Herpetology Notes, volume 14: 15-20 (2021) (published online on 08 January 2021)

Preliminary data on the reproductive biology for almae Franco and Ferreira, 2002, (Dipsadidae: : Tachymenini) in a Caatinga- Atlantic Forest transitional area of Bahia,

Sandro P. de Faria1,5,*, Felipe Camurugi2, Julio C. de Moura-Leite3,4, and Ricardo Marques2

The tribe Tachymenini comprises five genera of biology of T. almae Franco and Ferreira, 2002 from the , being Thamnodynastes Wagler, 1830 the most Caatinga biome. diverse with 20 described , distributed Behavioural and ecological data of T. almae are scarce from northern to Central . and it is mostly restricted to its endemic geographic Five of the 12 species that are currently reported for distribution in the Caatinga biome, occurring in open Brazil (Bailey et al., 2005; Costa and Bérnils, 2018; and forested habitats of xeric vegetation and rocky Uetz et al., 2019) occur in the Caatinga region within soils in the Brazilian states of Piauí, Ceará, Rio northeastern Brazil (Guedes et al., 2014; Franco et al., Grande do Norte,Paraíba, Pernambuco, Alagoas, and 2017). The genus is viviparous and reproductive data is Bahia (Roberto et al., 2009; Guedes, 2010; Jorge and available for T. chaquensis Bergna and Alvarez, 1993, T. Freire, 2011; Coelhoet al., 2013; Guedes et al., 2014; hypoconia (Cope, 1860), T. pallidus (Linnaeus, 1758), Barbosa et al., 2020). Thamnodynastes almae differs T. phoenix Franco, Trevine, Montingelli and Zaher, from other species of the genus by presenting a large 2017, T. sertanejo Bailey, Thomas and Silva Jr., 2005, size (maximum snout-vent length = 657 mm for males and T. strigatus (Günther, 1858) (Vitt and Vangilder and 595 mm for females), light brownish-yellowish 1983; Barbosa et al. 2006; Bellini et al., 2014; Marques colouration, immaculate gular region, 144–157 ventral et al., 2014; Rebelato et al., 2016; Marques et al., 2016; scales, 55–65 subcaudal scales, 8 supralabials, 9 Santana et al., 2017; Silva et al., 2019). However, there infralabials, 2–4 longitudinal ventral stripes, dorsal is a lack of information regarding the reproductive keeled scales and 19 middorsal scale rows (Franco and Ferreira, 2002; Coelho et al., 2013; Franco et al., 2017). This species is mainly active during night time and feeds on lizards and frogs in terrestrial or arboreal 1 Universidade Federal do Paraná–UFPR. Departamento strata (Guedes et al., 2014; Marques et al., 2017). When de Zoologia, Setor de Ciências Biológicas. Av. Coronel threatened, T. almae displays many defensive behaviours Francisco H. Santos, 100, CEP 81530-001, Curitiba, Paraná, (e.g. bite, dorsoventral flattening, triangulation of the Brazil. 2 Universidade Federal da Paraíba–UFPB, Centro de Ciências head and cloacal discharge; Marques et al., 2017). Exatas e da Natureza, Cidade Universitária,CEP 58000-000, However, further information on its biology are João Pessoa, Paraíba, Brazil. unknown. Herein, we provide the first contribution 3 Laboratório de Herpetologia, Museu de História Natural on T. almae reproduction with information of clutch Capão da Imbuia. Rua Prof. Nivaldo Braga 1378, CEP size and offspring’s morphology and pholidosis data. 82590-300, Curitiba, Paraná, Brazil. Additionally, we provide a new record for the species 4 Escola de Educação e Humanidades, Curso de Biologia, distribution and an updated distribution map. Pontificia Universidade Católica do Paraná, Rua Imaculada Conceição, 1500, CEP 82590-300, Curitiba, Paraná, Brazil. On 27 November 2018 at 19:30 h during the dry season, 5 Mapinguari – Laboratório de Sistemática e Biogeografia de SPF manually captured one specimen of T. almae at the Répteis e Anfíbios, Universidade Federal de Mato Grosso do rural area of Feira de Santana municipality (12.1245 S, Sul, Cidade Universitária, CEP 79070-900 Campo Grande, 38.9274 W, 230 m above sea level), Bahia state, Brazil. MS, Brazil. This transition area is defined as the “Agreste” ecotone * Corresponding author. E-mail: [email protected] of the Caatinga and the Atlantic Forest biomes (Silva © 2020 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. et al., 2017). The was found active within an 16 Sandro P. de Faria et al. area of rock outcrop covered by bromeliads (Bromelia Capão da Imbuia at the Universidade Federal do Paraná laciniosa Mart. ex Schult). The specimen was an adult (adult female: MHNCI 17551; offspring: 17552–17561, female with 539 mm snout-vent length and 153 mm tail Figure 1). length. The was transported to the laboratory in a We observed a clutch size of 10 snakes. The pregnancy plastic bag and killed with a thiopental injection (Hawk was in its final stage since the offspring were fully et al., 2005), fixed in 10% formalin, and preserved in formed. Offspring presented an average body mass of 70% alcohol. In order to obtain a tissue sample, we 2.02 g ± 0.27 standard deviation (SD) (1.46–2.34 g, n made a small incision on the venter and noticed the = 10). The smallest offspring measured 179 mm total individual was gravid. We proceeded by removing all length (female) and the largest 195 mm total length the embryos from the oviducts and sexed the specimens (male), corresponding to 33.3% and 36.2% of the by performing a longitudinal subcaudal incision on the mother’s total length size, respectively. Males (n = 6) tail to check for hemipenial tubes, observed through showed an average SVL size of 184.98 mm ± 5.05 SD a Leica Zoom 2000 stereo microscope. We measured (176.7–190.6 mm), and an average tail length of 43.5 specimens with a Mitutoyo calliper and used a 10g mm ± 0.1 SD (42–45 mm). Females (n = 4) presented Pesola dynamometer to obtain the body mass. The an average SVL size of 177.5 mm ± 2.21 SD (175–180 following variables were measured: mass (in grams), mm), and an average tail length of 42.5 mm ± 0.26 SD snout-vent length, tail length, head length, head height, (40–46 mm). All other measurement values were very head width, ocular diameter, distance between eyes and similar and overlapped between males and females ventral-symphysial distance. Additionally, we counted (Table 1). ventral and subcaudal scales starting counting after the Offspring data is available for few species of cloacal plate, and observed the umbilical scar position Thamnodynastes, but require some taxonomic within the ventrals. Vouchers were deposited in the clarification. Vitt and Vangilder (1983) report T. herpetological collection of Museu de História Natural pallidus and T. strigilis, with reproductive data for the

Figure 1. Voucher offspring of Thamnodynastes almae deposited at MHNCI (17552–17561). Preliminary data on the reproductive biology for Thamnodynastes almae 17

Table 1. Morphological measurements and meristic information of Thamnodynastes almae offspring. Abbreviate measurements are: US = umbilical scar position in relation to ventral scale count, SVL = Snout-vent length, TL = tail length, HL = Head length, TableHH =1. Head Morphological height, HW measurements = Head width, and meristic OD = information Ocular diameter, of Thamnodynastes DBE = Distance almae offspring. between Abbreviate eyes, VSD measurements = Ventral-symphysial are: US = umbilical distance. scar position in relation to ventral scale count,Size SVLmeasurements = Snout-vent are length, in millimetres TL = tail length, and HL body = Head mass length, in grams. HH = Head We provideheight, HW the = meanHead width, and standardOD = Ocular deviation diameter, (SD) DBE for = Distance males andbetween eyes, VSD = Ventral-symphysial distance. Size measurements are in millimetres and body mass in grams. We provide the mean and standard deviation (SD) for males and females. females.

Voucher Sex Ventrals Subcaudals US SVL TL HL HH HW OD DBE VSD Mass MHNCI 17552 M 155 64/64 26 190.6 44 12 4.1 5.9 2 3.1 8.7 2.34 MHNCI 17553 M 156 65/65 27 184.6 44 12 4.5 6.8 2.5 3.3 8.1 1.84 MHNCI 17554 M 160 66/66 28 185.5 45 11.2 4.1 5.7 2.1 3.8 8 1.82 MHNCI 17555 F 151 60/60 24 174.7 43 13.1 4.2 6.6 2.2 3.6 7.3 2.13 MHNCI 17556 F 151 63/63 24 178.4 46 11.7 4.2 6.3 2.4 3.9 7.4 2.15 MHNCI 17557 F 150 61/61 22 177 40 11.3 3.9 6.8 2.3 3.1 7 2.26 MHNCI 17558 M 155 66/66 25 189.7 43 11.8 4.3 5.2 2.6 3.3 8.3 2.01 MHNCI 17559 F 153 63/63 28 179.9 41 12.2 4.1 6.4 2.7 3.3 7.7 2.1 MHNCI 17560 M 156 64/64 25 176.7 43 11.7 4.2 5.6 2.7 3 7.4 1.46 MHNCI 17561 M 156 67/67 23 182.8 42 11.1 4.4 5.6 2.3 4 7.2 1.94 Mean M 156.3 65.3 25.7 185 43.5 11.6 4.3 5.8 2.4 3.4 8 1.9 SD M 1.9 1.2 1.8 5.1 1 0.4 0.2 0.5 0.3 0.4 0.6 0.3 Mean F 151.3 61.8 24.5 177.5 42.5 12.1 4.1 6.5 2.4 3.5 7.4 2.2 SD F 1.3 1.5 2.5 2.2 2.6 0.8 0.1 0.2 0.2 0.4 0.3 0.1

later. However, they identified species based on Peters Though males were slightly bigger than females, our and Orejas-Miranda (1970) and these were the closest sample size is limited to attest for sexual size dimorphism species at the time. Bailey at al. (2005) showed later the in neonates of T. almae. Unsexed offspring of T. pallidus specimen of T. pallidus from Vitt and Vangilder (1983) were smaller than T. almae and T. sertanejo, while was actually T. sertanejo. In addition, Vanzolini et al. offspring of T. sertanejo showed larger females than (1980) also report T. strigilis in Caatinga, based on Peters males (Table 2; Barbosa et al., 2006; Santana et al., 2017; and Orejas-Miranda (1970), but with the morphological Silva et al., 2019). Adult female snakes usually show description they provided, we now can allocate this larger body sizes, probably related to increase fecundity species as the recently described T. phoenix (see Franco (Pizzatto and Marques, 2002; Pizzatto, 2005; Cox et et al., 2017). Therefore, based on the same reference al., 2007; Bertona and Chiaraviglio, 2008; Pizzatto et used by both studies to identify species, we believe that al., 2008; Prieto et al., 2012). However, male offspring T. strigilis from Vitt and Vangilder (1983) refers to T. of T. almae presented larger SVL sizes, a pattern also phoenix. Likewise, Barbosa et al. (2006) report captive observed in T. hypoconia (Bellini et al., 2014). We also breeding of an unidentified species of Thamnodynastes. observed males with longer tails compared to females, a Despite their morphological description report 19 feature of sexual size dimorphism commonly presented middorsal scale rows, the photographs in the study in other species of snakes (Pizzatto et al., 2008; Orofino clearly show specimens of T. sertanejo (17 middorsal et al., 2010; Bellini et al., 2014). Some authors suggest scale rows; see Franco et al., 2017). that longer tails in males is an anatomical constraint in Following this background, T. almae is the species order to hold the hemipenis and the retractor muscles with more offspring so far. The clutch size of T. phoenix accommodation in the basal portion of the tail (Zug et varies from 3–8 snakes, and females present enlarged al., 1979; Lewis et al., 2013). However, longer tails in follicles in August (Vitt and Vangilder, 1983; Franco et males may also result as an advantage in species that al., 2017). For T. pallidus, there is record of 10 follicles have male-male ‘tail-wrestling’ competition (King, in July and two reports of five offspring each in January 1989; Madsen and Shine, 1993). Offspring of T. almae and February (Marques et al. 2014; Santana et al., and T. pallidus are proportionally similar compared to 2017; Silva et al., 2019) and 9 snakes for T. sertanejo in the mother’s size, while T. sertanejo show the smallest January (Barbosa et al., 2006). offspring. Silva et al. (2019) pointed out that the smallest 18 Sandro P. de Faria et al.

Table 2. Morphological comparison between offspring of Thamnodynastes species. Abbreviations are: M = male, F = female, I = indeterminate, CS = clutch size, SVL = Snout-vent length, TL = tail length, HL = Head length, HW = Head width, OD = Ocular Table 2. Morphological comparison between offspring of Thamnodynastes species. Abbreviations are: M = male, F = female, I = indeterminate, CS = clutch size, SVL = Snout-vent length, TL diameter,= tail length, and HL =DBE Head =length, Distance HW = Headbetween width, eyes. OD = OcularSize measurements diameter, and DBE are = Distance in millimetres between eyes. and Size body measurements mass in aregrams. in millimetres Values and are body provided mass in grams. Values are asprovided mean as ± mean standard ± standard deviation deviation (min-max). (min-max).

Species Source Sex CS SVL TL HL HW OD DBE Mass T. almae This study M 6 185 ± 5.1 43.5 ± 0.1 11.6 ± 0.4 5.8 ± 0.5 2.4 ± 0.3 3.4 ± 0.4 1.9 ± 0.3 (176.7–190.6) (4.2–4.5) (11.1–12.0) (5.2–6.8) (2–2.7) (3–4) (1.46–2.34) This study F 4 177.5 ± 2.2 42.5 ± 0.3 12.1 ± 0.8 6.5 ± 0.2 2.4 ± 0.2 3.475 ± 0.35 2.2± 0.1 (174.7–179.9) (4–4.6) (11.3–13.1) (6.3–6.8) (2.2–2.7) (3.1–3.9) (2.1–2.26) T. pallidus Santana et al. 2017 I 5 157.1 ± 6.8 61.5 ± 2.4 9.1 ± 0.1 5 ± 0.2 5.3 ± 0.2 - 1.6 ± 0.2 (149.7–164.2) (58–64) (8.9–9.2) (4.7–5.3) (5–5.5) (1.4–1.8) T. pallidus Silva et al. 2019 I 5 133.8 ± 4.5 53.2 ± 3.4 7.6 ± 0.2 4.4 ± 0.6 3 ± 0.5 3 ± 0.5 1 ± 0 (1–1) (126–137) (49–56) (7.4–7.9) (3.7–5) (2.2–3.4) (2.1–3.4) T. sertanejo Barbosa et al. 2006 M 6 173.7 ± 2.0 59.5 ± 1.9 11.2 ± 0.8 6.2 ± 0.4 - - - (171–176) (58–63) (10–12) (6–7) Barbosa et al. 2006 F 3 179.3 ± 2.3 65.3 ± 1.5 11.3 ± 0.6 6.3 ± 0.6 - - - (178–182) (64–67) (11–12) (6–7)

and largest offspring of T. pallidus were respectively The offspring’s morphometric and meristic data are 175 mm and 192 mm total length, corresponding to 31% poorly explored (Norval et al., 2007; Santana et al., 2017; and 35% of the mother’s size. Santana et al. (2017) also Morais et al., 2018; Silva et al., 2019), but are extremely showed a similar relation for T. pallidus, as the smallest relevant to the knowledge of phenotypic plasticity and and largest offspring correspond to 35% and 39% of natural history of these organisms. Also, the increasing the mother’s size. As for T. sertanejo, the smallest and natural history data in a highly biodiverse region as largest offspring were 28% (171 mm) and 29.8% (182 the Neotropics, allow us to answer future comparative mm) from the mother’s size, respectively (Barbosa et al., questions on ecology and evolution of viviparity in 2006). Such similar sizes can be a result of the shared snakes. This is a compelling aspect for the Caatinga, evolutionary history of closely related species.

Male offspring of T. almae presented higher values in pholidosis variables for ventral and subcaudal scales, with an average of 156.33 ± 1.86 SD ventrals (range 155–160 ventrals) and 65.33 ± 1.21 SD subcaudal scales (range 64–67 subcaudal scales), respectively. As for females, the average values for the same variables were 151.25 ± 1.26 SD ventrals (range 150–153 ventrals) and 61.75 ± 1.5 SD subcaudal scales (60–63 subcaudal scales) (Table 1). Sexual dimorphism related to pholidosis variables, with males having more ventral and subcaudal scales besides T. almae, was also found for T. chaquensis, T. hypoconia and T. strigatus (Bellini et al., 2014). However, Barbosa et al. (2006) found females of T. sertanejo with more ventrals than males. In this study we also present the southernmost record of T. almae. Our observation extends the species known distribution 187 km Southeast from the closest record in Itiúba, Bahia state (Guedes, 2010) and 908 km Southeast from the farthest record in José de Freitas, Piauí state Figure 2. Distribution map of T. almae in Caatinga biome. (Barbosa et al., 2020) (Table 3, Figure 2). The polygon Blue dots show literature records and the red star show our of occurrence provided by all records result in an area new record in Feira de Santana municipality, Bahia state. of 307,764 km2, which represents nearly 30% of the Dashed grey line show the Caatinga biome boundaries. See Caatinga territory (Silva et al., 2017). Table 3 for localities details. Preliminary data on the reproductive biology for Thamnodynastes almae 19

Table 3. Localities from the geographic distribution of T. almae updated from Coelho et al. (2013). Brazilian states acronyms are: AL = Alagoas,Table BA 3: = Localities Bahia, CE from = Ceará, the geographic PB = Paraíba, distribution PE = of Pernambuco, T. almae updated PI = from Piauí, Coelho RN = et Rio al. (2013).Grande Brazilian do Norte. states (*) approximate Brazilian statesacronyms coordinate, are: AL not = Alagoas, provided BA in = the Bahia, original CE = paper.Ceará, PB = Paraíba, PE = Pernambuco, PI = Piauí, RN = Rio Grande do Norte. (*) approximate Brazilian states coordinate, not provided in the original paper.

Municipality (Locality) State Latitute Longitude Reference 1. Feira de Santana BA -38,9275 -12,1245 This study 2. Itiúba BA -39,85 -10,7 Guedes, 2010 3. Piranhas AL -37,75 -9,6167 Guedes, 2010 4. Paulo Afonso BA -38,2167 -9,4 Guedes, 2010 5. Petrolândia PE -38,3 -9,1333 Guedes, 2010 6. Rodelas (Usina Hidrelétrica Luiz Gonzaga) BA -38,5777 -9,0027 Franco and Ferreira, 2002 7. Floresta PE -38,2665 -8,6788 Coelho et al., 2013 8. Arcoverde PE -37,0585 -8,4184 Freitas et al., 2019 9. Custódia PE -37,5275 -8,1611 Coelho et al., 2013 10. Sertânia PE -37,5115 -8,1569 Coelho et al., 2013 11. Cabrobó PE -39,4607 -8,5172 Coelho et al., 2013 12. Salgueiro PE -39,122 -7,982 Coelho et al., 2013 13. Cabaceiras (Fazenda Bravo) PB -36,2833 -7,4833 Guedes, 2010 14. Picuí PB -36,3457 -6,5039 Guedes et al., 2014 15. São José de Piranhas PB -38,4995 -7,1196 Coelho et al., 2013 16. Santa Maria (Fazenda Tanques) RN -35,701 -5,854 Jorge and Freire, 2011 17. Milagres CE -38,9764 -7,2536 Roberto et al., 2009 18. José de Freitas PI -42,6136 -4,7993 Barbosa et al., 2020

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