Research Article |
Corresponding author: Frederico de Alcântara Menezes ( fred-alcantara@hotmail.com ) Academic editor: Robert Jadin
© 2018 Frederico de Alcântara Menezes, Arthur Diesel Abegg, Bruno Rocha da Silva, Francisco Luís Franco, Renato Neves Feio.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Menezes FA, Abegg AD, Silva BR, Franco FL, Feio RN (2018) Composition and natural history of the snakes from the Parque Estadual da Serra do Papagaio, southern Minas Gerais, Serra da Mantiqueira, Brazil. ZooKeys 797: 117-160. https://doi.org/10.3897/zookeys.797.24549
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The Serra da Mantiqueira is one of the least inventoried physiographic areas of southeastern Brazil. There is great potential for detection of endemic species for which little or nothing is known about basic aspects of natural history. The Parque Estadual da Serra do Papagaio (PESP) within the Serra da Mantiqueira is an area of extreme biological importance because it houses mixed formations of grasslands, ombrophilous forests, and enclaves of Araucaria forests (mixed ombrophilous forest). Currently, the mixed ombrophilous forest covers less than 5% of its original range and areas occupied by this forest type, and associated ecosystems constitute refuges, housing several endemic, high altitude species. Between September 2015 and April 2016, field samplings were performed in the PESP using four distinct methods. The objective was to determine the composition and natural history of snakes from an isolated, high altitude area of the Serra da Mantiqueira. In PESP and surrounding areas, 80 individuals representing 24 species, 19 genera, and three families were recorded. Data are presented on abundance, habitat, daily activity, diet, reproduction, and defense. Comparison of the PESP snake assemblage with 30 other Atlantic Forest areas in southeastern Brazil indicate the Serra da Mantiqueira presents particular characteristics regarding snake composition.
Araucaria Forests, Atlantic Rainforest, cluster analysis, Serpentes , southeastern Brazil
The structure of a snake community can be influenced by historical factors, such as biogeography (
Snake fauna of tropical areas is typically characterized by high species richness, low abundance, and complex ecological interactions (
The Serra da Mantiqueira is one of the least well-understood physiographic areas of southeastern Brazil. There is considerable potential for the record of remarkable and endemic species, for which basic natural history has yet to be described. To date only a single study reports on a snake assemblage in this region (
This study was conducted in the Parque Estadual da Serra do Papagaio (PESP) (22°8'33"S, 44°43'43"W, ca. 22.900 ha) located at Serra da Mantiqueira, southern Minas Gerais State, southeastern Brazil. The PESP overlaps the municipalities of Aiuruoca, Alagoa, Baependi, Itamonte and Pouso Alto within MG, and has an altitudinal range from 1200–2359 m above sea level (
Snake sampling lasted eight months, from September 2015 to April 2016. We sampled between altitudes 1600 and 2359 m a.s.l. across all of the available vegetation types. Field trips of four to seven days at a time were made monthly for a total of 40 days of field observation. Four different sampling methods were used to capture snakes:
1) Pitfall traps with drift fences (
2) Time-constrained search was also employed (
3) Accidental encounter (
4) Records made by local people (
For each specimen, we recorded: date and time of observation, habitat, microhabitat, mass (g), sex, diet, reproduction, activity, and defensive behavior. Snake size was categorized according to
We compared the snake assemblage from Parque Estadual da Serra do Papagaio with those of 30 other localities in the Atlantic forest, in southeastern Brazil (Fig.
With this data set, we generated a binary presence/absence matrix of 120 species. To compare snake assemblages, we used this matrix to run a Cluster Analysis, using the Jaccard’s similarity index, and the Pair Group Average Method (UPGMA) as the grouping method. We also calculated the cophenetic correlation coefficient to indicate the similarity matrix degree of representation in the dendrogram. In this index, values greater than or equal to 0.8 allow considering the dendrogram as adequate to the similarity matrix (
Finally, a Nonmetric Multidimensional Scaling (NMDS) was used for another view of the Jaccard index clusters. The stress value was used as a representative measure of the groupings, and values <0.20 were considered acceptable (Clarke and Warnick 1994). Multivariate analyzes were performed in R software (R Core Team 2014), using the vegan package (
Areas used for the analysis of similarity between snake assemblages.The following snake assemblages were included in the analyzes: São Paulo State: 1 Parque Estadual Turístico do Alto Ribeira (
Several nomenclature changes have been proposed to the taxonomy of Neotropical snakes in recent years. To provide consistency during similarity analysis we present a brief list of species that have had names changes and identify the name used in this report. The name “Taeniophallus gr. occipitalis” was used for specimens traditionally referred to as Taeniophallus occipitalis (Jan, 1863) because more than one species often fall under this designation (Santos-Jr. 2008). Specimens identified as Mussurana montana (Franco, Marques & Puorto, 1997) by
We recorded 80 snakes during the eight month period of fieldwork through all sampling methods. In this group we separated 24 species of 19 genera, of which 67% are dipsadids, 19% viperids, and 14% colubrids (Table
List of species found in the Parque Estadual da Serra do Papagaio, Minas Gerais, Brazil. Abbreviations: abundance (N), relative frequency of each species (f%), environment (1 - High altitude grassland, 2 - Rocky Field, 3 - Dense Montane Ombrophilous Forest, 4 - Mixed Ombrophilous Forest), habitat (aa - open area, bf - forest edge, da – disturbed areas, fl - forest, lo - lotic environment, le - lentic environment), and habits (F-fossorial, C-cryptozoic, SAQ-sub-aquatic, T-terrestrial, SA-sub-arboreal, and A-arboreal). Species registered outside the park area (*).
Family/ Species | N | F% | Environments | Habitat | Habit | Altitudinal variation (m) | New record altitudinal (m) |
---|---|---|---|---|---|---|---|
COLUBRIDAE | |||||||
Chironius bicarinatus (Wied, 1820) | 1 | 1.2 | 1,3 | bf, at | SA | 0–1850 | – |
Chironius brazili Hamdan & Fernandes, 2015 | 1 | 1,2 | 1,3 | Bf, fl, at | SA | 200–2030 | – |
Spílotes pullatus (Linnaeus, 1758) | 2 | 2.5 | 3 | fl | SA | 0–1100 | 1630 |
DIPSADIDAE | |||||||
Apostolepis assimilis (Reinhardt, 1861)* | 1 | 1.2 | 1 | Aa, at | F, T | 170–1610 | – |
Atractus zebrinus (Jan, 1962) | 15 | 18.7 | 1,2,3 | Aa, fl, at | F, T | 20–1610 | 1730 |
Boiruna maculata (Boulenger, 1896) | 1 | 1.2 | 3 | fl | T | 0–1880 | – |
Echinanthera cephalostriata Di-Bernardo, 1996 | 2 | 2.5 | 3 | Bf, fl | C, T | 0–1610 | 1730 |
Erythrolamprus miliaris (Linnaeus, 1758) | 1 | 1.2 | 3 | Fl, le | SAQ | – | 1643 |
Gomesophis brasiliensis (Gomes, 1918) | 8 | 10 | 1 | Aa, lo, le, at | SAQ | 430–1650 | 1750 |
Mussurana montana (Franco, Marques & Puorto, 1997)* | 1 | 1.2 | 4 | fl | T | 750–1610 | 1740 |
Oxyrhopus clathratus Duméril, Bibron & Duméril, 1854* | 1 | 1.2 | 1 | Aa, at | T | 0–1610 | – |
Oxyrhopus rhombifer Duméril, Bibron & Duméril, 1854 | 1 | 1.2 | 1 | Aa, at | T | 0–1330 | 1730 |
Philodryas aestiva (Duméril, Bibron & Duméril, 1854) | 2 | 2.5 | 1,2 | aa | T | 0–1730 | 1800 |
Philodryas patagoniensis (Girard, 1858) | 9 | 11.2 | 1,2 | Aa, at | T | 0–1600 | 2200 |
Sibynomorphus mikanii (Schlegel, 1837) | 1 | 1.2 | 1 | Aa, at | T | 110–1350 | 1630 |
Taeniophallus affinis (Günther, 1858) | 3 | 3.7 | 1,3 | aa | T, C | 0–1600 | 1760 |
Taeniophallus gr. occipitalis* | 1 | 1.2 | 1 | bf | T, C | – | 1600 |
Thamnodynastes strigatus (Günther, 1858) | 15 | 18.7 | 1,3,4 | Aa, fl, bf, at | SAQ | 0–2450 | – |
Tomodon dorsatus Duméril, Bibron & Duméril, 1854 | 2 | 2.5 | 3,4 | fl | T | 0–1610 | 1730 |
Xenodon merremii (Wagler in Spix, 1824) | 1 | 1.2 | 1,2 | aa | T | 0–1300 | 1610 |
VIPERIDAE | |||||||
Bothrops fonsecai Hoge & Belluomini, 1959 | 7 | 8.7 | 1,2,3,4 | Aa, bf, fl | T | 440–1730 | 2175 |
Bothrops jararaca (Wied, 1824) | 2 | 2.5 | 1,2,3,4 | Bf, fl | T | 0–1640 | 2150 |
Bothrops neuwiedi Wagler in Spix, 1824 | 1 | 1.2 | 1,2 | Aa, bf | T | 0–1600 | 2150 |
Crotalus durissus Linnaeus, 1758 | 1 | 1.2 | 1,2 | Aa, bf | T | 0–1400 | 1950 |
Total | 80 | 100 |
The cluster analysis (cophenetic correlation coefficient = 0.8381) based on 120 snake species recorded at 31 localities, including Parque Estadual da Serra do Papagaio, resulted in three main groupings (Fig.
Group 1 was composed generally by localities of mid to high altitudes (> 600 m): the coastal forests of Serra do Mar and Bahia Interior Forests (
Group 2 consisted of low areas (< 400 m a.s.l.) of the Coastal Forests of Serra do Mar, comprising island and continental regions in the coastal strip of São Paulo and Rio de Janeiro states. In this group, three subgroups can also be observed. The first (7) consists of two continental islands on the south coast of São Paulo state (Ilha do Cardoso and Ilha Comprida). The second (8) composed by the other insular and lowland locations of the Atlantic Forest in São Paulo state, in addition to an island in Rio de Janeiro State (Ilha Grande). Geographically, this subgroup is close to the Parque Estadual da Serra do Mar (Picinguaba, São Paulo State), with which it shares several snake species. The third (9) is composed of lowland locations in the state of Rio de Janeiro, east of Serra dos Órgãos, for which similarities have already been described by
Group 3 is composed of only two high altitude localities (> 1100 m a.s.l.) in the southern Mantiqueira Mountains: the Parque Estadual da Serra do Papagaio and Munhoz.
The NMDS ordering analysis (stress 0.1547) graphically depicts the relative position of localities in a two-dimensional space (Fig.
Medium-sized snake (n = 1), diurnal and semi-arboreal (
From sea level, from the northern coast of Rio Grande do Sul to Bahia, to a maximum altitude of 1610 m in Campos do Jordão, SP (
Northeast, central-west, southeast and south of Brazil (Bahia, Goiás, Mato Grosso do Sul, Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina and Rio Grande do Sul), Argentina, and Uruguay (
Medium-size species (n = 1), diurnal and semi-arboreal (
The maximum altitudinal record for the species was at 2030 m a.s.l. at Pico do Inficionado, Catas Altas, MG (
Central-west, southeast and south of Brazil (Federal District, Goiás, Minas Gerais, São Paulo and Rio Grande do Sul) (Hamdan and Fernandes 2015). This species is thought to live in habitats similar to those of C. flavolineatus and inhabits riverine forests and forest borders near open areas (Hamdan and Fernandes 2015).
Snakes from the Serra do Papagaio. A Chironius brazili B Apostolepis assimilis C Gomesophis brasiliensis D Philodryas aestiva E Philodryas patagoniensis F Thamnodynastes strigatus G Tomodon dorsatus H Xenodon merremii I Bothrops fonsecai J Bothrops jararaca K Bothrops neuwiedi L Crotalus durissus. Photographs by Mário Sacramento, Frederico de Alcântara Menezes, Arthur Diesel Abegg, and Leonardo Chaves.
A large species (n = 1), with semi-arboreal habits, and diurnal activity (
In Brazil, range spans at minimum of sea level from the coast of Santa Catarina to Bahia to a maximum of 1100 m a.s.l., Brasília, Federal District (
This species can be found in all Brazilian states, and in Argentina and Paraguay (
Species of small size (n = 1), with nocturnal activity and cryptozoic or fossorial habits (
This species was found at a minimum of 170 m a.s.l., in Cuiabá, Mato Grosso and maximum of 1610 m a.s.l. in Campos do Jordão, SP (
Northeast, central-west, southeast and southern Brazil (Bahia, Mato Grosso, Mato Grosso do Sul, Goiás, Distrito Federal, Minas Gerais, São Paulo and Paraná), southeast Paraguay, northern Argentina, and Bolivia (
Species of small size (n = 15), nocturnal, with fossorial habits (
Found at a minimum of 20 m a.s.l. in Itaboraí, RJ and maximum of 1610 m a.s.l. in Campos do Jordão, SP (
South and southeast of Brazil (Santa Catarina, Paraná, São Paulo, Rio de Janeiro, Minas Gerais and Espírito Santo) (
A large species (n = 1), terrestrial (
Species records indicate a minimum of sea level in Rio Grande do Sul and maximum of 1240 m a.s.l. in Serra do Salitre, MG (
North, central-west, southeast and south of Brazil (Amazonas, Distrito Federal, Goiás, Mato Grosso do Sul, Minas Gerais, Rio Grande do Sul, São Paulo), Argentina, Paraguay, and Uruguay (
Species of small size (n = 2), predominantly diurnal, terrestrial and cryptozoic (
Found at altitude minimum of sea level from Santa Catarina coast to Rio de Janeiro and maximum of 1610 m a.s.l. in Campos do Jordão, SP (
Northeast, southeast, and southern Brazil (Bahia, Espírito Santo, Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina and São Paulo) (
Species of medium size (n = 1), predominantly diurnal and semi-aquatic (
We found no information in the literature regarding the altitudinal variation of the species. In this study, the maximum altitudinal record was at 1643 m a.s.l., in Baependi-MG.
Northern, northeast, central-west, southeast and southern Brazil (Alagoas, Amapá, Amazonas, Bahia, Ceará, Espírito Santo, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, Pernambuco, Piauí, Rio de Janeiro, Rio Grande do Norte, Rio Grande Do Sul, Rondônia, Santa Catarina, São Paulo, Sergipe) (
Species of small size (n = 8). This species is considered nocturnal and aquatic, associating with lentic watercourses (
Found at a minimum of 430 m a.s.l. in Encruzilhada do Sul, RS and maximum of 1650 m a.s.l. in the Parque Estadual da Serra do Papagaio, Alagoa, MG. The maximum altitudinal record derives from the same are in this study, where most individuals were recorded at 1750 m altitude, in the PESP, Baependi, MG. The toponyms obtained for this species occur in two altitudinal ranges: 51% are located between 430 and 800 meters and 49% above this range (
This species occurs in natural field areas (
A large species (n = 1), terrestrial (
This species was found at a minimum of 750 m a.s.l. in Guaratinguetá, SP and maximum of 1610 m a.s.l. in Campos do Jordão, SP (
Southeast Brazil (Minas Gerais, Rio de Janeiro, and São Paulo) (
Species of medium size (n = 1), terrestrial and nocturnal (
The species is found at a minimum of sea level from the coast of Rio Grande do Sul to Rio de Janeiro, and a maximum of 1610 m a.s.l. in Campos do Jordão, SP (
Northeast and southeast Brazil (Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo), and Argentina (
Species of medium size (n = 1), nocturnal and terrestrial (
This species is found at a minimum of sea level in Argentina, Uruguay, Rio Grande do Sul, and south of Santa Catarina, and at a maximum of 1330 m a.s.l. in Liberdade, MG (
Northeast, central-west, southeast and southern Brazil (Bahia, Ceará, Distrito Federal, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Rondônia, Santa Catarina and São Paulo), Argentina, Bolivia, Paraguay and Uruguay (
Species of medium size (n = 2), diurnal, often found on the ground in open areas (
This species was found at a minimum of sea level from the coast of Uruguay to Santa Catarina and maximum of 1730 m a.s.l. in Campos do Jordão, SP (
Northeast, central-west, southeast and southern Brazil (Bahia, Distrito Federal, Goiás, Mato Grosso do Sul, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, and São Paulo), Argentina, Bolivia, Paraguay and Uruguay (
Species of medium size (n = 9), diurnal and terrestrial (
In September, an adult was observed at 14:00 h near a ravine, while it was being attacked by two different birds (Poospiza sp. and an unidentified Passeriformes), possibly in defense of a nearby nest. In July, an adult was observed at 15:00 h, while it was ingesting a rodent. Out of the four examined specimens, two presented rodents in their stomach.
This species was found at a minimum of sea level from the coast of Argentina to the state of Espírito Santo and maximum of 1660 m a.s.l. in Umuarama, Campos do Jordão, SP (
North, northeast, central-west, southeast and southern Brazil (Bahia, Distrito Federal, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Rondônia, and São Paulo), Argentina, Bolivia, Chile, Paraguay and Uruguay (
Species of small size (n = 1), nocturnal and terrestrial (
This species was found at a minimum of 110 m a.s.l., in Puerto Bemberg, Iguazú, Argentina and maximum of 1350 m a.s.l. in the Serra do Ouro Branco, Ouro Branco, MG (
Northeast, central-west, southeast and southern Brazil (Bahia, Goiás, Maranhão, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, Norte and São Paulo), Argentina and Paraguay (
Species of small size (n = 3), diurnal, terrestrial and cryptozoic (
This species is found at a minimum of sea level from the coast of Rio Grande do Sul to Rio de Janeiro and maximum at 1600 m a.s.l. in Parque Estadual Ibitipoca, Lima Duarte, MG (
Northeast, southeast and southern Brazil (Alagoas, Bahia, Ceará, Espírito Santo, SE Minas Gerais, Paraná, Rio de Janeiro and Rio Grande do Sul, Santa Catarina, São Paulo) (
Species of small size (n = 1), diurnal and cryptozoic (
In this study, the maximum record was at 1600 m a.s.l., in the Aiuruoca, MG.
North, northeast, central-west, southeast and southern Brazil (Bahia, Ceará, Distrito Federal, Goiás, Pará, Paraíba, Paraná, Piauí, Rio Grande do Sul, Rondônia, São Paulo and Sergipe), Argentina, Bolivia, Paraguay, Peru and Uruguay (
Species of medium size (n = 15), nocturnal and terrestrial (
This species was found at a minimum of sea level and maximum of 2450 a.s.l. in Itatiaia National Park, state of Rio de Janeiro, Brazil (
Southern, southeast, and northern Brazil (Espírito Santo, Minas Gerais, Pará, Paraná, Rio de Janeiro, Santa Catarina, Rio Grande do Sul, Roraima and São Paulo), Paraguay, Uruguay and Argentina (
Species of medium size (n = 2), diurnal and terrestrial (
This species is found at a minimum of sea level from the coast of Rio Grande do Sul to Rio de Janeiro, and a maximum of 1610 m a.s.l. in Campos do Jordão, SP (
Central-west, southeast and southern Brazil (Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina, and São Paulo), Argentina, Paraguay and Uruguay (
Species of medium size (n = 1), diurnal and terrestrial (
This species is found at a minimum of sea level from the northern coast of Rio Grande do Sul to the extreme south of Santa Catarina, São Paulo, Rio de Janeiro, Espírito Santo, and Bahia. Maximum altitude recorded is 1300 m a.s.l. in the Parque Estadual de Itacolomi, Ouro Preto, MG (
North, northeast, central-west, southeast, and southern Brazil (Bahia, Brasília, Ceará, Goiás, Mato Grosso, Pará, Paraiba, Paraná, Pernambuco, Rondônia, São Paulo, and Tocantins), Bolivia and Paraguay (
Species of medium size (n = 7), nocturnal and terrestrial (
This species is found at a minimum of 400 m a.s.l. in Barra Mansa, RJ and a maximum of 1730 m a.s.l. in Campos do Jordão, SP (
Southeastern Brazil (Minas Gerais, Rio de Janeiro, and São Paulo) (
A species of medium size (n = 2), semi-arboreal and mainly nocturnal (
This species was found at a minimum of sea level between Rio Grande do Sul and Bahia with a maximum of 1640 m a.s.l. in Parque Nacional da Serra da Bocaina, SP (
Central-west, northeast, southeast, and southern Brazil (Bahia, Espírito Santo, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, São Paulo, and Santa Catarina), Paraguay and Argentina (
Medium-sized snake (n = 1), terrestrial and nocturnal (Marques et al. 2016). We spotted an adult during the day (12:50 h) in March, in a rocky field area. It was basking near a forest fragment at 2150 m a.s.l. When the observer approached, it fled into the forest. The available data in the literature indicates it was found mainly in fields and other open formations (Borges and Araújo 1998,
The neuwiedi complex species was found at a minimum of sea level in the coast of Rio de Janeiro State, and a maximum of 1600 m a.s.l. in Parque Estadual do Ibitipoca, MG (
Northeast, central-west, southeast and southern Brazil (Bahia, Goiás, Minas Gerais, Paraiba, Paraná, Rio de Janeiro, São Paulo and Santa Catarina) (
Species of medium size (n = 1), with terrestrial and nocturnal habits (
This species is found at a minimum of sea level for the coasts of Argentina, Uruguay, Rio Grande do Sul and Bahia and maximum of 1400 m a.s.l. at Taquaral Farm, Paraty, RJ (
Southern and southeastern Brazil (Amapá, Amazonas, Bahia, Ceará, Goiás, Mato Grosso, Mato Grosso do Sul, Maranhão, Minas Gerais, Pará, Paraíba, Paraná, Pernambuco, Piauí, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina, São Paulo), Netherlands Antilles (Aruba), Guyana, Suriname, French Guiana, Peru, Colombia, Venezuela, Uruguay, Bolivia and Paraguay (
The 24 snake species recorded in this study correspond to approximately 11% of the 219 species known from the phytogeographical domain of the Atlantic Forest (
In any study, the relative frequency of snakes may be influenced by the sampling method (
In contrast to other Neotropical snake community studies (
The high species richness registered for the PESP might be related to environmental heterogeneity and the mix of habitats allowing more species of reptiles to coexist (
The species composition of the Parque Estadual da Serra do Papagaio is very similar to the Munhoz, which is also in southern Serra da Mantiqueira. These areas have very similar vegetation types. Both are present in the Alto Paraná Atlantic Forest, which contains rocky fields, savannahs, and montane forests (sensu
The other two main groups are composed of plateau areas (1) in the states of São Paulo and Minas Gerais (except localities in southern Serra da Mantiqueira); and (2) lowland areas and in São Paulo and Rio de Janeiro states. The formation of a cluster containing low coastal localities has already been described for anurans (
The Serra da Mantiqueria is defined as an area of particular biological importance, and it stands out in the region as one of the most important areas for biodiversity conservation in the state of Minas Gerais. This status is justified by the high abundance of rare and threatened species (
The dipsadid Mussurana montana is another species of interest found in this area. This species is endemic to a region that covers about 16,000 km2 in the mountainous portions of São Paulo, Minas Gerais, and Rio de Janeiro states (
The present study provides data on the occurrence and natural history of snakes in the Parque Estadual da Serra do Papagaio. However, additional field research is encouraged. New inventories could contribute additional knowledge about the natural history of the species of this region, especially those of which ecological, biological, and morphological data are scarce, such as Mussurana montana. Our ordination analysis shows a remarkable difference between the compositions of snake faunas between different altitudinal gradients in the Brazilian Atlantic Rainforest. The Serra da Mantiqueira, where the Parque Estadual da Serra do Papagaio is located, seems to present a very unique composition of snake species, most of which are not shared with others localities from the southeast of Brazil. The diverse vegetation types found in this region permits the existence and maintenance of diverse species of animals. Additionally, the protected areas belonging to the Atlantic Forest affords greater potential to find rare and endemic species. These facts together make an argument for prioritizing the conservation of this park. Nevertheless, even if recognized as a conservation unit, the park continues to suffer depredation of its flora and fauna, mainly through illegal hunting. Regrettably, this alarming scenario is widely observable in Brazil due to the scientific scrapping and the neglect of the Brazilian government to control its natural resources (
1 | Loreal pit present; solenoglyphous dentition | Viperidae |
– | Loreal pit absent, aglyphous or opisthoglyphous | 2 |
2 | Aglyphous dentition; even number of dorsal scale rows | Colubridae |
– | Aglyphous or opisthoglyphous dentition; odd number of dorsal scale rows | Dipsadidae |
1 | Rattle at the tip of the tail; with some enlarged shields on top of the head | Crotalus durissus |
– | No rattle at the tip of the tail; tiny shields on top of the head | 2 |
2 | Inverted v-shaped spots along the dorsum; second supralabial fusioned with the prelaculal | Bothrops jararaca |
– | Trapezoid spots along the dorsum; second supralabial not fused with the prelaculal | 3 |
3 | Non-fragmented trapezoid spots along the dorsum; black venter | Bothrops fonsecai |
– | Trapezoid spots are fragmented at the midline to ventrals; venter cream, with several tiny brown spots | Bothrops neuwiedi |
1 | 12 scale rows at mid-body | 2 |
– | More than 12 scale rows at mid-body | Spilotes pullatus |
2 | 2-4 rows of keeled dorsal scales; first third of body black or dark gray, vertebral stripe yellowish or cream; top of the head tan to brown (distinct from the background color of the body); venter cream, with black bordered scales | Chironius brazili |
– | Two rows of keeled dorsal; olive green at the first third of the body; head color similar to the body; venter yellow, without black bordered scales | Chironius bicarinatus |
1 | 17 or fewer scale rows at mid-body | 2 |
– | 19 scale rows at mid-body | 10 |
2 | 15 scale rows at mid-body | 3 |
– | 17 scale rows at mid-body | 5 |
3 | Internasal shields absent; body almost uniformly red, with black tail | Apostolepis assimilis |
– | Internasal shields present; without red color on the dorsum | 4 |
4 | Black eyes, indistinguishable pupil; venter white, with black spots | Sibynomorphus mikanii |
– | Eyes with brown background color and round pupil, easily distinguishable; venter immaculate yellow | Taeniophallus gr. occipitalis |
5 | Dark lining, loreal shield absent | Tomodon dorsatus |
– | Light lining, loreal shield present | 6 |
6 | Dorsal color brown, with a lighter longitudinal line on each side; three supralabials in contact with the eyeball | Gomesophis brasiliensis |
– | Coloration not as above; two supralabials in contact with the eyeball | 7 |
7 | Dorsal scales with black borders and light centers; venter cream yellow, base of ventral scales with black edges | Erythrolamprus miliaris |
– | Coloration not as above | 8 |
8 | Stout and short body; small eyes | Atractus zebrinus |
– | Slender and elongate body; large eyes | 9 |
9 | Fourth row of dorsal scales with white dots that, together, form a continuous line along the body; brown top of the head and body; ventral scales with transverse band | Echinanthera cephalostriata |
– | The fourth row of dorsals without white dots; top of the head black, contrasting with the brown body; ventral scales without transverse band | Taeniophallus affinis |
10 | Two apical pits | 11 |
– | Single apical pit | 14 |
11 | Red iris | 12 |
– | Black or dark brown irish, never red | 13 |
12 | Juveniles are brick red, with a dark brown longitudinal vertebral stripe; adults are entirely dark brown | Mussurana montana |
– | Coral color pattern, with black triangular spots, bordered with white, background color red | Oxyrhopus rhombifer |
13 | Loreal shield usually absent; contact between frontal and preocular absent | Oxyrhopus clathratus |
– | Loreal shield present; contact between frontal and preocular present | Boiruna maculata |
14 | Keeled dorsal scales; dorsum uniform green | Philodryas aestiva |
– | Smooth dorsal scales; dorsum never green | 15 |
15 | Canthus rostralis very evident; without postocular stripe | Philodryas patagoniensis |
– | Canthus rostralis not evident; conspicuous postocular stripe | 16 |
16 | With two large post-diastemal fangs, aglyphous; venter cream, without longitudinal lines | Xenodon merremii |
– | Without two large post-diastemal fangs, but opisthoglyphous; venter cream, with two to four longitudinal lines | Thamnodynastes strigatus |
The authors are grateful to Instituto Butantan for the infrastructure offered, Instituto Estadual de Florestas (IEF) for permission to work at the park, Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis (IBAMA) for authorizing snake capture (47749–3), CAPES and Christiane Sarkis de Alcântara for financial support; Emmanuel M. J. Landroz, Marcos Lopez Menezes and all his family, Roseli, Darci, Rita and Marcilei Lopes Menezes for fieldwork assistance; Renato S. Bérnils, for the valuable comments throughout the development of the study.