Constrictor constrictor constrictor Boa constrictor constrictor Constrictor constrictor constrictor Charina bottae Corallus hortulana Epicrates cenchria Eunectes murinus B . occidentalis B . atlantica B . constrictor B . amarali amarali constrictor constrictor B . atlantica B . occidentalis Boa constrictor Boa constrictor constrictor Boa constrictor Boa constrictor constrictor Boa constrictor Boa constrictor constrictor Boa constrictor The good, the bad and the boa: An unexpected new species of a true boa revealed by morphological and molecular evidence Gonzalez, Rodrigo Castellari Lima, Lorena Corina Bezerra de Passos, Paulo Silva, Maria José J. PLoS ONE 2024 e 0298159 2024-04-17 19 4 1 29 Gonzalez & Lima & Passos & Silva 2024 [565,712,1664,1688] Squamata Boidae Boa Animalia 9 10 Chordata species atlantica sp. nov.    urn:lsid:zoobank.org: act: 242E539F-CF4C-4C0B-A5E1-D4E143BE82A5.     Constrictor constrictor constrictor–Ihering, 1911 [ 63] (part.); Amaral, 1930 [ 64] (part.); Stull, 1932 [ 65] (part.); Stull, 1935 [ 66] (part.);    Boa constrictor constrictor–Forcart, 1951 [ 27] (part.); Stimson, 1969 [ 67] (part.); Peters & Orejas-Miranda, 1970 [ 68] (part.);    Constrictor constrictor constrictor–Amaral, 1977 [ 69] (part.);   Fig 2. Phylogenetic tree inferred by using Maximum Likelihood and general time reversible model.The matrix is composed of 39 samples from South America, being 35 samples with the four concatenated markers (cyt- b, ND4, NTF3, and ODC resulting in 2305 positions in the final dataset) and the outgroup (  Charina bottae,  Corallus hortulana,  Epicrates cenchria, and  Eunectes murinus) with cyt bonly). The percentage of trees in which the associated taxa clustered together is shown above the branches (bootstrap). The tree is drawn to scale, with branch lengths measured in the number of substitutions per site. Group colours: Black: Outgroup; blue:  B. occidentalis; green:  B. atlantica  sp. nov.; orange:  B. constrictorand red:  B. amarali. Photographs: A) B. c.  amarali, no locality; B. c.  constrictor, no locality, Brazil; C) B. c.  constrictor, from Cuiabá, Mato Grosso, Brazil;  B. atlantica sp. nov., from Rio de Janeiro, Brazil;  B. occidentalisfrom Tucumán, Argentina. https://doi.org/10.1371/journal.pone.0298159.g002    Boa constrictor–Henderson et al., 1995 [ 6] (part.);    Boa constrictor constrictor–McDiarmid et al., 1999 [ 25] (part.);    Boa constrictor–Marques et al., 2001 [ 70]; Argôlo, 2004 [ 71]; Tipton, 2005 (part.) [ 72];  Boa constrictor constrictor–Bonny, 2007 (part.) [ 3];    Boa constrictor–Pontes & Rocha, 2008 (part.) [ 73]; Hynkovà et al. 2009 [ 30] (part.); Reynolds et al., 2014 [ 35] (part.); Wallach et al., 2014 [ 74] (part.); Card et al., 2016 [ 11] (part.);  Boa constrictor constrictor–Reynolds & Henderson, 2018 [ 34] (part.);  Boa constrictor–Nogueira et al., 2019 [ 9] (part.).     Holotype. Brazil: Rio de Janeiro, Fundação Osório, Rio Comprido, Rua Paula Ramos, 52( 22˚56’03.0"S 43˚12’36.9"W, datum WGS84;  93 mabove sea level, hereafter asl), MNRJ 27242, adult male, collected by Sergeant Marco Aurélio da Silvaon  13 September 2019. The holotype was collected on a tree, while copulating with the paratype MNRJ 27243.   Paratypes(n = 47).All from Brazil:  Alagoas State: Passo do Camaragibe( 9˚16’28.5"S, 35˚ 28’04.8"W),  44 masl, MNRJ 3940, female, collected by H. Silvaat Passo on  18 January 1988;  Bahia State: Ilhéus, CEPEC-CEPLAC( 14˚45’21.6"S, 39˚13’53.4"W),  58 masl, CZGB 4862, female, collected by G.A. Costaon  10 July 1996;  Salobrinho, Ilhéus( 14˚47’S, 39˚10’W),  29 masl, MNRJ 6362, male, collected by S. Rangelin  September 1987;  Ituaçú( 13˚48’S, 41˚18’W),  539.95 masl, MNRJ 6361, female, collected by U. Caramaschi& H. Niemeyeron  07 August 1997;  Espírito Santo state: Cachoeiro do Itapemerim( 20˚51’00.4"S, 41˚06’42.9"W),  35 masl, IBSP 79063, female, collector unknown, collected on  10 January 2011;  Setiba, Guarapari( 20˚ 37’S, 40˚26’W),  20 masl, MNRJ 23361, female, collected by T.S. Soares, date of collection unknown and  MNRJ 24903, male, collected by C.F.D. Rochaon  15 November 1999;  Reserva Biológica de Comboios, Linhares( 19˚33’S, 40˚03’W),  13 masl, MNRJ 23879, male, collected by A.P. Almeidaon  31 August 2007;  São Mateus, Campus CEUNES/UFES( 18˚40’S, 39˚51’W),  8 masl, MNRJ 23882, male, collected by R.S. Bérnilson  15 March 2011;  Vitória, Morro da Gamela( 20˚17’50.1"S, 40˚18’05.5"W),  17 masl, MNRJ 9565, male, collected by J.L. Gasparini, & F. Campagnollion  14 January 2002;  Rio de Janeiro state: Cabo Frio( 22˚50’S, 41˚59’W),  6 masl, MNRJ 22936, female, collected by R.R. Pintoet al on  03 May 2012;  Carapebus, Parque Nacional da Restinga de Jurubatiba( 22˚15’S, 41˚39’W),  8 masl, MNRJ 26802, male, collected by D.S. Fernandeset al. on the  8 March 2018;  Guapimirim( 22˚34’S, 43˚0’W)  23 masl, MNRJ 14238, female, collector unknown, collected in  May 2001;  Guapimirim( 22˚34’S, 43˚0’W)  23 masl, MNRJ 14250, female, collector unknown, collected in  May 2001;  Iguaba Grande( 22˚50’S, 42˚10’W),  37 masl, MNRJ 17353, female, date of collection and collector unknown;  Itaboraí, Complexo Petroquímico do Rio de Janeiro (COMPERJ)( 22˚39’S, 42˚51’W),  21 masl, MNRJ 25057, female, collected by J. Creusenon  03 June 2013and  MNRJ 26324, male, collected on  28 July 2016;  Itaboraí( 22˚39’S, 42˚51’W),  134 masl, MNRJ 25413, female, date of collection and collector unknown;  Maricá( 22˚55’S, 42˚51’W),  12 masl, MNRJ 13111, female, collected by C. L. Prataon  12 May 2005;  Itaipu, Niterói( 22˚57’S, 43˚02’W),  7 masl, MNRJ 11205, female, collected by F. Vieira, J.V. Camargo, & M.A. Gonçalvesin  May 2004;  Pendotiba, Niterói( 22˚54’S, 43˚04’W),  111 masl, MNRJ 23573, female, collected by M.S.C. Mesquitaon  19 January 2013;  Niterói( 22˚57’S, 43˚02’W),  2 masl, MNRJ 16922, female, collected on  17 July 2008, by R.W. Kisling;  Nova Iguaçu( 22˚42’S, 43˚28’W),  34 masl, MNRJ 24860, male, collected by A. Antuneson  13 June 2013; and MNRJ 26213, female, collector unknown, collected in 2014;  Porciúncula, Fazenda Vargem Alegre( 20˚58’S, 42˚02’W),  231 masl, MNRJ 14172, female, collected by B. Pimentaon  08 March 2006;  Rio das Ostras( 22˚30’S, 41˚56’W),  31 masl, MNRJ 10117, female, collected by D.S. Fernandeset al. on  17 November 2003;  Rio de Janeiro, Aeroporto Internacional Tom Jobim( 22˚48’S, 43˚15’W),  7 masl, MNRJ 25952, female, collected by J.T. Baldinein  November 2015and  MNRJ 26350, female, collector unknown, collected on  9 May 2016;  Água Santa( 22˚54’19.0"S, 43˚18’26.1"W),  221 masl, MNRJ 25950, female, collected by D.S. Fernandes, B. Miranda, & P. Pinna  09 December 2015;  Bairro Jardim Botânico, Horto Grotão( 22˚57’S, 43˚14’W),  650 masl, MNRJ 19740, male, collected by L. Caetanoon  13 July 2010;  Cosme velho( 22˚56’S, 43˚11’W),  18 masl, MNRJ 19412, male, collected by J.P. Pombalon  05 April 2010and  MNRJ 19564, male, collected on  18 May 2010;  Del Castilho( 22˚52’S 43˚16’W),  22 masl, MNRJ 25953, female, collected by R. Baptistaon  30 January 2016;  Estrada do Rio Morto( 23˚00’S, 43˚29’W),  3 masl, MNRJ 23144, female, by R. L. Santos, date of collection unknown;  Estrada dos Mananciais( 22˚55’S, 43˚22’W),  21 masl, MNRJ 26589, female, collected by D. B. Santoson  21 April 2017;  Ilha do Governador( 14˚47’52.6"S, 39˚10’35.3"W),  15 masl, MNRJ 9449, female, collector unknown, collected on  21 March 2001and  MNRJ 26585, female, collected by A. Carneiroon  6 June 2017;  Parque Nacional da Tijuca( 22˚56’S, 43˚ 17’W),  360 masl, MNRJ 26886, female, collected by C. Buenoon  13 March 2018and  MNRJ 27262, female, same collector, collected on  29 February 2019;  Recreio dos Bandeirantes( 23˚ 01’S, 43˚28’W),  14 masl, MNRJ 14200, female, collected on by J.R. Gomes  27 May 2006and  MNRJ 14201, female, same collector, collected on  13 May 2006;  Serra do Mendanha( 22˚50’S, 43˚29’W),  190 masl, MNRJ 17547, female, collected by J.A.L. Pontes, date of collection unknown;  Rio de Janeiro( 22˚57’S, 43˚18’W),  10 masl, MNRJ 10092, male, collected by M. Mocelin, date of collection unknown and  MNRJ 26796, male, collector unknown, collected on  22 August 2017;  São Francisco de Itabapoana( 21˚14’03.0"S, 41˚07’19.0"W),  4 masl, MBML 2097, male, collected by G.L. Forrequeon  10 October 2006;  Teresópolis, road BR- 116 km86.5( 22˚24’S, 42˚58’W),  972 masl, MNRJ 20700, female, collector unknown, collected on  25 July 2011;  Fundação Osório, Rio Comprido, Rua Paula Ramos, 52( 22˚56’03.0"S 43˚12’36.9"W, WGS84),  93 masl, MNRJ 27243, adult female, collected by Sgt. Marco Aurélio da Silvaon  13 September 2019[copulating with the holotype MNRJ 27242].   Diagnosis.  Boa atlantica sp. nov.can be distinguished from the other congeners by the following unique combination of characters: (i) ventrals 228–243; (ii) subcaudals 47–58 inmales, and 31–56 infemales; (iii) anterior dorsal scale rows 51–66; (iv) midbody dorsal scale rows 69–90; (v) posterior dorsal scale rows 42–54; (vi) dorsal body spots 17–23; (vii) tail spots 4–6 inmales, 2–6 infemales; (viii) longitudinal head stripe usually continuous; (ix) head stripe without lateral projections; (x) elliptical, circular or double-oval dorsal interspots; (xi) posterior dorsal interspots not blotched; (xii) no change in dorsal spots towards the tail; (xiii) lateral ocelli dark brown, black or faint reddish, white bordered; (xiv) belly cream with tones of orange, brown and black, scattered of black dots and large groups of black spots; (xv) belly with progressive darkening towards the tail; (xvi) black spots on ventral surface of tail.   Etymology.The species epithet  atlanticais a Latin adjective that refers to the Atlantic coast but mainly the Atlantic Forest ecoregion, the homeland of several endemic species, including this new  Boa. The preservation of this biome is crucial for conservation, nonetheless, it has been suffering deforestation since the colonial times and only 12% of it remains standing. This situation is under serious threat since the past Brazilian policies lack empathy for conservation issues and seem to foster deforestation even further.   Description of the holotype.MNRJ 27242 ( Figs 3and 4), adult male, SVL 2016 mm, TL 294 mm, head width 27.5 mm, head length 77.9 mm, head height 27.5 mm, distance between eyes 22.9 mm, eye-rostral distance 26.3 mm, eye-mouth distance 8.2 mm, distance between nostrils 12.3 mm, cloacal spur length 7.3/ 6.8 mm, eye height 6.2 mm, eye length 6.5 mm, scales over the preocular stripe 9/7, circumorbitals 17/15, suboculars 1/2, supralabials 22/21, infralabials 26/26, scales between nostrils 6, intrasupraoculars 17, gulars 18/17, dorsal scale rows 64/ 90/50, tail dorsal scale rows 24, preventrals 2, ventrals 245, subcaudals 57. Dorsum of head light brown light brown with discrete dark grey speckles; longitudinal head stripe brown, bordered by dark brown and black, extending from internasal region, breaking in occipital region, and continued up to cervical region; lateral head surface light brown; preocular stripe faded brown; and subocular stripe brown marginally reaching supralabials; postocular stripe dark brown, connecting with first dorsal saddle; some supralabials with dark brown dots; head ventral surface cream, with grey and salmon speckles and black dots; black dots unite forming two larger pairs of infralabial spots, and another three pairs of gular spots; dorsal background light brown progressively darkens towards tail; saddle-shaped dorsal spots, 22, laterally connected to each other, delimiting oval spots, dark brown in 2/3 of body, closer to each other and brownish-red before tail; body, lateral surface 2/3 of body greyish cream with salmon speckles and dark brown ocelli, encircling creamish white spots; last body third cream, with black speckles and irregular black ocelli encircling brick-red spots; ventral body surface cream, with black speckles and larger black blotches united on the sides, almost in alternate way; salmon tones in the midline of belly; dorsal background of tail yellow with five brick-red spots, bordered by black; first three spots with a pair of yellowish ocelli; ventral surface of tail cream, with alternate black spots among lateral sides of subcaudals not contacting each other.   Fig 3.Dorsal (A), ventral (B), and lateral (C) views of the head of the holotype of  Boa atlantica  sp. nov.(MNRJ 27242) from Rio de Janeiro, Atlantic coast of Brazil. https://doi.org/10.1371/journal.pone.0298159.g003 Hemipenis of the holotype: fully everted and maximally expanded hemipenis renders a bilobed, non-capitate, and non-calyculated organ; lobes relatively short and sub-cylindrical with rounded apices, similar size and oriented centrifugally; lobes covered by 6 transversal flounces on its basal portion; lobes naked from median to apical region; sulcus spermaticus divides on distal portion of organ; sulcus spermaticus branches centrifugally oriented running to tip of lobes; margins of sulcus spermaticus bordered by flounces on basal to most of distal portion of lobes; sulcus spermaticus expanded at apices of lobes; hemipenial body subcylindrical; distal region of hemipenial body defined by transversal series of 4 transversal flounces connected to the sulcus spermaticus; proximal region of hemipenis naked ( Fig 5).   Fig 4.Dorsal (A) and ventral (B) views of the holotype of  Boa atlantica  sp. nov.(MNRJ 27242) from Rio de Janeiro, Atlantic coast of Brazil. https://doi.org/10.1371/journal.pone.0298159.g004   Fig 5.Sulcate (A) and asulcate (B) sides of hemipenis of the holotype of  Boa atlantica  sp. nov.(MNRJ 27242) from Rio de Janeiro, Atlantic coast of Brazil. https://doi.org/10.1371/journal.pone.0298159.g005  Variation.We refer to Tables 3and S 1for the synthesis of quantitative variation of  Boaatlantica. Dorsal ground colour from light to pinkish brown; flanks on the first two thirds of body lighter than dorsum, usually with grey tones; flanks darker on last third of body, usually with tones of blackish brown; longitudinal head stripe darker than background colouration, usually with brown; head stripe usually continuous, with a brown blotch of the same colour as the body; longitudinal head stripe without lateral projections, from internasal region, usually extending to occipital region; preocular and subocular pigmented with tiny brown dots; last two stripes lighter than postocular or with same colour; postocular stripe extends to quadratemandibular articulation or connects to first dorsal saddle; usually no gular blotch; dorsal spots 17–23, elliptical, circular or double-oval, not blotched; dorsal spots (13–28 scales long, 8–17 scales wide) light brown to pinkish brown, bordered by brown; posterior dorsal spots do not change in colour or form; saddles dark brown or black (4–13 scales long), bordered by light brown to pinkish brown, usually connected to each other; dorsal saddles usually with brown on last body third; lateral ocelli from dark red, brownish orange to blackish brown, bordered by cream; belly cream with pinkish tones and black blotches; ventral black blotches progressively increase in number and size towards last body third (= posterior ventral darkening); dorsal tail colour light brown, with 3–6 dorsal brown or black spots, without tail interspots; ventral surface of tail cream with spots generally uniformly black or light brown, bordered by black; cloacal blotch, if present, brown or black.  Table 3. Variations in scale counts of  Boa atlantica sp. nov.     Min Max mean SD n 95%-IC  Circumorbitals 13 20 15.9 1.4 122 0.25   Suboculars 0 2 0.4 0.6 123 0.11   Supralabials 18 25 20.8 1.5 122 0.27   Intrasupraoculars 13 21 16.7 1.5 123 0.27   Infralabials 20 29 24.5 1.5 123 0.27   Gulars 12 22 16.2 1.6 121 0.29   Preventrals 0 6 1.3 1.1 120 0.20   Ventrals 225 248 235.2 4.1 115 0.75   Subcaudals 31 59 51.5 4 120 0.72   Anterior dorsal scale rows 48 70 58.7 4 119 0.72   Midbody dorsal scale rows 67 93 79.5 5 120 0.89   Posterior dorsal scale rows 38 58 47.9 3.4 124 0.60   Tail dorsal scale rows 17 26 20.9 2 122 0.35  95%-CI = 95% confidence interval; max = maximum; min = minimum; n = number of samples; and SD = standard deviation.  https://doi.org/10.1371/journal.pone.0298159.t003  Sexual secondary dimorphism. Despitethe size of the lateral cloacal spurs, which is larger in males than in females, we found evidence of sexual dimorphism in  Boa atlantica sp. nov., as confirmed by the MANOVA (F = 1038; p<0.03). Thevariables in which we found sexual differences were: number of subcaudal rows (F 1,28= 6.49; p<0.02), number of tail spots (F 1,28= 6.19; p<0.02), eye-mouth distance (F 1,28= 4.88; p<0.04), head width (F 1,2= 4.24; p<0.05). Comparingthe central tendency measures and 95% confidence intervals ( Table 4), females have broader heads and have larger eye-mouth distance than males, and males have more subcaudals and more tail spots than females. Thelargest male is from São Fidelis, state of Rio de Janeiro( IBSP 4620) 2114 mmSVL, 290 mmTL; largest female from Floresta da Tijuca, Rio de Janeiro, state of Rio de Janeiro( MNRJ 26886) 1.808 mmSVL, 245 mmTL.  Comparison with South American mainland congeners.  Boa atlantica  sp. nov.differs from  B. constrictor amarali(in parenthesis) by having belly cream with tones of orange, brown and black, scattered of black dots and large groups of black spots (vs. heavily pigmented with black and brown); posterior dorsal spots not blotched (vs. blotched); and tail interspots absent (vs. fuzzy) ( Fig 6).  Boa atlantica  sp. nov.differs from B. c.  constrictor(in parenthesis) by having posterior dorsal saddle spots shape similar to the anterior spots (vs. last dorsal saddles comprising large red polygons different from the anterior in B. c.  constrictor); last lateral ocelli dark brown, black or dark red (vs. blood-red in B. c.  constrictor) ( Fig 6); tail spots black (vs. blood-red in B. c.  constrictor) ( Fig 6).  Boa atlantica  sp. nov.differs from  B. nebulosa(in parenthesis) by having 228–243 ventrals (vs. 256–269 in  B. nebulosa), 69–90 midbody dorsal scale rows (vs. 59–69 in  B. nebulosa), subocular stripe present (vs. absent in  B. nebulosa), posterior dorsal spots and saddles do not change shape (vs. posterior saddles fuse with lateral ocelli forming brown bands in  B. nebulosa), and belly with shades of brown and black with and randomly distributed black dots, and larger spots concentrated on the paraventral region (vs. belly heavily pigmented with black, brown dots and shades in  B. nebulosa).  Boa atlantica  sp. nov.differs from  B. orophias(in parenthesis) by having 228–243 ventrals (vs. 262–280 in  B. orophias), 44– 59 subcaudals (vs. 63–69 in  B. orophias), 17–23 dorsal spots (vs. 25–30 in  B. orophias), 3–6 tail spots (vs. 6–9 in  B. orophias), longitudinal head stripe usually continuous with regular borders and no lateral projections (vs. longitudinal head stripe usually fragmented, with carved borders and none, two or multiple lateral projections in  Boa orophias).  Boa atlantica  sp. nov.differs from B. c.  occidentalis(in parenthesis) by having 17–23 dorsal spots (vs. 23–29 in B. c.  occidentalis); general colour light to dark brown (vs. dark brown in B. c.  occidentalis); longitudinal head stripe with no lateral projections (vs. two or more projections in B. c.  occidentalis); lateral ocelli dark brown, black or faint reddish (vs. cream in B. c.  occidentalis); belly cream with tones of orange, brown and black, scattered of black dots and large groups of black spots (vs. heavily variegated with brown, yellow, black and white blotches in B. c.  occidentalis); belly progressively darker towards tail (vs. no changes in belly pattern in B. c.  occidentalis); Tail spots black (vs. cream in B. c.  occidentalis) ( Fig 6). Additionally,  B. occidentaliswas always recovered as monophyletic on the molecular analyses, although grouped either as the sister group to the South American representatives or as the sister group to the Mexico+ Central America Clade, depending on the markers and the dataset used. We refer to S1–S4 Tables for the meristic and morphometric data on  B. atlantica,  B. occidentalis,  B. orophiasand  B. nebulosa(respectively).  Table 4. Basic descriptive statistics for the sexual dimorphic characters of  Boa atlantica. max = maximum; min = minimum; n = number of samples; SD = standard deviation.     Sex Min max Mean SD n  Head width (mm) M 22.9 57.6 35.3 8.1 21  F 29.8 52.7 39.7 6.6 26  Distance Eye-Mouth (mm) M 3.5 7.4 5.3 1.3 20  F 4.3 8.8 6.5 1.4 26  Number of subcaudal rows M 47 58 52.9 2.8 26  F 31 56 48.0 5.0 25  Number of tail spots M 4 6 5.0 0.6 26  F 2 6 4.3 0.8 25  Abbreviations: F = female; M = male.  https://doi.org/10.1371/journal.pone.0298159.t004  Hemipenial morphology of the sympatric congeners.The hemipenes of the sympatric Boasare similar in several aspects: organ bilobed, non-capitate, and non-calyculate; lobes relatively short and sub-cylindrical with rounded apices, similar in size and oriented centrifugally; sulcus spermaticus divides on distal portion of hemipenial body below sulk bifurcation; sulcus spermaticus branches centrifugally oriented running to tip of lobes; margins of sulcus spermaticus bordered by flounces on basal to most of distal portion of lobes; sulcus spermaticus expanded at apices of lobes; hemipenial body subcylindrical; proximal region of hemipenis naked. Nevertheless, the hemipenes of  Boa atlantica  sp. nov.( Fig 5) differ from those of B. c.  constrictorand B. c.  amarali(in parentheses) by having lobes bifurcated at about 70–74% of the total length (vs. at 54–79% in B. c.  constrictorand 65–76% in B. c.  amarali), lobes covered by 4–6 transversal flounces on its basal portion (vs. 6–8 flounces in B. c.  amarali), lobes naked from median to apical region (vs. with shallow flounces in B. c.  constrictorand B. c.  amarali); distal region of hemipenial body with 5–7 transversal flounces connected to the sulcus spermaticus (vs. 4–6 in B. c.  constrictorand B. c.  amarali).   Distribution.  Boa atlantica  sp. nov.is found along the coastal Atlantic Forest along eastern Brazilfrom Caicó (6˚27’23.2"S 37˚06’05.8"W; Fig 7and S 5Table, point 1) in Rio Grande do Norte Stateto IlhaGrande (23˚08’48.4"S, 44˚13’40.3"W; Fig 7and S 5Table, point 156) in Rio de Janeiro State, which is also de westernmost point in the southern distribution. The westernmost point in the northern distribution is Ituaçú (13˚48’26.7"S 41˚18’39.5"W; Fig 7and S 5Table point 26) in Bahia State.   Fig 6.Pre-cloacal region and tails of mainland South-American boas.  B. atlantica sp. nov.(MNRJ 27243): a) superior view, b) lateral view;  B. constrictor(USNM 566533): c) superior view, d) lateral view;  B. amarali(UFG 134): e) superior view, f) lateral view;  B. occidentalis(FML28405): g) superior view, h) lateral view. https://doi.org/10.1371/journal.pone.0298159.g006   Fig 7.Distribution of  Boa atlantica sp. nov.based on the examined vouchers and tissue samples. A) Detailment in Rio de Janeiro City, including type locality. B) records in northern Rio de Janeiro and Espírito Santo States, C) records in Rio de Janeiro State. See also S4 Table. https://doi.org/10.1371/journal.pone.0298159.g007 In this heterogeneous environment,  B. atlantica  sp. nov.is found from the sea level up to 906 ma.s.l. ( Fig 7). 4993332327 2019-09-13 WGS, MNRJ Sergeant Marco Aurelio da Silva Brazil Fundacao Osorio 93 11 12 -22.934 Rio Comprido 75 -43.210 Rio Comprido 10 11 MNRJ 27242 1 1 Rio de Janeiro holotype 4993332311 1988-01-18 MNRJ H. Silva Brazil 44 -9.274 Passo do Camaragibe -35.468 11 12 MNRJ 3940 1 1 Alagoas State paratype 4993332333 1996-07-10 S, W G. A. Costa Brazil 58 -14.756 CEPEC-CEPLAC -39.231 Ilheus 11 12 CZGB 4862 1 1 Bahia State paratype 4993332329 1987-09 MNRJ S. Rangel Brazil 29 -14.783 Ilheus -39.166 Salobrinho 11 12 MNRJ 6362 1 1 Bahia paratype 4993332338 1997-08-07 MNRJ U. Caramaschi & H. Niemeyer Brazil 540 -13.800 Ituacu -41.300 Salobrinho 11 12 MNRJ 6361 1 1 Bahia paratype 4993332330 2011-01-10 MNRJ G. A. Costa Brazil 35 -20.8501 Cachoeiro do Itapemerim -41.111 Salobrinho 11 12 IBSP 79063 1 1 Espirito Santo state paratype 4993332346 1996-07-10 MNRJ Soares Brazil 20 -20.617 Guarapari -40.43333333 Setiba 11 12 MNRJ 23361 1 1 Bahia paratype 4993332325 [688,1422,589,613] 1999-11-15 MNRJ C. F. D. Rocha Brazil 29 Ilheus Salobrinho 11 12 MNRJ 24903 1 1 Bahia paratype 4993332345 2007-08-31 MNRJ A. P. Almeida Brazil 13 -19.55 Linhares -40.05 Reserva Biologica de Comboios 11 12 MNRJ 23879 1 1 Bahia paratype 4993332305 2011-03-15 MNRJ R. S. Bernils Brazil 8 -18.66666666 Campus CEUNES / UFES -39.85 Sao Mateus 11 12 MNRJ 23882 1 1 Bahia paratype 4993332306 2002-01-14 MNRJ J. L. Gasparini & F. Campagnolli Brazil 17 -20.297 Morro da Gamela -40.301 Vitoria 11 12 MNRJ 9565 1 1 Espirito Santo paratype 4993332307 2012-05-03 MNRJ R. R. Pinto Brazil 6 -22.833 Cabo Frio -41.983 Vitoria 11 12 MNRJ 22936 1 1 Rio de Janeiro state paratype 4993332320 2018-03-08 MNRJ D. S. Fernandes Brazil 8 -22.25 Parque Nacional da Restinga de Jurubatiba -41.65 Carapebus 11 12 MNRJ 26802 1 1 Espirito Santo paratype 4993332331 2001-05 MNRJ S. Bernils Brazil 23 -22.567 Guapimirim -43 Vitoria 11 12 MNRJ 14238 1 1 Espirito Santo paratype 4993332313 2001-05 MNRJ S. Bernils Brazil 23 -22.567 Guapimirim -43 Vitoria 11 12 MNRJ 14250 1 1 Espirito Santo paratype 4993332340 2011-03-15 MNRJ S. Bernils Brazil 37 -22.833 Iguaba Grande 42.167 Vitoria 11 12 MNRJ 17353 1 1 Espirito Santo paratype 4993332334 2013-06-03 MNRJ J. Creusen Brazil 21 -22.65 Complexo Petroquimico do Rio de Janeiro (COMPERJ) -42.85 Itaborai 11 12 MNRJ 25057 1 1 Espirito Santo paratype 4993332303 2016-07-28 MNRJ S. Bernils Brazil 8 Morro da Gamela Vitoria 11 12 MNRJ 26324 1 1 Espirito Santo paratype 4993332339 2011-03-15 MNRJ S. Bernils Brazil 134 -22.65 Itaborai -42.85 Vitoria 11 12 MNRJ 25413 1 1 Espirito Santo paratype 4993332347 2005-05-12 MNRJ C. L. Prata Brazil 12 22.9167 Marica -42.85 Vitoria 11 12 MNRJ 13111 1 1 Espirito Santo paratype 4993332312 2004-05 MNRJ F. Vieira & J. V. Camargo & M. A. Goncalves Brazil 7 -22.95 Niteroi -43.033 Itaipu 11 12 MNRJ 11205 1 1 Espirito Santo paratype 4993332321 2013-01-19 MNRJ M. S. C. Mesquita Brazil 111 -22.9 Niteroi -43.066 Pendotiba 11 12 MNRJ 23573 1 1 Espirito Santo paratype 4993332316 2008-07-17 MNRJ R. W. Kisling Brazil 2 -22.95 Niteroi -43.033 Vitoria 11 12 MNRJ 16922 1 1 Espirito Santo paratype 4993332332 2013-06-13 MNRJ A. Antunes Brazil 34 -22.7 Nova Iguacu -43.46666667 Porciuncula 11 12 MNRJ 24860, MNRJ 26213 2 Rio de Janeiro paratype 4993332319 2006-03-08 MNRJ B. Pimenta Brazil 231 -20.967 Fazenda Vargem Alegre -42.033 Porciuncula 11 12 MNRJ 14172 1 Rio de Janeiro paratype 4993332314 2003-11-17 MNRJ D. S. Fernandes Brazil 31 -22.5 Rio das Ostras -41.933 Porciuncula 11 12 MNRJ 10117 1 Rio de Janeiro paratype 4993332310 2015-11 2016-05-09 2015-11 MNRJ Baldine Brazil 7 -22.8 Aeroporto Internacional Tom Jobim -43.25 Porciuncula 11 12 MNRJ 25952, MNRJ 26350 2 Rio de Janeiro paratype 4993332342 [755,1462,1490,1514] 2015-11 2016-05-09 2015-11 MNRJ Baldine Brazil 7 Aeroporto Internacional Tom Jobim Porciuncula 11 12 MNRJ 25952, MNRJ 26350 2 Rio de Janeiro paratype 4993332326 2015-12-09 MNRJ D. S. Fernandes & B. Miranda & P. Pinna Brazil 221 -22.905 Agua Santa -43.307 Porciuncula 11 12 MNRJ 25950 1 Rio de Janeiro paratype 4993332328 2010-07-13 MNRJ L. Caetano Brazil 650 -22.95 Horto Grotao -43.233 Bairro Jardim Botanico 11 12 MNRJ 19740 1 Rio de Janeiro paratype 4993332308 2010-04-05 2010-05-18 2010-04-05 MNRJ J. P. Pombal Brazil 18 -22.933 Cosme velho 43.183 Porciuncula 11 12 MNRJ 19412, MNRJ 19564 1 Rio de Janeiro paratype 4993332336 [698,1186,1664,1688] 2010-04-05 2010-05-18 2010-04-05 MNRJ J. P. Pombal Brazil 18 Cosme velho Porciuncula 11 12 MNRJ 19412, MNRJ 19564 1 Rio de Janeiro paratype 4993332337 2016-01-30 MNRJ Baptista Brazil 22 -22.866 Del Castilho -43.266 Porciuncula 11 12 MNRJ 25953 1 Rio de Janeiro paratype 4993332335 2013-06-13 MNRJ L. Santos Brazil 3 -23 Estrada do Rio Morto -43.483 Porciuncula 11 12 MNRJ 23144 1 Rio de Janeiro paratype 4993332324 2017-04-21 MNRJ D. B. Santos Brazil 21 22.916 Estrada dos Mananciais -43.366 Porciuncula 11 12 MNRJ 26589 1 Rio de Janeiro paratype 4993332304 2001-03-21 MNRJ W. Kisling & Nova Iguacu Brazil 15 -14.79 Ilha do Governador -39.17 Porciuncula 11 12 MNRJ 9449 1 Rio de Janeiro paratype 4993332341 2017-06-06 MNRJ A. Carneiro Brazil 34 Fazenda Vargem Alegre Porciuncula 11 12 MNRJ 26585 1 Rio de Janeiro paratype 4993332315 2018-03-13 MNRJ C. Bueno Brazil 360 12 13 -22.933 Parque Nacional da Tijuca -43.283 Porciuncula 11 12 MNRJ 26886 1 Rio de Janeiro paratype 4993332309 2019-02-29 MNRJ W. Kisling & Nova Iguacu Brazil 34 Fazenda Vargem Alegre Porciuncula 12 13 MNRJ 27262 1 Rio de Janeiro paratype 4993332322 2006-05-27 MNRJ J. R. Gomes Brazil 14 -23.017 Recreio dos Bandeirantes -43.467 Porciuncula 12 13 MNRJ 14200 1 Rio de Janeiro paratype 4993332323 [533,1205,312,336] 2006-05-13 MNRJ W. Kisling & Nova Iguacu Brazil 34 Fazenda Vargem Alegre Porciuncula 12 13 MNRJ 14201 1 Rio de Janeiro paratype 4993332317 2013-06-13 MNRJ J. A. L. Pontes Brazil 190 -22.833 Serra do Mendanha -43.483 Porciuncula 12 13 MNRJ 17547 1 Rio de Janeiro paratype 4993332343 2013-06-13 2017-08-22 2006-10-10 MNRJ M. Mocelin Brazil 10 -22.95 Rio de Janeiro -43.3 Porciuncula 12 13 MNRJ 10092 1 Rio de Janeiro paratype 4993332301 2017-08-22 2017-08-22 2006-10-10 MNRJ W. Kisling & Nova Iguacu Brazil 34 Fazenda Vargem Alegre Porciuncula 12 13 MNRJ 26796 1 Rio de Janeiro paratype 4993332344 2006-10-10 2017-08-22 2006-10-10 MNRJ G. L. Forreque Brazil 4 -21.234 Sao Francisco de Itabapoana -41.122 Sao Francisco de Itabapoana 12 13 MBML 2097 1 Rio de Janeiro paratype 4993332302 2011-07-25 2017-08-22 2006-10-10 MNRJ W. Kisling & Nova Iguacu Brazil Fundacao Osorio 972 -22.4 Teresopolis -42.967 Porciuncula 12 13 MNRJ 20700 1 Rio de Janeiro paratype 4993332318 2019-09-13 2017-08-22 2006-10-10 MNRJ Sgt. Marco Aurelio da Silva Brazil Fundacao Osorio 93 -22.934 Rio Comprido -43.210 Rio Comprido 12 13 WGS84, MNRJ 27243 1 Rio de Janeiro paratype