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Activity Pattern of the Black-And-White Tegu, Salvator Merianae (Squamata, Teiidae), in an Atlantic Forest Remnant in Southeastern Brazil

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Activity Pattern of the Black-And-White Tegu, Salvator Merianae (Squamata, Teiidae), in an Atlantic Forest Remnant in Southeastern Brazil Herpetology Notes, volume 13: 93-99 (2020) (published online on 05 February 2020) Activity pattern of the Black-and-White Tegu, Salvator merianae (Squamata, Teiidae), in an Atlantic Forest remnant in southeastern Brazil Ana Carolina Srbek-Araujo1,2,3,*, Lillian Jardim Guimarães1, and David Costa-Braga1 Abstract. Temperature is an important factor for metabolism, and lizards dispend part of their daily activity in interactions with the thermal environment. Here we aimed to determine the activity pattern (monthly and daily activity) of Salvator merianae in a remnant of Atlantic Forest in southeastern Brazil, and to analyse the association between activity (records/month) and local weather conditions (temperature and rainfall). The records were collected by camera traps between October/2015 and July/2017 (22 months). We obtained 162 records of S. merianae. The daily activity was between 8 h and 16 h (~9 hours). The species was inactive from April to September 2016 (hibernation period), with two active seasons during our sampling period. In the first activity season (October/2015 to March/2016), the local temperature varied between 26.4ºC and 27.8ºC, and the activity pattern was bimodal. In the second season (October/2016 to April/2017), ambient temperature varied between 23.8ºC and 26.8ºC, and the activity pattern was unimodal. The daily distribution of records tended to be different between activity seasons (W=5.251; p=0.072). The number of records/month was related to monthly mean temperature (rs=0.616; p=0.002; 22 pairs) and not related to total monthly rainfall (rs=0.128; p=0.570; 22 pairs). Teiid lizards need high body temperatures to perform their activities, which may explain their unimodal activity pattern. The bimodal pattern differs from that previously described to S. merianae and can be attributed to behavioural responses to higher temperatures during the first activity season aiming to avoid thermal stress. The activity of S. merianae was influenced by environment temperature, and the difference between activity seasons (bimodal or unimodal pattern) suggests behavioural plasticity in response to differences in annual temperatures. Keywords. behavioural plasticity, circadian rhythm, hibernation, Macroteiid, Reptilia, thermoregulation, time of activity. Introduction body temperature. The body temperature varies from values that allow the beginning of their activities to the Lizards are ectothermic vertebrates that use behavioural maximum physiologically tolerated values, establishing mechanisms, such as alternating movements between an optimal range in which they can develop their daily warm and cooler thermal microenvironments (e.g. activities (Brattstrom, 1965). The body temperature Duran et al., 2018), and physiological mechanisms, influences several biological processes, such as such as enzymes regulated by hormones (e.g. John- antipredator responses (Cury de Barros et al., 2010), Alder, 1990), to maintain metabolic rates and regulate oxygen consumption (Beyer and Spotilat, 1994) and immunological function (Mondal and Rai, 2001). As presented above, the temperature is a physical factor of great importance for metabolism, and lizards dispend a large part of their daily activity in interactions 1 Laboratório de Ecologia e Conservação de Biodiversidade, with the thermal environment (Rocha et al., 2009). The Programa de Pós-graduação em Ecologia de Ecossistemas, thermal environment influences their cycle of daily and Universidade Vila Velha - UVV, Rua Comissário José Dantas seasonal activity (Pianka and Vitt, 2003), and in some de Melo, n° 21, Bairro Boa Vista, CEP 29.102-920, Vila cases leads to hibernation during the months with lower Velha, Espírito Santo, Brazil. temperatures (Winck and Cechin, 2008). In general, the 2 Programa de Pós-graduação em Ciência Animal, Universidade availability of sunlight is the most important factor for Vila Velha - UVV, Vila Velha, Espírito Santo, Brazil. 3 Instituto SerraDiCal de Pesquisa e Conservação, Belo the activity of lizards, mainly the heliothermic ones, Horizonte, Minas Gerais, Brazil. usually found in warm microhabitats where direct * Corresponding author. E-mail: srbekaraujo�hotmail.com sunlight is available (Vitt and Caldwell, 2009). 94 Ana Carolina Srbek-Araujo et al. The Teiidae Family comprises 18 living genus of Material and Methods lizards (Uetz et al., 2019) distributed throughout the Study area.—The study was carried out in Reserva new world (Harvey et al., 2012). They are all terrestrial Natural Vale (RNV - Vale Natural Reserve; -19.1094°S, with diurnal activity (heliophiles), mainly carnivorous -39.9238°W; 22.711 ha) and in Reserva Biológica de or insectivorous, and most of them are active foragers Sooretama (RBS - Sooretama Biological Reserve; - performing their activities within an optimal thermal 19.0044°S, -40.1474°W; 27.859 ha), in the northern range (Bauer, 1998). Most of them are lizards with large region of the state of Espírito Santo, southeastern Brazil. snout-vent length (SVL), as macroteiids of the genus These areas, together with other small private fragments, Salvator (mean SVL = 416.0 ± 3.5 mm; e.g. Winck et form an important remnant of native vegetation al., 2011), but some teiids have smaller representatives, (>53,000 ha, Bloco Florestal Linhares-Sooretama: BLS such as Ameivula spp. (mean SVL = 56.1 ± 4.5 mm; - Linhares-Sooretama Forest Block) that represents e.g. Menezes et al., 2008). Larger animals have more approximately 11% of the forest remaining in the state body mass to heat and lowest relative body surface (based on data available in FSOSMA and INPE, 2019). (smaller surface area-to-volume ratio), than smaller The BLS is composed by a mosaic of habitats, with ones, resulting in a lower heat loss rate to the external predominance of dense lowland forest (Tabuleiro environment (Vitt and Caldwell, 2009). Thermal inertia Forest), classified as Perennial Seasonal Forest (Jesus is therefore different and may influence the activity and Rolim, 2005). The regional climate is tropical with pattern because larger mass can maintain body heat for dry winter, type Aw, according to Köppen classification a longer time compared to smaller animals. (Alvares et al., 2014). The region is characterized by The Black-and-White Tegu, Salvator merianae two climatic periods throughout the year (dry and rainy (Duméril and Bibron, 1839), is heliothermic (Winck et season), with the dry season extending from April to al., 2011) and has unimodal daily activity pattern (Van- September, and the rainy season from October to March Sluys and Rocha, 1999; Ferreguetti et al., 2018), with (Kierulff et al., 2014). The mean annual temperature in seasonal variation in activity (Van-Sluys and Rocha, the region is 24.3 ± 2.1°C, ranging from 18.7 ± 0.6°C 1999) and seasonal reproductive pattern (Naretto et al., to 29.9 ± 0.9°C; and the mean annual rainfall is 1,215 2015). Its activity is more intense in warmest months, ± 260 mm, with high interannual variability (Kierulff with hibernation period in the colder months (Winck et et al., 2014). al., 2011). Thermal preferences are similar between body sizes, sexes and reproductive condition in S. merianae, Data collection.—Data were collected from October within a narrow range of temperature variation in thermal 2015 to July 2017, totalling ~22 months of sampling. For gradient, suggesting similar thermal requirements for a data collection, Bushnell Trophy Cam HD Aggressor broad range of physiological processes (Cechetto and Low-Glow digital camera traps (Bushnell Inc., Overland Naretto, 2015). Contradictorily, the activity pattern can Park, USA) were used. The camera traps were installed be differentiated between adults and subadults, and on trails inside the forest or on unpaved roads located between sexes, with males emerging from hibernation within the study area, totalling 14 sampling points. The before females to establish their territories (Winck equipment was attached to tree trunks (~45 cm above et al., 2011). Despite that, the reproductive cycle is the ground) and operated 24 hours/day. Solar time was considered throughout the sampling period. No baits synchronised, and both male and female concentrate were used during the study. their reproductive activities between October and December (Naretto et al., 2015). Species distribution Data analysis.—To ensure independent counts of models indicate mean temperature of the coldest quarter lizard’s detections, all records obtained within the of the year, followed by annual precipitation, as the same interval of one hour and at the same sampling most important variables for S. merianae (Jarnevich et point were considered as a single detection event (= al., 2018). independent record; Srbek-Araujo and Chiarello, Here we aimed to determine the activity pattern of S. 2013). The sampling effort (number of camera traps x merianae in an important remnant of Atlantic Forest in number of sampling days) and capture success (number southeastern Brazil, considering the species monthly of independent records per sampling effort × 100) were and daily activity pattern, and to analyse the association calculated according to Srbek-Araujo and Chiarello between activity and local weather conditions (2005). The number of sampling days correspond to (temperature and precipitation). the time between first and last records in each month of Activity pattern of the Black-and-White Tegu in an Atlantic Forest remnant 95 sampling for each sampling point. Circular statistical analyses were done in the Oriana
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