown. Tadpole. Unknown., Published as part of Rojas, Rommel R., Peña, Pedro Perez, Ávila, Robson W., Carvalho, Vinicius Tadeu De, Perez, Renata, Farias, Izeni Pires, Gordo, Marcelo & Hrbek, Tomas, 2022, Two new surprising species of leaf-litter toad of the Rhinella margaritifera species group (Anura: Bufonidae) from the Peruvian Amazon, Loreto-Peru, pp. 487-515 in Zootaxa 5150 (4) on pages 498-504, DOI: 10.11646/zootaxa.5150.4.2, http://zenodo.org/record/6626813, {"references":["Gordo, M. (2006) s. n. In: Vriesendorp, C., Pitman, N., Rojas, M., Pawlak, B. A., Rivera, C., Calixto, M., Vela, C. P. & Fasabi, R. (Eds.), Peru: Matses. Rapid Biological Inventories. Report 16. The Field Museum, Chicago, Illinois, pp. 1 - 336.","Perez, P., Bodmer, R. & Puertas, P. (2006) In: Fang, T., Bodmer, R., Aquino, R. & Valqui, M. (Ed.), Manejo de Fauna Silvestre en la Amazonia. Quito, Ecuador. Anuros y Saurios del Interfluvio Yavari- Tahuayo y su comparacion con las areas naturales protegidas en la region Loreto, Peru, pp. 1 - 15."]}

Rhinella unapensis sp. nov. Synonyms Bufo typhonius (Rivera-Gonzales et al. 2003) Bufo typhonius gr. margaritifera Ribeyro & Layche (2008) Holotype. CRBIIAP-2125 adult male collected at Puerto Almendra (3° 49’ 58’’ S, 73° 22’ 28’’ W), Department of Loreto, Province of Maynas, District of San Juan, Nina-Rumi community, Peru by Rommel R. Rojas on December 7, 2017, at 10 h. (Figure 7 A–F). Paratypes. CRBIIAP-2114, CRBIIAP-2115, CHUFC-A 9201, CHUFC-A 9202, adult females collected at Puerto Almendra, (3° 49’ 27’’ S, 73° 22’ 59’’ W), Department of Loreto, Province of Maynas, District of San Juan, Nina-Rumi community, Peru by Rommel R. Rojas Zamora. CRBIIAP-2116, CRBIIAP-2117, CRBIIAP-2123, CRBIIAP-2124, adult males collected at Puerto Almendra, (3° 49’ 52’’ S, 73° 22’ 29’’ W), Department of Loreto, Province of Maynas, District of San Juan, Nina-Rumi community, Peru by Rommel R. Rojas Zamora. CRBIIAP-2118, CRBIIAP-2119, CRBIIAP-2120, CRBIIAP-2121, CRBIIAP-2122, juveniles collected at Km 70.0 by the Iquitos-Nauta highway (04° 21’ 15’’ S, 73° 30’ 53’’ W) Department of Loreto, Province of Maynas, District of San Juan, Nuevo San Juan community, Peru by Rommel R. Rojas. MZUNAP-AN 509, MZUNAP-AN 540 adult males collected at Fundo UNAP km 31.5 by the Iquitos-Nauta highway (03º 56’ 36’’S, 72º 24’ 06’’W), Department of Loreto, Province of Maynas, District of San Juan, Iquitos-Nauta highway by Percy Mejia (Figure 8 A–L). Diagnosis. Rhinella unapensis sp. nov. is diagnosed by: (1) small-size (snout-vent length range 30.1–42.5 in adult males, 36.3–46.1 mm in adult females); (2) snout pointed in dorsal view, nearly acute in lateral view; (3) canthal and pre-orbital crests absent; (4) low supra-orbital, parietal and supra-tympanic crests; (5) small bony protrusion at the angle of jaws; (6) tympanum evident; (7) dorsolateral row of tubercles present; (8) parotoid glands small, elongate and evident; (9) skin on dorsum bearing a mixture of warts. Holotype measurements (mm). SVL 36.0; HL 12.3; HW 11.85; IND 2.3; END 4.1; ED 3.2; UEW 3.4; IOD 4.0; POCH 1.0; POCL 3.1; HTD 2.1; VTD 1.8; PGL 4.6; HAL 10.4; FAL 7.6; THL 14.8.; TL 13.2; TAL 8.4; FL 10.4. Comparison with other species. By the absence of canthal and pre-orbital crests Rhinella unapensis sp. nov. is promptly distinguished from R. alata, R. cristinae, R. exostosica, R. hoogmoedi, R. iserni, R. margaritifera, R. ocellata, R. roqueana, R. scitula, R. sebbeni, R. sclerocephala, and R. stanlaii. Rhinella unapensis sp. nov. It also differs from the Amazonian species cited above by (characteristics of other species in parenthesis): From R. alata by presence of bone protrusion at angle of jaw (absent), snout pointed in dorsal view, nearly acute in lateral view (sub acuminate in dorsal view, rounded to protruding in lateral view), palmar tubercle ovoid (oval). From R. cristinae by the acute snout in lateral view (rounded), tympanum distinct (indistinct), larger males, with 30.1–45.5 mm SVL (30.7–34.3 mm), smaller females, with 36.3–46.1 mm SVL (55.1 mm). From R. exostosica by being smaller in SVL (64.2 vs. 37.7 mm R. unapensis sp. nov.), palmar tubercle shape ovoid (tear-shaped), parotoid glands small and elongate (prominent), cephalic crest poorly developed (developed). From R. iserni by the presence of external tympanum (not distinguish), parotoid gland lacking lateral line of tubercles (presence), smaller SVL (35.4 mm in R. unapensis sp. nov. vs. 58.5 mm in adult males). From R. margaritifera by the small vertebral apophyses (protruding), low cranial crests (vs. well developed), snout nearly acute in lateral view (mucronate), smaller SVL (30.1–45.5 mm in adult males in R. unapensis sp. nov. vs. 58 mm in adult males). From R. roqueana by the small SVL (30.1– 45.5 in adult males of R. unapensis sp. nov. vs 70–79 mm in adult males of R. roqueana), absence of protruding vertebral apophyses (present), tympanum evident (hidden), the snout nearly acute in lateral view (pointed). From R. sclerocephala by snout nearly acute in lateral view (truncate), by smaller SVL (35.4 mm in R. unapensis sp. nov. vs. 61.2 mm in adult males), absence or very small vertebral apophyses (hypertrophied). The new species can also be distinguished from the species of the Rhinella margaritifera group from outside the Amazonia as follows. From R. hoogmoedi by the absence of a rounded tubercle at the posterior corner of mouth (present); dorsolateral line of pointed tubercles not well marked (well-marked). From R. ocellata by the absence of dorsal ocelli colour pattern. From R. scitula by the parotoid gland small (larger); absence of bony knobs at the angle of jaws (present); absence of a line of spinulose tubercles on parotoid gland (present). From R. sebbeni by supratympanic crest poorly developed (well-developed), and snout pointed in dorsal view (mucronate). From R. stanlaii by the smaller SVL (35.44 mm in R. unapensis sp. nov. vs. 45.8 mm in adult males), small parotoid gland (paratoid gland developed). In addition to the absence of canthal and pre-orbital crests, eight species of the Rhinella margaritifera group are similar to the new species for their low supra-orbital, parietal and supra-tympanic crests: R. acutirostris, R. castaneotica, R. dapsilis, R. lescurei, R. magnussoni, R. parecis, R. proboscidea and R. angeli. The new species is distinguished from R. acutirostris by absence of tubercles on the lateral of parotoid gland (presence), the snout nearly acute in lateral view (pointed), and dorsolateral line of tubercles weak (marked dorsolateral line of tubercles). From R. castaneotica by the presence of a small bone protrusion at angle of jaw (absent), presence of external tympanum (not distinguish), the snout nearly acute in lateral view (pointed). From R. dapsilis by smaller SVL (30.1–45.5 in adult males, 36.3–46.1 in adult females in R. unapensis sp. nov. vs. 50–59 in males and 63–94 mm SVL in adult females of R. dapsilis, snout lacking a developed proboscis (present), granular skin (smooth) and by the dorsolateral line of tubercles weak (marked dorsolateral line of tubercles). From R. lescurei by the elongate parotoid gland (triangular parotoid gland), snout nearly acute in lateral view (pointed). From R. magnussoni by skin on dorsum bearing a mixture of warts (skin of dorsum with spicules), pointed snout (rounded), nostril not protuberant (nostril protuberant) and distinct tympanic membrane (barely distinct). From R. parecis by the snout nearly acute in lateral view (rounded), presence of bone protrusion at angle of jaw (absent), elongate and distinct parotoid glands (small and not well distinct). From R. proboscidea by dorsal skin texture granular (smooth), less protruding snout (snout distinctly prominent). Finally, from R. angeli sp. nov. the new species is differentiated by the absence of a fleshy proboscis (present even in juveniles), presence of bone protrusion at angle of jaw (absent), snout profile nearly acute in lateral view (pointed), head as longer as wide (wider than long), postocular crest longer with 2.9–4.4mm (1.2–1.5mm) and lower 0.8–1.3mm (3.2–3.7mm), forearm length larger than hand length (equal) and thigh and tibia length almost equal (thigh longer than tibia). Both species are also geographically separated by the Ucayali and Amazon Rivers, with R. angeli sp. nov. distributed on the left bank and R. unapensis sp. nov. limited to the right bank. Description of holotype. CRBIIAP-2125 adult male. Body robust. SVL 36.0 mm; head wider (12.3 mm) than longer (11.8 mm). Snout pointed in dorsal view, nearly acute in lateral view. Canthal crest not raised; supraorbital and postorbital crests not developed with low supratympanic crest. Small parotoid gland, not well developed, slightly triangular in dorsal view, expanding laterally. Lateral row of tubercles from posterior margin of parotoid gland to base of the leg. Small bony protrusion at the angle of jaws. Canthus rostralis concave; interorbital and internarial region flat, nostrils acute directed dorsolaterally. Eye to nostril distance longer than eye diameter, larger than internarial distance. Upper eyelid width about 91.7% of eye diameter. Tip of the snout without developed proboscis. Tympanum distinct, rounded; its horizontal diameter representing 95.5% of the vertical diameter and smaller than internarinal distance. Triangular corner of mouth, presence of bony knobs at the lower angle of jaws. Paratoid glands distinguished, elongate. Vocal slits absent; small choanae, ovoid, lateral, widely separated; tongue free. Dorsolateral line of small white pointed granules extends from the posterior border of the parotoid gland to the groin. Very small vertebral apophyses. Skin on dorsal and ventral surfaces highly granular. Presence of rounded granules in dorsal and ventral surfaces. Small tubercles covering lips, eyelids, interorbital and loreal region. Slender forearms, robust forelimbs, covered with small and scattered tubercles around the edges. Free fingers, with the tips slightly enlarged; length of fingers 2 Color in life. Dark brown dorsal coloration. A light-brown vertebral line from the interorbital region to cloaca, surrounded by light brown spots. Small light brown spots scattered distributed at head, midbody and shanks. Dark brown bands on arms and legs and dark brown blotches on flanks and tympanic region. White-cream venter with pointed dark brown blotches on chest and belly. Loreal and tympanic region with dark brown blotches. Cream color throat; arms and legs white cream with dark brown points. Gold iris, with black reticulations (Figure 9 A–H). Morphological variation. The type series includes 17 specimens (8 adult males, 4 adult females and 5 juveniles). Dorsal skin texture not variable between sexes, being covered with rounded tubercles in males and females. Sexual dimorphism is present (Table 5). Presence of low cranial crests in both sexes, without differences between male and female, in juveniles the crest is indistinct. Dorsal coloration is variable, from dark brown to brown (Figure 9 A–H). There is presence of dark-brown spots in the head, neck, midbody and shanks region. Some individuals present a light brown vertebral line. Gular and chest region may present a brown coloration or cream coloration. Ventral coloration variable from almost immaculate cream to heavily pigmented with dark blotches or points. Distribution and natural history. Rhinella unapensis sp. nov. is known from five localities from Loreto department, Peru, western Amazonia (Figure 5). Fundo Zamora, Fundo UNAP, Allpahuayo Mishana National Reserve, the vicinity of Nauta and Puerto Almendra in the Nanay river basin. Its distribution is in the northwest of Iquitos (left bank of the Amazon, around the Nanay and Marañon Rivers), where it spans the length of the Iquitos-Nauta highway; further north, its distribution is poorly known but is possibly delimited by the Napo and Marañon Rivers. This species has been recorded from white sand forests, locally known as “ varillales ” (in Portuguese “ campinaranas ”), from around Iquitos city (Figure 10 A). They have been found in the leaf litter of primary and secondary forests, and occur between 103 to 135 m a.s.l. They are diurnal species. During night surveys, the species was seen resting in stems and leaves of low shrubs (~ 1.20 m) (Figure 10 B–C). Colour in preservative. Similar to that in life. In 70% alcohol, the body coloration presents a slightly faded mainly in dark brown blotches (Figure 8). Etymology. The specific epithet refers to the Universidad Nacional de la Amazonia Peruana-UNAP in honor of the legacy of all professors who passed away due to COVID-19 and other diseases. Tadpole. Unknown. Vocalization. Unknown

Rhinella unapensis sp. nov. Synonyms Bufo typhonius (Rivera-Gonzales et al. 2003) Bufo typhonius gr. margaritifera Ribeyro & Layche (2008) Holotype. CRBIIAP-2125 adult male collected at Puerto Almendra (3° 49’ 58’’ S, 73° 22’ 28’’ W), Department of Loreto, Province of Maynas, District of San Juan, Nina-Rumi community, Peru by Rommel R. Rojas on December 7, 2017, at 10 h. (Figure 7 A–F). Paratypes. CRBIIAP-2114, CRBIIAP-2115, CHUFC-A 9201, CHUFC-A 9202, adult females collected at Puerto Almendra, (3° 49’ 27’’ S, 73° 22’ 59’’ W), Department of Loreto, Province of Maynas, District of San Juan, Nina-Rumi community, Peru by Rommel R. Rojas Zamora. CRBIIAP-2116, CRBIIAP-2117, CRBIIAP-2123, CRBIIAP-2124, adult males collected at Puerto Almendra, (3° 49’ 52’’ S, 73° 22’ 29’’ W), Department of Loreto, Province of Maynas, District of San Juan, Nina-Rumi community, Peru by Rommel R. Rojas Zamora. CRBIIAP-2118, CRBIIAP-2119, CRBIIAP-2120, CRBIIAP-2121, CRBIIAP-2122, juveniles collected at Km 70.0 by the Iquitos-Nauta highway (04° 21’ 15’’ S, 73° 30’ 53’’ W) Department of Loreto, Province of Maynas, District of San Juan, Nuevo San Juan community, Peru by Rommel R. Rojas. MZUNAP-AN 509, MZUNAP-AN 540 adult males collected at Fundo UNAP km 31.5 by the Iquitos-Nauta highway (03º 56’ 36’’S, 72º 24’ 06’’W), Department of Loreto, Province of Maynas, District of San Juan, Iquitos-Nauta highway by Percy Mejia (Figure 8 A–L). Diagnosis. Rhinella unapensis sp. nov. is diagnosed by: (1) small-size (snout-vent length range 30.1–42.5 in adult males, 36.3–46.1 mm in adult females); (2) snout pointed in dorsal view, nearly acute in lateral view; (3) canthal and pre-orbital crests absent; (4) low supra-orbital, parietal and supra-tympanic crests; (5) small bony protrusion at the angle of jaws; (6) tympanum evident; (7) dorsolateral row of tubercles present; (8) parotoid glands small, elongate and evident; (9) skin on dorsum bearing a mixture of warts. Holotype measurements (mm). SVL 36.0; HL 12.3; HW 11.85; IND 2.3; END 4.1; ED 3.2; UEW 3.4; IOD 4.0; POCH 1.0; POCL 3.1; HTD 2.1; VTD 1.8; PGL 4.6; HAL 10.4; FAL 7.6; THL 14.8.; TL 13.2; TAL 8.4; FL 10.4. Comparison with other species. By the absence of canthal and pre-orbital crests Rhinella unapensis sp. nov. is promptly distinguished from R. alata, R. cristinae, R. exostosica, R. hoogmoedi, R. iserni, R. margaritifera, R. ocellata, R. roqueana, R. scitula, R. sebbeni, R. sclerocephala, and R. stanlaii. Rhinella unapensis sp. nov. It also differs from the Amazonian species cited above by (characteristics of other species in parenthesis): From R. alata by presence of bone protrusion at angle of jaw (absent), snout pointed in dorsal view, nearly acute in lateral view (sub acuminate in dorsal view, rounded to protruding in lateral view), palmar tubercle ovoid (oval). From R. cristinae by the acute snout in lateral view (rounded), tympanum distinct (indistinct), larger males, with 30.1–45.5 mm SVL (30.7–34.3 mm), smaller females, with 36.3–46.1 mm SVL (55.1 mm). From R. exostosica by being smaller in SVL (64.2 vs. 37.7 mm R. unapensis sp. nov.), palmar tubercle shape ovoid (tear-shaped), parotoid glands small and elongate (prominent), cephalic crest poorly developed (developed). From R. iserni by the presence of external tympanum (not distinguish), parotoid gland lacking lateral line of tubercles (presence), smaller SVL (35.4 mm in R. unapensis sp. nov. vs. 58.5 mm in adult males). From R. margaritifera by the small vertebral apophyses (protruding), low cranial crests (vs. well developed), snout nearly acute in lateral view (mucronate), smaller SVL (30.1–45.5 mm in adult males in R. unapensis sp. nov. vs. 58 mm in adult males). From R. roqueana by the small SVL (30.1– 45.5 in adult males of R. unapensis sp. nov. vs 70–79 mm in adult males of R. roqueana), absence of protruding vertebral apophyses (present), tympanum evident (hidden), the snout nearly acute in lateral view (pointed). From R. sclerocephala by snout nearly acute in lateral view (truncate), by smaller SVL (35.4 mm in R. unapensis sp. nov. vs. 61.2 mm in adult males), absence or very small vertebral apophyses (hypertrophied). The new species can also be distinguished from the species of the Rhinella margaritifera group from outside the Amazonia as follows. From R. hoogmoedi by the absence of a rounded tubercle at the posterior corner of mouth (present); dorsolateral line of pointed tubercles not well marked (well-marked). From R. ocellata by the absence of dorsal ocelli colour pattern. From R. scitula by the parotoid gland small (larger); absence of bony knobs at the angle of jaws (present); absence of a line of spinulose tubercles on parotoid gland (present). From R. sebbeni by supratympanic crest poorly developed (well-developed), and snout pointed in dorsal view (mucronate). From R. stanlaii by the smaller SVL (35.44 mm in R. unapensis sp. nov. vs. 45.8 mm in adult males), small parotoid gland (paratoid gland developed). In addition to the absence of canthal and pre-orbital crests, eight species of the Rhinella margaritifera group are similar to the new species for their low supra-orbital, parietal and supra-tympanic crests: R. acutirostris, R. castaneotica, R. dapsilis, R. lescurei, R. magnussoni, R. parecis, R. proboscidea and R. angeli. The new species is distinguished from R. acutirostris by absence of tubercles on the lateral of parotoid gland (presence), the snout nearly acute in lateral view (pointed), and dorsolateral line of tubercles weak (marked dorsolateral line of tubercles). From R. castaneotica by the presence of a small bone protrusion at angle of jaw (absent), presence of external tympanum (not distinguish), the snout nearly acute in lateral view (pointed). From R. dapsilis by smaller SVL (30.1–45.5 in adult males, 36.3–46.1 in adult females in R. unapensis sp. nov. vs. 50–59 in males and 63–94 mm SVL in adult females of R. dapsilis, snout lacking a developed proboscis (present), granular skin (smooth) and by the dorsolateral line of tubercles weak (marked dorsolateral line of tubercles). From R. lescurei by the elongate parotoid gland (triangular parotoid gland), snout nearly acute in lateral view (pointed). From R. magnussoni by skin on dorsum bearing a mixture of warts (skin of dorsum with spicules), pointed snout (rounded), nostril not protuberant (nostril protuberant) and distinct tympanic membrane (barely distinct). From R. parecis by the snout nearly acute in lateral view (rounded), presence of bone protrusion at angle of jaw (absent), elongate and distinct parotoid glands (small and not well distinct). From R. proboscidea by dorsal skin texture granular (smooth), less protruding snout (snout distinctly prominent). Finally, from R. angeli sp. nov. the new species is differentiated by the absence of a fleshy proboscis (present even in juveniles), presence of bone protrusion at angle of jaw (absent), snout profile nearly acute in lateral view (pointed), head as longer as wide (wider than long), postocular crest longer with 2.9–4.4mm (1.2–1.5mm) and lower 0.8–1.3mm (3.2–3.7mm), forearm length larger than hand length (equal) and thigh and tibia length almost equal (thigh longer than tibia). Both species are also geographically separated by the Ucayali and Amazon Rivers, with R. angeli sp. nov. distributed on the left bank and R. unapensis sp. nov. limited to the right bank. Description of holotype. CRBIIAP-2125 adult male. Body robust. SVL 36.0 mm; head wider (12.3 mm) than longer (11.8 mm). Snout pointed in dorsal view, nearly acute in lateral view. Canthal crest not raised; supraorbital and postorbital crests not developed with low supratympanic crest. Small parotoid gland, not well developed, slightly triangular in dorsal view, expanding laterally. Lateral row of tubercles from posterior margin of parotoid gland to base of the leg. Small bony protrusion at the angle of jaws. Canthus rostralis concave; interorbital and internarial region flat, nostrils acute directed dorsolaterally. Eye to nostril distance longer than eye diameter, larger than internarial distance. Upper eyelid width about 91.7% of eye diameter. Tip of the snout without developed proboscis. Tympanum distinct, rounded; its horizontal diameter representing 95.5% of the vertical diameter and smaller than internarinal distance. Triangular corner of mouth, presence of bony knobs at the lower angle of jaws. Paratoid glands distinguished, elongate. Vocal slits absent; small choanae, ovoid, lateral, widely separated; tongue free. Dorsolateral line of small white pointed granules extends from the posterior border of the parotoid gland to the groin. Very small vertebral apophyses. Skin on dorsal and ventral surfaces highly granular. Presence of rounded granules in dorsal and ventral surfaces. Small tubercles covering lips, eyelids, interorbital and loreal region. Slender forearms, robust forelimbs, covered with small and scattered tubercles around the edges. Free fingers, with the tips slightly enlarged; length of fingers 2Color in life. Dark brown dorsal coloration. A light-brown vertebral line from the interorbital region to cloaca, surrounded by light brown spots. Small light brown spots scattered distributed at head, midbody and shanks. Dark brown bands on arms and legs and dark brown blotches on flanks and tympanic region. White-cream venter with pointed dark brown blotches on chest and belly. Loreal and tympanic region with dark brown blotches. Cream color throat; arms and legs white cream with dark brown points. Gold iris, with black reticulations (Figure 9 A–H). Morphological variation. The type series includes 17 specimens (8 adult males, 4 adult females and 5 juveniles). Dorsal skin texture not variable between sexes, being covered with rounded tubercles in males and females. Sexual dimorphism is present (Table 5). Presence of low cranial crests in both sexes, without differences between male and female, in juveniles the crest is indistinct. Dorsal coloration is variable, from dark brown to brown (Figure 9 A–H). There is presence of dark-brown spots in the head, neck, midbody and shanks region. Some individuals present a light brown vertebral line. Gular and chest region may present a brown coloration or cream coloration. Ventral coloration variable from almost immaculate cream to heavily pigmented with dark blotches or points. Distribution and natural history. Rhinella unapensis sp. nov. is known from five localities from Loreto department, Peru, western Amazonia (Figure 5). Fundo Zamora, Fundo UNAP, Allpahuayo Mishana National Reserve, the vicinity of Nauta and Puerto Almendra in the Nanay river basin. Its distribution is in the northwest of Iquitos (left bank of the Amazon, around the Nanay and Marañon Rivers), where it spans the length of the Iquitos-Nauta highway; further north, its distribution is poorly known but is possibly delimited by the Napo and Marañon Rivers. This species has been recorded from white sand forests, locally known as “ varillales ” (in Portuguese “ campinaranas ”), from around Iquitos city (Figure 10 A). They have been found in the leaf litter of primary and secondary forests, and occur between 103 to 135 m a.s.l. They are diurnal species. During night surveys, the species was seen resting in stems and leaves of low shrubs (~ 1.20 m) (Figure 10 B–C). Colour in preservative. Similar to that in life. In 70% alcohol, the body coloration presents a slightly faded mainly in dark brown blotches (Figure 8). Etymology. The specific epithet refers to the Universidad Nacional de la Amazonia Peruana-UNAP in honor of the legacy of all professors who passed away due to COVID-19 and other diseases. Tadpole. Unknown. Vocalization. Unknown, Published as part of Rojas, Rommel R., Peña, Pedro Perez, Ávila, Robson W., Carvalho, Vinicius Tadeu De, Perez, Renata, Farias, Izeni Pires, Gordo, Marcelo & Hrbek, Tomas, 2022, Two new surprising species of leaf-litter toad of the Rhinella margaritifera species group (Anura: Bufonidae) from the Peruvian Amazon, Loreto-Peru, pp. 487-515 in Zootaxa 5150 (4) on pages 504-510, DOI: 10.11646/zootaxa.5150.4.2, http://zenodo.org/record/6626813, {"references":["Ribeyro, S. & Layche, J. (2008) Herpetofauna en el bosque de varillal del centro de ensenanza forestal (CIEFOR) - Puerto Almendra, Iquitos-Peru. Bachelor dissertation. Universidad Nacional de la Amazonia Peruana-UNAP, Iquitos. [unknown pagination]"]}

Rhinella angeli sp. nov. Synonyms Bufo sp. nov. (pinocho) (Rodrigues & Knel, 2003) Bufo sp. nov. (pinocho) (Gordo et al. 2006) Bufo sp. nov. (Pérez et al. 2006) Holotype. CRBIIAP -2126 adult male collected at Sacacaca, Yavari-Mirin basin (4° 32’ 3’’ S, 72° 30’ 17’’ W) Department of Loreto, Province of Ramon Castilla, District of Yavari, Peru by Pedro Perez Peña on 22 November 2011 at 9:00 am (Figure 2 A–E). Paratypes. MZUNAP-AN 551 adult female collected at Sierra del Divisor National Reserve (6° 21’ 59” S, 73° 27’ 43” W) Department of Loreto, Province of Requena, Perú by Jose Luis Ramirez. CRBIIAP-2127 adult male collected at Miricillo locality (4° 22’ 42” S, 73° 6’ 38” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña. CRBIIAP-2128 adult female collected at Panguana locality (4° 22’ 38” S, 73° 6’ 21” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña. CRBIIAP-2129 juvenile collected at Nueva Esperanza locality (4° 22’ 31” S, 73° 6’ 3” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña. CRBIIAP -2130 juvenile collected at Taguado locality (4° 22’ 12” S, 73° 5’ 42” W) Department of Loreto, Province of Ramon Castilla, Perú by Pedro Perez Peña (Figure 3 A-L). Diagnosis. Rhinella angeli sp. nov. is diagnosed by: (1) presence of a very long fleshy proboscis (20% of HL) on its snout that can hang flaccid or stand stiffly straight, (2) snout-vent length 36.2-37.5 mm in males and 40.1-48.6 mm in females, (3) cephalic crests poorly developed, (4) vertebral apophyses absent, (5) absence of bony knobs at the lower angle of jaws, (6) dorsolateral row of tubercles present, (7) texture of dorsum highly granular (Figure 2 A–H). Holotype measurements (mm). SVL 37.4; HL 11.8; HW 12.3; IND 2.3; END 3.4; ED 4.1; UEW 3.4; IOD 6.6; POCH 1.4; POCL 3.6; HTD 2.6; VTD 2.5; PGL 7.8; HAL 8.8; FAL 8.5; THL 16.5; TL 10.9; TAL 8.6; FL 13.2. Comparison with other species. Rhinella angeli sp. nov. differs from all other species belonging to the R. margaritifera species group by their pronounced fleshy proboscis in the nose (Table 3, Figure 4 A–H). From similar species (characteristics of other species in parentheses), two have been reported as having proboscis (less pronounced than the new species): R. dapsilis and R. proboscidea. The new species is additionally distinguished from R. dapsilis by snout-vent length smaller in males (36.2–37.5 mm) and females (40.1–48.6 mm) (vs. 50–59 mm in males and 63–94 mm in females) and poorly developed cephalic crests (well developed). From R. proboscidea the new species is promptly distinguished by the presence of pronounced fleshy proboscis (snout acute referred as a proboscis), by the granulose dorsal skin in males (smooth dorsal skin in males), and poorly developed crests (developed). Also, from its phylogenetically most closely related described species R. magnussoni the new species is distinguished by the absence of a dorsolateral line of tubercles that extends from behind the eye alongside the base of the legs (present), absence of bony knobs at angle of jaws (presence), presence of pronounced fleshy proboscis (absent). Finally, small-bodied toads from R. margaritifera group have been confused with R. castaneotica, from which the new species is also distinguished by the presence of external tympanum (not distinguished), the snout strongly acute in lateral view (protruding) and the presence of a long fleshy proboscis (absent). Description of holotype. CRBIIAP-2126, adult male. SVL 37.4 mm. Body robust. Head wider (12.3 mm) than longer (11.8 mm). Snout strongly acute in dorsal and lateral view, strongly acute with fleshy ridge extending to tip of snout in lateral view. Cephalic crests not developed, granulose with keratinized borders. Supraorbital crest not developed, granulose with a few coalescent granules. Supraorbital crest low, granulose, occasionally with a few coalescent granules. Preorbital and postorbital crest low. Infraorbital crest low, curved, granulose. Supratympanic crest low and short, straight, with granules of irregular size and form. Parietal crest absent. Canthus rostralis without crests, concave. Interorbital and inter nostril region flat, nostrils strongly protuberant directed dorsolaterally. Eye to nostril distance equal to eye diameter, larger than inter nostril distance. Upper eyelid width about 62% of eye diameter. Tympanum distinct, rounded; its horizontal diameter representing 88% of the vertical diameter and equal to inter nostril distance. Corner of mouth triangular. Bony knobs at the lower angle of jaws absent. Parotoid elongate. Vocal slits absent; choanae small, ovoid, lateral, widely separated; tongue free. Dorsolateral line of small-pointed granules extends from the posterior border of the parotoid gland to the groin. Vertebral apophyses absent. Skin on dorsum and limbs highly granular, with many small, rounded tubercles, irregularly distributed. Small tubercles covering lips, eyelids, interorbital and loreal region. Forearms slender, covered with small, scattered tubercles on the edges. Fingers free, with tips slightly enlarged; length of fingers 2 Color in life (Figure 4 A–H). Doral coloration light brown. A light brown vertebral line from the interorbital region to the cloaca, surrounded by a dark brown spot. Black small spots distributed at midbody. Dark brown bands on arms and legs, and dark brown blotches on flanks and tympanic region. Venter cream with scattered dark brown blotches on chest. Loreal and tympanic region with dark brown blotches. Dark brown throat, arms and legs cream in ventral view. Iris gold, with black reticulations. Colour in preservative. Similar to that in life. In alcohol 70% the body coloration is slightly faded, mainly in dark brown blotches (Figure 3 A–L). Morphological variation. The type series includes 6 specimens (2 adult males, 2 adult females and 2 juveniles). The new species is phenotypically variable, although all of the specimens present the characteristic fleshy proboscis. Sexual dimorphism is present (Table 4). Patterning on the dorsum varies, with specimens presenting brown chevrons extending from the head to the vent. Dorsal coloration varies from dark to light brown (Figure 2–3). Specimens can present dark brown spots in the midbody. Dorsal tubercles present variable sizes, conical with keratinized apices in larger individuals. Dorsal skin texture does not vary between sexes, being covered with scattered conical tubercles in males and females. Some individuals have a thin clear brown line that extends from the rostrum to the cloaca along the central dorsum. Dorsolateral region may vary in color, from dark brown to light brown. Color of the gular and chest region varies from dark brown to creamy brown. Ventral coloration also varies from almost immaculate light brown to heavily pigmented with dark brown blotches. Tubercles on vent can vary and depend on individual size. Younger specimens present crests with highly scattered granules and reduced amount of keratin. Distribution and natural history. Rhinella angeli sp. nov. is known from the Loreto Department, Peru, western Amazonia (Figure 5) in the localities of Sacacaca, Miricillo, Panguana and Nueva Esperanza in the Yavari-Mirin river basin, as well as the localities of Taguado, Corrientes and Cauchuyo in the Tahuayo-Quebrada blanco river basin and the Sierra del Divisor Natural Reserve. We also reported specimens from the Matsés National Reserve, right bank of the Yaquerana River (reported by Gordo et al. 2006). Individuals were found in the primary forest, in upland (= terra firme) forests away from rivers, within root mass of bushes or in leaf litter. Specimens were visualized and collected in diurnal surveys. The species are found between 75 to 180 m a.s.l. When individuals were disturbed during surveys, they tried to camouflage in the litter by expanding their forelimbs and hindlimbs and becoming immobile (Figure 6 A–B). The color of the secretion of the parotoid glands is yellowish (Figure 6 C). Etymology. The specific epithet is honorific to Angel Ruiz Frias, professor of Biological Sciences at Universidad Nacional de la Amazonia Peruana-UNAP that left this world due to COVID 19. Vocalization. Unknown. Tadpole. Unknown.

Rhinella parecis sp. nov. (Figs. 4–6) Holotype. UFMT-A 13716 (adult male) collected at Vilhena Municipality, Rondônia State, Brazil (12º42’54”S, 60º15’27” W; 560m a.s.l.), on October 10, 2007 by D.H. Morais (Figure 4). Paratypes. Eleven males, four females and two juveniles of undetermined sex: UFMT-A 13698, and 13711 (adult males) and UFMT-A 13821 (juvenile) collected with the holotype. UFMT-A 7496 (adult male) collected at Santa Rita Farm, Alto Alegre do Parecis Municipality, Rondônia State, Brazil (12º07’41”S, 61º52’02” W; 370m a.s.l.), on December 12, 2007 by J.A.S. Rodrigues-Filho. MNRJ 84743 (adult male) collected at Parecis Municipality, Rondônia State, Brazil, on February 02, 2013 by F.B.S. Teles; G.B.B. Silva. UFMT-A 18706, 18715, and 18723 (adult males), UFMT-A 18718, 18720, 18729, and 18735 (adult females) collected at Guaporé district, Chupinguaia Municipality, Rondônia State, Brazil (12º13’49”S, 60º43’40” W; 330m a.s.l.), on March 08, 2014 by R. W. Ávila; P.H.B. Dávila. MZUSP 148801–02 (adult males) and MZUSP 148803 (juvenile) collected at Comodoro Municipality, Mato Grosso State, Brazil (13º04’28”S, 59º40’08” W; 560m a.s.l.), on November 27–28, 2009 by biologists of JGP Consultoria Ambiental. MNRJ 85730 and 85740 (adult males) collected at Comodoro Municipality, Mato Grosso State, Brazil on February 9, 2013 by F.B.S. Teles; G.B.B. Silva. Referred specimens. MTR 25661 (adult male) and MTR 25809 (juvenile) from Parque Nacional dos PacaásNovos, Campo Novo de Rondônia municipality, Rondônia state, Brazil. Etymology. The specific name parecis, a noun in apposition, refers to the Chapada dos Parecis, a plateau occupying large portions of the Brazilian states of Mato Grosso and Rondônia, Brazil. Diagnosis. Rhinella parecis sp. nov. is diagnosed by the following combination of characters: (1) snout-vent length range 40.7–53.5 mm in males and 44.9–54.8 mm in females; (2) snout rounded in dorsal view, acute with fleshy ridge extending to the tip of the snout in lateral view; (3) canthal and pre-orbital crests absent; (4) supra-orbital, parietal and supra-tympanic crests present and low; (5) dorsolateral row of tubercles present; (6) bony protrusion at the angle of jaws absent; (7) tympanum evident; (8) vertebral apophyses absent; (9) toes half-webbed. Comparison. Among the species of the Rhinella margaritifera species group (characters in parenthesis), the absence of canthal and pre-orbital crests distinguish Rhinella parecis sp. nov. from R. alata, R. gildae, R. hoogmoedi, R. margaritifera, R. martyi, R. ocellata, R. paraguayensis, R. roqueana, R. scitula, R. sebbeni, R. sclerocephala, R. stanlaii and R. yunga (canthal and pre-orbital crests present). Rhinella parecis sp. nov. is distinguished from R. alata by larger snout-vent length (SVL), dorsolateral line of tubercles weak, and triangular paratoid glands (SVL 37.54–39.20 mm in males, 37.78–49.69 mm in females, marked dorsolateral line of tubercles and elongated paratoid glands). Rhinella parecis sp. nov. is differentiated from R. gildae by having smaller SVL, dorsolateral line of tubercles weak, absence of both vertebral apophyses and bony knobs at the angle of jaws (SVL 46.3–64.5 mm in males and 67.3–70.6 mm in females, dorsolateral line of conical tubercles well marked, vertebral apophyses present especially in females and a prominent bony protrusion at the angle of jaws present). From Rhinella hoogmoedi, R. parecis sp. nov. is differentiated by smaller size, dorsolateral line of tubercles weak and absence of bony protrusion at the angle of jaws (SVL 38.3–63.9 mm in males and 40.9–69.7 mm in females, dorsolateral line of pointed tubercles well marked and a rounded tubercle at the posterior corner of mouth present). From Rhinella margaritifera and R. roqueana the new species is distinguished by the absence of both vertebral apophyses and bony knobs at the angle of jaws and dorsolateral line of tubercles weak (vertebral apophyses and a bony protrusion at the angle of jaws present and spinose dorsolateral line of tubercles). Rhinella parecis sp. nov. is distinguished from R. martyi by smaller size, dorsolateral line of tubercles weak, absence of both vertebral apophyses and bony knobs at the angle of jaws (SVL 55.3 ± 5.8mm in males and 64.7 ± 3.4mm in females, dorsolateral line of tubercles well marked, vertebral apophyses present and protruding bony knob at the angle of jaws present). From Rhinella paraguayensis the new species is differentiated by the absence of bony knobs at the angle of jaws, less marked dorsolateral line of tubercles, and snout acute in lateral view (bony protrusion at the angle of jaws present, marked dorsolateral line of tubercles and snout rounded in lateral view). From Rhinella scitula, R. parecis sp. nov. is distinguished by larger size, dorsolateral line of tubercles weak, and absence of bony knobs at the angle of jaws (33.8–46.1 mm in males, 45.9–50.5 mm in females, marked dorsolateral line of tubercles and a bony protrusion at the angle of jaws). The absence of bony protrusions at the angle of jaws, slightly smaller size, and rounded snout in dorsal view differs the new species from R. sebbeni (bony protrusion present, SVL 48.5–59.7 mm in males and 54.7–66.7 mm in females, and mucronate snout in dorsal view). The new species can be separated from R. sclerocephala by toes half-webbed, smaller size, absence of bony protrusions at the angle of jaws, and dorsolateral line of tubercles weak (toes fully webbed, SVL 55.4–67.3mm in males, 68.5–77.4 mm in females, bony knobs in the angle of jaws present and marked dorsolateral line of tubercles). From R. stanlaii, the new species is differentiated by the rounded snout in dorsal view, smaller paratoid gland, and absence of bony protrusions at the angle of jaws (pointed snout in dorsal view, paratoid gland developed, bony knobs in the angle of jaws present). From R. yunga, the new species is distinguished by its smaller size, granulose dorsal skin and presence of vocal slits (SVL 57.5–59.5 mm in males, 53.5–65.5 mm in females, skin on dorsum smooth with scattered tubercles in males, vocal slits absent). From the seven species of R. margaritifera group with low cranial crests, R. parecis sp. nov. is distinguished from R. acutirostris by rounded snout in dorsal view, absence of bony protrusions at the angle of jaws, and by the dorsolateral line of tubercles weak (snout pointed, presence of bony knobs at the angle of jaws, and marked dorsolateral line of tubercles). Rhinella parecis sp. nov. can be differentiated from R. dapsilis by smaller size, snout lacking a developed proboscis, and by the dorsolateral line of tubercles weak (SVL 52.1–56.2 mm in males, 66.4–79.8mm in females, proboscis well developed, and marked dorsolateral line of tubercles). Rhinella parecis sp. nov. is distinguishable from R. lescurei by the triangular paratoid gland, and larger size (elongated paratoid, SVL 34.6 ± 4.3 mm in males, 43.7 ± 0.8 mm in females). The absence of dorsal ocelli colour pattern and acute snout with fleshy ridge extending to the tip of the snout in lateral view differentiates R. parecis sp. nov. from R. ocellata (dorsal ocelli colour pattern present and rounded snout in lateral view). The larger size, distinct tympanum, granulose skin on the dorsum, and upper eyelid width significantly smaller (5.5–8.6% of SVL in males—Tukey Test pRhinella parecis sp. nov. from R. castaneotica (SVL 32.4–40.4 mm in males, 34.8–41.6 mm in females, tympanum indistinct, smoother skin on dorsum and larger upper eyelid width: 6.5–11.7% of SVL in males and 8.1–10.9% of SVL in females). From R. magnussoni the new species differs by the rounded snout in dorsal view, absence of bony knobs, and significantly smaller internarial distance in females (4.2–4.6% of SVL—Tukey Test p=0.025) and significantly larger head length in males (28.7–37.5% of SVL—Tukey Test p=0.041) (snout pointed in dorsal view, presence of bony protrusion at the angle of jaws, larger internarial distance in females: 4.7–5.9% of SVL, and smaller head length in males: 30.6–34.5% of SVL). From R. proboscidea, R. parecis sp. nov. is promptly distinguished by the snout lacking a developed proboscis, granulose dorsal skin in males and significantly smaller tibia length in males 34.1–39.3% of SVL—Tukey Test p Description of the holotype. UFMT 13716, Adult male (Figure 4). SVL 42.9 mm; head wider (16.5 mm) than long (15.2 mm) at the angle of jaws. Snout rounded in dorsal view, acute with fleshy ridge extending to tip of snout in lateral view. Cephalic crests weakly developed, only the supraorbital, parietal and supratympanic crests present; canthus rostralis without crests, concave; interorbital and internarial region flat, nostrils protuberant directed dorsolaterally; eye to nostril distance equal to eye diameter, larger than internarial distance; upper eyelid width about 80% of eye diameter. Tympanum distinct, large, almost rounded; its horizontal diameter representing 93% of the vertical diameter and equal to internarinal distance. Corner of mouth angular, without bony knobs at the lower angle of jaws. Vocal slits present, lateral to the tongue; choanae small, ovoid, lateral, widely separated; tongue free, not notched behind. Paratoid glands small, triangular. A dorsolateral line of small tubercles extends from the posterior border of the paratoid gland to the groin. Vertebral apophyses absent. Skin on dorsal and ventral surfaces granulose, with scattered conical tubercles. Small tubercles covering lips, eyelids, interorbital and loreal region. Forearms and forelimbs robust covered with small tubercles of the same size of those in the dorsum and scattered larger tubercles on the edges. Fingers free, with tips slightly enlarged; length of fingers 2 Measurement of the holotype (mm). SVL 42.9; HL 15.2; HW 16.5; IND 2.6; END 4.3; ED 4.2; UEW 3.4; IOD 6.6; POCH 1.0; POCL 4.9; HTD 2.6; VTD 2.8; PGL 5.2; HAL 11; FAL 12; THL 18.8; TL 15.8; TAL 10.4; FL 14.4. Colour in life. Dorsum light brown, with dark brown bands on arms and legs and dark brown blotches on flanks and tympanic region (Figure 5a). A gray vertebral line from the interorbital region to cloaca, surrounded by dark brown bands and black spots at midbody. Venter light brown with scattered dark brown blotches. Loreal and tympanic region with dark brown blotches. Throat dark brown, arms and legs with dark brown bands. Iris gold, with black reticulations. Colour in preservative. Similar to that in life, although slightly faded mainly in dark brown blotches (Figure 4). Tadpole. Unknown. Variation among paratypes. The type series includes 17 specimens (11 adult males, 4 adult females and 2 juveniles). Females are slightly larger than males and a summary of variation in measurements between sexes can be found in Table 3. Low cranial crests in both sexes, without differences between males and females. Dorsal skin texture not variable between sexes, being covered with scattered conical tubercles in males and females. Dorsal coloration is variable, from dark brown to cream (Figure 6), sometimes with dark brown spots in midbody. Vertebral line is present in about 60% of the individuals. Ventral coloration is also variable from almost immaculate light brown to heavily pigmented with dark brown blotches. Males may present gular and chest brown coloration with a cream midventral stripe. Geographic distribution. Rhinella parecis sp. nov. is known from seven localities from Mato Grosso and Rondônia states, western Brazil (Figure 7). The species is also known from the Pacaás-Novos National Park. Most of these localities are in the Chapada dos Parecis (~ 650m a.s.l.), which separate the Paraguay and Amazon basins and present transition zones between Cerrado and Amazonia domains. Individuals of Rhinella parecis sp. nov. were found inside rainforest fragments, since several areas in this region have been deforested and transformed for cattle ranching and agricultural farming.

Rhinella parecis sp. nov. (Figs. 4���6) Holotype. UFMT-A 13716 (adult male) collected at Vilhena Municipality, Rond��nia State, Brazil (12��42���54���S, 60��15���27��� W; 560m a.s.l.), on October 10, 2007 by D.H. Morais (Figure 4). Paratypes. Eleven males, four females and two juveniles of undetermined sex: UFMT-A 13698, and 13711 (adult males) and UFMT-A 13821 (juvenile) collected with the holotype. UFMT-A 7496 (adult male) collected at Santa Rita Farm, Alto Alegre do Parecis Municipality, Rond��nia State, Brazil (12��07���41���S, 61��52���02��� W; 370m a.s.l.), on December 12, 2007 by J.A.S. Rodrigues-Filho. MNRJ 84743 (adult male) collected at Parecis Municipality, Rond��nia State, Brazil, on February 02, 2013 by F.B.S. Teles; G.B.B. Silva. UFMT-A 18706, 18715, and 18723 (adult males), UFMT-A 18718, 18720, 18729, and 18735 (adult females) collected at Guapor�� district, Chupinguaia Municipality, Rond��nia State, Brazil (12��13���49���S, 60��43���40��� W; 330m a.s.l.), on March 08, 2014 by R. W. ��vila; P.H.B. D��vila. MZUSP 148801���02 (adult males) and MZUSP 148803 (juvenile) collected at Comodoro Municipality, Mato Grosso State, Brazil (13��04���28���S, 59��40���08��� W; 560m a.s.l.), on November 27���28, 2009 by biologists of JGP Consultoria Ambiental. MNRJ 85730 and 85740 (adult males) collected at Comodoro Municipality, Mato Grosso State, Brazil on February 9, 2013 by F.B.S. Teles; G.B.B. Silva. Referred specimens. MTR 25661 (adult male) and MTR 25809 (juvenile) from Parque Nacional dos Paca��sNovos, Campo Novo de Rond��nia municipality, Rond��nia state, Brazil. Etymology. The specific name parecis, a noun in apposition, refers to the Chapada dos Parecis, a plateau occupying large portions of the Brazilian states of Mato Grosso and Rond��nia, Brazil. Diagnosis. Rhinella parecis sp. nov. is diagnosed by the following combination of characters: (1) snout-vent length range 40.7���53.5 mm in males and 44.9���54.8 mm in females; (2) snout rounded in dorsal view, acute with fleshy ridge extending to the tip of the snout in lateral view; (3) canthal and pre-orbital crests absent; (4) supra-orbital, parietal and supra-tympanic crests present and low; (5) dorsolateral row of tubercles present; (6) bony protrusion at the angle of jaws absent; (7) tympanum evident; (8) vertebral apophyses absent; (9) toes half-webbed. Comparison. Among the species of the Rhinella margaritifera species group (characters in parenthesis), the absence of canthal and pre-orbital crests distinguish Rhinella parecis sp. nov. from R. alata, R. gildae, R. hoogmoedi, R. margaritifera, R. martyi, R. ocellata, R. paraguayensis, R. roqueana, R. scitula, R. sebbeni, R. sclerocephala, R. stanlaii and R. yunga (canthal and pre-orbital crests present). Rhinella parecis sp. nov. is distinguished from R. alata by larger snout-vent length (SVL), dorsolateral line of tubercles weak, and triangular paratoid glands (SVL 37.54���39.20 mm in males, 37.78���49.69 mm in females, marked dorsolateral line of tubercles and elongated paratoid glands). Rhinella parecis sp. nov. is differentiated from R. gildae by having smaller SVL, dorsolateral line of tubercles weak, absence of both vertebral apophyses and bony knobs at the angle of jaws (SVL 46.3���64.5 mm in males and 67.3���70.6 mm in females, dorsolateral line of conical tubercles well marked, vertebral apophyses present especially in females and a prominent bony protrusion at the angle of jaws present). From Rhinella hoogmoedi, R. parecis sp. nov. is differentiated by smaller size, dorsolateral line of tubercles weak and absence of bony protrusion at the angle of jaws (SVL 38.3���63.9 mm in males and 40.9���69.7 mm in females, dorsolateral line of pointed tubercles well marked and a rounded tubercle at the posterior corner of mouth present). From Rhinella margaritifera and R. roqueana the new species is distinguished by the absence of both vertebral apophyses and bony knobs at the angle of jaws and dorsolateral line of tubercles weak (vertebral apophyses and a bony protrusion at the angle of jaws present and spinose dorsolateral line of tubercles). Rhinella parecis sp. nov. is distinguished from R. martyi by smaller size, dorsolateral line of tubercles weak, absence of both vertebral apophyses and bony knobs at the angle of jaws (SVL 55.3 �� 5.8mm in males and 64.7 �� 3.4mm in females, dorsolateral line of tubercles well marked, vertebral apophyses present and protruding bony knob at the angle of jaws present). From Rhinella paraguayensis the new species is differentiated by the absence of bony knobs at the angle of jaws, less marked dorsolateral line of tubercles, and snout acute in lateral view (bony protrusion at the angle of jaws present, marked dorsolateral line of tubercles and snout rounded in lateral view). From Rhinella scitula, R. parecis sp. nov. is distinguished by larger size, dorsolateral line of tubercles weak, and absence of bony knobs at the angle of jaws (33.8���46.1 mm in males, 45.9���50.5 mm in females, marked dorsolateral line of tubercles and a bony protrusion at the angle of jaws). The absence of bony protrusions at the angle of jaws, slightly smaller size, and rounded snout in dorsal view differs the new species from R. sebbeni (bony protrusion present, SVL 48.5���59.7 mm in males and 54.7���66.7 mm in females, and mucronate snout in dorsal view). The new species can be separated from R. sclerocephala by toes half-webbed, smaller size, absence of bony protrusions at the angle of jaws, and dorsolateral line of tubercles weak (toes fully webbed, SVL 55.4���67.3mm in males, 68.5���77.4 mm in females, bony knobs in the angle of jaws present and marked dorsolateral line of tubercles). From R. stanlaii, the new species is differentiated by the rounded snout in dorsal view, smaller paratoid gland, and absence of bony protrusions at the angle of jaws (pointed snout in dorsal view, paratoid gland developed, bony knobs in the angle of jaws present). From R. yunga, the new species is distinguished by its smaller size, granulose dorsal skin and presence of vocal slits (SVL 57.5���59.5 mm in males, 53.5���65.5 mm in females, skin on dorsum smooth with scattered tubercles in males, vocal slits absent). From the seven species of R. margaritifera group with low cranial crests, R. parecis sp. nov. is distinguished from R. acutirostris by rounded snout in dorsal view, absence of bony protrusions at the angle of jaws, and by the dorsolateral line of tubercles weak (snout pointed, presence of bony knobs at the angle of jaws, and marked dorsolateral line of tubercles). Rhinella parecis sp. nov. can be differentiated from R. dapsilis by smaller size, snout lacking a developed proboscis, and by the dorsolateral line of tubercles weak (SVL 52.1���56.2 mm in males, 66.4���79.8mm in females, proboscis well developed, and marked dorsolateral line of tubercles). Rhinella parecis sp. nov. is distinguishable from R. lescurei by the triangular paratoid gland, and larger size (elongated paratoid, SVL 34.6 �� 4.3 mm in males, 43.7 �� 0.8 mm in females). The absence of dorsal ocelli colour pattern and acute snout with fleshy ridge extending to the tip of the snout in lateral view differentiates R. parecis sp. nov. from R. ocellata (dorsal ocelli colour pattern present and rounded snout in lateral view). The larger size, distinct tympanum, granulose skin on the dorsum, and upper eyelid width significantly smaller (5.5���8.6% of SVL in males���Tukey Test pRhinella parecis sp. nov. from R. castaneotica (SVL 32.4���40.4 mm in males, 34.8���41.6 mm in females, tympanum indistinct, smoother skin on dorsum and larger upper eyelid width: 6.5���11.7% of SVL in males and 8.1���10.9% of SVL in females). From R. magnussoni the new species differs by the rounded snout in dorsal view, absence of bony knobs, and significantly smaller internarial distance in females (4.2���4.6% of SVL���Tukey Test p=0.025) and significantly larger head length in males (28.7���37.5% of SVL���Tukey Test p=0.041) (snout pointed in dorsal view, presence of bony protrusion at the angle of jaws, larger internarial distance in females: 4.7���5.9% of SVL, and smaller head length in males: 30.6���34.5% of SVL). From R. proboscidea, R. parecis sp. nov. is promptly distinguished by the snout lacking a developed proboscis, granulose dorsal skin in males and significantly smaller tibia length in males 34.1���39.3% of SVL���Tukey Test pDescription of the holotype. UFMT 13716, Adult male (Figure 4). SVL 42.9 mm; head wider (16.5 mm) than long (15.2 mm) at the angle of jaws. Snout rounded in dorsal view, acute with fleshy ridge extending to tip of snout in lateral view. Cephalic crests weakly developed, only the supraorbital, parietal and supratympanic crests present; canthus rostralis without crests, concave; interorbital and internarial region flat, nostrils protuberant directed dorsolaterally; eye to nostril distance equal to eye diameter, larger than internarial distance; upper eyelid width about 80% of eye diameter. Tympanum distinct, large, almost rounded; its horizontal diameter representing 93% of the vertical diameter and equal to internarinal distance. Corner of mouth angular, without bony knobs at the lower angle of jaws. Vocal slits present, lateral to the tongue; choanae small, ovoid, lateral, widely separated; tongue free, not notched behind. Paratoid glands small, triangular. A dorsolateral line of small tubercles extends from the posterior border of the paratoid gland to the groin. Vertebral apophyses absent. Skin on dorsal and ventral surfaces granulose, with scattered conical tubercles. Small tubercles covering lips, eyelids, interorbital and loreal region. Forearms and forelimbs robust covered with small tubercles of the same size of those in the dorsum and scattered larger tubercles on the edges. Fingers free, with tips slightly enlarged; length of fingers 2Measurement of the holotype (mm). SVL 42.9; HL 15.2; HW 16.5; IND 2.6; END 4.3; ED 4.2; UEW 3.4; IOD 6.6; POCH 1.0; POCL 4.9; HTD 2.6; VTD 2.8; PGL 5.2; HAL 11; FAL 12; THL 18.8; TL 15.8; TAL 10.4; FL 14.4. Colour in life. Dorsum light brown, with dark brown bands on arms and legs and dark brown blotches on flanks and tympanic region (Figure 5a). A gray vertebral line from the interorbital region to cloaca, surrounded by dark brown bands and black spots at midbody. Venter light brown with scattered dark brown blotches. Loreal and tympanic region with dark brown blotches. Throat dark brown, arms and legs with dark brown bands. Iris gold, with black reticulations. Colour in preservative. Similar to that in life, although slightly faded mainly in dark brown blotches (Figure 4). Tadpole. Unknown. Variation among paratypes. The type series includes 17 specimens (11 adult males, 4 adult females and 2 juveniles). Females are slightly larger than males and a summary of variation in measurements between sexes can be found in Table 3. Low cranial crests in both sexes, without differences between males and females. Dorsal skin texture not variable between sexes, being covered with scattered conical tubercles in males and females. Dorsal coloration is variable, from dark brown to cream (Figure 6), sometimes with dark brown spots in midbody. Vertebral line is present in about 60% of the individuals. Ventral coloration is also variable from almost immaculate light brown to heavily pigmented with dark brown blotches. Males may present gular and chest brown coloration with a cream midventral stripe. Geographic distribution. Rhinella parecis sp. nov. is known from seven localities from Mato Grosso and Rond��nia states, western Brazil (Figure 7). The species is also known from the Paca��s-Novos National Park. Most of these localities are in the Chapada dos Parecis (~ 650m a.s.l.), which separate the Paraguay and Amazon basins and present transition zones between Cerrado and Amazonia domains. Individuals of Rhinella parecis sp. nov. were found inside rainforest fragments, since several areas in this region have been deforested and transformed for cattle ranching and agricultural farming., Published as part of ��vila, Robson Waldemar, Morais, Drausio Honorio, Perez, Renata, Pansonato, Andr��, Carvalho, Vinicius Tadeu De, Rojas, Rommel R., Gordo, Marcelo & Farias, Izeni Pires, 2020, A new species of the Rhinella margaritifera (Laurenti 1768) species group (Anura Bufonidae) from southern Brazilian Amazonia, pp. 368-388 in Zootaxa 4868 (3) on pages 378-381, DOI: 10.11646/zootaxa.4868.3.3, http://zenodo.org/record/4417812

Neacomys xingu, new species Xingu Spiny Mouse Figures 11, 12 HOLOTYPE: The holotype (UFMT 1268) is an adult female (age class 5), collected on 28 August 2009 by Cleuton Lima Miranda (original field number PSA 242) in a pitfall trap. The specimen is preserved as a skin, skull, and skeleton in good condition; additionally, a tissue is preserved in alcohol, and a partial cytochrome b sequence that we obtained from it has been deposited in Genbank with accession number MT 462060. TYPE LOCALITY: Flona Tapirapé-Aquiri, Marabá, state of Pará, Brazil (5°46′S, 50°32′W, fig. 5). DIAGNOSIS: Neacomys xingu is a small species (table 3) that differs from congeneric taxa by the following combination of craniodental traits: skull delicate; interorbital region narrow; nasal bones expanded anteriorly; supraorbital margins convergent anteriorly; subsquamosal fenestra small (about ¼ the size of the postglenoid foramen on each side of the skull); paraoccipital process separated from the auditory bullae; sphenopalatine foramen small; carotid circulation usually pattern 1 (sensu Voss, 1988); maxillary part of incisive septum (between the incisive foramina) wide; M1 usually with flat and undivided anterocone; and M1 anteroloph fused with the anterolabial conule. MORPHOLOGICAL DESCRIPTION: Dorsal pelage orange-brown, sprinkled with black (fig. 11); ventral pelage varying from pure white to buffy white and separated from the dorsal pelage by a thin orange lateral line. Superciliary, genal, and mystacial vibrissae blackish and long (extending behind ears when laid back alongside the head); submental vibrissae absent; interramal vibrissae short and white; Ears small and rounded; postauricular hairs gray based with orange tips, forming an orange tuft behind each pinna. Ungual tufts white, longer than claws in length; fore- and hind feet covered dorsally with buffy-cream hairs; hind feet narrow and elongate with small interdigital membranes present. Tail about the same length as head and body, usually unicolored (except in PSA 069, MPEG 39901, and MPEG 42019, in which the tail is dark above and paler below), and covered by short, spiny, and clearly visible hairs; tail tip with very short (1–2 mm) terminal tuft; caudal scales small, arranged in annular series; each caudal scale with three subequal hairs inserted along its posterior margin. Skull small and delicate in dorsal view (fig. 12); with anteriorly expanded nasal margins; notably broad rostrum and shallow zygomatic notches; posterior nasal terminus slightly pointed, extending beyond the maxillary-frontal suture; premaxillaries terminating slightly anterior to nasals; lacrimal bone small and visible in dorsal view, equally contacting the maxillary and frontal bones; supraorbital margins convergent anteriorly; interorbital region narrow; supraorbital beads developed as projecting shelves; lateral expansion of the parietal restricted to the dorsal cranial surface (except in MCN-M 1404 and MPEG 39901, in which the parietal is slightly expanded ventrally near the squamosal root of the zygomatic arch). Incisive foramina small with subparallel lateral margins, not extending posteriorly to level of M1s; maxillary portion of incisive septum (dividing the left and right foramina) usually wide. Zygomatic plate varying from broad to narrow. Palate with two posterolateral pits on each side. Auditory bullae small and usually globular, with short and narrow eustachian tubes; periotic bone extends anteriorly to the internal carotid canal but does not enter it (except in MPEG 42715 and MCN-M 1404, in which the periotic does not reach the internal carotid canal). Subsquamosal fenestra small (about ¼ the size of the postglenoid foramen); hamular process of the squamosal long. Paraoccipital process narrow and small and usually separated from the auditory bullae (Sánchez-Vendizú et al., 2018: fig. 3C). Sphenopalatine foramen small (except in MPEG 41991, whose sphenopalatine foramen is large); alisphenoid strut absent; carotid circulation pattern usually primitive 8 (pattern 1, as identified by retaining a well-developed squamosal-alisphenoid groove and sphenofrontal foramen, both indicative of the presence of the supraorbital branch of the stapedial artery; Voss, 1988: fig. 18A, B). First upper molar (M1) usually with slightly flat and undivided anterocone; anteroloph usually fused with the anterolabial conule (such that the anteroflexus is not distinguishable); M3 small; labial cusps (paracone, metacone) usually taller than lingual cusps (protocone, hypocone); m1 anteroconid undivided. Mandible with mental foramen opening laterally; capsular process of lower incisor alveolus present, but indistinct (reduced to a slight rounded elevation), approximately at same height of coronoid process. TAXONOMIC COMPARISONS: Neacomys xingu differs from N. dubosti in dorsal pelage color (orange-brown sprinkled with black versus light to dark brown finely sprinkled with orange), and by its narrower interorbital region, globular auditory bullae (the bullae are usually flask shaped in N. dubosti), and carotid circulation usually pattern 1 (versus always pattern 1 in N. dubosti). Karyotypically, Neacomys xingu differs from all other members of the Dubosti Group by having a diploid chromosomal complement of 58 (versus 2 n = 64 in N. dubosti and 2 n = 54 in “species 2”), a uniquely small FN of 64 autosomal arms (versus FN = 66– 60 in other species), and submetacentric sex chromosomes (at least one sex chromosome is acrocentric in N. dubosti and “species 2”). DISTRIBUTION: Neacomys xingu has been collected on the right bank of the lower Xingu River and in the region of Serra de Carajás, in southeastern Pará state (fig. 5). According to cytogenetic data (Di-Nizo et al., 2017; Oliveira da Silva et al., 2019), the species also ranges southward into Vila Rica in northeastern Mato Grosso state. These records suggest that the species is restricted to the Xingu center of endemism (Silva et al., 2005). ETYMOLOGY: The specific epithet xingu is to be treated as a noun in apposition. The species name refers to the Xingu center of endemism, delimited by the Xingu and Tapajós rivers, where the species occurs. REMARKS: Neacomys xingu was previously reported in the literature as “ Neacomys clade 7” by Patton et al. (2000), “ Neacomys sp.” (in part) by Silva at al. (2015), “ Neacomys sp.” by Di-Nizo et al. (2017) and Brandão et al. (2019), and “ Neacomys sp. C” by Oliveira da Silva et al. (2019).

Neacomys vossi, new species Voss’s Spiny Mouse Figures 8, 9 HOLOTYPE: The holotype (UFPA 1583) is an adult female (age class 2) collected on 2 April 2013, by Ana Cristina Mendes Oliveira (original field number JB 031). The specimen consists of a stuffed skin, skull, and skeleton, all in good condition; additionally, a tissue sample is preserved in ethanol, and a partial cytochrome b sequence that we obtained from it has been deposited in Genbank with accession number MT 462028. TYPE LOCALITY: Boca do Rato, on the right bank of the Rio Tapajós, Itaituba municipality, state of Pará, Brazil (5°14′S, 56°56′W, fig. 5). DIAGNOSIS: Neacomys vossi is a small species (table 3) that differs from congeneric taxa by the following combination of craniodental traits: skull delicate; interorbital region narrow; nasal bones expanded anteriorly; supraorbital margins convergent anteriorly; subsquamosal fenestra usually small (about ¼ the size of the postglenoid foramen on each side of the skull); paroccipital processes close to the auditory bullae; sphenopalatine foramen large; carotid circulation pattern 1 (sensu Voss, 1988); maxillary part of incisive septum (between the incisive foramina) usually wide; M1 usually with slightly flat, narrow, and undivided anterocone; and M1 anteroloph usually distinct (not fused with the anterolabial conule). MORPHOLOGICAL DESCRIPTION: Dorsal pelage brown sprinkled with orange and/or black (fig. 8); ventral pelage varying from pure white to buffy white; very thin orange lateral line. Superciliary, genal, and mystacial vibrissae blackish and long (extending behind ears when laid back alongside the head); submental vibrissae absent; interramal vibrissae short and white. Ears small and rounded; post-auricular hairs gray-based with orange tips, forming an orange tuft behind each pinna. Ungual tufts white, longer than claws; fore- and hind feet covered dorsally with buffycream hairs; hind feet narrow and elongate with small interdigital membranes. Tail about the same length as head and body, usually slightly bicolored (except in UFPA 1277, 1417, 1444, 1654, and 1736 which have unicolored tail) and covered by spiny and clearly visible hairs; tail tip of the tail with very short (approximately 1.4 mm long) terminal tuft; caudal scales small, arranged in annular series; each caudal scale with three subequal hairs inserted along its posterior margin. Skull small and delicate in dorsal view (fig. 9); with expanded anterior nasal margins; notably broad rostrum and shallow zygomatic notches; posterior nasal terminus usually flat, extending beyond the maxillary-frontal suture; premaxillaries terminating slightly anterior to nasals; lacrimal bone small and visible in dorsal view, equally contacting maxillary and frontal bones; supraorbital margins convergent anteriorly; interorbital region narrow; supraorbital beads developed as projecting shelves; lateral expansion of the parietal restricted to the dorsal surface (except in UFPA 1227 and UFPA 1577, in which the parietal is slightly expanded ventrally near the squamosal root of the zygomatic arch). Incisive foramina small with subparallel lateral margins, not extending posteriorly to the level of M1s; maxillary portion of incisive septum (dividing the left and right foramina) usually wide. Zygomatic plate narrow. Palate with two posterolateral pits on each side of the palate. Auditory bullae small and globular, with short and narrow eustachian tubes; periotic bone extending anteriorly to internal carotid canal, but usually not entering into it (except in UFPA 1227, 1417, 1530, 1654, 1417, and 1736, in which the periotic does enter the internal carotid canal). Subsquamosal fenestra usually small (about ¼ the size of the postglenoid foramen); hamular process of the squamosal long. Paraoccipital process narrow, small, and close to the auditory bullae (Sánchez-Vendizú et al., 2018: fig. 3C). Sphenopalatine foramen large; alisphenoid strut absent; carotid circulation primitive (pattern 1, as identified by retaining a well-developed squamosal-alisphenoid groove and sphenofrontal foramen, both indicative of the presence of the supraorbital branch of the stapedial artery; Voss et al., 1988). First upper molar (M1) usually with slightly flat and undivided anterocone; anteroloph usually distinct (not fused with the anterolabial conule); M3 small; labial cusps (paracone, metacone) usually taller than lingual cusps (protocone, hypocone); m1 anteroconid undivided. Mandible with mental foramen opening laterally; capsular process of lower incisor alveolus present, but indistinct (reduced as a slight, rounded elevation), approximately at same height as coronoid process. TAXONOMIC COMPARISONS: Neacomys vossi differs from N. dubosti in dorsal pelage color (brown sprinkled with orange and/or black versus dark brown finely sprinkled with orange in N. dubosti), and by its narrower interorbital region, and globular auditory bullae (the bullae are usually flask shaped in N. dubosti). Neacomys vossi differs from N. xingu in dorsal pelage color (brown sprinkled with orange and/or black versus orange-brown sprinkled with black in N. xingu), larger sphenopalatine foramen, and an M1 anteroloph that is usually distinct (versus fused in N. xingu). Karyotypically, Neacomys vossi differs from other species in the Dubosti Group by having a diploid chromosome complement of 58 (versus 2 n = 64 in N. dubosti and 2 n = 64 in “species 2”), an FN of 68 autosomal arms (versus FN = 70 in N. marajoara, FN = 64 in N. xingu, and FN = 66 in “species 2”), and submetacentric sex chromosomes (at least one of the sex chromosomes is acrocentric in N. dubosti and “species 2”) DISTRIBUTION: Neacomys vossi has been collected on the right bank of the upper and middle Tapajós River and on the left bank of lower Xingu (fig. 5). According to our records, the species appears to be endemic to the Tapajós center of endemism (Silva et al., 2005). ETYMOLOGY: Named in honor of Robert S. Voss (fig. 10), curator of mammals at the American Museum of Natural History, New York, for his extensive contributions to our knowledge of Neotropical mammals, especially the taxonomy of Neacomys in northeastern Amazonia. . FIELD NOTES: Among the specimens we examined for which trapping information is available, all 17 were captured in pitfall traps, two (UFPA 1277 and 1284) in upland (terra firme) forests and three (UFPA 1444, 1487, and 1583) in primary forest of unknown character. REMARKS: Neacomys vossi was previously reported in the literature as “ Neacomys sp. A” by Oliveira da Silva et al. (2017, 2019)., Published as part of Semedo, Thiago Borges Fernandes, Silva, Maria Nazareth Ferreira Da, Gutiérrez, Eliécer E., Ferreira, Daniela Cristina, Nunes, Mario Da Silva, Mendes-Oliveira, Ana Cristina, Farias, Izeni Pires & Rossi, Rogério Vieira, 2020, Systematics of Neotropical Spiny Mice, Genus Neacomys Thomas, 1900 (Rodentia: Cricetidae), from Southeastern Amazonia, with Descriptions of Three New Species, pp. 1-43 in American Museum Novitates 2020 (3958) on pages 26-30, DOI: 10.1206/3958.1, http://zenodo.org/record/5056873, {"references":["Voss, R. S. 1988. Systematics and ecology of ichthyomyine rodents (Muroidea): patterns of morphological evolution in a small adaptive radiation. Bulletin of the American Museum of Natural History 188 (2): 259 - 493.","Sanchez-Vendizu, P., V. Pacheco and D. Vivas-Ruiz. 2018. An introduction to the systematics of smallbodied Neacomys (Rodentia: Cricetidae) from Peru with descriptions of two new species. American Museum Novitates 3913: 1 - 38.","Silva, J. M. C., A. B. Rylands and G. A. B. Fonseca 2005. O destino das areas de endemismo da Amazonia. Megadiversidade 1: 124 - 131.","Oliveira da Silva, W., et al. 2017. Chromosomal diversity and molecular divergence among three undescribed species of Neacomys (Rodentia, Sigmodontinae) separated by Amazonian rivers. PLoS ONE 12: 1 - 19.","Oliveira da Silva, W., et al. 2019. Chromosomal phylogeny and comparative chromosome painting among Neacomys species (Rodentia, Sigmodontinae) from eastern Amazonia. Bmc Evolutionary Biology 19: 1 - 13."]}

Neacomys marajoara, new species Marajoara Spiny Mouse Figures 4, 7 HOLOTYPE: The holotype (MPEG 40432) is an adult male (age class 3), collected on 19 January 2009, by R. V. Rossi (field number MAJ 23; fig. 4) in a pitfall trap. The specimen consists of a stuffed skin (missing the tip of the tail), skull, and skeleton; a tissue sample of this specimen is preserved in ethanol, and a partial cytochrome b sequence that we obtained from it has been deposited in Genbank with accession number MT 462070. TYPE LOCALITY: Tauari Farm, municipality of Chaves, Marajó Island, state of Pará, Brazil (0°39′S, 50°11′W, figs. 5, 6). DIAGNOSIS: Neacomys marajoara is a small species (table 3) that differs from congeneric taxa by the following combination of craniodental traits: skull delicate; interorbital region relatively broad; nasal bones straight anteriorly; supraorbital margins slightly convergent anteriorly; subsquamosal fenestra large (almost half the size of the postglenoid foramen on each side of the skull); paraoccipital processes separated from the auditory bullae; sphenopalatine foramen large; carotid circulation usually pattern 2 (sensu Voss, 1988); maxillary part of incisive septum (between the incisive foramina) narrow; M1 usually with flat and undivided anterocone; and M1 anteroloph usually fused with anterolabial conule. MORPHOLOGICAL DESCRIPTION: Dorsal pelage dark brown finely sprinkled with orange (fig. 4); ventral pelage varying from pure white to yellowish white, separated from the dorsal pelage by a very thin orange lateral line. Superciliary, genal, and mystacial vibrissae blackish and long (extending behind ears when laid back alongside the head); submental vibrissae absent; interramal vibrissae short and white. Ears small and rounded; postauricular hairs gray based with orange tips, forming an orange tuft behind each pinna. Ungual tufts white, longer than claws in most specimens examined (except MPEG 40435 and MPEG 40446, in which ungual tufts are as long as the claws); fore- and hind feet covered dorsally with buffy-cream hairs; hind feet narrow and elongate with small interdigital membranes present. Tail about the same length as head and body, bicolored, and covered by short, spiny, and clearly visible hairs; white hairs present on ventral caudal surface from base to midlength; tail tip with very short (1 mm) terminal tuft; caudal scales small, arranged in annular series; each caudal scale with three subequal hairs inserted along its posterior margin. Skull small and delicate in dorsal view (fig. 7), with straight anterior nasal margins, notably broad rostrum, and shallow zygomatic notches; posterior nasal terminus usually slightly pointed, extending beyond maxillary-frontal suture; premaxillaries terminating anterior to nasals; lacrimal bone small and visible in dorsal view, usually in broad contact with frontal bones; supraorbital margins slightly convergent anteriorly; interorbital region relatively broad; supraorbital beads developed as projecting shelves; lateral expansion of the parietal restricted to the dorsal cranial surface. Incisive foramina small and usually teardrop shaped, not extending posteriorly to level of M1s; maxillary portion of incisive septum (dividing the left and right foramina) narrow. Zygomatic plate narrow. Palate with two posterolateral pits on each side. Auditory bullae small and globular, with short and narrow eustachian tubes; periotic bone extends anteriorly to internal carotid canal (except MPEG 40435, MPEG 40443, and MPEG 40439, in which the periotic does not reach the internal carotid canal), but does not enter the canal. Subsquamosal fenestra large (almost half the size of the postglenoid foramen); hamular process of the squamosal long. Paraoccipital process narrow and small and separated from the auditory bullae. Sphenopalatine foramen large; alisphenoid strut absent; carotid circulation pattern usually derived 7 (pattern 2, as identified by the presence of a large stapedial foramen and a large posterior opening of the alisphenoid canal, and by the absence of a squamosalalisphenoid groove and a sphenofrontal foramen; Voss, 1988). First upper molar (M1) usually with broad, flat, and undivided anterocone; anteroloph usually fused with anterolabial conule (such that the anteroflexus is not distinguishable); M3 small; labial cusps (paracone and metacone) usually smaller than lingual cusps (protocone and hypocone); m1 anterocone undivided. Mandible with mental foramen opening laterally; capsular process of lower incisor alveolus present, but indistinct (reduced to a slight rounded elevation), approximately at same height as coronoid process. KARYOTYPES: Karyotypes of specimens from the type locality were obtained by Silva et al. (2015), and karyotypes from specimens collected elsewhere on Marajó Island were obtained by Oliveira da Silva et al. (2019), who described a chromosomal complement of 2 n = 58 and a fundamental number (FN) = 70 (table 5). 7 Most specimens of Neacomys marajoara (seven out of nine) exhibit carotid circulation pattern 2, but one specimen (MPEG 40446) exhibits circulatory pattern 1, and another (MPEG 40431) exhibits both patterns (on opposite sides of the skull). TAXONOMIC COMPARISONS: Neacomys marajoara differs from N. dubosti in dorsal pelage color (dark brown finely sprinkled with orange versus light to dark brown finely sprinkled with orange in N. dubosti), and by its bicolored tail (the tail is usually unicolored in N. dubosti), narrower interorbital region, teardrop-shaped incisive foramina (the lateral margins of the incisive foramina are usually subparallel in N. dubosti), globular auditory bullae (the bullae are usually flask-shaped in N. dubosti), and carotid circulation usually pattern 2 (versus pattern 1 in N. dubosti). a We found variation in this character, please see Neacomys xingu description. b We found variation in this character, please see Neacomys marajoara description. Neacomys marajoara differs from N. xingu in dorsal pelage color (dark brown finely sprinkled with orange versus orange-brown sprinkled with black in N. xingu), and by its bicolored tail (the tail is usually unicolored in N. xingu), anteriorly straight (versus anteriorly expanded) nasal bones, broader interorbital region, narrower maxillary septum of the incisive foramina, larger subsquamosal fenestrae, and carotid circulation usually pattern 2 (versus usually pattern 1). Neacomys marajoara differs from N. vossi in dorsal pelage color (dark brown finely sprinkled with orange versus brown sprinkled with orange and/or black in N. vossi), and by its anteriorly straight (versus anteriorly expanded) nasal bones, broader interorbital region, paraoccipital processes separated from the auditory bullae (versus processes close to the auditory bullae), larger subsquamosal fenestrae, carotid circulation usually pattern 2 (versus pattern 1), and narrower maxillary septum of the incisive foramina. Karyotypically, Neacomys marajoara differs from other species in the Dubosti Group by having a diploid chromosomal complement of 58 (versus 2 n = 64 in N. dubosti; table 5), a uniquely large FN of 70 autosomal arms (versus FN = 64–68 in other group members), and submetacentric sex chromosomes (at least one of the sex chromosomes is acrocentric in N. dubosti and “species 2”). DISTRIBUTION: Neacomys marajoara has been recorded only on Marajó Island, state of Pará, Brazil (fig. 5). ETYMOLOGY: The specific epithet marajoara is to be treated as a noun in apposition. It is derived from Tupi Guarani (a language family that comprises many different indigenous Amazonian dialects) and denotes a native of Marajó Island (the type locality). FIELD NOTES: Of the nine specimens for which trapping information is available, eight were captured in pitfall traps and one was taken in a Sherman trap placed on the ground. One paratype (MPEG 40435) was pregnant with three fetuses. REMARKS: Neacomys marajoara was previously reported in the literature as “ Neacomys sp.” (in part) by Silva at al. (2015) and as “ Neacomys sp. D” by Oliveira da Silva et al. (2019).

Neacomys marajoara, new species Marajoara Spiny Mouse Figures 4, 7 HOLOTYPE: The holotype (MPEG 40432) is an adult male (age class 3), collected on 19 January 2009, by R. V. Rossi (field number MAJ 23; fig. 4) in a pitfall trap. The specimen consists of a stuffed skin (missing the tip of the tail), skull, and skeleton; a tissue sample of this specimen is preserved in ethanol, and a partial cytochrome b sequence that we obtained from it has been deposited in Genbank with accession number MT 462070. TYPE LOCALITY: Tauari Farm, municipality of Chaves, Marajó Island, state of Pará, Brazil (0°39′S, 50°11′W, figs. 5, 6). DIAGNOSIS: Neacomys marajoara is a small species (table 3) that differs from congeneric taxa by the following combination of craniodental traits: skull delicate; interorbital region relatively broad; nasal bones straight anteriorly; supraorbital margins slightly convergent anteriorly; subsquamosal fenestra large (almost half the size of the postglenoid foramen on each side of the skull); paraoccipital processes separated from the auditory bullae; sphenopalatine foramen large; carotid circulation usually pattern 2 (sensu Voss, 1988); maxillary part of incisive septum (between the incisive foramina) narrow; M1 usually with flat and undivided anterocone; and M1 anteroloph usually fused with anterolabial conule. MORPHOLOGICAL DESCRIPTION: Dorsal pelage dark brown finely sprinkled with orange (fig. 4); ventral pelage varying from pure white to yellowish white, separated from the dorsal pelage by a very thin orange lateral line. Superciliary, genal, and mystacial vibrissae blackish and long (extending behind ears when laid back alongside the head); submental vibrissae absent; interramal vibrissae short and white. Ears small and rounded; postauricular hairs gray based with orange tips, forming an orange tuft behind each pinna. Ungual tufts white, longer than claws in most specimens examined (except MPEG 40435 and MPEG 40446, in which ungual tufts are as long as the claws); fore- and hind feet covered dorsally with buffy-cream hairs; hind feet narrow and elongate with small interdigital membranes present. Tail about the same length as head and body, bicolored, and covered by short, spiny, and clearly visible hairs; white hairs present on ventral caudal surface from base to midlength; tail tip with very short (1 mm) terminal tuft; caudal scales small, arranged in annular series; each caudal scale with three subequal hairs inserted along its posterior margin. Skull small and delicate in dorsal view (fig. 7), with straight anterior nasal margins, notably broad rostrum, and shallow zygomatic notches; posterior nasal terminus usually slightly pointed, extending beyond maxillary-frontal suture; premaxillaries terminating anterior to nasals; lacrimal bone small and visible in dorsal view, usually in broad contact with frontal bones; supraorbital margins slightly convergent anteriorly; interorbital region relatively broad; supraorbital beads developed as projecting shelves; lateral expansion of the parietal restricted to the dorsal cranial surface. Incisive foramina small and usually teardrop shaped, not extending posteriorly to level of M1s; maxillary portion of incisive septum (dividing the left and right foramina) narrow. Zygomatic plate narrow. Palate with two posterolateral pits on each side. Auditory bullae small and globular, with short and narrow eustachian tubes; periotic bone extends anteriorly to internal carotid canal (except MPEG 40435, MPEG 40443, and MPEG 40439, in which the periotic does not reach the internal carotid canal), but does not enter the canal. Subsquamosal fenestra large (almost half the size of the postglenoid foramen); hamular process of the squamosal long. Paraoccipital process narrow and small and separated from the auditory bullae. Sphenopalatine foramen large; alisphenoid strut absent; carotid circulation pattern usually derived 7 (pattern 2, as identified by the presence of a large stapedial foramen and a large posterior opening of the alisphenoid canal, and by the absence of a squamosalalisphenoid groove and a sphenofrontal foramen; Voss, 1988). First upper molar (M1) usually with broad, flat, and undivided anterocone; anteroloph usually fused with anterolabial conule (such that the anteroflexus is not distinguishable); M3 small; labial cusps (paracone and metacone) usually smaller than lingual cusps (protocone and hypocone); m1 anterocone undivided. Mandible with mental foramen opening laterally; capsular process of lower incisor alveolus present, but indistinct (reduced to a slight rounded elevation), approximately at same height as coronoid process. KARYOTYPES: Karyotypes of specimens from the type locality were obtained by Silva et al. (2015), and karyotypes from specimens collected elsewhere on Marajó Island were obtained by Oliveira da Silva et al. (2019), who described a chromosomal complement of 2 n = 58 and a fundamental number (FN) = 70 (table 5). 7 Most specimens of Neacomys marajoara (seven out of nine) exhibit carotid circulation pattern 2, but one specimen (MPEG 40446) exhibits circulatory pattern 1, and another (MPEG 40431) exhibits both patterns (on opposite sides of the skull). TAXONOMIC COMPARISONS: Neacomys marajoara differs from N. dubosti in dorsal pelage color (dark brown finely sprinkled with orange versus light to dark brown finely sprinkled with orange in N. dubosti), and by its bicolored tail (the tail is usually unicolored in N. dubosti), narrower interorbital region, teardrop-shaped incisive foramina (the lateral margins of the incisive foramina are usually subparallel in N. dubosti), globular auditory bullae (the bullae are usually flask-shaped in N. dubosti), and carotid circulation usually pattern 2 (versus pattern 1 in N. dubosti). a We found variation in this character, please see Neacomys xingu description. b We found variation in this character, please see Neacomys marajoara description. Neacomys marajoara differs from N. xingu in dorsal pelage color (dark brown finely sprinkled with orange versus orange-brown sprinkled with black in N. xingu), and by its bicolored tail (the tail is usually unicolored in N. xingu), anteriorly straight (versus anteriorly expanded) nasal bones, broader interorbital region, narrower maxillary septum of the incisive foramina, larger subsquamosal fenestrae, and carotid circulation usually pattern 2 (versus usually pattern 1). Neacomys marajoara differs from N. vossi in dorsal pelage color (dark brown finely sprinkled with orange versus brown sprinkled with orange and/or black in N. vossi), and by its anteriorly straight (versus anteriorly expanded) nasal bones, broader interorbital region, paraoccipital processes separated from the auditory bullae (versus processes close to the auditory bullae), larger subsquamosal fenestrae, carotid circulation usually pattern 2 (versus pattern 1), and narrower maxillary septum of the incisive foramina. Karyotypically, Neacomys marajoara differs from other species in the Dubosti Group by having a diploid chromosomal complement of 58 (versus 2 n = 64 in N. dubosti; table 5), a uniquely large FN of 70 autosomal arms (versus FN = 64–68 in other group members), and submetacentric sex chromosomes (at least one of the sex chromosomes is acrocentric in N. dubosti and “species 2”). DISTRIBUTION: Neacomys marajoara has been recorded only on Marajó Island, state of Pará, Brazil (fig. 5). ETYMOLOGY: The specific epithet marajoara is to be treated as a noun in apposition. It is derived from Tupi Guarani (a language family that comprises many different indigenous Amazonian dialects) and denotes a native of Marajó Island (the type locality). FIELD NOTES: Of the nine specimens for which trapping information is available, eight were captured in pitfall traps and one was taken in a Sherman trap placed on the ground. One paratype (MPEG 40435) was pregnant with three fetuses. REMARKS: Neacomys marajoara was previously reported in the literature as “ Neacomys sp.” (in part) by Silva at al. (2015) and as “ Neacomys sp. D” by Oliveira da Silva et al. (2019)., Published as part of Semedo, Thiago Borges Fernandes, Silva, Maria Nazareth Ferreira Da, Gutiérrez, Eliécer E., Ferreira, Daniela Cristina, Nunes, Mario Da Silva, Mendes-Oliveira, Ana Cristina, Farias, Izeni Pires & Rossi, Rogério Vieira, 2020, Systematics of Neotropical Spiny Mice, Genus Neacomys Thomas, 1900 (Rodentia: Cricetidae), from Southeastern Amazonia, with Descriptions of Three New Species, pp. 1-43 in American Museum Novitates 2020 (3958) on pages 19-26, DOI: 10.1206/3958.1, http://zenodo.org/record/5056873, {"references":["Voss, R. S. 1988. Systematics and ecology of ichthyomyine rodents (Muroidea): patterns of morphological evolution in a small adaptive radiation. Bulletin of the American Museum of Natural History 188 (2): 259 - 493.","Silva, W. O., et al. 2015. Diversity and karyotypic evolution in the genus Neacomys (Rodentia, Sigmodontinae). Cytogenetic and Genome Research 146: 1 - 10.","Oliveira da Silva, W., et al. 2019. Chromosomal phylogeny and comparative chromosome painting among Neacomys species (Rodentia, Sigmodontinae) from eastern Amazonia. Bmc Evolutionary Biology 19: 1 - 13."]}

Neacomys xingu, new species Xingu Spiny Mouse Figures 11, 12 HOLOTYPE: The holotype (UFMT 1268) is an adult female (age class 5), collected on 28 August 2009 by Cleuton Lima Miranda (original field number PSA 242) in a pitfall trap. The specimen is preserved as a skin, skull, and skeleton in good condition; additionally, a tissue is preserved in alcohol, and a partial cytochrome b sequence that we obtained from it has been deposited in Genbank with accession number MT 462060. TYPE LOCALITY: Flona Tapirapé-Aquiri, Marabá, state of Pará, Brazil (5°46′S, 50°32′W, fig. 5). DIAGNOSIS: Neacomys xingu is a small species (table 3) that differs from congeneric taxa by the following combination of craniodental traits: skull delicate; interorbital region narrow; nasal bones expanded anteriorly; supraorbital margins convergent anteriorly; subsquamosal fenestra small (about ¼ the size of the postglenoid foramen on each side of the skull); paraoccipital process separated from the auditory bullae; sphenopalatine foramen small; carotid circulation usually pattern 1 (sensu Voss, 1988); maxillary part of incisive septum (between the incisive foramina) wide; M1 usually with flat and undivided anterocone; and M1 anteroloph fused with the anterolabial conule. MORPHOLOGICAL DESCRIPTION: Dorsal pelage orange-brown, sprinkled with black (fig. 11); ventral pelage varying from pure white to buffy white and separated from the dorsal pelage by a thin orange lateral line. Superciliary, genal, and mystacial vibrissae blackish and long (extending behind ears when laid back alongside the head); submental vibrissae absent; interramal vibrissae short and white; Ears small and rounded; postauricular hairs gray based with orange tips, forming an orange tuft behind each pinna. Ungual tufts white, longer than claws in length; fore- and hind feet covered dorsally with buffy-cream hairs; hind feet narrow and elongate with small interdigital membranes present. Tail about the same length as head and body, usually unicolored (except in PSA 069, MPEG 39901, and MPEG 42019, in which the tail is dark above and paler below), and covered by short, spiny, and clearly visible hairs; tail tip with very short (1–2 mm) terminal tuft; caudal scales small, arranged in annular series; each caudal scale with three subequal hairs inserted along its posterior margin. Skull small and delicate in dorsal view (fig. 12); with anteriorly expanded nasal margins; notably broad rostrum and shallow zygomatic notches; posterior nasal terminus slightly pointed, extending beyond the maxillary-frontal suture; premaxillaries terminating slightly anterior to nasals; lacrimal bone small and visible in dorsal view, equally contacting the maxillary and frontal bones; supraorbital margins convergent anteriorly; interorbital region narrow; supraorbital beads developed as projecting shelves; lateral expansion of the parietal restricted to the dorsal cranial surface (except in MCN-M 1404 and MPEG 39901, in which the parietal is slightly expanded ventrally near the squamosal root of the zygomatic arch). Incisive foramina small with subparallel lateral margins, not extending posteriorly to level of M1s; maxillary portion of incisive septum (dividing the left and right foramina) usually wide. Zygomatic plate varying from broad to narrow. Palate with two posterolateral pits on each side. Auditory bullae small and usually globular, with short and narrow eustachian tubes; periotic bone extends anteriorly to the internal carotid canal but does not enter it (except in MPEG 42715 and MCN-M 1404, in which the periotic does not reach the internal carotid canal). Subsquamosal fenestra small (about ¼ the size of the postglenoid foramen); hamular process of the squamosal long. Paraoccipital process narrow and small and usually separated from the auditory bullae (Sánchez-Vendizú et al., 2018: fig. 3C). Sphenopalatine foramen small (except in MPEG 41991, whose sphenopalatine foramen is large); alisphenoid strut absent; carotid circulation pattern usually primitive 8 (pattern 1, as identified by retaining a well-developed squamosal-alisphenoid groove and sphenofrontal foramen, both indicative of the presence of the supraorbital branch of the stapedial artery; Voss, 1988: fig. 18A, B). First upper molar (M1) usually with slightly flat and undivided anterocone; anteroloph usually fused with the anterolabial conule (such that the anteroflexus is not distinguishable); M3 small; labial cusps (paracone, metacone) usually taller than lingual cusps (protocone, hypocone); m1 anteroconid undivided. Mandible with mental foramen opening laterally; capsular process of lower incisor alveolus present, but indistinct (reduced to a slight rounded elevation), approximately at same height of coronoid process. TAXONOMIC COMPARISONS: Neacomys xingu differs from N. dubosti in dorsal pelage color (orange-brown sprinkled with black versus light to dark brown finely sprinkled with orange), and by its narrower interorbital region, globular auditory bullae (the bullae are usually flask shaped in N. dubosti), and carotid circulation usually pattern 1 (versus always pattern 1 in N. dubosti). Karyotypically, Neacomys xingu differs from all other members of the Dubosti Group by having a diploid chromosomal complement of 58 (versus 2 n = 64 in N. dubosti and 2 n = 54 in “species 2”), a uniquely small FN of 64 autosomal arms (versus FN = 66– 60 in other species), and submetacentric sex chromosomes (at least one sex chromosome is acrocentric in N. dubosti and “species 2”). DISTRIBUTION: Neacomys xingu has been collected on the right bank of the lower Xingu River and in the region of Serra de Carajás, in southeastern Pará state (fig. 5). According to cytogenetic data (Di-Nizo et al., 2017; Oliveira da Silva et al., 2019), the species also ranges southward into Vila Rica in northeastern Mato Grosso state. These records suggest that the species is restricted to the Xingu center of endemism (Silva et al., 2005). ETYMOLOGY: The specific epithet xingu is to be treated as a noun in apposition. The species name refers to the Xingu center of endemism, delimited by the Xingu and Tapajós rivers, where the species occurs. REMARKS: Neacomys xingu was previously reported in the literature as “ Neacomys clade 7” by Patton et al. (2000), “ Neacomys sp.” (in part) by Silva at al. (2015), “ Neacomys sp.” by Di-Nizo et al. (2017) and Brandão et al. (2019), and “ Neacomys sp. C” by Oliveira da Silva et al. (2019)., Published as part of Semedo, Thiago Borges Fernandes, Silva, Maria Nazareth Ferreira Da, Gutiérrez, Eliécer E., Ferreira, Daniela Cristina, Nunes, Mario Da Silva, Mendes-Oliveira, Ana Cristina, Farias, Izeni Pires & Rossi, Rogério Vieira, 2020, Systematics of Neotropical Spiny Mice, Genus Neacomys Thomas, 1900 (Rodentia: Cricetidae), from Southeastern Amazonia, with Descriptions of Three New Species, pp. 1-43 in American Museum Novitates 2020 (3958) on pages 30-33, DOI: 10.1206/3958.1, http://zenodo.org/record/5056873, {"references":["Voss, R. S. 1988. Systematics and ecology of ichthyomyine rodents (Muroidea): patterns of morphological evolution in a small adaptive radiation. Bulletin of the American Museum of Natural History 188 (2): 259 - 493.","Sanchez-Vendizu, P., V. Pacheco and D. Vivas-Ruiz. 2018. An introduction to the systematics of smallbodied Neacomys (Rodentia: Cricetidae) from Peru with descriptions of two new species. American Museum Novitates 3913: 1 - 38.","Di-Nizo, C. B., K. R. S. Banci, Y. Sato-Kuwabara, and M. J. J. Silva. 2017. Advances in cytogenetics of Brazilian rodents: cytotaxonomy, chromosome evolution and new karyotypic data. Comparative Cytogenetics 11: 833 - 89.","Oliveira da Silva, W., et al. 2019. Chromosomal phylogeny and comparative chromosome painting among Neacomys species (Rodentia, Sigmodontinae) from eastern Amazonia. Bmc Evolutionary Biology 19: 1 - 13.","Silva, J. M. C., A. B. Rylands and G. A. B. Fonseca 2005. O destino das areas de endemismo da Amazonia. Megadiversidade 1: 124 - 131.","Patton, J. L., M. N. F. da Silva, and J. R. Malcolm. 2000. Mammals of the rio Jurua and the evolutionary and ecological diversification of Amazonia. Bulletin of the American Museum of Natural History 244: 1 - 306.","Brandao, M. B. et al. 2019. Mammals of Mato Grosso, Brazil: Annotated species list and historical review. Mastozoologia Neotropical, 26: 263 - 307."]}

Semedo, Thiago Borges Fernandes, Silva, Maria Nazareth Ferreira Da, Gutiérrez, Eliécer E., Ferreira, Daniela Cristina, Nunes, Mario Da Silva, Mendes-Oliveira, Ana Cristina, Farias, Izeni Pires, Rossi, Rogério Vieira (2020): Systematics of Neotropical Spiny Mice, Genus Neacomys Thomas, 1900 (Rodentia: Cricetidae), from Southeastern Amazonia, with Descriptions of Three New Species. American Museum Novitates 2020 (3958): 1-43, DOI: https://doi.org/10.1206/3958.1, URL: http://dx.doi.org/10.1206/3958.1

Neacomys vossi, new species Voss’s Spiny Mouse Figures 8, 9 HOLOTYPE: The holotype (UFPA 1583) is an adult female (age class 2) collected on 2 April 2013, by Ana Cristina Mendes Oliveira (original field number JB 031). The specimen consists of a stuffed skin, skull, and skeleton, all in good condition; additionally, a tissue sample is preserved in ethanol, and a partial cytochrome b sequence that we obtained from it has been deposited in Genbank with accession number MT 462028. TYPE LOCALITY: Boca do Rato, on the right bank of the Rio Tapajós, Itaituba municipality, state of Pará, Brazil (5°14′S, 56°56′W, fig. 5). DIAGNOSIS: Neacomys vossi is a small species (table 3) that differs from congeneric taxa by the following combination of craniodental traits: skull delicate; interorbital region narrow; nasal bones expanded anteriorly; supraorbital margins convergent anteriorly; subsquamosal fenestra usually small (about ¼ the size of the postglenoid foramen on each side of the skull); paroccipital processes close to the auditory bullae; sphenopalatine foramen large; carotid circulation pattern 1 (sensu Voss, 1988); maxillary part of incisive septum (between the incisive foramina) usually wide; M1 usually with slightly flat, narrow, and undivided anterocone; and M1 anteroloph usually distinct (not fused with the anterolabial conule). MORPHOLOGICAL DESCRIPTION: Dorsal pelage brown sprinkled with orange and/or black (fig. 8); ventral pelage varying from pure white to buffy white; very thin orange lateral line. Superciliary, genal, and mystacial vibrissae blackish and long (extending behind ears when laid back alongside the head); submental vibrissae absent; interramal vibrissae short and white. Ears small and rounded; post-auricular hairs gray-based with orange tips, forming an orange tuft behind each pinna. Ungual tufts white, longer than claws; fore- and hind feet covered dorsally with buffycream hairs; hind feet narrow and elongate with small interdigital membranes. Tail about the same length as head and body, usually slightly bicolored (except in UFPA 1277, 1417, 1444, 1654, and 1736 which have unicolored tail) and covered by spiny and clearly visible hairs; tail tip of the tail with very short (approximately 1.4 mm long) terminal tuft; caudal scales small, arranged in annular series; each caudal scale with three subequal hairs inserted along its posterior margin. Skull small and delicate in dorsal view (fig. 9); with expanded anterior nasal margins; notably broad rostrum and shallow zygomatic notches; posterior nasal terminus usually flat, extending beyond the maxillary-frontal suture; premaxillaries terminating slightly anterior to nasals; lacrimal bone small and visible in dorsal view, equally contacting maxillary and frontal bones; supraorbital margins convergent anteriorly; interorbital region narrow; supraorbital beads developed as projecting shelves; lateral expansion of the parietal restricted to the dorsal surface (except in UFPA 1227 and UFPA 1577, in which the parietal is slightly expanded ventrally near the squamosal root of the zygomatic arch). Incisive foramina small with subparallel lateral margins, not extending posteriorly to the level of M1s; maxillary portion of incisive septum (dividing the left and right foramina) usually wide. Zygomatic plate narrow. Palate with two posterolateral pits on each side of the palate. Auditory bullae small and globular, with short and narrow eustachian tubes; periotic bone extending anteriorly to internal carotid canal, but usually not entering into it (except in UFPA 1227, 1417, 1530, 1654, 1417, and 1736, in which the periotic does enter the internal carotid canal). Subsquamosal fenestra usually small (about ¼ the size of the postglenoid foramen); hamular process of the squamosal long. Paraoccipital process narrow, small, and close to the auditory bullae (Sánchez-Vendizú et al., 2018: fig. 3C). Sphenopalatine foramen large; alisphenoid strut absent; carotid circulation primitive (pattern 1, as identified by retaining a well-developed squamosal-alisphenoid groove and sphenofrontal foramen, both indicative of the presence of the supraorbital branch of the stapedial artery; Voss et al., 1988). First upper molar (M1) usually with slightly flat and undivided anterocone; anteroloph usually distinct (not fused with the anterolabial conule); M3 small; labial cusps (paracone, metacone) usually taller than lingual cusps (protocone, hypocone); m1 anteroconid undivided. Mandible with mental foramen opening laterally; capsular process of lower incisor alveolus present, but indistinct (reduced as a slight, rounded elevation), approximately at same height as coronoid process. TAXONOMIC COMPARISONS: Neacomys vossi differs from N. dubosti in dorsal pelage color (brown sprinkled with orange and/or black versus dark brown finely sprinkled with orange in N. dubosti), and by its narrower interorbital region, and globular auditory bullae (the bullae are usually flask shaped in N. dubosti). Neacomys vossi differs from N. xingu in dorsal pelage color (brown sprinkled with orange and/or black versus orange-brown sprinkled with black in N. xingu), larger sphenopalatine foramen, and an M1 anteroloph that is usually distinct (versus fused in N. xingu). Karyotypically, Neacomys vossi differs from other species in the Dubosti Group by having a diploid chromosome complement of 58 (versus 2 n = 64 in N. dubosti and 2 n = 64 in “species 2”), an FN of 68 autosomal arms (versus FN = 70 in N. marajoara, FN = 64 in N. xingu, and FN = 66 in “species 2”), and submetacentric sex chromosomes (at least one of the sex chromosomes is acrocentric in N. dubosti and “species 2”) DISTRIBUTION: Neacomys vossi has been collected on the right bank of the upper and middle Tapajós River and on the left bank of lower Xingu (fig. 5). According to our records, the species appears to be endemic to the Tapajós center of endemism (Silva et al., 2005). ETYMOLOGY: Named in honor of Robert S. Voss (fig. 10), curator of mammals at the American Museum of Natural History, New York, for his extensive contributions to our knowledge of Neotropical mammals, especially the taxonomy of Neacomys in northeastern Amazonia. . FIELD NOTES: Among the specimens we examined for which trapping information is available, all 17 were captured in pitfall traps, two (UFPA 1277 and 1284) in upland (terra firme) forests and three (UFPA 1444, 1487, and 1583) in primary forest of unknown character. REMARKS: Neacomys vossi was previously reported in the literature as “ Neacomys sp. A” by Oliveira da Silva et al. (2017, 2019).

Amazophrynella bilinguis sp. nov. Kaefer, Rojas, Ferrão & Lima urn:lsid:zoobank.org:act: 6E3730AD-CC35-47B1-99F6-E7FEC5382F9F Holotype. INPA-H 39784 (field number APL 18244), adult male (Fig. 1 A–B, 2A–C, 3A, 3C, 4A, 5B) collected on February 2012 by I.L. Kaefer, A.P. Lima and L. Vasconcelos in the farm Fazenda Taperinha (2°34’48.4” S; 54°22’16.6” W, 7 m a.s.l.) on the south margin of the Amazon River, Municipality of Santarém, Pará State, Brazil. Paratype s . Eleven specimens. Nine males: INPA-H 39782, INPA-H 39780, INPA-H 39774, INPA-H 39776, INPA-H 39783, INPA-H 39778, INPA-H 39781, INPA-H 39775, INPA-H 39785 (field numbers APL 18240, APL 18241, APL 18243, APL 18245, APL 18246, APL 18247, APL 18248, APL 18249, and APL 18250, respectively); two females: INPA-H 39779 and INPA-H 39777 (field numbers APL 18242, APL 18251, respectively); all collected by I.L. Kaefer, A.P. Lima and L. Vasconcelos between February 3–5 2012 at the same locality of holotype. Female INPA-H 39777 is designated as the allotype (Fig. 1 C–D, 2D–F, 3B, 3D, 4B, 5A). Diagnosis. A species of Amazophrynella characterized by: (1) SVL 13–14.5 mm in males and 19.6–20.4 mm in females; (2) Finger I Ż Finger II; (3) palmar tubercle rounded; (4) dorsal surfaces highly granular; (5) whitish belly with black tiny spots in life; (6) call type 1 composed by one note with duration of 0.206– 0.292 s and 69–77 pulses; (7) call type 2 with duration of 0.779– 3.191 s and constituted by 6–22 multipulsed notes (9–14 pulses/note). Comparison with other species. We compared the new species with all currently recognized species of Amazophrynella: A. amazonicola; A. bokermanni; A. javierbustamantei Rojas, Chaparro, Carvalho, Ávila, Farias, Hrbek, and Gordo, 2016; A. manaos Rojas, Carvalho, Ávila, Farias, and Hrbek, 2014; A. matses Rojas, Carvalho, Ávila, Farias, Gordo, and Hrbek, 2015; A. minuta; A. moisesii Rojas, Fouquet, Ron, Hernández-Ruz, Melo- Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, Ávila, Farias, Gordo, and Hrbek, 2018; A. siona; A. teko Rojas, Fouquet, Ron, Hernández-Ruz, Melo-Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, Ávila, Farias, Gordo, and Hrbek, 2018; A. vote Ávila, Carvalho, Gordo, Kawashita-Ribeiro, and Morais, 2012; A. xinguensis Rojas, Fouquet, Ron, Hernández-Ruz, Melo-Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, Ávila, Farias, Gordo, and Hrbek, 2018. Amazophrynella bilinguis sp. nov. can be distinguished from all other Amazophrynella species by the combination of morphological and bioacoustic characters. In addition, the new species differs genetically from all other species in the genus. Characteristics of compared species are presented in parentheses. The whitish belly in life distinguishes Amazophrynella bilinguis sp. nov. from A. minuta (yellow to orange: Rojas et al. 2018c), A. amazonicola (yellow to reddish orange: Rojas et al. 2015), A. matses (yellow to yellowish orange: Rojas et al. 2015), A. siona (yellow to reddish brown: Rojas et al. 2018a), A. javierbustamantei (pale yellow: Rojas et al. 2016), A. moisesi (pale yellow: Rojas et al. 2018a), A. teko (pale yellow to cream: Rojas et al. 2018a). The Finger I Ż Finger II in A. bilinguis sp. nov. differs from those of all above cited species (Finger I smaller than Finger II: Ávila et al. 2012, Rojas et al. 2014, 2015, 2016, 2018a, 2018c). Additionally, by emitting call type 1 with duration of 0.206– 0.292 s and 69–77 pulses, the new species can be distinguished from A. minuta (0.132– 0.143 s and 43–48 pulses: present study), and A. teko (0.150– 0.190 s and 10–30 pulses: Rojas et al. 2018a). In addition, A. bilinguis sp. nov. emits call type 2 with multipulsed notes that differs from those of A. amazonicola, A. minuta, and A. siona (non-pulsed: Rojas et al. 2018a, b, present study; see Discussion section). The whitish belly of the new species is similar to A. manaos, A. vote, A. xinguensis and A. bokermanni. The new species differs from A. manaos by having Finger I Ż Finger II (Finger I shorter than Finger II: Rojas et al. 2014), whitish belly with black tiny spots (whitish belly with black blotches: Rojas et al. 2014), and advertisement call type 1 with duration of 0.206– 0.292 s (0.133– 0.156 s: Rojas et al. 2018b). Amazophrynella bilinguis sp. nov. differs from A. vote by reaching 14.5 mm SVL in males and 20.4 mm in females (maximum SVL 19.3 mm in males, and 25.7 mm in females: Ávila et al. 2012) and Finger I Ż Finger II (Finger I et al. 2012). Moreover, A. bilinguis sp. nov. differs of A. vote by having call type 1 with duration of 0.206– 0.292 s and 69–77 pulses (0.098– 0.150 s and 41–60 pulses: Rojas et al. 2018b). According to the genetic analyses (see section Phylogenetic relationships), A. xinguensis and A. bokermanni are the most closely-related nominal species in relation to Amazophrynella bilinguis sp. nov. The new species can be distinguished from A. xinguensis by having SVL 13–14.5 mm in males and 19.6–20.4 mm in females (SVL males 17.7–20.0 mm; SVL females 22.4–26.3 mm: Rojas et al. 2018a), palmar tubercle rounded (elliptical: Rojas et al. 2018a). The advertisement call of A. xinguensis is unknown. Amazophrynella bilinguis sp. nov. is differentiated from A. bokermanni by SVL 13–14.5 mm in males (SVL = 15.9–16.5 mm: Rojas et al. 2018a), dorsal surfaces highly granular (granular), black tiny spots on venter (small black dots). Moreover, the advertisement call type 1 of A. bilinguis sp. nov. can be distinguished from those of A. bokermanni by duration of notes 0.206– 0.292 s (0.125– 0.163 s: Rojas et al. 2018b). Description of holotype. Body small, elongate. Head triangular in dorsal and lateral view. Head longer than wide. HL 38.04% and HW 30.43% of SVL. Snout acuminate in lateral view and triangular in dorsal view. SL 44.4% of HL. Nostrils protuberant, closer to snout than eyes. Cantus rostralis straight in dorsal view. Internarinal distance smaller than eye diameter. IND 28.6% of HW. Upper eyelid covered by small granules. Eye prominent, 29.0% of HL. Tympanum not visible through the skin. Texture of skin on tympanum covered with granules. Texture of dorsal skin highly granular. Texture of dorsolateral skin granular. Abundance of granules on arms insertion. Forelimbs thick. Upper arms robust. UAL 39.1% of SVL. Presence of granules on upper arm and arm. HAL 20.9% of UAL. Fingers slender. Tips unexpanded. Fingers basally webbed. Relative length of Fingers: I Ż II Measurement of the holotype (in mm) . SVL: 13.8; HW: 4.2; HL: 5.2; SL: 2.3; ED: 1.5; IND: 1.2; UAL: 5.4; HAL: 2.9; FI: 1.5; FII:1.5; THL: 8.1; TAL: 7.5; TL: 5.1; FL: 4.9. Coloration of the holotype. In life, head brown in dorsal view. Dorsum brown with brown chevrons. Flanks light brown. Dorsal surfaces of upper arm, arm and hand light brown. Dorsal surfaces of thighs, tibia, tarsus and foot light brown. Gular region cream with brown spots. Belly whitish with brown spots and white granules (Fig. 5B). Ventral surfaces of upper arm and arm creamy. Palm reddish. Ventral surfaces of thighs, tarsus and tibia creamy, sole black. Iris golden and pupil black. In preservative, the coloration is almost the same than the one in life. The coloration of the dorsum becomes light brown. Gular region and venter become cream. The iris loses its coloration. Fingers and toes become cream (Fig. 6). Variation. There is little variation among specimens of the type series (Table 2). Sexual dimorphism was detected in SVL, with 13.0– 14.5 mm (13.65 ± 0.43 mm, n = 10) in males and 19.56–20.4 mm (19.98 ± 0.59 mm, n =2) in females. Three specimens (INPA-H 39775, INPA-H 39780, INPA-H 39776) have few granules on posterior region of head. The specimen INPA-H 39779 present a white line from head to cloaca. There is variation in the size (mm) of the FI and FII between individuals, while some (e.g. INPA-H 39778, INPA-H 39774, INPA-H 39783) present Finger I> Finger II and other present Finger I Ż Finger II (e.g. INPA-H 39782, INPA-H 39784, INPA-H 39775). Subarticular tubercles more protruding and swollen in females. The specimen INPA-H39777 shows small spots on dorsolateral surfaces. Dorsum with different tonalities of brown (light brown to brown). Spots on venter vary in sizes (small to medium size). Thighs, shanks and tarsus between cream and whitish coloration, in ventral view. Palm and sole present different tonalities of cream, in ventral view. In preserved specimens, the palmar tubercle is more flattened. Advertisement call. Two different call types were recorded (Fig. 7 A–D), both of them from the same two individuals. Additional males, not recorded, also emitted both call types. We did not categorize these vocalizations as a single composed advertisement call because each one of the call types can be repeated for several minutes. The calls of type 1 (Fig. 7A) consist of one multipulsed note (Fig. 7C) with average duration of 0.248 ± 0.020 s (0.206– 0.292 s) and are composed by 72 ± 2 pulses (69–77 pulses). Calls type 1 are emitted in series (Fig. 7A) and are interleaved by silent intervals of 1.076 ± 0.162 s (0.621– 3.779 s). Calls type 1 have a mean dominant frequency of 3526 ± 170 Hz (3338–4264 Hz), low frequency of 3058 ± 119 Hz (2674–3234 Hz) and high frequency of 4478 ± 173 Hz (4277–4744 Hz). Different from calls of type 1, the calls of type 2 are arranged in bouts (Fig. 7B) of notes and have a duration of 1.9 ± 0.77 s (0.779– 3.191 s). The mean number of notes per call is 13 ± 5 (6–22 notes) and notes are 0.034 ± 0.001 s (0.010– 0.065 s) long in average. The duration of the silent interval between notes in call type 2 is 0.125 ± 0.028 s (0.085– 0.206 s). Notes are composed of 12 ± 1 pulses (9–14 pulses) (Fig. 7D). The mean dominant frequency of calls type 2 is 3450 ± 68 Hz (3220–4680 Hz), while the mean low frequency is 3200 ± 18 Hz (2961–3410 Hz) and high frequency is 4002 ± 91 Hz (3345–4680 Hz). Phylogenetic relationships. Our phylogenetic tree inferred from 559 aligned sites of 16S rRNA (Fig. 8) was partially concordant with Rojas et al. (2018a). All nominal species of Amazophrynella in our reconstruction formed well supported clades (PP Ż 0.95), excepted A. siona, and A. teko. Amazophrynella bilinguis sp. nov. was recovered as the sister taxon of Amazophrynella sp. 3 (sensu Rojas et al. 2018a) from Parque de Desenvolvimento Jatobá (Tapajós River, Pará, Brazil) with low support (PP = 0.74). Genetic distance between A. bilinguis sp. nov. and Amazophrynella sp. 3 was relatively low (p and K2P = 2%) compared to distances calculated between A. bilinguis sp. nov. and other Amazophrynella species that ranged from 3% (p) and 4% (K2P) (Amazophrynella sp. 2) to 14% (p) and 16% (K2P) (A. siona). Amazophrynella bilinguis sp. nov. presented 4% (p and K2P) genetic distance from samples of A. bokermanni, the nominal species that was placed closest to it in the phylogenetic tree. See Table 3 for genetic distances among all analyzed species. Distribution and natural history. Amazophrynella bilinguis sp. nov. is known only from Fazenda Taperinha (2°34’48.4” S; 54°22’16.6” W, 7 m a.s.l.) on south margin of the Amazon River, Municipality of Santarém, Pará State, Brazil (Fig. 9). However, it is likely that sampling efforts, which are historically scarce at that region, might expand the known distribution of this taxon. Like in other species within the genus, males of this species were found calling during the day, usually perched up to 10–40 cm above the ground on vegetation, fallen logs, branches or on leaf litter on the forest floor. Females and vocally active males were found near a stream and associated shallow ponds, where egg deposition and tadpole development may occur. After collection, a pair (female = INPA-H 39779, SVL = 20.8 mm; male = INPA-H 39774; SVL = 15.4 mm) engaged in axillary amplexus inside a plastic bag in the improvised field laboratory (Fig. 4A and 4B). Etymology. The epithet " bilinguis " is Latin and means "bilingual". It refers to the two distinct advertisement calls emitted by the male individuals.

Amazophrynella bilinguis sp. nov. Kaefer, Rojas, Ferr��o & Lima urn:lsid:zoobank.org:act: 6E3730AD-CC35-47B1-99F6-E7FEC5382F9F Holotype. INPA-H 39784 (field number APL 18244), adult male (Fig. 1 A���B, 2A���C, 3A, 3C, 4A, 5B) collected on February 2012 by I.L. Kaefer, A.P. Lima and L. Vasconcelos in the farm Fazenda Taperinha (2��34���48.4��� S; 54��22���16.6��� W, 7 m a.s.l.) on the south margin of the Amazon River, Municipality of Santar��m, Par�� State, Brazil. Paratype s . Eleven specimens. Nine males: INPA-H 39782, INPA-H 39780, INPA-H 39774, INPA-H 39776, INPA-H 39783, INPA-H 39778, INPA-H 39781, INPA-H 39775, INPA-H 39785 (field numbers APL 18240, APL 18241, APL 18243, APL 18245, APL 18246, APL 18247, APL 18248, APL 18249, and APL 18250, respectively); two females: INPA-H 39779 and INPA-H 39777 (field numbers APL 18242, APL 18251, respectively); all collected by I.L. Kaefer, A.P. Lima and L. Vasconcelos between February 3���5 2012 at the same locality of holotype. Female INPA-H 39777 is designated as the allotype (Fig. 1 C���D, 2D���F, 3B, 3D, 4B, 5A). Diagnosis. A species of Amazophrynella characterized by: (1) SVL 13���14.5 mm in males and 19.6���20.4 mm in females; (2) Finger I Ż Finger II; (3) palmar tubercle rounded; (4) dorsal surfaces highly granular; (5) whitish belly with black tiny spots in life; (6) call type 1 composed by one note with duration of 0.206��� 0.292 s and 69���77 pulses; (7) call type 2 with duration of 0.779��� 3.191 s and constituted by 6���22 multipulsed notes (9���14 pulses/note). Comparison with other species. We compared the new species with all currently recognized species of Amazophrynella: A. amazonicola; A. bokermanni; A. javierbustamantei Rojas, Chaparro, Carvalho, ��vila, Farias, Hrbek, and Gordo, 2016; A. manaos Rojas, Carvalho, ��vila, Farias, and Hrbek, 2014; A. matses Rojas, Carvalho, ��vila, Farias, Gordo, and Hrbek, 2015; A. minuta; A. moisesii Rojas, Fouquet, Ron, Hern��ndez-Ruz, Melo- Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, ��vila, Farias, Gordo, and Hrbek, 2018; A. siona; A. teko Rojas, Fouquet, Ron, Hern��ndez-Ruz, Melo-Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, ��vila, Farias, Gordo, and Hrbek, 2018; A. vote ��vila, Carvalho, Gordo, Kawashita-Ribeiro, and Morais, 2012; A. xinguensis Rojas, Fouquet, Ron, Hern��ndez-Ruz, Melo-Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, ��vila, Farias, Gordo, and Hrbek, 2018. Amazophrynella bilinguis sp. nov. can be distinguished from all other Amazophrynella species by the combination of morphological and bioacoustic characters. In addition, the new species differs genetically from all other species in the genus. Characteristics of compared species are presented in parentheses. The whitish belly in life distinguishes Amazophrynella bilinguis sp. nov. from A. minuta (yellow to orange: Rojas et al. 2018c), A. amazonicola (yellow to reddish orange: Rojas et al. 2015), A. matses (yellow to yellowish orange: Rojas et al. 2015), A. siona (yellow to reddish brown: Rojas et al. 2018a), A. javierbustamantei (pale yellow: Rojas et al. 2016), A. moisesi (pale yellow: Rojas et al. 2018a), A. teko (pale yellow to cream: Rojas et al. 2018a). The Finger I Ż Finger II in A. bilinguis sp. nov. differs from those of all above cited species (Finger I smaller than Finger II: ��vila et al. 2012, Rojas et al. 2014, 2015, 2016, 2018a, 2018c). Additionally, by emitting call type 1 with duration of 0.206��� 0.292 s and 69���77 pulses, the new species can be distinguished from A. minuta (0.132��� 0.143 s and 43���48 pulses: present study), and A. teko (0.150��� 0.190 s and 10���30 pulses: Rojas et al. 2018a). In addition, A. bilinguis sp. nov. emits call type 2 with multipulsed notes that differs from those of A. amazonicola, A. minuta, and A. siona (non-pulsed: Rojas et al. 2018a, b, present study; see Discussion section). The whitish belly of the new species is similar to A. manaos, A. vote, A. xinguensis and A. bokermanni. The new species differs from A. manaos by having Finger I Ż Finger II (Finger I shorter than Finger II: Rojas et al. 2014), whitish belly with black tiny spots (whitish belly with black blotches: Rojas et al. 2014), and advertisement call type 1 with duration of 0.206��� 0.292 s (0.133��� 0.156 s: Rojas et al. 2018b). Amazophrynella bilinguis sp. nov. differs from A. vote by reaching 14.5 mm SVL in males and 20.4 mm in females (maximum SVL 19.3 mm in males, and 25.7 mm in females: ��vila et al. 2012) and Finger I Ż Finger II (Finger I et al. 2012). Moreover, A. bilinguis sp. nov. differs of A. vote by having call type 1 with duration of 0.206��� 0.292 s and 69���77 pulses (0.098��� 0.150 s and 41���60 pulses: Rojas et al. 2018b). According to the genetic analyses (see section Phylogenetic relationships), A. xinguensis and A. bokermanni are the most closely-related nominal species in relation to Amazophrynella bilinguis sp. nov. The new species can be distinguished from A. xinguensis by having SVL 13���14.5 mm in males and 19.6���20.4 mm in females (SVL males 17.7���20.0 mm; SVL females 22.4���26.3 mm: Rojas et al. 2018a), palmar tubercle rounded (elliptical: Rojas et al. 2018a). The advertisement call of A. xinguensis is unknown. Amazophrynella bilinguis sp. nov. is differentiated from A. bokermanni by SVL 13���14.5 mm in males (SVL = 15.9���16.5 mm: Rojas et al. 2018a), dorsal surfaces highly granular (granular), black tiny spots on venter (small black dots). Moreover, the advertisement call type 1 of A. bilinguis sp. nov. can be distinguished from those of A. bokermanni by duration of notes 0.206��� 0.292 s (0.125��� 0.163 s: Rojas et al. 2018b). Description of holotype. Body small, elongate. Head triangular in dorsal and lateral view. Head longer than wide. HL 38.04% and HW 30.43% of SVL. Snout acuminate in lateral view and triangular in dorsal view. SL 44.4% of HL. Nostrils protuberant, closer to snout than eyes. Cantus rostralis straight in dorsal view. Internarinal distance smaller than eye diameter. IND 28.6% of HW. Upper eyelid covered by small granules. Eye prominent, 29.0% of HL. Tympanum not visible through the skin. Texture of skin on tympanum covered with granules. Texture of dorsal skin highly granular. Texture of dorsolateral skin granular. Abundance of granules on arms insertion. Forelimbs thick. Upper arms robust. UAL 39.1% of SVL. Presence of granules on upper arm and arm. HAL 20.9% of UAL. Fingers slender. Tips unexpanded. Fingers basally webbed. Relative length of Fingers: I Ż II Measurement of the holotype (in mm) . SVL: 13.8; HW: 4.2; HL: 5.2; SL: 2.3; ED: 1.5; IND: 1.2; UAL: 5.4; HAL: 2.9; FI: 1.5; FII:1.5; THL: 8.1; TAL: 7.5; TL: 5.1; FL: 4.9. Coloration of the holotype. In life, head brown in dorsal view. Dorsum brown with brown chevrons. Flanks light brown. Dorsal surfaces of upper arm, arm and hand light brown. Dorsal surfaces of thighs, tibia, tarsus and foot light brown. Gular region cream with brown spots. Belly whitish with brown spots and white granules (Fig. 5B). Ventral surfaces of upper arm and arm creamy. Palm reddish. Ventral surfaces of thighs, tarsus and tibia creamy, sole black. Iris golden and pupil black. In preservative, the coloration is almost the same than the one in life. The coloration of the dorsum becomes light brown. Gular region and venter become cream. The iris loses its coloration. Fingers and toes become cream (Fig. 6). Variation. There is little variation among specimens of the type series (Table 2). Sexual dimorphism was detected in SVL, with 13.0��� 14.5 mm (13.65 �� 0.43 mm, n = 10) in males and 19.56���20.4 mm (19.98 �� 0.59 mm, n =2) in females. Three specimens (INPA-H 39775, INPA-H 39780, INPA-H 39776) have few granules on posterior region of head. The specimen INPA-H 39779 present a white line from head to cloaca. There is variation in the size (mm) of the FI and FII between individuals, while some (e.g. INPA-H 39778, INPA-H 39774, INPA-H 39783) present Finger I> Finger II and other present Finger I Ż Finger II (e.g. INPA-H 39782, INPA-H 39784, INPA-H 39775). Subarticular tubercles more protruding and swollen in females. The specimen INPA-H39777 shows small spots on dorsolateral surfaces. Dorsum with different tonalities of brown (light brown to brown). Spots on venter vary in sizes (small to medium size). Thighs, shanks and tarsus between cream and whitish coloration, in ventral view. Palm and sole present different tonalities of cream, in ventral view. In preserved specimens, the palmar tubercle is more flattened. Advertisement call. Two different call types were recorded (Fig. 7 A���D), both of them from the same two individuals. Additional males, not recorded, also emitted both call types. We did not categorize these vocalizations as a single composed advertisement call because each one of the call types can be repeated for several minutes. The calls of type 1 (Fig. 7A) consist of one multipulsed note (Fig. 7C) with average duration of 0.248 �� 0.020 s (0.206��� 0.292 s) and are composed by 72 �� 2 pulses (69���77 pulses). Calls type 1 are emitted in series (Fig. 7A) and are interleaved by silent intervals of 1.076 �� 0.162 s (0.621��� 3.779 s). Calls type 1 have a mean dominant frequency of 3526 �� 170 Hz (3338���4264 Hz), low frequency of 3058 �� 119 Hz (2674���3234 Hz) and high frequency of 4478 �� 173 Hz (4277���4744 Hz). Different from calls of type 1, the calls of type 2 are arranged in bouts (Fig. 7B) of notes and have a duration of 1.9 �� 0.77 s (0.779��� 3.191 s). The mean number of notes per call is 13 �� 5 (6���22 notes) and notes are 0.034 �� 0.001 s (0.010��� 0.065 s) long in average. The duration of the silent interval between notes in call type 2 is 0.125 �� 0.028 s (0.085��� 0.206 s). Notes are composed of 12 �� 1 pulses (9���14 pulses) (Fig. 7D). The mean dominant frequency of calls type 2 is 3450 �� 68 Hz (3220���4680 Hz), while the mean low frequency is 3200 �� 18 Hz (2961���3410 Hz) and high frequency is 4002 �� 91 Hz (3345���4680 Hz). Phylogenetic relationships. Our phylogenetic tree inferred from 559 aligned sites of 16S rRNA (Fig. 8) was partially concordant with Rojas et al. (2018a). All nominal species of Amazophrynella in our reconstruction formed well supported clades (PP Ż 0.95), excepted A. siona, and A. teko. Amazophrynella bilinguis sp. nov. was recovered as the sister taxon of Amazophrynella sp. 3 (sensu Rojas et al. 2018a) from Parque de Desenvolvimento Jatob�� (Tapaj��s River, Par��, Brazil) with low support (PP = 0.74). Genetic distance between A. bilinguis sp. nov. and Amazophrynella sp. 3 was relatively low (p and K2P = 2%) compared to distances calculated between A. bilinguis sp. nov. and other Amazophrynella species that ranged from 3% (p) and 4% (K2P) (Amazophrynella sp. 2) to 14% (p) and 16% (K2P) (A. siona). Amazophrynella bilinguis sp. nov. presented 4% (p and K2P) genetic distance from samples of A. bokermanni, the nominal species that was placed closest to it in the phylogenetic tree. See Table 3 for genetic distances among all analyzed species. Distribution and natural history. Amazophrynella bilinguis sp. nov. is known only from Fazenda Taperinha (2��34���48.4��� S; 54��22���16.6��� W, 7 m a.s.l.) on south margin of the Amazon River, Municipality of Santar��m, Par�� State, Brazil (Fig. 9). However, it is likely that sampling efforts, which are historically scarce at that region, might expand the known distribution of this taxon. Like in other species within the genus, males of this species were found calling during the day, usually perched up to 10���40 cm above the ground on vegetation, fallen logs, branches or on leaf litter on the forest floor. Females and vocally active males were found near a stream and associated shallow ponds, where egg deposition and tadpole development may occur. After collection, a pair (female = INPA-H 39779, SVL = 20.8 mm; male = INPA-H 39774; SVL = 15.4 mm) engaged in axillary amplexus inside a plastic bag in the improvised field laboratory (Fig. 4A and 4B). Etymology. The epithet " bilinguis " is Latin and means "bilingual". It refers to the two distinct advertisement calls emitted by the male individuals., Published as part of Kaefer, Igor Luis, Rojas, Rommel R., Ferr��o, Miqu��ias, Farias, Izeni Pires & Lima, Albertina Pimentel, 2019, A new species of Amazophrynella (Anura: Bufonidae) with two distinct advertisement calls, pp. 316-334 in Zootaxa 4577 (2) on pages 319-329, DOI: 10.11646/zootaxa.4577.2.5, http://zenodo.org/record/2629715, {"references":["Rojas, R. R., Chaparro, J. C., Carvalho, V. T., Avila, R. W., Farias, I. P., Hrbek, T. & Gordo, M. (2016) Uncovering the diversity inside the Amazophrynella minuta complex: integrative taxonomy reveals a new species of Amazophrynella (Anura: Bufonidae) from southern Peru. ZooKeys, 71, 43 - 71.","Rojas, R. R., Carvalho, V. T., Avila, R. W., Farias, I. P., Gordo, M. & Hrbek, T. (2015) Two new species of Amazophrynella (Amphibia: Anura: Bufonidae) from Loreto, Peru. Zootaxa, 3946 (1), 79 - 103. https: // doi. org / 10.11646 / zootaxa. 3946.1.3","Rojas, R. R., Fouquet, A., Carvalho, V. T., Ron, S., Chaparro, J. C., Vogt, R. C., Avila, R. W., Farias, I. P., Gordo, M. & Hrbek, T. (2018 c) Redescription of the Amazonian tiny tree toad Amazophrynella minuta (Melin, 1941) (Anura: Bufonidae) from its type locality. Zootaxa, 4482 (3), 511 - 526. https: // doi. org / 10.11646 / zootaxa. 4482.3.4","Rojas, R. R., Fouquet, A., Ron, S. R., Hernandez-Ruz, E. J., Melo-Sampaio, P. R., Chaparro, J. C., Vogt, R. C., Carvalho, V. T., Pinheiro, L. C., Avila, R. W., Farias, I. P., Gordo, M. & Hrbek, T. (2018 a) A Pan-Amazonian species delimitation: high species diversity within the genus Amazophrynella (Anura: Bufonidae). PeerJ, 6, e 4941. https: // doi. org / 10.7717 / peerj. 4941","Avila, R. W., Carvalho, V. T. D., Gordo, M., Kawashita-Ribeiro, R. A. & Morais, D. H. (2012) A new species of Amazophrynella (Anura: Bufonidae) from southern Amazonia. Zootaxa, 3484, 65 - 74.","Rojas, R. R., Carvalho, V. T., Gordo, M. & Avila, R. W. (2014) A new species of Amazophrynella (Anura: Bufonidae) from the Brazilian Guiana Shield. Zootaxa, 3753, 79 - 95.","Rojas, R. R., Carvalho, V. T., Avila, R. W., Kawashita-Ribeiro, R. A., Hrbek, T. & Gordo, M. (2018 b) Description of the advertisement calls of four species of Amazophrynella (Anura: Bufonidae). Zootaxa, 4459 (1), 193 - 196. https: // doi. org / 10.11646 / zootaxa. 4459.1.11"]}

Amazophrynella minuta (Melin, 1941) Synonymy Atelopus minutus: Melin (1941: 40, Brazil; Amazonas Taracuá) Dendrophryniscus minutus: McDiarmid (1971: 39, in part) Lectotype (designated herewithin). NaTURHISTORISka MUSEET, GöTEbORg, SWEdEN, NHMG 462, SVL 13.40, adUlT malE, cOllEcTEd aT TaRacUá (0.10°S, 68.46°W) 100 m a.S.l., AmaZONaS STaTE, BRaZIl bY DOUglaS MElIN ON 1924. Paralectotypes (FIgURE 4). NHMG 463, NHMG 465 (adUlT malES), NHMG 464 (adUlT fEmalE) cOllEcTEd aT THE SamE PlacE aS THE lEcTOTYPE. Topotypical specimens. INPA – H 32721 –34, adUlT malES aNd INPA – H 32735 –40, adUlT fEmalES cOllEcTEd bY ROmmEl R. ROjaS aT THE SamE PlacE aS THE lEcTOTYPE ON AUgUST 19 aNd 20, 2013. Diagnosis. A SPEcIES Of Amazophrynella WITH: (1) MEdIUm bOdY SIZE fOR THE gENUS (SEE TablE 1), adUlT fEmalES 15.0–19.0 mm SVL (N=11), adUlT malES 12.2–14.2 mm SVL (N=20); (2) SNOUT POINTEd IN laTERal VIEW; UPPER jaW, IN laTERal VIEW, PROTRUdINg bEYONd lOWER; (3) TYmPaNUm, VOcal Sac, PaROTId glaNd aNd cRaNIal cRESTS INcONSPIcUOUS; (4) TEXTURE Of dORSal SkIN HIgHlY gRaNUlaR; (5) abUNdaNT Small ROUNdEd (IN PaRT cONIcal) WaRTS PRESENT ON dORSal SURfacES Of fORElImbS aNd HINdlImbS; (6) TEXTURE Of VENTRal SkIN HIgHlY gRaNUlaR; (7) fINgERS SlENdER, baSallY WEbbEd; (8) fINgER I SHORTER THaN fINgER II; (9) fINgER III RElaTIVElY lONg (HAL / SVL 0.2–0.3 mm, N=31); (10) PalmaR TUbERclE EllIPTIcal; (11) SUPERNUmERaRY TUbERclES ROUNdEd; (12) lONg HINd lImbS (TAL / SVL 0.4–0.5, N=31); (13) TOES SlENdER, baSallY WEbbEd; (14) TOES lackINg laTERal fRINgES; (15) PlaNTaR SURfacES Of fEET bEaRINg ONE mETaTaRSal TUbERclE, THE INNER 2.0X laRgER THaN THE OUTER, OUTER SUbcONIcal; SUPERNUmERaRY PlaNTaR TUbERclES ROUNdEd; (16) IN lIfE, VENTER cOlORaTION YEllOW–ORaNgE WITH laRgE TO mEdIUm SIZE black blOTcHES ON VENTER. Comparisons with other species. Amazophrynella minuta IS mOST SImIlaR TO A. amazonicola fROm WHIcH IT caN bE dISTINgUISHEd bY (cHaRacTERISTIcS Of cOmPaREd SPEcIES IN PaRENTHESES) abSENcE Of Small TRIaNgUlaR PROTRUSION aT THE TIP Of THE SNOUT; TEXTURE Of dORSal SkIN HIgHlY gRaNUlaR (gRaNUlaR); PalmaR TUbERclE EllIPTIcal (ROUNdEd). FROm A. matses bY SNOUT POINTEd IN laTERal VIEW; TEXTURE Of dORSal SkIN HIgHlY gRaNUlaR (SPIcUlaTE); VENTER cOlORaTION YEllOW–ORaNgE (PalE YEllOW; FIgURE 5A VS. 5C). FROm A. javierbustamantei bY TEXTURE Of dORSal SkIN HIgHlY gRaNUlaR (TUbERcUlaTE); PalmaR TUbERclE EllIPTIcal (ROUNdEd); THROaT aNd cHEST cOlORaTION lIgHT bROWN (gRaYISH cREam); laRgE blOTcHES ON VENTER (Small POINTS). FROm A. siona bY SNOUT PROfIlE POINTEd (acUTE); WaRTS ON dORSUm (gRaNUlES); TEXTURE Of dORSal SkIN HIgHlY gRaNUlaR (fINElY gRaNUlaR). FROm A. moisesii bY SNOUT PROfIlE POINTEd (acUmINaTE); TEXTURE Of dORSal SkIN HIgHlY gRaNUlaR (TUbERcUlaTE); laRgE blOTcHES ON VENTER (TINY POINTS). FROm A. bokermanni, A. manaos, A. vote, A. xinguensis aNd A. manaos THE maIN dIffERENcE IS IN VENTER cOlORaTION, YEllOWISH–ORaNgE (WHITE IN A. manaos, A. bokermanni aNd A. teko, REd bROWN IN A. vote, bROWN IN A. xinguensis; FIgURE 5A VS. 5F–J); FI FII IN A. bokermanni, aNd FI ≥ FII IN A. xinguensis). Redescription. BOdY SlENdER, SlIgHTlY ENlaRgEd POSTERIORlY. SEXUal dImORPHISm ObSERVEd IN SVL, WITH 12.2– 14.7 mm (13.6 ± 0.6 mm, N=20, TablE 4) IN adUlT malES aNd 14.5–19.4 mm (17.5 ± 1.5 mm, N=11) IN adUlT fEmalES. HEad POINTEd, lONgER THaN WIdE: HL 4.6–6.8 mm (5.4 ± 0.5 mm), 36.0% Of SVL; HW 3.8–5.8 mm (4.6 ± 0.1 mm), 30.0% Of SVL. SNOUT PROfIlE POINTEd IN laTERal VIEW; SL 2.1–3.2 mm (2.5 ± 0.3 mm), 46.0% Of HL. Canthus rostralis STRaIgHT aNd lOREal REgION VERTIcal. NOSTRIlS PROTUbERaNT, clOSER TO SNOUT THaN TO EYES. IND 1.0– 1.5 mm (1.2 ± 0.1 mm), 27.0% Of HW. INTERORbITal aNd OccIPITal REgIONS flaT. TYmPaNIc mEmbRaNE aNd TYmPaNIc aNNUlUS NOT aPPaRENT. PaROTOId glaNdS NOT VISIblE. VOcal Sac NOT VISIblE IN malES. EYES PROmINENT. EYE dIamETER 1.2–2.0 mm (1.5 ± 0.2 mm), 29.0% Of HL. EYElId WITH Small TUbERclES ON bORdERS. Choanae Small aNd cIRcUlaR. DORSal SkIN TEXTURE HIgHlY gRaNUlaR. Small ROUNdEd WaRTS (IN PaRT cONIcal) ON dORSOlaTERal aNd laTERal SURfacES POSTERIOR TO EYE, ON POSTERIOR dORSUm, aNd ON dORSal SURfacES Of ENTIRE aRm. TEXTURE Of gUlaR REgION, cHEST aNd bEllY HIgHlY gRaNUlaR. FORElImbS SlENdER. FOREaRmS RObUST, ESPEcIallY IN malES. UAL 3.5–5.5 mm (4.4 ± 0.7 mm); 29.3% Of SVL. FINgERS SlENdER. HAL 2.4–4.5 mm (3.1 ± 0.5 mm); 70.0% Of UAL. EdgES Of fORElImbS WITH ScaTTEREd gRaNUlES, IN dORSal aNd VENTRal VIEW. FINgERS baSallY WEbbEd. RElaTIVE lENgTH Of fINgERS: I Variation. VaRIaTION IN mEaSUREmENTS IS PRESENTEd IN TablE 1. SPEcImENS fROm JaPURa RIVER (INPA–H 32731, INPA–H 32724, INPA–H 32734) aRE lONgER (SL abOUT 1.0 mm) THaN INdIVIdUalS fROm THE TYPE lOcalITY. SOmE INdIVIdUalS fROm SÃO GabRIEl da CacHOEIRa PRESENT RElaTIVElY fEW WaRTS ON dORSal SURfacE (INPA–H32731, INPA–H 32724 aNd INPA–H 32734). MOTTlEd WaRTS dISSEmINaTEd ON cHEST aNd bEllY aRE fOUNd IN SPEcImENS INPA–H32731 (SÃO GabRIEl da CacHOEIRa) aNd INPA–35496 (JaPURa RIVER). SOmE SPEcImENS fROm THE TYPE lOcalITY (INPA–H 32728, INPA–H 32735) aNd fROm THE JaPURa RIVER (INPA–H 35512), PRESENT lOWER abUNdaNcE Of WaRTS ON aRmS. Coloration in life (FIgURE 6). HEad bROWN. DORSUm REddISH–bROWN. FlaNkS lIgHT bROWN. DORSal SURfacES Of THE UPPER aRm aNd aRm REddISH–bROWN. HaNdS aNd fINgERS lIgHT bROWN, IN dORSal VIEW. INSERTION Of THE aRmS PRESENT YEllOW blOTcHES. DORSal SURfacES Of THE THIgHS, TIbIa, TaRSUS aNd fEET REddISH–bROWN. VENTRal SURfacES Of UPPER aRm aNd aRm cREam. Palm lIgHT REd. VENTRal SURfacES Of THIgHS mOTTlEd cREam WITH Small black blOTcHES. THIgHS WITH a TRaNSVERSE laTERal black baR. POSTERIOR REgION Of THE THIgH aNd clOaca YEllOW, cOVEREd bY Small black blOTcHES. TaRSUS WITH TRaNSVERSE bROWN baRS. SOlE REddISH–bROWN. GUlaR REgION aNd cHEST gRaYISH–bROWN. VENTER cOlORaTION bETWEEN YEllOW aNd YEllOW–ORaNgE. VENTER cOVEREd bY laRgE TO mEdIUm SIZE black blOTcHES. IN VENTRal VIEW, SOmE SPEcImENS PRESENT YEllOW blOTcHES aT THE INSERTION Of THE aRmS. THE SPEcImENS INPA–H 32725 (TaRacUá) aNd INPA–H 35494 (JaPURa RIVER), dISPlaY YEllOW cOlORaTION WITH Small black blOTcHES ON POSTERIOR REgION Of THE THIgH NEaR THE clOaca. IRIS gOldEN aNd PUPIl black. Color in preservative (FIgURE 7). THE cOlOR IN PRESERVaTIVE fadEd. WE NOTEd THE PROgRESSIVE lOSS Of THE VENTRal cOlORaTION THaT bEcamE PalE YEllOW. THE YEllOW cOlORaTION ON aXIllaRY SURfacE dISaPPEaREd. IN VENTRal VIEW, THE cOlOR Of fINgERS aNd TOES lOST THEIR REd cOlOR aNd bEcamE cREam. Distribution (FIgURE 8). Amazophrynella minuta IS dISTRIbUTEd THROUgHOUT THE EaSTERN bORdER REgION Of THE AmaZONaS STaTE IN BRaZIl, a REgION alSO kNOWN aS “CabEÇa dO CacHORRO” (“DOg’S HEad”, ENglISH fREE TRaNSlaTION). THE SPEcIES IS fOUNd IN THE fOllOWINg lOcalITIES: TaRacUá (0.10°N, 68.46°W), SÃO GabRIEl da CacHOEIRa (0.16°S, 66.98°W), CUcUÍ (1.07°N, 66.88°W) aNd JaPURa RIVER SOUTH-EaST Of VIla BITTENcOURT (1.81°S, 68.98°W), aT ElEVaTIONS bETWEEN 90–105 m a.S.l. IN COlOmbIa IT IS REPORTEd IN THE DEPaRTmENT Of CaqUETá (0.92°N, 75.67°W) aNd VaUPéS (0.90°S, 69.67°W) (LYNcH, 2006) aT ElEVaTIONS bETWEEN 100–200 m a.S.l. (AcOSTa– GalVIS, 2017). IN VENEZUEla, THE SPEcIES WaS REPORTEd fROm RaUdal dE DaNTO, RÍO CUaO (4.53°N, 67.18°W) AmaZONaS STaTE, aT ElEVaTION Of 105 m a.S.l. (ROjaS–RUNjaIc et al., 2013). Ecology. WE fOUNd all INdIVIdUalS IN THE mORNINg, NEaR a bOdY Of WaTER. SPEcImENS WERE cOllEcTEd IN PRImaRY “ terra firme ” fOREST (“fIRm laNd” – NON–flOOdEd fOREST, ENglISH fREE TRaNSlaTION) SOmE IN THE lEaf lITTER aNd OTHERS amONg TREE ROOTS. SPEcImENS fROm SÃO GabRIEl da CacHOEIRa WERE cOllEcTEd IN a fRagmENT Of fOREST NEaR THE aIRPORT; THE INdIVIdUalS WERE fOUNd aROUNd a STREam WITH abUNdaNT lEaf lITTER. AmPlEXUS IS aXIllaR (FIgURE 9). Phylogenetic analysis. THE cOmPlETE PHYlOgENETIc RElaTIONSHIPS Of Amazophrynella, THE PHYlOgENETIc POSITION Of A. minuta aNd gENETIc dISTaNcE bETWEEN SPEcIES Of Amazophrynella aRE REPORTEd IN ROjaS et al. (2018, FIgURE 1 aNd TablE 1). Advertisement call (FIgURE 10). THE cHaRacTERIZaTION Of THE adVERTISEmENT call Of Amazophrynella minuta WaS baSEd ON 14 callS fROm ONE INdIVIdUal fROm SÃO GabRIEl da CacHOEIRa (INPA–H 32730), SVL: 12.2 mm, REcORdEd bETWEEN 15:30–16:00 H. AIR TEmPERaTURE aT THE TImE Of REcORdINg VaRIEd bETWEEN 26.0 TO 27.0°C. THE call cONSISTEd Of a TRIllEd NOTE EmITTEd bETWEEN REgUlaR SIlENT INTERValS. NOTES Had aN UPWaRd amPlITUdE mOdUlaTION, REacHINg THEIR maXImUm INTENSITY NEaR THE mIddlE Of THE NOTE. NOTE dURaTION RaNgEd bETWEEN 0.310 aNd 0.108 S (0.223 ± 0.068 S). NUmbER Of PUlSES PER NOTE RaNgEd fROm 18 TO 35 PUlSES/NOTE (26.286 ± 6.557 PUlSES/NOTE). THE INTER–NOTE INTERVal WaS 0.663 TO 1.050 S (0.827 ± 0.120 S). THE dOmINaNT fREqUENcY RaNgEd fROm 4242.790 TO 4562.011 HZ (4419.213 ± 125.642 HZ). THE fUNdamENTal fREqUENcY RaNgEd fROm 3201.010 TO 3788.520 HZ (3489.211 ± 235.791 HZ). THE RISE TImE RaNgEd fROm 0.045 TO 0.083 S (0.059 ± 0.015 S). NUmbER Of HaRmONIcS VaRIEd fROm 3 TO 5 (4.150 ± 0.587). Osteology. A dETaIlEd REVIEW aNd dEScRIPTION Of THE SkElETON Of A. minuta WERE PROVIdEd bY McDIaRmId (1971). IN THE SamE PUblIcaTION, McDIaRmINd (1971) REPORTEd cHaRacTERS THaT cOUld bE TENTaTIVElY cONSIdEREd aS PUTaTIVE gENUS lEVEl SYNaPOmORPHIES (SEE FOUqUET et al., 2012b)., Published as part of Rojas, Rommel R., Fouquet, Antoine, Carvalho, Vinícius Tadeu De, Ron, Santiago, Chaparro, Juan Carlos, Vogt, Richard C., Ávila, Robson W., Farias, Izeni Pires, Gordo, Marcelo & Hrbek, Tomas, 2018, Redescription of the Amazonian tiny tree toad Amazophrynella minuta (Melin, 1941) (Anura: Bufonidae) from its type locality, pp. 511-526 in Zootaxa 4482 (3) on pages 515-522, DOI: 10.11646/zootaxa.4482.3.4, http://zenodo.org/record/1440763, {"references":["Melin, D. E. (1941) Contributions to the knowledge of the Amphibia of South America. Goteborgs Kungl. Uetenskaps - och Uitterhets - samhalles. Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1 - 71","McDiarmid, R. w. (1971) Comparative morphology and evolution of frogs of the Neotropical genera Atelopus, Dendrophryniscus, Melanophryniscus and Oreophrynella. Bulletin of the Los Angeles County Museum of Natural History, 12, 1 - 66.","Lynch, J. D. (2006) The amphibian fauna in the Villavicencio region of eastern Colombia. Caldasia, 28, 135 - 155.","Acosta-Galvis, A. R. (2017) Lista de los Anfibios de Colombia: Referencia en linea V. 07.2017.0 (21 June 2017). Batrachia, Villa de Leyva, Boyaca. Available from: http: // www. batrachia. com (accessed 13 August 2018)"]}

Amazophrynella matses sp. nov. (Figure 9) Synonyms Dendrophryniscus minutus (Gordo et al. 2006). Holotype. MZUNAP 921 (adult male), collected at Nuevo Salvador locality (5 ° 2 ' 33 '' S, 73 ° 50 ' 02'' W), district of Requena, Province of Requena, Loreto, Peru, by Ian Pool Medina, Richard Curto and Rommel R. Rojas on February 5, 2013, at 16 h. Paratopotypes. MZUNAP 934, MZUNAP 955 MZUNAP 940, MZUNAP 948 MZUNAP 943, MZUNAP 952, MZUNAP 953 (adult males), collected by Richard Curto on January 6, 2013. MZUNAP 932, MZUNAP 945, MZUNAP 950, (adult females), by Ian Pool Medina on January 7, 2013. MZUNAP 958, MZUNAP 922, MZUNAP 923, MZUNAP 925, MZUNAP 926 (adult males), MZUNAP 927, MZUNAP 944, MZUNAP 938, MZUNAP 936 (adult females) by Ian Pool Medina, Richard Curto and Rommel R. Rojas on February 5, 2013 (Figure 10). Paratypes. MZUNAP 928, MZUNAP 929, MZUNAP 930, MZUNAP 931, MZUNAP 933, MZUNAP 955, MZUNAP 935, MZUNAP 950, MZUNAP 937, MZUNAP 939, MZUNAP 941, MZUNAP 942, MZUNAP 946, MZUNAP 947, MZUNAP 949, (adult females) collected near Jenaro Herrera community, district of Requena (4 ° 53 ' 54 '' S, 73 ° 37 ' 05'' W) on January 6, 2013, by Rommel R. Rojas. Referred specimens. USNM 332894-332907 collected by Lily O. Rodriguez on November 27, 1989 in Jenaro Herrera (4 ° 55 ' S, 74 ° 45 ' W), on eastern bank of Río Ucayali, district of Requena, Loreto, Peru. Diagnosis. Amazophrynella matses sp. nov. is assigned to the genus Amazophrynella by the combination of the following synapomorphies (sensu Fouquet et al. 2012 b): small size with maximum SVL of 26.5 mm, elongated snout and slender body, eyelids smaller than eye diameter, cranial crests not evident, parotid glands and external tympanum not visible, texture of the skin granular or finely granular, ventrum with pattern of blotches or spots. The new species is readily diagnosed by the combination of the following characters: 1) smallest size of the genus: females 17.1 ± 0.7 mm (16.5–17.7 mm), males 12 ± 0.6 mm (11.5–13.5 mm); 2) canthus rostralis straight in dorsal view; 3) snout slightly truncate, with edges of nasal protrusion dilated in ventral view; 4) palmar tubercles elliptical; 5) finger tips expanded, 6) presence of medium-sized black ocelli and/or streaks covering venter; 7) belly yellow with undefined blotches; and 8) ten diagnostic characters in the 16 S rDNA gene fragment (Figure 1, Table 5). Comparisons with other species. Amazophrynella matses sp. nov. can be distinguished from the other species of Amazophrynella by the following characters (characters of compared species in parentheses). Compared to A. minuta sensu stricto, SVL is smaller in males (larger SVL; Table 2), and snout slightly truncate, with edges of nasal protrusion dilated in ventral view (snout pointed, with no dilatation on the nasal protrusion) (Figure 5). Compared to A. bokermanni, relative size of fingers differs, with finger II larger than I (I>II), and belly yellow-orange with black ocelli and/or streaks (white belly with small black dots). Compared to A. vote, snout is truncate (rounded) and belly yellow-orange with black ocelli and/or streaks (ventral color pattern reddish brown, with presence of small white dots and brown spots and absence of large blotches). Relative to A. manaos, finger tips are expanded (unexpanded), texture of body skin spiculate (granulated body surfaces), belly yellow-orange with black ocelli and/ or streaks (white abdomen with black spots or stripes). Compared to A. amazonicola sp. nov., fingers are unwebbed (fingers I and II webbed at base), snout truncate with edges of nasal protrusion dilated in ventral view (snout pointed with no dilated nasal protrusion), palmar tubercles elliptical (rounded) and greenish-white granules on body absent (present). Description of the holotype. Body robust; head triangular, as long as wide; head length 33.4 % of SVL; head width 28.8 % of SVL. Snout truncate in dorsal view, with a triangular nasal protrusion with edges dilated in ventral view; snout pointed in lateral view; snout length 48.8 % of head length; tympanum and vocal sac indistinct; circular eyes, diameter about 24.4 % of head length; nostril closer to the tip of snout than to eyes. Dorsal skin texture spiculate and covered by several scattered small granules; few spine granules on the upper arm; ventral surfaces covered by granular tubercles larger than that of dorsum; forelimbs slender, length of upper arm 29.6 % of SVL; edges of lower arms and upper arms covered by few small spines; fingers slender, relative length of fingers I Measurement of the holotype (mm). SVL: 13.5; HW: 3.9; HL: 4.5; SL: 2.2; ED: 1.1; IND: 1.1; UAL: 4.0; HAL: 2.8; THL: 5.9; TAL: 5.7; TL: 4.2; FL: 5.3. Coloration. In males and females the dorsal coloration light brown with dark brown transverse bars. A continuous or barely white line visible on upper jaw, extending from the tip of the snout to the corner of the mouth. Golden iris and black pupil (Figure 11). Throat dark gray, chest grayish brown; belly yellow-orange with black ocelli and/or streaks covering the abdomen, reaching the chest and throat. Ventral skin of thighs grayish brown with scattered black dots. Reddish brown granules covering dorsal and ventral surfaces. Palms and soles reddish brown. In alcohol, the dorsal coloration becomes darker. Ventral coloration fades, with yellow colors becoming white. Variation. The species shows little variation in dorsal pattern, with few specimens presenting a thin brown vertebral line from the tip of snout to the vent. However, ventral coloration is highly variable among individuals (Figure 12). The black ocelli on the throat, chest and belly vary in abundance. Some specimens exhibited black streaks on the belly, which are the result of union of ocelli. The yellow surface may extend from thighs to the chest. In juveniles the edges of nasal protrusion in less perceptible. The elliptical shape of the palmar tubercle became flattened in preserved specimens. In juveniles specimens the protrusion on the edges of the nose is more visible. Distribution and natural history. Amazophrynella matses sp. nov. was found at four localities on the right bank of the Ucayali River in the state of Loreto, southeastern Peru: Requena, around Jenaro Herrera community, Nuevo Salvador and the Matses Indian Reserve. Amazophrynella matses sp. nov. can be found in upland (= terra firme) forests away from rivers, within root mass of bushes or in leaf litter. Reproduction is coincident with the rainy season, where oviposition occurs in puddles with abundant herbaceous and shrubby vegetation. Etymology. The specific epithet matses, a substantive from a polysynthetic language, is a reference to the Matsés ethnic group, which inhabits the region where we found the new species.

Rojas, Rommel R., Carvalho, Vinícius Tadeu De, Ávila, Robson W., Farias, Izeni Pires, Gordo, Marcelo, Hrbek, Tomas (2015): Two new species of Amazophrynella (Amphibia: Anura: Bufonidae) from Loreto, Peru. Zootaxa 3946 (1): 79-103, DOI: http://dx.doi.org/10.11646/zootaxa.3946.1.3

Amazophrynella matses sp. nov. (Figure 9) Synonyms Dendrophryniscus minutus (Gordo et al. 2006). Holotype. MZUNAP 921 (adult male), collected at Nuevo Salvador locality (5 �� 2 ' 33 '' S, 73 �� 50 ' 02'' W), district of Requena, Province of Requena, Loreto, Peru, by Ian Pool Medina, Richard Curto and Rommel R. Rojas on February 5, 2013, at 16 h. Paratopotypes. MZUNAP 934, MZUNAP 955 MZUNAP 940, MZUNAP 948 MZUNAP 943, MZUNAP 952, MZUNAP 953 (adult males), collected by Richard Curto on January 6, 2013. MZUNAP 932, MZUNAP 945, MZUNAP 950, (adult females), by Ian Pool Medina on January 7, 2013. MZUNAP 958, MZUNAP 922, MZUNAP 923, MZUNAP 925, MZUNAP 926 (adult males), MZUNAP 927, MZUNAP 944, MZUNAP 938, MZUNAP 936 (adult females) by Ian Pool Medina, Richard Curto and Rommel R. Rojas on February 5, 2013 (Figure 10). Paratypes. MZUNAP 928, MZUNAP 929, MZUNAP 930, MZUNAP 931, MZUNAP 933, MZUNAP 955, MZUNAP 935, MZUNAP 950, MZUNAP 937, MZUNAP 939, MZUNAP 941, MZUNAP 942, MZUNAP 946, MZUNAP 947, MZUNAP 949, (adult females) collected near Jenaro Herrera community, district of Requena (4 �� 53 ' 54 '' S, 73 �� 37 ' 05'' W) on January 6, 2013, by Rommel R. Rojas. Referred specimens. USNM 332894-332907 collected by Lily O. Rodriguez on November 27, 1989 in Jenaro Herrera (4 �� 55 ' S, 74 �� 45 ' W), on eastern bank of R��o Ucayali, district of Requena, Loreto, Peru. Diagnosis. Amazophrynella matses sp. nov. is assigned to the genus Amazophrynella by the combination of the following synapomorphies (sensu Fouquet et al. 2012 b): small size with maximum SVL of 26.5 mm, elongated snout and slender body, eyelids smaller than eye diameter, cranial crests not evident, parotid glands and external tympanum not visible, texture of the skin granular or finely granular, ventrum with pattern of blotches or spots. The new species is readily diagnosed by the combination of the following characters: 1) smallest size of the genus: females 17.1 �� 0.7 mm (16.5���17.7 mm), males 12 �� 0.6 mm (11.5���13.5 mm); 2) canthus rostralis straight in dorsal view; 3) snout slightly truncate, with edges of nasal protrusion dilated in ventral view; 4) palmar tubercles elliptical; 5) finger tips expanded, 6) presence of medium-sized black ocelli and/or streaks covering venter; 7) belly yellow with undefined blotches; and 8) ten diagnostic characters in the 16 S rDNA gene fragment (Figure 1, Table 5). Comparisons with other species. Amazophrynella matses sp. nov. can be distinguished from the other species of Amazophrynella by the following characters (characters of compared species in parentheses). Compared to A. minuta sensu stricto, SVL is smaller in males (larger SVL; Table 2), and snout slightly truncate, with edges of nasal protrusion dilated in ventral view (snout pointed, with no dilatation on the nasal protrusion) (Figure 5). Compared to A. bokermanni, relative size of fingers differs, with finger II larger than I (I>II), and belly yellow-orange with black ocelli and/or streaks (white belly with small black dots). Compared to A. vote, snout is truncate (rounded) and belly yellow-orange with black ocelli and/or streaks (ventral color pattern reddish brown, with presence of small white dots and brown spots and absence of large blotches). Relative to A. manaos, finger tips are expanded (unexpanded), texture of body skin spiculate (granulated body surfaces), belly yellow-orange with black ocelli and/ or streaks (white abdomen with black spots or stripes). Compared to A. amazonicola sp. nov., fingers are unwebbed (fingers I and II webbed at base), snout truncate with edges of nasal protrusion dilated in ventral view (snout pointed with no dilated nasal protrusion), palmar tubercles elliptical (rounded) and greenish-white granules on body absent (present). Description of the holotype. Body robust; head triangular, as long as wide; head length 33.4 % of SVL; head width 28.8 % of SVL. Snout truncate in dorsal view, with a triangular nasal protrusion with edges dilated in ventral view; snout pointed in lateral view; snout length 48.8 % of head length; tympanum and vocal sac indistinct; circular eyes, diameter about 24.4 % of head length; nostril closer to the tip of snout than to eyes. Dorsal skin texture spiculate and covered by several scattered small granules; few spine granules on the upper arm; ventral surfaces covered by granular tubercles larger than that of dorsum; forelimbs slender, length of upper arm 29.6 % of SVL; edges of lower arms and upper arms covered by few small spines; fingers slender, relative length of fingers IMeasurement of the holotype (mm). SVL: 13.5; HW: 3.9; HL: 4.5; SL: 2.2; ED: 1.1; IND: 1.1; UAL: 4.0; HAL: 2.8; THL: 5.9; TAL: 5.7; TL: 4.2; FL: 5.3. Coloration. In males and females the dorsal coloration light brown with dark brown transverse bars. A continuous or barely white line visible on upper jaw, extending from the tip of the snout to the corner of the mouth. Golden iris and black pupil (Figure 11). Throat dark gray, chest grayish brown; belly yellow-orange with black ocelli and/or streaks covering the abdomen, reaching the chest and throat. Ventral skin of thighs grayish brown with scattered black dots. Reddish brown granules covering dorsal and ventral surfaces. Palms and soles reddish brown. In alcohol, the dorsal coloration becomes darker. Ventral coloration fades, with yellow colors becoming white. Variation. The species shows little variation in dorsal pattern, with few specimens presenting a thin brown vertebral line from the tip of snout to the vent. However, ventral coloration is highly variable among individuals (Figure 12). The black ocelli on the throat, chest and belly vary in abundance. Some specimens exhibited black streaks on the belly, which are the result of union of ocelli. The yellow surface may extend from thighs to the chest. In juveniles the edges of nasal protrusion in less perceptible. The elliptical shape of the palmar tubercle became flattened in preserved specimens. In juveniles specimens the protrusion on the edges of the nose is more visible. Distribution and natural history. Amazophrynella matses sp. nov. was found at four localities on the right bank of the Ucayali River in the state of Loreto, southeastern Peru: Requena, around Jenaro Herrera community, Nuevo Salvador and the Matses Indian Reserve. Amazophrynella matses sp. nov. can be found in upland (= terra firme) forests away from rivers, within root mass of bushes or in leaf litter. Reproduction is coincident with the rainy season, where oviposition occurs in puddles with abundant herbaceous and shrubby vegetation. Etymology. The specific epithet matses, a substantive from a polysynthetic language, is a reference to the Mats��s ethnic group, which inhabits the region where we found the new species., Published as part of Rojas, Rommel R., Carvalho, Vin��cius Tadeu De, ��vila, Robson W., Farias, Izeni Pires, Gordo, Marcelo & Hrbek, Tomas, 2015, Two new species of Amazophrynella (Amphibia: Anura: Bufonidae) from Loreto, Peru, pp. 79-103 in Zootaxa 3946 (1) on pages 93-100, DOI: 10.11646/zootaxa.3946.1.3, http://zenodo.org/record/236987, {"references":["Gordo, M. (2006) In: Vriesendorp, C., Pitman, N., Rojas, M., Pawlak, B. A., Rivera, C., Calixto M., Vela C. P. & Fasabi, R. (Eds.), 2006. Peru: Matses. Rapid Biological Inventories Report 16. The Field Museum, Chicago, Illinois, 336 pp.","Fouquet, A., Recoder, R., Teixeira Jr., M., Cassimiro, J., Amaro, R. C., Camacho, A., Damasceno, R., Carnaval, A. C., Moritz, C. & Rodrigues, M. T. (2012 b) Amazonella Fouquet et al. 2012 (Anura: Bufonidae) junior homonym of Amazonella Lundblad, 1931 (Acari: Unionicolidae): proposed replacement by Amazophrynella nom. nov. Zootaxa, 3244, 68."]}

Amazophrynella amazonicola sp. nov. (Figure 2) Synonyms Dendrophryniscus minutus (Rodriguez & Duellman, 1994). Holotype. MZUNAP 901 adult male, collected in Puerto Almendra (3 ° 49 ' 41 '' S, 73 ° 22 ' 07'' W), 10 km from the municipality of Iquitos, Province of Maynas, District of San Juan, Nina-Rumi community, Peru, collected by Ian Pool Medina and Rommel R. Rojas on January 15, 2013, at 10 h. Paratopotypes. MZUNAP 906; MZUNAP 915; MZUNAP 110; MZUNAP 907, MZUNAP 917; MZUNAP 889; MZUNAP 910; MZUNAP 911; MZUNAP 916; MZUNAP 913; MZUNAP 914, adult females collected in Puerto Almendra on January 15, 2013, by Ian Pool Medina and Richard Curto (Figure 3). Paratypes. MZUNAP 908, MZUNAP 924, MZUNAP 886, MZUNAP 900, MZUNAP 888, MZUNAP 919, MZUNAP 902, MZUNAP 887 (adult males), MZUNAP 905, MZUNAP 920 (adult females) collected at Fundo Zamora, (03° 57 ' 58 '' S, 73 ° 25 ' 07'' W), San Juan Bautista district, 58 km of Iquitos—Nauta highway, by Richard Curto and Ian Pool Medina on January 17, 2013. MZUNAP 918, MZUNAP 909 (adult females) collected on Nauta locality (04° 34 ' 09'' S, 73 ° 43 ' 25 '' W), Maynas province, 100 km from the municipality of Iquitos, by Ian Pool Medina on June 22, 2012; MZUNAP 242 collected on Fundo UNAP (04° 00' 11 '' S; 73 ° 26 ' 04'' W), Maynas province, 31 km from the municipality of Iquitos, by the Iquitos-Nauta highway. Diagnosis. Amazophrynella amazonicola sp. nov. is assigned to the genus Amazophrynella by the combination of the following synapomorphies (sensu Fouquet et al. 2012 b): small size with maximum SVL of 26.5 mm, elongated snout and slender body, eyelids smaller than eye diameter, cranial crests not evident, parotid glands and external tympanum not visible, texture of the skin granular or finely granular, ventrum with pattern of blotches or spots. The new species is diagnosable by: 1) snout pointed (in dorsal view); 2) tip of the snout with a small triangular protrusion (in both dorsal and ventral views); 3) snout profile pointed (in lateral view); 4) body surface sparsely granular, covered with medium-sized granules scattered irregularly; (5) base of fingers I and II webbed; (6) ventral coloration yellow-orange with dark brown blotches; and 7) brown dorsum with greenish white granules; and 8) four diagnostic characters in the 16 S rDNA gene fragment (Figure 1, Table 4). Comparisons with other species. Amazophrynella amazonicola sp. nov. can be distinguished from the four nominal species of Amazophrynella by the following characters (characters of compared species in parentheses). In contrast to A. minuta sensu stricto, dorsal surfaces are finely granular (dorsal surfaces roughly granular), fingers I and II webbed at the base (unwebbed fingers) (Figure 4 A), palmar tubercle rounded (palmar tubercle elliptical), fingers expanded (unexpanded), and spiny granules on axillary region absent (prickly warty skin on axillary region). Compared to A. bokermanni, the main difference is the relative size of fingers, with finger II being smaller than I (I>II); body size is smaller in males and females (22 mm SVL in males and 28 mm SVL in females) (Table 2), with belly yellow-orange with dark brown blotches (belly white with small black dots). In contrast to A. manaos, snout is pointed (triangular snout with a slightly truncate culmination) (Figure 5 C), fingers I and II basally at the base (unwebbed fingers), dorsal skin finely granular (dorsal surfaces granular), and belly yellow-orange with dark brown blotches (white belly with black spots or stripes). Compared to A. vote, snout is pointed (rounded), subrostral crest converging anteriorly (not converging), and belly yellow-orange with dark brown blotches (ventral color pattern reddish brown, with presence of small white dots and brown spots and absence of large blotches). Description of the holotype. Body slender; head triangular, as long as wide; head length 37.3 % of SVL; head width 31.7 % of SVL. Snout pointed in lateral view; head rounded in dorsal view; tip of snout with a small triangular protrusion; snout length 44.1 % of head length; tympanum not visible; vocal sac not externally visible; eyes prominent, 21.4 % of head length; nostril is closer to the tip of snout than to eyes; internarinal distance equal to eye diameter; presence of a line of small spiny granules from the outer edge of the mouth to upper arm. Dorsal skin spiculate, covered with several small rounded tubercles; scattered spiny granules on upper arm; texture of ventral skin granular, covered by small rounded granules. Forelimbs slender, upper arm length 34.4 % of SVL; edges of lower arm and upper arm with small spiny tubercles; hand length 66 % of upper arm length; fingers slender, with tips expanded; relative length of fingers I Measurement of the holotype (mm). SVL: 14.5; HW: 4.6; HL: 5.5; SL: 2.4; ED: 1.2; IND: 1.2; UAL: 5.0; HAL: 3.3; THL: 8.1; TAL: 7.2; TL: 4.5; FL: 5.3. Coloration. In life, dorsal coloration light brown with greenish-white granules, ventral coloration yelloworange with dark brown blotches. In alcohol, dorsal surfaces become dark brown, and the ventral coloration fades, with yellow color becoming pale brown. Variation. Coloration of belly is highly variable (Figure 6), with some specimens displaying an intense yellow-orange belly instead of yellow. During the breeding season, coloration of throat and chest of males and females becomes red. Tubercles on thighs of some specimens are yellow instead of brown, and a yellow spots cover the belly in some specimens. In some males the inner surface of the thighs can vary from clear yellow to grayish reddish color. In some individuals, the palmar tubercle shape may vary depending on the size (SVL), being much more perceptible to the eye in larger-sized individuals. In preserved individuals, the form of palmar tubercle looses its shape, becoming more flattened. The nasal bump on the tip of the nose is more visible in adults males and females, while in juveniles this character is less visible. Distribution and natural history. Amazophrynella amazonicola sp. nov. is known to occur from five localities in the Peruvian department of Loreto: Fundo Zamora, Fundo UNAP, Allpahuayo Mishana National Reserve, the vicinity of Nauta and Puerto Almendra. This species was recorded in white sand forests, locally known as “ varillales ” (in Portuguese “ campinaranas ”), from around Iquitos. It occurs in northwest of Iquitos (left bank of the Amazon, Nanay and Marañon Rivers), where its distribution spans the length of the Iquitos-Nauta highway; further north, its distribution is poorly known but is possibly delimited by the Napo and Marañon Rivers (Figure 7). Amazophrynella amazonicola sp. nov. breeds between December and March (Figure 8). The species is diurnal, and can be found amidst leaf litter, where its coloration makes it difficult to locate. Etymology. The specific epithet refers to the Amazon River, the world’s largest river and one of the new seven wonders of nature.

Amazophrynella amazonicola sp. nov. (Figure 2) Synonyms Dendrophryniscus minutus (Rodriguez & Duellman, 1994). Holotype. MZUNAP 901 adult male, collected in Puerto Almendra (3 �� 49 ' 41 '' S, 73 �� 22 ' 07'' W), 10 km from the municipality of Iquitos, Province of Maynas, District of San Juan, Nina-Rumi community, Peru, collected by Ian Pool Medina and Rommel R. Rojas on January 15, 2013, at 10 h. Paratopotypes. MZUNAP 906; MZUNAP 915; MZUNAP 110; MZUNAP 907, MZUNAP 917; MZUNAP 889; MZUNAP 910; MZUNAP 911; MZUNAP 916; MZUNAP 913; MZUNAP 914, adult females collected in Puerto Almendra on January 15, 2013, by Ian Pool Medina and Richard Curto (Figure 3). Paratypes. MZUNAP 908, MZUNAP 924, MZUNAP 886, MZUNAP 900, MZUNAP 888, MZUNAP 919, MZUNAP 902, MZUNAP 887 (adult males), MZUNAP 905, MZUNAP 920 (adult females) collected at Fundo Zamora, (03�� 57 ' 58 '' S, 73 �� 25 ' 07'' W), San Juan Bautista district, 58 km of Iquitos���Nauta highway, by Richard Curto and Ian Pool Medina on January 17, 2013. MZUNAP 918, MZUNAP 909 (adult females) collected on Nauta locality (04�� 34 ' 09'' S, 73 �� 43 ' 25 '' W), Maynas province, 100 km from the municipality of Iquitos, by Ian Pool Medina on June 22, 2012; MZUNAP 242 collected on Fundo UNAP (04�� 00' 11 '' S; 73 �� 26 ' 04'' W), Maynas province, 31 km from the municipality of Iquitos, by the Iquitos-Nauta highway. Diagnosis. Amazophrynella amazonicola sp. nov. is assigned to the genus Amazophrynella by the combination of the following synapomorphies (sensu Fouquet et al. 2012 b): small size with maximum SVL of 26.5 mm, elongated snout and slender body, eyelids smaller than eye diameter, cranial crests not evident, parotid glands and external tympanum not visible, texture of the skin granular or finely granular, ventrum with pattern of blotches or spots. The new species is diagnosable by: 1) snout pointed (in dorsal view); 2) tip of the snout with a small triangular protrusion (in both dorsal and ventral views); 3) snout profile pointed (in lateral view); 4) body surface sparsely granular, covered with medium-sized granules scattered irregularly; (5) base of fingers I and II webbed; (6) ventral coloration yellow-orange with dark brown blotches; and 7) brown dorsum with greenish white granules; and 8) four diagnostic characters in the 16 S rDNA gene fragment (Figure 1, Table 4). Comparisons with other species. Amazophrynella amazonicola sp. nov. can be distinguished from the four nominal species of Amazophrynella by the following characters (characters of compared species in parentheses). In contrast to A. minuta sensu stricto, dorsal surfaces are finely granular (dorsal surfaces roughly granular), fingers I and II webbed at the base (unwebbed fingers) (Figure 4 A), palmar tubercle rounded (palmar tubercle elliptical), fingers expanded (unexpanded), and spiny granules on axillary region absent (prickly warty skin on axillary region). Compared to A. bokermanni, the main difference is the relative size of fingers, with finger II being smaller than I (I>II); body size is smaller in males and females (22 mm SVL in males and 28 mm SVL in females) (Table 2), with belly yellow-orange with dark brown blotches (belly white with small black dots). In contrast to A. manaos, snout is pointed (triangular snout with a slightly truncate culmination) (Figure 5 C), fingers I and II basally at the base (unwebbed fingers), dorsal skin finely granular (dorsal surfaces granular), and belly yellow-orange with dark brown blotches (white belly with black spots or stripes). Compared to A. vote, snout is pointed (rounded), subrostral crest converging anteriorly (not converging), and belly yellow-orange with dark brown blotches (ventral color pattern reddish brown, with presence of small white dots and brown spots and absence of large blotches). Description of the holotype. Body slender; head triangular, as long as wide; head length 37.3 % of SVL; head width 31.7 % of SVL. Snout pointed in lateral view; head rounded in dorsal view; tip of snout with a small triangular protrusion; snout length 44.1 % of head length; tympanum not visible; vocal sac not externally visible; eyes prominent, 21.4 % of head length; nostril is closer to the tip of snout than to eyes; internarinal distance equal to eye diameter; presence of a line of small spiny granules from the outer edge of the mouth to upper arm. Dorsal skin spiculate, covered with several small rounded tubercles; scattered spiny granules on upper arm; texture of ventral skin granular, covered by small rounded granules. Forelimbs slender, upper arm length 34.4 % of SVL; edges of lower arm and upper arm with small spiny tubercles; hand length 66 % of upper arm length; fingers slender, with tips expanded; relative length of fingers IMeasurement of the holotype (mm). SVL: 14.5; HW: 4.6; HL: 5.5; SL: 2.4; ED: 1.2; IND: 1.2; UAL: 5.0; HAL: 3.3; THL: 8.1; TAL: 7.2; TL: 4.5; FL: 5.3. Coloration. In life, dorsal coloration light brown with greenish-white granules, ventral coloration yelloworange with dark brown blotches. In alcohol, dorsal surfaces become dark brown, and the ventral coloration fades, with yellow color becoming pale brown. Variation. Coloration of belly is highly variable (Figure 6), with some specimens displaying an intense yellow-orange belly instead of yellow. During the breeding season, coloration of throat and chest of males and females becomes red. Tubercles on thighs of some specimens are yellow instead of brown, and a yellow spots cover the belly in some specimens. In some males the inner surface of the thighs can vary from clear yellow to grayish reddish color. In some individuals, the palmar tubercle shape may vary depending on the size (SVL), being much more perceptible to the eye in larger-sized individuals. In preserved individuals, the form of palmar tubercle looses its shape, becoming more flattened. The nasal bump on the tip of the nose is more visible in adults males and females, while in juveniles this character is less visible. Distribution and natural history. Amazophrynella amazonicola sp. nov. is known to occur from five localities in the Peruvian department of Loreto: Fundo Zamora, Fundo UNAP, Allpahuayo Mishana National Reserve, the vicinity of Nauta and Puerto Almendra. This species was recorded in white sand forests, locally known as ��� varillales ��� (in Portuguese ��� campinaranas ���), from around Iquitos. It occurs in northwest of Iquitos (left bank of the Amazon, Nanay and Mara��on Rivers), where its distribution spans the length of the Iquitos-Nauta highway; further north, its distribution is poorly known but is possibly delimited by the Napo and Mara��on Rivers (Figure 7). Amazophrynella amazonicola sp. nov. breeds between December and March (Figure 8). The species is diurnal, and can be found amidst leaf litter, where its coloration makes it difficult to locate. Etymology. The specific epithet refers to the Amazon River, the world���s largest river and one of the new seven wonders of nature., Published as part of Rojas, Rommel R., Carvalho, Vin��cius Tadeu De, ��vila, Robson W., Farias, Izeni Pires, Gordo, Marcelo & Hrbek, Tomas, 2015, Two new species of Amazophrynella (Amphibia: Anura: Bufonidae) from Loreto, Peru, pp. 79-103 in Zootaxa 3946 (1) on pages 86-92, DOI: 10.11646/zootaxa.3946.1.3, http://zenodo.org/record/236987, {"references":["Rodriguez, L. O. & Duellman, W. E. (1994) Guide to the Frogs of the Iquitos Region. Amazonian Peru. University of Kansas Natural History Museum, special Publications, 22, 1 - 80.","Fouquet, A., Recoder, R., Teixeira Jr., M., Cassimiro, J., Amaro, R. C., Camacho, A., Damasceno, R., Carnaval, A. C., Moritz, C. & Rodrigues, M. T. (2012 b) Amazonella Fouquet et al. 2012 (Anura: Bufonidae) junior homonym of Amazonella Lundblad, 1931 (Acari: Unionicolidae): proposed replacement by Amazophrynella nom. nov. Zootaxa, 3244, 68."]}

Amazophrynella manaos sp. nov. (Figure 1) Dendrophryniscus minutus (McDiarmid, 1971; Zimmerman & Rodrigues, 1990; Magnusson & Hero, 1991; Lima et al., 2006) Amazonella minuta (Fouquet et al., 2012 a) Amazophrynella minuta (Fouquet et al., 2012 b) Holotype. INPA-H 31866, adult male, collected in forest remnants of the campus of the Universidade Federal do Amazonas—UFAM (03° 05' 56 "S, 59 ° 58 ' 09" W), municipality of Manaus, Amazonas State, by Vinicius Tadeu de Carvalho and Rommel Roberto Rojas, on June 26, 2012 at 16: 35 h. Paratypes. INPA-H 29568, INPA-H 29571, INPA-H 29570, INPA-H 29572, INPA-H 29569 (adult males), collected in Mineração Taboca (0° 42 ' 59 " S, 60 ° 10 ' 24 " W), municipality of Presidente Figueiredo, Amazonas State, by Vinicius Tadeu de Carvalho, on December 17, 2011; INPA-H 6983, INPA-H 6984, INPA-H 6987 (adult males), collected on the campus of the Universidade Federal do Amazonas—UFAM (03° 05' 37 " S, 59 ° 58 ' 26 " W), Amazonas State, by Ana Cristina de Oliveira Cordeiro, on February 10, 2010; INPA-H 20986; INPA-H 21217 (adult females), INPA-H 21028, INPA-H 21170, INPA-H 21060 (adult males), collected in the Reserva Florestal Adolpho Ducke (02° 55 ' 37 " S, 59 ° 58 ' 14 " W), by Albertina P. Lima, in December 1992; INPA-H 30577, INPA-H 30575, INPA-H 30573, INPA-H 30572, INPA-H 30576 (adult females), collected at the municipality of Presidente Figueiredo (01° 56 ' 20 " S, 60 ° 02' 14 " W), Amazonas State, by André Luiz Ferreira da Silva, on January 5, 2012; INPA-H 21442, INPA-H 21400 (adult females), INPA-H 21451, INPA-H 21398, INPA-H 21821 (adult males), collected at Reserve ZF- 2 (02° 35 ' 20 "S, 60 °06' 55 "W), municipality of Presidente Figueiredo, Amazonas State, by Albertina P. Lima, in December 1997; INPA-H 1859 (adult male), collected at Parque Estadual Rio Negro Setor Sul, Cuieiras River (02° 43 ' 27 " S, 60 ° 24 ' 22 " W), municipality of Manaus, Amazonas State, by Vinicius Tadeu de Carvalho and Lucéia Bonora, on January 10, 2007. Diagnosis. The new species is assigned to the genus Amazophrynella by the combination of the following generic characters: small size, depressed body, eardrum and parotid glands not visible, presence of granulate bodies, presence of subarticular tubercles on hands, elongated snout, and via phylogenetically nested position within the genus Amazophrynella. The new species is characterized by: 1) triangular snout with a slightly truncate culmination (in ventral view); 2) ventral surface covered with high density fine granules; 3) white abdomen; 4) black patches or stripes on the abdomen; 5) fifteen molecular autapomorphies in the 16 S rDNA (Table 2). Divergence from its nominal sister taxon is 9 %. Molecular phylogeny and genetic distance. Phylogenetic analysis of nominal and hypothetical taxa of the genus Amazophrynella indicates the existence of six lineages (Figure 2). Basal divergence is between the clade composed of an undescribed Amazophrynella species from western Amazônia and A. minuta specimens sampled from the type locality, and a clade comprising all other species and lineages. The A. minuta + A. aff. minuta “western Amazonia” clade is supported by 97 % bootstrap value while the other clade is supported by 80 % bootstrap value. The new species, A. manaos is sister to a possibly a new species of Amazophrynella from the Guiana Shield (bootstrap support 70 %) and both are sister to A. bokermanni (bootstrap support 75 %). The recently described A. vote (italicize) is sister to A. bokemanni + (A. manaos + A. sp. “Guianas”) with a bootstrap support of 80 %. Smallest uncorrected 16 S rDNA p -distances (4 %) was observed between A. manaos sp. nov. and A. sp. “Guianas”. Greatest intrageneric distance (14 %) was observed between A. manaos sp. nov. and A. minuta, and A. sp. “Guianas” and A. minuta (see Table 3). Characterization of topotypic material of Amazophrynella minuta. Topotypic material of A. minuta was collected in mission Taracuá, right bank of the middle Uaupés River in the last week of August 2013. Adult males measured 13.5 ± 0.57 mm (12.4–14.2 mm), females 17.5 ± 0.61 mm (17.1–18.9 mm) in snout vent length. The specimens (Figure 3) were characterized by a pointed head (in ventral view); pointed snout (in lateral view); roughly granular dorsal and ventral skin; a scattering of abundant pointed spicules on the body; big warts on the dorso-lateral region; dark gray brown throat and chest; intense yellow-orange coloration of the abdomen; and by irregular black blotches or ocelli on the abdomen. Fingers are unwebbed. Formula of the fingers: I Comparisons with other species. The new species can be differentiated from the three other nominal species of Amazophrynella by the following characters (characters of compared species in parentheses): a) from A. minuta (Melin, 1941) by the triangular snout with a slightly truncate culmination (snout pointed), absence of spiny tubercles on dorsum (prickly warty skin on dorsum), skin of axillary region and forearm finely granular (rough granular skin on forearm and axillary region, especially on the neck), venter white with black spots and darkbrown chest in life (venter bright reddish-orange, with small black blotches and throat light brown) (Figure 4 A-C and Figure 5 A, Table 4); b) from A. bokermanni (Izecksohn, 1993), by finger I shorter than finger II (finger I longer than finger II), smaller SVL, with males reaching 15.8 mm and females 24.7 mm (maximum SVL of males and females 22 mm and 28 mm, respectively) (Figure 4 G–I, Figure 5 B); and c) from A. vote Ávila, Carvalho, Gordo, Kawashita & Morais, 2012, by snout triangular with a slight culmination (rounded), slightly smaller SVL in males (14.7–15.8 mm vs. 16.4 mm—Ávila et al., 2012), and white abdomen with black spots or stripes and blackish to dark-brown chest and throat in life (abdomen reddish-brown with small white dots and brown spots) and subrostral crest converging anteriorly (subrostral crest not converging anteriorly) (Figure 4 D–F, Figure 5 C). Description of the holotype. Small species; body slender; head longer than wide; head triangular; head length 36 % SVL; upper eyelids close to 60 % of interorbital distance; snout elongate; snout profile slightly truncated in lateral view; eyes prominent, 31 % of head width; nostril is closer to the tip of the snout than to the eyes; tympanums not visible; parotid gland absent. Body covered by abundant rounded granules; texture of dorsal skin granular; abundant granules grouped in the axillar region; texture of the ventral skin granular. Forelimbs slender; upper arm covered by tiny conical tubercles. Fingers unwebbed; formula of the fingers: I Measurement of the holotype (in mm). SVL: 13.9; HL: 5.1; HW: 4.1; EW: 1.1; ED: 1.3; SL: 2.3; END: 1.4; IND: 1.0; IOD: 1.8; HAL: 3.0; UAL: 4.1; THL: 7.2; TL: 6.9; TAL: 4.4 and FL: 4.9. Coloration of the holotype. In life, dorsal color pattern of the holotype dark-brown with transversal black and light-brown bars, more evident on limbs; thin cream medial line extending from head to cloaca, a white longitudinal stripe on upper jaw extending from nostril to forearm; arms white ventrally; throat and chest region blackish; belly white with large black spots, thighs and tibiae predominantly grey ventrally with few small black dots; palms and soles reddish (Figure 5 D). Coloration of preserved specimens is nearly identical to that described in life but black spots on the belly are less evident. Variation. Variation exists among individuals within the same locality as well as between localities. In some individuals the coloration of the throat extends onto the chest and the belly has few black spots; other specimens show a ventral pattern of black stripes that extend dorsally to the upper part of the lateral region. Juveniles have a less pronounced pattern on the belly, with few black spots. Individuals from some localities have a thin clear brown line that extends from the rostrum to the cloaca along the central dorsum (Figure 6). Distribution and natural history. Amazophrynella manaos sp. nov. is distributed throughout the southwestern part of the Brazilian Guiana region. It was collected in eight localities in the state of Amazonas, Brazil: Mineração Taboca, Campus of the Universidade Federal do Amazonas, Reserva Florestal Adolpho Ducke, Presidente Figueiredo, Reserva ZF- 2, REBIO Uatumã, RDS Uatumã and Parque Estadual Rio Negro Setor Sul, Cuieiras River. To the south and west its distribution is likely to be delimited by the Amazon and Negro Rivers, respectively. It thus appears to be restricted to the south-central portion of the Guiana Shield (Figure 7). It occurs in primary forests and forest fragments, and is found in leaf litter—often fallen fronds of the buriti palm (Mauritia flexuosa)—generally close to creeks. The species is diurnal, is clearly sexually dimophic in size (Figure 9, Table 5, online supplement 1) with females being distinctly larger than males. The amplexus is cephalic (Figure 9). Reproductive period is from November to April (Magnusson & Hero, 1991). Etymology. We name the species in honor of the Manaos Amerindian tribe that inhabited the region of the present day city of Manaus, Amazonas, Brazil, where the species is distributed.

Amazophrynella manaos sp. nov. (Figure 1) Dendrophryniscus minutus (McDiarmid, 1971; Zimmerman & Rodrigues, 1990; Magnusson & Hero, 1991; Lima et al., 2006) Amazonella minuta (Fouquet et al., 2012 a) Amazophrynella minuta (Fouquet et al., 2012 b) Holotype. INPA-H 31866, adult male, collected in forest remnants of the campus of the Universidade Federal do Amazonas���UFAM (03�� 05' 56 "S, 59 �� 58 ' 09" W), municipality of Manaus, Amazonas State, by Vinicius Tadeu de Carvalho and Rommel Roberto Rojas, on June 26, 2012 at 16: 35 h. Paratypes. INPA-H 29568, INPA-H 29571, INPA-H 29570, INPA-H 29572, INPA-H 29569 (adult males), collected in Minera����o Taboca (0�� 42 ' 59 " S, 60 �� 10 ' 24 " W), municipality of Presidente Figueiredo, Amazonas State, by Vinicius Tadeu de Carvalho, on December 17, 2011; INPA-H 6983, INPA-H 6984, INPA-H 6987 (adult males), collected on the campus of the Universidade Federal do Amazonas���UFAM (03�� 05' 37 " S, 59 �� 58 ' 26 " W), Amazonas State, by Ana Cristina de Oliveira Cordeiro, on February 10, 2010; INPA-H 20986; INPA-H 21217 (adult females), INPA-H 21028, INPA-H 21170, INPA-H 21060 (adult males), collected in the Reserva Florestal Adolpho Ducke (02�� 55 ' 37 " S, 59 �� 58 ' 14 " W), by Albertina P. Lima, in December 1992; INPA-H 30577, INPA-H 30575, INPA-H 30573, INPA-H 30572, INPA-H 30576 (adult females), collected at the municipality of Presidente Figueiredo (01�� 56 ' 20 " S, 60 �� 02' 14 " W), Amazonas State, by Andr�� Luiz Ferreira da Silva, on January 5, 2012; INPA-H 21442, INPA-H 21400 (adult females), INPA-H 21451, INPA-H 21398, INPA-H 21821 (adult males), collected at Reserve ZF- 2 (02�� 35 ' 20 "S, 60 ��06' 55 "W), municipality of Presidente Figueiredo, Amazonas State, by Albertina P. Lima, in December 1997; INPA-H 1859 (adult male), collected at Parque Estadual Rio Negro Setor Sul, Cuieiras River (02�� 43 ' 27 " S, 60 �� 24 ' 22 " W), municipality of Manaus, Amazonas State, by Vinicius Tadeu de Carvalho and Luc��ia Bonora, on January 10, 2007. Diagnosis. The new species is assigned to the genus Amazophrynella by the combination of the following generic characters: small size, depressed body, eardrum and parotid glands not visible, presence of granulate bodies, presence of subarticular tubercles on hands, elongated snout, and via phylogenetically nested position within the genus Amazophrynella. The new species is characterized by: 1) triangular snout with a slightly truncate culmination (in ventral view); 2) ventral surface covered with high density fine granules; 3) white abdomen; 4) black patches or stripes on the abdomen; 5) fifteen molecular autapomorphies in the 16 S rDNA (Table 2). Divergence from its nominal sister taxon is 9 %. Molecular phylogeny and genetic distance. Phylogenetic analysis of nominal and hypothetical taxa of the genus Amazophrynella indicates the existence of six lineages (Figure 2). Basal divergence is between the clade composed of an undescribed Amazophrynella species from western Amaz��nia and A. minuta specimens sampled from the type locality, and a clade comprising all other species and lineages. The A. minuta + A. aff. minuta ���western Amazonia��� clade is supported by 97 % bootstrap value while the other clade is supported by 80 % bootstrap value. The new species, A. manaos is sister to a possibly a new species of Amazophrynella from the Guiana Shield (bootstrap support 70 %) and both are sister to A. bokermanni (bootstrap support 75 %). The recently described A. vote (italicize) is sister to A. bokemanni + (A. manaos + A. sp. ���Guianas���) with a bootstrap support of 80 %. Smallest uncorrected 16 S rDNA p -distances (4 %) was observed between A. manaos sp. nov. and A. sp. ���Guianas���. Greatest intrageneric distance (14 %) was observed between A. manaos sp. nov. and A. minuta, and A. sp. ���Guianas��� and A. minuta (see Table 3). Characterization of topotypic material of Amazophrynella minuta. Topotypic material of A. minuta was collected in mission Taracu��, right bank of the middle Uaup��s River in the last week of August 2013. Adult males measured 13.5 �� 0.57 mm (12.4���14.2 mm), females 17.5 �� 0.61 mm (17.1���18.9 mm) in snout vent length. The specimens (Figure 3) were characterized by a pointed head (in ventral view); pointed snout (in lateral view); roughly granular dorsal and ventral skin; a scattering of abundant pointed spicules on the body; big warts on the dorso-lateral region; dark gray brown throat and chest; intense yellow-orange coloration of the abdomen; and by irregular black blotches or ocelli on the abdomen. Fingers are unwebbed. Formula of the fingers: IComparisons with other species. The new species can be differentiated from the three other nominal species of Amazophrynella by the following characters (characters of compared species in parentheses): a) from A. minuta (Melin, 1941) by the triangular snout with a slightly truncate culmination (snout pointed), absence of spiny tubercles on dorsum (prickly warty skin on dorsum), skin of axillary region and forearm finely granular (rough granular skin on forearm and axillary region, especially on the neck), venter white with black spots and darkbrown chest in life (venter bright reddish-orange, with small black blotches and throat light brown) (Figure 4 A-C and Figure 5 A, Table 4); b) from A. bokermanni (Izecksohn, 1993), by finger I shorter than finger II (finger I longer than finger II), smaller SVL, with males reaching 15.8 mm and females 24.7 mm (maximum SVL of males and females 22 mm and 28 mm, respectively) (Figure 4 G���I, Figure 5 B); and c) from A. vote ��vila, Carvalho, Gordo, Kawashita & Morais, 2012, by snout triangular with a slight culmination (rounded), slightly smaller SVL in males (14.7���15.8 mm vs. 16.4 mm�����vila et al., 2012), and white abdomen with black spots or stripes and blackish to dark-brown chest and throat in life (abdomen reddish-brown with small white dots and brown spots) and subrostral crest converging anteriorly (subrostral crest not converging anteriorly) (Figure 4 D���F, Figure 5 C). Description of the holotype. Small species; body slender; head longer than wide; head triangular; head length 36 % SVL; upper eyelids close to 60 % of interorbital distance; snout elongate; snout profile slightly truncated in lateral view; eyes prominent, 31 % of head width; nostril is closer to the tip of the snout than to the eyes; tympanums not visible; parotid gland absent. Body covered by abundant rounded granules; texture of dorsal skin granular; abundant granules grouped in the axillar region; texture of the ventral skin granular. Forelimbs slender; upper arm covered by tiny conical tubercles. Fingers unwebbed; formula of the fingers: IMeasurement of the holotype (in mm). SVL: 13.9; HL: 5.1; HW: 4.1; EW: 1.1; ED: 1.3; SL: 2.3; END: 1.4; IND: 1.0; IOD: 1.8; HAL: 3.0; UAL: 4.1; THL: 7.2; TL: 6.9; TAL: 4.4 and FL: 4.9. Coloration of the holotype. In life, dorsal color pattern of the holotype dark-brown with transversal black and light-brown bars, more evident on limbs; thin cream medial line extending from head to cloaca, a white longitudinal stripe on upper jaw extending from nostril to forearm; arms white ventrally; throat and chest region blackish; belly white with large black spots, thighs and tibiae predominantly grey ventrally with few small black dots; palms and soles reddish (Figure 5 D). Coloration of preserved specimens is nearly identical to that described in life but black spots on the belly are less evident. Variation. Variation exists among individuals within the same locality as well as between localities. In some individuals the coloration of the throat extends onto the chest and the belly has few black spots; other specimens show a ventral pattern of black stripes that extend dorsally to the upper part of the lateral region. Juveniles have a less pronounced pattern on the belly, with few black spots. Individuals from some localities have a thin clear brown line that extends from the rostrum to the cloaca along the central dorsum (Figure 6). Distribution and natural history. Amazophrynella manaos sp. nov. is distributed throughout the southwestern part of the Brazilian Guiana region. It was collected in eight localities in the state of Amazonas, Brazil: Minera����o Taboca, Campus of the Universidade Federal do Amazonas, Reserva Florestal Adolpho Ducke, Presidente Figueiredo, Reserva ZF- 2, REBIO Uatum��, RDS Uatum�� and Parque Estadual Rio Negro Setor Sul, Cuieiras River. To the south and west its distribution is likely to be delimited by the Amazon and Negro Rivers, respectively. It thus appears to be restricted to the south-central portion of the Guiana Shield (Figure 7). It occurs in primary forests and forest fragments, and is found in leaf litter���often fallen fronds of the buriti palm (Mauritia flexuosa)���generally close to creeks. The species is diurnal, is clearly sexually dimophic in size (Figure 9, Table 5, online supplement 1) with females being distinctly larger than males. The amplexus is cephalic (Figure 9). Reproductive period is from November to April (Magnusson & Hero, 1991). Etymology. We name the species in honor of the Manaos Amerindian tribe that inhabited the region of the present day city of Manaus, Amazonas, Brazil, where the species is distributed., Published as part of Rojas, Rommel R., Carvalho, Vin��cius Tadeu De, Gordo, Marcelo, ��vila, Robson W., Farias, Izeni Pires & Hrbek, Tomas, 2014, A new species of Amazophrynella (Anura: Bufonidae) from the southwestern part of the Brazilian Guiana Shield, pp. 79-95 in Zootaxa 3753 (1) on pages 82-90, DOI: 10.11646/zootaxa.3753.1.7, http://zenodo.org/record/285395, {"references":["McDiarmid, R. W. (1971) Comparative morphology and evolution of frogs of the Neotropical genera Atelopus, Dendrophryniscus, Melanophryniscus and Oreophrynella. Bulletin of the Los Angeles County Museum of Natural History, 12, 1 - 66.","Zimmerman, B. L. & Rodrigues, M. T. (1990) Frogs, snakes, and lizards of INPA-WWF reserves near Manaus, Brazil. In: Gentry, A. H. (Eds.), Four Neotropical Rainforests. Yale University Press, Connecticut, USA, pp. 426 - 454.","Magnusson, W. E. & Hero, J. H. (1991) Predation and the evolution of complex oviposition behaviour in Amazon rainforest frogs. Oecologia, 86, 310 - 318. http: // dx. doi. org / 10.1007 / bf 00317595","Lima, A. P., Magnusson, W. E., Menin, M., Erdtmann, L. K., Rodrigues, D. J., Keller, C. & Hodl, W. (2006) Guia de sapos da Reserva Adolpho Ducke, Amazonia Central - Guide to the frogs of Reserva Adolpho Ducke, Central Amazonian. Attema Design Editorial, Manaus, 168 pp.","Fouquet, A., Recoder, R., Teixeira Jr., M., Cassimiro, J., Amaro, R. C., Camacho, A., Damasceno, R., Carnaval, A. C., Moritz, C. & Rodrigues, M. T. (2012 a) Molecular phylogeny and morphometric analyses reveal deep divergence between Amazonian and Atlantic Forest species of Dendrophryniscus. Molecular Phylogenetics and Evolution, 62, 823 - 838. http: // dx. doi. org / 10.1016 / j. ympev. 2011.11.023","Fouquet, A., Recoder, R., Teixeira Jr., M., Cassimiro, J., Amaro, R. C., Camacho, A., Damasceno, R., Carnaval, A. C., Moritz, C. & Rodrigues, M. T. (2012 b) Amazonella Fouquet et al., 2012 (Anura: Bufonidae) junior homonym of Amazonella Lundblad, 1931 (Acari: Unionicolidae): proposed replacement by Amazophrynella nom. nov. Zootaxa, 3244, 68.","Melin, D. E. (1941) Contributions to the knowledge of the Amphibia of South America. Goteborgs Kungl. Vetenskaps - och Vitterhets - samhalles. Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1 - 71.","Izecksohn, E. (1993) Nova especie de Dendrophryniscus da regiao amazonica (Amphibia, Anura, Bufonidae). Revista Brasileira de Zoologia, 10, 407 - 412. http: // dx. doi. org / 10.1590 / s 0101 - 81751993000300006","Avila, R. W., Carvalho, V. T., Gordo, M., Ribeiro, K. & Morais, D. (2012) A new species of Amazophrynella (Anura: Bufonidae) from southern Amazonia. Zootaxa, 3484, 65 - 74."]}

Semedo, Thiago Borges Fernandes, Brandão, Marcus Vinicius, Carmignotto, Ana Paula, Nunes, Mario Da Silva, Farias, Izeni Pires, Silva, Maria Nazareth Ferreira Da, Rossi, Rogério Vieira (2015): Taxonomic status and phylogenetic relationships of Marmosa agilis peruana Tate, 1931 (Didelphimorphia: Didelphidae), with comments on the morphological variation of Gracilinanus from central-western Brazil. Zoological Journal of the Linnean Society 173 (1): 190-216, DOI: 10.1111/zoj.12203, URL: http://dx.doi.org/10.1111/zoj.12203

Allobates hodli sp. nov Figures 2–5. Epipedobates femoralis Hödl et al. 2004 p. 823, Catuaba, Acre population (partim). Allobates femoralis Amézquita et al. 2006 p. 1877, Catuaba, Acre population (partim); Lötters et al. 2007 p. 307, Fig. 379; Simões et al. 2008 p. 610, Fig. 2 B. (partim); Amézquita et al. 2009, Fig. 1, Catuaba pattern (partim). Holotype. INPA – H 16555 (original field number APL 2014). Adult male, collected by P. I. Simões and A. P. Lima after recording of advertisement calls at 07: 55 h, 25 th of November 2004, at Cachoeira do Jirau, on the left bank of the upper Madeira River (09.3347° S, 64.7375 ° W), approximately 125 km upstream from the city of Porto Velho, Estado de Rondônia, Brazil. Paratopotypes. INPA-H 16541–16554, INPA-H 16556–16569 (original field numbers APL 2000 –2013, 2015–2018, 2022–2030, 2032), 6 females, 22 males. Collected in the same locality as holotype, 23-25 November 2004 by P. I. Simões and A. P. Lima. Paratypes. All from Brazil. Acre: INPA-H 11621–11640,, 4 females, 17 males, Fazenda Catuaba, Municipality of Rio Branco, 10.0742 ° S, 67.6249 ° W, collected in February 2004 by A. P. Lima. Rondônia: INPA-H 16578, 16584–16587, 16589, 16591 –16592, 16597, 16602 –16603, 16605–16607, 16611–16614, 16620–16624, 16626, 16628, 16631, 16633, 16636–16637, 16639–16641, 16643, 16645 –16646, 16648, 13 females, 26 males, collected on the left bank of the upper Madeira River, across the river from the village of Fortaleza do Abunã, 160 km upstream from the city of Porto Velho, 72 km upstream from Cachoeira do Jirau, 9.5160 ° S, 65.3249 ° W, collected 05-08 January 2005 by P.I. Simões and A.P. Lima. INPA-H 16596, 16730, 16739, 16756, 16758, 16767, 16771, 16777–16778, 16788, 16805, 16818–16819, 2 females, 11 males, collected on the left bank of the upper Madeira River, across the river from the village of Mutum-Paraná, 121 km upstream from the city of Porto Velho, 34 km upstream from Cachoeira do Jirau, 9.5732 ° S, 64.9211 ° W, collected 10–13 January 2005 by P. I. Simões and A. P. Lima. Etymology. The specific epithet is a patronym for Dr. Walter Hödl, an Austrian biologist and professor who pioneered research on behavior and acoustic communication in anurans. For the past two decades, Walter and his students have dedicated special attention to the Allobates femoralis complex. Diagnosis. The new species is assigned to the genus Allobates by the combination of the following characters: presence of a pale dorsolateral stripe, dorsal skin texture granular posteriorly, basal webbing present only between Toes III and IV, Finger I longer than Finger II, finger discs generally weakly expanded (moderately expanded on Finger I), median lingual process absent, testes not pigmented, dark collar absent on throat, oral disc of tadpoles emarginate, not umbelliform. Allobates hodli is distinguished in life from all other species of Allobates (except Allobates femoralis, Allobates myersi and Allobates zaparo) for presenting relatively large body-size (average SVL = 24.76 ± 1.08 mm, males and females pooled), by the lack of brown or light-brown colors or patterning on dorsum and lateral surface of body, and by presenting dark and white marbling on anterior ventral surface of body, replaced by solid reddish-orange color on the ventral surface of hind limbs. Allobates hodli is distinguished from other taxa and morphotypes that form the A. femoralis complex by presenting advertisement calls consisting of groups of two notes repeated in series or bouts (instead of groups of one, three or four notes), and by presenting a conspicuous reddish-orange coloration on the ventral surface of legs, instead of an exclusively black and white reticulated pattern, observed in A. femoralis. Allobates hodli also has diffuse reddish-orange and black patches on dorsal surface of thighs, as opposed to regular, pale (yellowish to red) longitudinal flash marks extending onto the entire dorsal surface of thighs, generally margined by dark patches, observed in typical A. femoralis. A. hodli is distinguished from A. zaparo and A. myersi by the color of dorsum, which is uniformly black/ dark-brown in A. hodli (Fig. 2), but reddish in A. zaparo and brown to light-brown in A. myersi. Allobates myersi also lacks a pale dorsolateral stripe. A. hodli is largely sympatric to Ameerega picta, a dendrobatid frog that presents similar body size and color pattern. However, A. hodli can be distinguished from Ameerega picta by lacking a bright (orange to red) flash mark on calf region. Description of holotype. Morphological measurements of holotype are presented in Table 1. Body robust, head slightly wider than long (HL/HW = 0.94) (Fig. 3 A). Eye diameter slightly larger than distance from nostril to anterior corner of the eye. Nares located posterolaterally to tip of snout, directed posterolaterally, visible in ventral and anterior view. Center of nostril not visible dorsally. Canthus rostralis convex from tip of snout to nostril, straight from nostril to anterior corner of the eye. Loreal region vertical. Tympanum well visible, with maximum diameter horizontal, corresponding to 44 % the maximum diameter of the eye. Maxillary teeth present. Tongue length twice as large as wide, attached anteriorly on first third. Median lingual process absent. Choanae round. A single vocal sac is present, corresponding to most of the area of the medial and posterior subgular region. Vocal sac round when expanded. When retracted, vocal sac forms two lateral slits at the level of maxilla articulation (Fig. 3, B). Skin granular on dorsum and dorsal surface of legs. Granules round, more developed on dorsal surface of urostyle region and shanks. Skin smooth ventrally and laterally. Dermal flap above cloaca absent. Palmar tubercle slightly triangular. Thenar tubercle well-developed, oval to elliptic, maximum diameter 1.28 times smaller than maximum diameter of palmar tubercle. Subarticular tubercles of Fingers II, III and IV are round, small, never exceeding the width of phalanges. Subarticular tubercle of Finger I elliptic, 1.21 times larger than thenar tubercle in maximum diameter. Supernumerary tubercles absent. Carpal pad and metacarpal ridges absent on hands. No fringes or webbing on fingers. A distal tubercle on finger IV is weakly developed (Fig. 3 C). Finger I is slightly (1.08 times) longer than Finger II. Length of finger IV does not reach distal subarticular tubercle of finger III when fingers are pressed against each other. Relative lengths of fingers: IV Length of shank corresponding to 48 % of snout-to-vent length (Table 1). Tarsal keel is tubercle-like, strongly curved at its proximal end, flattening towards the metatarsal tubercle. Metatarsal fold evident (but not folding over itself) running from the base of Toe V towards metatarsal tubercle, but not reaching it. Preaxial edge of tarsus smooth, with no fringe. Basal webbing present only between Toes III and IV, and II and III. Relative lengths of toes: I Variation in type series. Morphological measurements of individuals constituting type series are presented in Table 1. Morphological characters described for the holotype apply to all individuals in type series, except for the following: Males slightly smaller (4.42 %, in average) than females. Head slightly longer than wide in males (HL/HW = 1.04) and females (HL/HW = 1.05) in average. Maximum diameter of tympanum corresponding to approximately half the maximum diameter of the eye in males and females (Table 1). Vocal sac and slits absent in females. Palmar tubercle round to slightly triangular. A distal tubercle on finger IV is present in 28 of a total 83 (34.1 %) inspected specimens, but is absent or weakly developed in the remaining 54 specimens (65.9 %). Color in life. Males and females do not present dimorphism in relation to color and color pattern. Dorsal surface of body solid black to solid dark-brown (Fig. 2 B). Lateral surface of body solid black. Dorsolateral line white, thinner than lateral line (Fig. 2 A, 2 E). When continuous with flash marks on thighs, dorsolateral line becomes reddish-orange on groin. Lateral line white. Gular region solid black to dark bluish-gray in males and females (Fig. 2 D). In males, vocal sacs usually with a paler bluish-gray color when inflated. Mid abdomen white with irregular black to dark-gray blotches or speckling, merging with solid dark color of gular region. Abdomen bright reddish-orange posteriorly, with dark irregular spots appearing marginally from lateral edges. Ventral surfaces of hind limbs also bright reddish-orange, sometimes with small marginal dark spots (Fig. 2 C, 2 D). Plantar surface of feet brown. Ventral surfaces of arms bright reddish-orange, with bright yellow flash marks extending from dorsal surface of upper arms. Black to dark-gray spot ventrally on upper arm, at the point of body insertion, continuous with gular region pattern. Dorsal surfaces of posterior and anterior limbs reddish to brick-brown (Fig. 2 B, 2 E). Dorsal and rear surfaces of thighs with irregular bright reddish-orange flash marks or patterning, same color as ventral surfaces of legs, with irregular black or darkbrown blotches or spots. Granules on dorsal surface of shanks usually darker than overall color of shanks. A yellow flash mark is present dorsally on upper arms, at the point of body insertion (Fig. 2 B, 2 E). The iris is evident, with metallic yellowish-brown pigmentation. Color in life of juveniles. Color of juveniles after metamorphosis is the same of adults. Dorsum and flanks are solid black to dark-brown, with dorsolateral and lateral lines white and conspicuous. Limbs generally reddish-brown. Bright yellow flash marks are present dorsally on the upper arms, and may reach the elbow. Dorsal surface of thighs with conspicuous longitudinal bright reddish-orange flash marks, lacking black or dark-brown blotches or spots (Fig. 2 F). Color in alcohol of holotype. Dorsum is solid black to dark-brown. A thin, pale white dorsolateral line is present, continuous from groin at hind limb insertion, over the orbit and nostril, to the tip of snout (Fig. 3 A). A pale white lateral line is present, broader than dorsolateral stripe, running from groin, over the insertion of arm, below nostril, to tip of upper lip. Lateral stripe is continuous on both sides of the body. Lateral surface between dorsolateral and lateral stripe solid black. Color of gular region and throat is solid black to darkbrown. Abdomen color is white with irregular black blotches or speckling. Abdomen color becomes solid black/dark-brown from chest towards the gular region. The black/dark-brown speckling over white background pattern is replaced posteriorly by a solid pale-tan pattern, continuous with the ventral color pattern of hind limbs (Fig. 3 B). Arms uniformly very pale-brown in dorsal view, paler on the axilla and carpal/metacarpal regions. Irregular dark blotches appear on dorsal surfaces of tarsus and fingers. Arms uniformly pale white to pale tan in ventral view, with a black patch (continuous with color pattern of gular region) on anterior surface of the arm. Surface of outer lateral edge of forearm and metacarpal region same color as dorsal surface of arms, extending laterally from elbow and reaching the palm of hands and ventral surface of fingers (Fig. 3 C). Legs are pale brown in dorsal view. Irregular pale, unpigmented patches, as well as irregular black blotches and spots are present on dorsal and rear surfaces of thighs (Fig. 3 A). The area immediately around vent is darker than the overall surface of thighs. Dorsal surfaces of shanks with darker granules. Inner dorsal surface of tarsal region is lighter than overall pattern of legs. Toes are generally darker than tarsal region in dorsal view. Ventral surface of legs is uniformly pale-tan with small brown spots appearing marginally from outer edges. Ventral surfaces of tarsal region and toes darker, same color as dorsal surface of legs (Fig. 3 B, 3 D). Tongue is cream-colored; large intestine (removed for the analysis of diet) is unpigmented. Testes are unpigmented. Color variation in type series. Color in alcohol described for the holotype apply to all individuals in type series, except for the following: Lateral stripe is usually continuous, but can be interrupted in some individuals, on one or on both sides of body. Gular region and throat solid black in females, solid black to dark-brown in males. Mature oocytes are pigmented, with black pigment concentrated on animal pole. Description of tadpoles. Tadpole measurements were obtained from four tadpoles in developmental stage 36 (Table 2). Tadpoles correspond to a lot under the same collection number, INPA-H 23693. The largest tadpole (TL = 24.7 mm, Fig. 4) was used for detailed description. Body is depressed, body width (6.0 mm) larger than body depth (4.8 mm), body length 16.1 mm. Snout nearly round, flattened anterodorsally in lateral view (Fig. 4 C). Tip of snout flattened anteriorly in dorsal view (Fig. 4 A). Nares small, directed anterolaterally, located 0.8 mm anterior to the eye, and 1.0 mm posterior to tip of snout. Nostrils narrowly spaced, distance between nostrils 0.9 mm. Eyes dorsal, directed dorsolaterally, 0.9 mm in maximum length, located 1.8 mm posterior to tip of snout. Distance between medial margins of the eyes is 1.4 mm. Spiracle single, sinistral, forming a free tube opening posterodorsally below body axis in lateral view, 5.0 mm posterior from tip of snout (Fig. 4 C). Vent tube medial, free, 0.9 mm in length, opening dextrally. Tail musculature reaches maximum depth (2.4 mm) approximately at the end of first third of tail length, and maximum width at body-tail insertion (2.6 mm). Ventral tail fin originates at body-tail insertion. Dorsal tail fin originates slightly posterior (0.8 mm) to body-tail insertion, and reaches maximum high 14.5 mm from tip of snout, corresponding to the region of maximum tail depth. At maximum depth of tail, depth of musculature is 1.4 mm, dorsal fin 1.5 mm and ventral fin 1.1 mm. Oral disc is positioned anteroventrally, emarginate laterally, transversely elliptical, 2.6 mm in transverse width. Anterior labium continuous with snout, 2.6 mm in length. Marginal papillae absent dorsally on anterior labium (gap 1.7 mm, 74 % of total anterior labium length), present only laterally, on its outer margins. Posterior labium free from body wall, 2.4 mm in length, with a single row of marginal papillae. All papillae with rounded tips (Fig. 5). Labial tooth row formula is 2 (2)/ 3 (1). Rows A- 1 and A- 2 with same length (2.1 mm), A- 2 with a large medial gap (0.6 mm). Rows P- 1, P- 2 and P- 3 with same length (2.0 mm), P- 1 presenting a very narrow medial gap ( Color in preservative is dark to light tan. Body is darker than tail, with scattered brown melanophores in higher densities on anterior dorsum. High concentrations of melanophores also appear posteriorly on dorsum at the top and on flanking regions of tail muscle insertion (Fig. 4 A). Melanophores are evenly distributed on anterior ventral surface of body. Posterior ventral surface of body is transparent, not pigmented, and intestines are clearly visible through skin (Fig. 4 B). Tail musculature is light tan, tail fins transparent, with scattered brown melanophores forming irregular blotches on tail surface (Fig. 4 C). Advertisement and courtship calls description and variation. Advertisement calls of Allobates hodli consist in trills of calls formed by two whistle-like notes with ascending frequency modulation (Fig. 6 A, 6 C). Measurements of advertisement call characteristics of holotype and average values for 60 males from type series are presented on Table 3. The average maximum frequency of calls within type series is 3425.0 ± 184.7 Hz, and average duration of calls (summed durations of first and second notes, and inter-note silent interval) 0.164 ± 0.011 s. First note is less modulated (average difference between lower and higher frequency = 470.7 ± 94.8 Hz) than the second note (average difference between lower and higher frequency = 740.7 ± 115.0 Hz) and shorter in duration (0.033 ± 0.004 s, in comparison to second note, 0.056 ± 0.007 s). Courtship calls of A. hodli are quite distinct from their advertisement calls and are constituted by a continuous pulsed tone (Fig. 6 D), emitted only in the presence of females near the male’s calling perch. Average maximum frequency and duration of courtship calls obtained from calls of four individuals were 3190.2 Hz and 0.571 s, respectively (Table 4). The average pulse emission rate between calls was 125.2 pulses per second. Individuals Molecular phylogeny and genetic distances. From an initial sequence database containing 203 16 S rDNA sequences (including outgroups), a total of 93 unique haplotypes were used in the phylogenetic analysis (Table 5). Phylogenetic reconstructions support the existence of two basal clades within the Allobates femoralis group, both forming the sister clade to Allobates zaparo (Fig. 7 A). One of the basal clades contains Allobates hodli and the second contains samples from areas nearby Allobates femoralis type-locality, which we refer to as Allobates femoralis sensu stricto (clade femo 0 4, Fig. 7 B). A. hodli is marginally paraphyletic to populations that occur in the southern reaches of the Brazilian State of Acre (clade Acre 0 1, Fig. 7 B), which present advertisement calls constituted by four notes and color pattern more similar to that of A. femoralis than to that of A. hodli. Samples from the locality Monte Alegre were not clearly positioned within A. hodli or Acre 0 1 clade. This locality probably corresponds to a relictual contact zone between these clades. Allobates hodli and clade

Allobates hodli sp. nov Figures 2���5. Epipedobates femoralis H��dl et al. 2004 p. 823, Catuaba, Acre population (partim). Allobates femoralis Am��zquita et al. 2006 p. 1877, Catuaba, Acre population (partim); L��tters et al. 2007 p. 307, Fig. 379; Sim��es et al. 2008 p. 610, Fig. 2 B. (partim); Am��zquita et al. 2009, Fig. 1, Catuaba pattern (partim). Holotype. INPA ��� H 16555 (original field number APL 2014). Adult male, collected by P. I. Sim��es and A. P. Lima after recording of advertisement calls at 07: 55 h, 25 th of November 2004, at Cachoeira do Jirau, on the left bank of the upper Madeira River (09.3347�� S, 64.7375 �� W), approximately 125 km upstream from the city of Porto Velho, Estado de Rond��nia, Brazil. Paratopotypes. INPA-H 16541���16554, INPA-H 16556���16569 (original field numbers APL 2000 ���2013, 2015���2018, 2022���2030, 2032), 6 females, 22 males. Collected in the same locality as holotype, 23-25 November 2004 by P. I. Sim��es and A. P. Lima. Paratypes. All from Brazil. Acre: INPA-H 11621���11640,, 4 females, 17 males, Fazenda Catuaba, Municipality of Rio Branco, 10.0742 �� S, 67.6249 �� W, collected in February 2004 by A. P. Lima. Rond��nia: INPA-H 16578, 16584���16587, 16589, 16591 ���16592, 16597, 16602 ���16603, 16605���16607, 16611���16614, 16620���16624, 16626, 16628, 16631, 16633, 16636���16637, 16639���16641, 16643, 16645 ���16646, 16648, 13 females, 26 males, collected on the left bank of the upper Madeira River, across the river from the village of Fortaleza do Abun��, 160 km upstream from the city of Porto Velho, 72 km upstream from Cachoeira do Jirau, 9.5160 �� S, 65.3249 �� W, collected 05-08 January 2005 by P.I. Sim��es and A.P. Lima. INPA-H 16596, 16730, 16739, 16756, 16758, 16767, 16771, 16777���16778, 16788, 16805, 16818���16819, 2 females, 11 males, collected on the left bank of the upper Madeira River, across the river from the village of Mutum-Paran��, 121 km upstream from the city of Porto Velho, 34 km upstream from Cachoeira do Jirau, 9.5732 �� S, 64.9211 �� W, collected 10���13 January 2005 by P. I. Sim��es and A. P. Lima. Etymology. The specific epithet is a patronym for Dr. Walter H��dl, an Austrian biologist and professor who pioneered research on behavior and acoustic communication in anurans. For the past two decades, Walter and his students have dedicated special attention to the Allobates femoralis complex. Diagnosis. The new species is assigned to the genus Allobates by the combination of the following characters: presence of a pale dorsolateral stripe, dorsal skin texture granular posteriorly, basal webbing present only between Toes III and IV, Finger I longer than Finger II, finger discs generally weakly expanded (moderately expanded on Finger I), median lingual process absent, testes not pigmented, dark collar absent on throat, oral disc of tadpoles emarginate, not umbelliform. Allobates hodli is distinguished in life from all other species of Allobates (except Allobates femoralis, Allobates myersi and Allobates zaparo) for presenting relatively large body-size (average SVL = 24.76 �� 1.08 mm, males and females pooled), by the lack of brown or light-brown colors or patterning on dorsum and lateral surface of body, and by presenting dark and white marbling on anterior ventral surface of body, replaced by solid reddish-orange color on the ventral surface of hind limbs. Allobates hodli is distinguished from other taxa and morphotypes that form the A. femoralis complex by presenting advertisement calls consisting of groups of two notes repeated in series or bouts (instead of groups of one, three or four notes), and by presenting a conspicuous reddish-orange coloration on the ventral surface of legs, instead of an exclusively black and white reticulated pattern, observed in A. femoralis. Allobates hodli also has diffuse reddish-orange and black patches on dorsal surface of thighs, as opposed to regular, pale (yellowish to red) longitudinal flash marks extending onto the entire dorsal surface of thighs, generally margined by dark patches, observed in typical A. femoralis. A. hodli is distinguished from A. zaparo and A. myersi by the color of dorsum, which is uniformly black/ dark-brown in A. hodli (Fig. 2), but reddish in A. zaparo and brown to light-brown in A. myersi. Allobates myersi also lacks a pale dorsolateral stripe. A. hodli is largely sympatric to Ameerega picta, a dendrobatid frog that presents similar body size and color pattern. However, A. hodli can be distinguished from Ameerega picta by lacking a bright (orange to red) flash mark on calf region. Description of holotype. Morphological measurements of holotype are presented in Table 1. Body robust, head slightly wider than long (HL/HW = 0.94) (Fig. 3 A). Eye diameter slightly larger than distance from nostril to anterior corner of the eye. Nares located posterolaterally to tip of snout, directed posterolaterally, visible in ventral and anterior view. Center of nostril not visible dorsally. Canthus rostralis convex from tip of snout to nostril, straight from nostril to anterior corner of the eye. Loreal region vertical. Tympanum well visible, with maximum diameter horizontal, corresponding to 44 % the maximum diameter of the eye. Maxillary teeth present. Tongue length twice as large as wide, attached anteriorly on first third. Median lingual process absent. Choanae round. A single vocal sac is present, corresponding to most of the area of the medial and posterior subgular region. Vocal sac round when expanded. When retracted, vocal sac forms two lateral slits at the level of maxilla articulation (Fig. 3, B). Skin granular on dorsum and dorsal surface of legs. Granules round, more developed on dorsal surface of urostyle region and shanks. Skin smooth ventrally and laterally. Dermal flap above cloaca absent. Palmar tubercle slightly triangular. Thenar tubercle well-developed, oval to elliptic, maximum diameter 1.28 times smaller than maximum diameter of palmar tubercle. Subarticular tubercles of Fingers II, III and IV are round, small, never exceeding the width of phalanges. Subarticular tubercle of Finger I elliptic, 1.21 times larger than thenar tubercle in maximum diameter. Supernumerary tubercles absent. Carpal pad and metacarpal ridges absent on hands. No fringes or webbing on fingers. A distal tubercle on finger IV is weakly developed (Fig. 3 C). Finger I is slightly (1.08 times) longer than Finger II. Length of finger IV does not reach distal subarticular tubercle of finger III when fingers are pressed against each other. Relative lengths of fingers: IV Variation in type series. Morphological measurements of individuals constituting type series are presented in Table 1. Morphological characters described for the holotype apply to all individuals in type series, except for the following: Males slightly smaller (4.42 %, in average) than females. Head slightly longer than wide in males (HL/HW = 1.04) and females (HL/HW = 1.05) in average. Maximum diameter of tympanum corresponding to approximately half the maximum diameter of the eye in males and females (Table 1). Vocal sac and slits absent in females. Palmar tubercle round to slightly triangular. A distal tubercle on finger IV is present in 28 of a total 83 (34.1 %) inspected specimens, but is absent or weakly developed in the remaining 54 specimens (65.9 %). Color in life. Males and females do not present dimorphism in relation to color and color pattern. Dorsal surface of body solid black to solid dark-brown (Fig. 2 B). Lateral surface of body solid black. Dorsolateral line white, thinner than lateral line (Fig. 2 A, 2 E). When continuous with flash marks on thighs, dorsolateral line becomes reddish-orange on groin. Lateral line white. Gular region solid black to dark bluish-gray in males and females (Fig. 2 D). In males, vocal sacs usually with a paler bluish-gray color when inflated. Mid abdomen white with irregular black to dark-gray blotches or speckling, merging with solid dark color of gular region. Abdomen bright reddish-orange posteriorly, with dark irregular spots appearing marginally from lateral edges. Ventral surfaces of hind limbs also bright reddish-orange, sometimes with small marginal dark spots (Fig. 2 C, 2 D). Plantar surface of feet brown. Ventral surfaces of arms bright reddish-orange, with bright yellow flash marks extending from dorsal surface of upper arms. Black to dark-gray spot ventrally on upper arm, at the point of body insertion, continuous with gular region pattern. Dorsal surfaces of posterior and anterior limbs reddish to brick-brown (Fig. 2 B, 2 E). Dorsal and rear surfaces of thighs with irregular bright reddish-orange flash marks or patterning, same color as ventral surfaces of legs, with irregular black or darkbrown blotches or spots. Granules on dorsal surface of shanks usually darker than overall color of shanks. A yellow flash mark is present dorsally on upper arms, at the point of body insertion (Fig. 2 B, 2 E). The iris is evident, with metallic yellowish-brown pigmentation. Color in life of juveniles. Color of juveniles after metamorphosis is the same of adults. Dorsum and flanks are solid black to dark-brown, with dorsolateral and lateral lines white and conspicuous. Limbs generally reddish-brown. Bright yellow flash marks are present dorsally on the upper arms, and may reach the elbow. Dorsal surface of thighs with conspicuous longitudinal bright reddish-orange flash marks, lacking black or dark-brown blotches or spots (Fig. 2 F). Color in alcohol of holotype. Dorsum is solid black to dark-brown. A thin, pale white dorsolateral line is present, continuous from groin at hind limb insertion, over the orbit and nostril, to the tip of snout (Fig. 3 A). A pale white lateral line is present, broader than dorsolateral stripe, running from groin, over the insertion of arm, below nostril, to tip of upper lip. Lateral stripe is continuous on both sides of the body. Lateral surface between dorsolateral and lateral stripe solid black. Color of gular region and throat is solid black to darkbrown. Abdomen color is white with irregular black blotches or speckling. Abdomen color becomes solid black/dark-brown from chest towards the gular region. The black/dark-brown speckling over white background pattern is replaced posteriorly by a solid pale-tan pattern, continuous with the ventral color pattern of hind limbs (Fig. 3 B). Arms uniformly very pale-brown in dorsal view, paler on the axilla and carpal/metacarpal regions. Irregular dark blotches appear on dorsal surfaces of tarsus and fingers. Arms uniformly pale white to pale tan in ventral view, with a black patch (continuous with color pattern of gular region) on anterior surface of the arm. Surface of outer lateral edge of forearm and metacarpal region same color as dorsal surface of arms, extending laterally from elbow and reaching the palm of hands and ventral surface of fingers (Fig. 3 C). Legs are pale brown in dorsal view. Irregular pale, unpigmented patches, as well as irregular black blotches and spots are present on dorsal and rear surfaces of thighs (Fig. 3 A). The area immediately around vent is darker than the overall surface of thighs. Dorsal surfaces of shanks with darker granules. Inner dorsal surface of tarsal region is lighter than overall pattern of legs. Toes are generally darker than tarsal region in dorsal view. Ventral surface of legs is uniformly pale-tan with small brown spots appearing marginally from outer edges. Ventral surfaces of tarsal region and toes darker, same color as dorsal surface of legs (Fig. 3 B, 3 D). Tongue is cream-colored; large intestine (removed for the analysis of diet) is unpigmented. Testes are unpigmented. Color variation in type series. Color in alcohol described for the holotype apply to all individuals in type series, except for the following: Lateral stripe is usually continuous, but can be interrupted in some individuals, on one or on both sides of body. Gular region and throat solid black in females, solid black to dark-brown in males. Mature oocytes are pigmented, with black pigment concentrated on animal pole. Description of tadpoles. Tadpole measurements were obtained from four tadpoles in developmental stage 36 (Table 2). Tadpoles correspond to a lot under the same collection number, INPA-H 23693. The largest tadpole (TL = 24.7 mm, Fig. 4) was used for detailed description. Body is depressed, body width (6.0 mm) larger than body depth (4.8 mm), body length 16.1 mm. Snout nearly round, flattened anterodorsally in lateral view (Fig. 4 C). Tip of snout flattened anteriorly in dorsal view (Fig. 4 A). Nares small, directed anterolaterally, located 0.8 mm anterior to the eye, and 1.0 mm posterior to tip of snout. Nostrils narrowly spaced, distance between nostrils 0.9 mm. Eyes dorsal, directed dorsolaterally, 0.9 mm in maximum length, located 1.8 mm posterior to tip of snout. Distance between medial margins of the eyes is 1.4 mm. Spiracle single, sinistral, forming a free tube opening posterodorsally below body axis in lateral view, 5.0 mm posterior from tip of snout (Fig. 4 C). Vent tube medial, free, 0.9 mm in length, opening dextrally. Tail musculature reaches maximum depth (2.4 mm) approximately at the end of first third of tail length, and maximum width at body-tail insertion (2.6 mm). Ventral tail fin originates at body-tail insertion. Dorsal tail fin originates slightly posterior (0.8 mm) to body-tail insertion, and reaches maximum high 14.5 mm from tip of snout, corresponding to the region of maximum tail depth. At maximum depth of tail, depth of musculature is 1.4 mm, dorsal fin 1.5 mm and ventral fin 1.1 mm. Oral disc is positioned anteroventrally, emarginate laterally, transversely elliptical, 2.6 mm in transverse width. Anterior labium continuous with snout, 2.6 mm in length. Marginal papillae absent dorsally on anterior labium (gap 1.7 mm, 74 % of total anterior labium length), present only laterally, on its outer margins. Posterior labium free from body wall, 2.4 mm in length, with a single row of marginal papillae. All papillae with rounded tips (Fig. 5). Labial tooth row formula is 2 (2)/ 3 (1). Rows A- 1 and A- 2 with same length (2.1 mm), A- 2 with a large medial gap (0.6 mm). Rows P- 1, P- 2 and P- 3 with same length (2.0 mm), P- 1 presenting a very narrow medial gap (Advertisement and courtship calls description and variation. Advertisement calls of Allobates hodli consist in trills of calls formed by two whistle-like notes with ascending frequency modulation (Fig. 6 A, 6 C). Measurements of advertisement call characteristics of holotype and average values for 60 males from type series are presented on Table 3. The average maximum frequency of calls within type series is 3425.0 �� 184.7 Hz, and average duration of calls (summed durations of first and second notes, and inter-note silent interval) 0.164 �� 0.011 s. First note is less modulated (average difference between lower and higher frequency = 470.7 �� 94.8 Hz) than the second note (average difference between lower and higher frequency = 740.7 �� 115.0 Hz) and shorter in duration (0.033 �� 0.004 s, in comparison to second note, 0.056 �� 0.007 s). Courtship calls of A. hodli are quite distinct from their advertisement calls and are constituted by a continuous pulsed tone (Fig. 6 D), emitted only in the presence of females near the male���s calling perch. Average maximum frequency and duration of courtship calls obtained from calls of four individuals were 3190.2 Hz and 0.571 s, respectively (Table 4). The average pulse emission rate between calls was 125.2 pulses per second. Individuals Molecular phylogeny and genetic distances. From an initial sequence database containing 203 16 S rDNA sequences (including outgroups), a total of 93 unique haplotypes were used in the phylogenetic analysis (Table 5). Phylogenetic reconstructions support the existence of two basal clades within the Allobates femoralis group, both forming the sister clade to Allobates zaparo (Fig. 7 A). One of the basal clades contains Allobates hodli and the second contains samples from areas nearby Allobates femoralis type-locality, which we refer to as Allobates femoralis sensu stricto (clade femo 0 4, Fig. 7 B). A. hodli is marginally paraphyletic to populations that occur in the southern reaches of the Brazilian State of Acre (clade Acre 0 1, Fig. 7 B), which present advertisement calls constituted by four notes and color pattern more similar to that of A. femoralis than to that of A. hodli. Samples from the locality Monte Alegre were not clearly positioned within A. hodli or Acre 0 1 clade. This locality probably corresponds to a relictual contact zone between these clades. Allobates hodli and clade, Published as part of Sim��es, Pedro Ivo, Lima, Albertina P. & Farias, Izeni Pires, 2010, The description of a cryptic species related to the pan-Amazonian frog Allobates femoralis (Boulenger 1883) (Anura: Aromobatidae), pp. 1-28 in Zootaxa 2406 on pages 5-25, DOI: 10.5281/zenodo.194212, {"references":["Hodl, W., Amezquita, A. & Narins, P. M. (2004) The role of call frequency and the auditory papillae in phonotactic behavior in male dart-poison frogs Epipedobates femoralis (Dendrobatidae). Journal of Comparative Physiology A, 190, 823 - 829.","Amezquita, A., Hodl, W., Lima, A. P., Castellanos, L., Erdtmann, L. & Araujo, M. C. (2006) Masking interference and the evolution of the acoustic communication system in the Amazonian dendrobatid frog Allobates femoralis. Evolution, 60 (9), 1874 - 1887.","Lotters, S., Jungfer, K. - H., Henkel, F. W. & Schmitz, W. (2007) Poison Frogs. 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