Microhabitat Selection of Ameiva Ameiva (Linnaeus , 1758), in the Brazilian Pantanal (Squamata: Sauria: Teiidae)

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Benício_etal_Microhabitat_selection_Ameiva_ameiva:HERPETOZOA.qxd 12.02.2019 14:53 Seite 1

heRPetozoa 31 (3/4): 211 - 218 211 Wien, 28. Februar 2019

microhabitat selection of Ameiva ameiva (Linnaeus , 1758), in the Brazilian Pantanal (squamata: sauria: teiidae)

mikrohabitatwahl von Ameiva ameiva (Linnaeus , 1758) im brasilianischen Pantanal (squamata: sauria: teiidae)

RoniLdo a. B eníCio & z aida oRteGa & aBRaham menCía & d anieL Cunha Passos

KuRzFassunG die autoren untersuchten den einfluß einiger umweltparameter auf die mikrohabitatwahl von Ameiva amei - va (Linnaeus , 1758) im brasilianischen Pantanal, einem Lebensraum mit markantem Wechsel von Überflutung und trockenfallen. die analyse der mikrohabitatwahl erfolgte im september mittels Ressourcenauswahlfunktion (RsF) und Bedingter Logistischer Regression (CLR) an neunundzwanzig adulten individuen von of A. ameiva . die mit - tlere temperatur an den aufenthaltsorten betrug 36 °C (spannweite: 27 °C – 51 °C; standardabweichung: 6,8 °C). im Verlauf der tagesaktivität waren die temperaturen des substrats auf dem A. ameiva beobachtet wurde, abwech - selnd sowohl bei sehr hohen als auch deutlich niedrigeren Werten gehäuft. das CLR modell zeigte, daß keiner der untersuchten umweltparameter substrattemperatur, substrattyp und sonnenexposition die Wahrscheinlichkeit des Vorkommens von A. ameiva an einem bestimmten aufenthaltsort signifikant beeinflußte (zugehörige P-Werte: 0,11; 0,69 und 0,87), wobei bei der sonnenexposition die Qualitäten volle Besonnung, schatten und halbschatten und beim substrattyp Gras, Laubstreu und blanker Boden unterschieden wurden. die ergebnisse legen nahe, daß A. ameiva zumindest während der trockenzeit im brasilianischen Pantanal kein mikrohabitat-spezialist ist.

aBstRaCt the authors studied the influence of three environmental traits on the microhabitat selection of Ameiva amei - va (Linnaeus , 1758), in the Brazilian Pantanal, an environment with a marked effect of seasonality, i.e., alternating periods of flooding and falling dry. microhabitat selection was assessed in the dry season using a Resource selection Function (RsF) and Conditional Logistic Regression (CLR) analysis. twenty-nine adult individuals of A. ameiva were observed. the average temperature in places where individuals occurred was 36 °C (ranging from 27 °C to 51 °C; sd = 6.8 °C). the pattern of the temperature of the substrate on which A. ameiva was observed showed peaks, occurring alternatively either at very high but also milder temperatures along its daily activity period. the CLR model revealed that none of the studied environmental variables (substrate temperature, type of substrate or sun exposure) significantly affected the probability of occurrence of Ameiva lizards at a particular point ( P = 0.11, P = 0.69, P = 0.87, respectively). Regarding the sun exposure situation the qualities full sun, shade and filtered sun were distinguished, the substrate type comprised the parameter values grass, leaflitter and exposed soil. the results suggest that A. ameiva is not a microhabitat specialist, at least during the dry season in the Pantanal of Brazil.

KeY WoRds Reptilia: squamata: sauria: teiidae; Ameiva ameiva ; ecology, behavior; foraging activity, microhabitat use, occurrence, substrate, wetlands, Pantanal, Brazil

intRoduCtion

Ameiva ameiva (Linnaeus , 1758), is a (VanzoLini et al. 1980) and occurring in medium-sized (snout-vent length up to 174 almost all Brazilian territory ( aViLa -P iRes mm after Vitt & C oLLi 1994 ; Fig. 1) ground- 1995). across its distribution, this species in- dwelling lizard that spends a large amount of habits a variety of environments ( Vitt & its activity period moving, a typical pattern CoLLi 1994), such as forest edges and open of wide/ active foragers ( RoCha et al. 2009). areas within rainforests ( CRuz -n eto & it has a vast geographical distribution, living GoRdo 1996; L ima et al. 2001), seasonally mainly in open areas of south america dry forests ( Vitt 1995; s aLes et al. 2011a), Benício_etal_Microhabitat_selection_Ameiva_ameiva:HERPETOZOA.qxd 12.02.2019 14:53 Seite 2

212 R. a. BeníCio & z. oRteGa & a. menCía & d. C. P assos

savannas ( CoLLi 1991) and coastal shrub - microhabitat use of several populations lands ( zaLuaR & R oCha 2000) as well as of A. ameiva was studied at different human disturbed areas ( saRtoRius et al. Brazilian biomes ( saRtoRius et al. 1999; 1999). Ameiva ameiva mainly feeds on inver - zaLuaR & R oCha 2000; s aLes et al. 2011a). tebrates (Vi tt & C oLLi 1994; z aLuaR & however, there is still a lack of knowledge RoCha 2000, s aLes et al. 2011b, FReitas et about A. ameiva lizards from the Pantanal, an al. 2012), despite sporadically including some area of flooded grasslands from western vertebrates and plant material in the diet Brazil ( JunK & d e Cunha 2005). Widely (RoCha & V RCiBRadiC 1998; saLes et al. distributed species often vary geographically 2010; moRaes & s antos 2012; siLVa et al. in the microhabitat use, and it is important to 2013). these lizards are mainly diurnal and study this intraspecific variability since it is concentrate their activity within the warmer related to niche specialization, evolution and hours of the day, usually between 10:00 h and adaptability to environmental changes ( BoR- 14:00 h ( Vitt & C oLLi 1994; saRtoRius et al. zee et al. 2016; BaRs -C LoseL et al. 2017). 1999; zaLuaR & R oCha 2000; saLes et al. thus, the authors’ present study aimed to 2011a). their life-history traits vary along the evaluate which environmental traits influ - broad geographical range, but reproduction ence the occurrence of A. ameiva in the Pan- begins and tends to be more intense during tanal, an environment with a marked effect of the wet period ( Vitt 1982; C oLLi 1991; V itt seasonality, i.e., alternating periods of flood - & C oLLi 1994; R oCha 2008). ing and falling dry ( meRCante et al. 2011).

mateRiaLs and methods

the study was conducted in the Pan - location were registered, measured with a tanal region of miranda-abrobal, near the laser infrared thermometer (Benetech ® Base de estudos do Pantanal (BeP) of the Gs320, precision 0.1 °C) from one meter Federal university of mato Grosso do sul maintaining a constant distance, to maxi - (Corumbá, mato Groso do sul, Brazil) at mize the comparability of the measurements. 19°34’37” s, 57°00’42” W, in the dry sea - in addition, the authors recorded the type of son, september 2017. the Pantanal domain substrate (‘grass’, ‘leaf litter’ and ‘exposed is embedded in one of the most distinct eco - soil’) and the sun exposure (‘full sun’, regions concerning orography: the flooded ‘shade’ and ‘filtered sun’) of the microhabi - grasslands and savannas ( PRado et al. 1992; tat where the individual was observed (= LouRiVaL et al. 2000). Located in the west - used point). to assess microhabitat selec - ern portion of the state of mato Grosso do tion, the same variables (t s, type of substrate sul, the Pantanal occupies approximately and sun exposure) were registered at four 71.500 km 2 of the state territory (FeRReiRa unused points related to each used point. et al. 2017). unused points were de fined one meter north, the Ameiva lizards were sampled south, west and east from each used point as within three consecutive days (september an objective criterion to standardize their 26-28) through active visual search ( CRumP location. these unused points represented & s Cott 1994) in the period from 07:00 h to the available microhabitats for each lizard in 17:00 h (local time) on two transects of one a given moment. Comparing environmental kilometer each, both in the surroundings of variables of the used and unused points the BeP. to avoid pseudo replication, each would allow for the study of the influence of transect was divided into parts, sampled the registered variables in the microhabitat sequentially in different days. in this way, selection of A. ameiva . each part of the transect was sampled only microhabitat selection was assessed once. the study was purely observational, using a Resource selection Function (RsF) without capture of any individual, to avoid (manLY et al. 2007). Resource selection animal stress. upon observation of a lizard, Functions calculate the odds ratio of an indi - the time of the day (hour in local time) and vidual to use a certain resource relative to its the substrate temperature (t s) of its exact availability in the environment. in the pres - Benício_etal_Microhabitat_selection_Ameiva_ameiva:HERPETOZOA.qxd 12.02.2019 14:53 Seite 3

microhabitat selection of Ameiva ameiva (Linnaeus , 1758) in the Brasilian Pantanal 213

Fig. 1: adult individual of Ameiva ameiva (L innaeus , 1758) , at the Base de estudos do Pantanal, Federal university of mato Grosso do sul, Corumbá, mato Grosso do sul, Brazil. abb. 1: adulte Ameiva ameiva (L innaeus , 1758) in der Base de estudos do Pantanal der universität von mato Grosso do sul, Corumbá, mato Grosso do sul, Brasilien.

ent case, the resources are the microhabitat presence of the lizard in a certain microhab - explicative variables of interest: t s, type of itat (recorded as 1 for the used points and 0 substrate and sun exposure. solving RsFs for the four associated unused points). the by means of Generalized Linear models CLR was conditioned by the identity of (GLms) allows modeling the probability of each individual lizard, to guarantee that val - occurrence of the lizard, given several ex- ues of the explicative variables of each used plicative variables and their interactions. in microhabitat were paired with those avail - addition, this approach can calculate the able to such individual ( duChesne et al. effect of each explicative variable in the 2010; LiedKe et al. 2018). hence, values of same probabilistic selection process ( Lied - the explicative variables of the microhabitat Ke et al. 2018). Finally, the RsF allows used by one lizard are compared with the determining the availability of the environ - simultaneously available values of these mental variables for each individual, simul - variables at its surroundings. an analogous taneously to the moment when the animal is approach, Paired Logistic Regression observed in the used point. this method (PLR), has been used for many years to provides a more mechanistic and powerful study habitat selection of reptiles ( ComPton approach to understand microhabitat selec - et al. 2002; oRteGa & P éRez -m eLLado tion than considering the general availabili - 2017). Conditional Logistic Regression has ty for all individuals of a population, thus, the advantage of having more than one needing fewer individuals to obtain repre - unused point paired to each used point, so sentative results than with traditional ap - the availability of microhabitats is better proaches ( LiedKe et al. 2018). represented for each individual. all analy - the authors assessed the RsF through ses were performed in the software R, ver - a Conditional Logistic Regression (CLR) sion 3.4 ( R C oRe team 2018) using the analysis. the response variable was the ‛lme4’ package ( Bates et al. 2015). Benício_etal_Microhabitat_selection_Ameiva_ameiva:HERPETOZOA.qxd 12.02.2019 14:53 Seite 4

214 R. a. BeníCio & z. oRteGa & a. menCía & d. C. P assos

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Fig. 2: absolute frequency during the hours of the day of 29 active adult individuals of Ameiva ameiva (L innaeus , 1758) , including the mean substrate temperature (■) at the respective observation points. study site: Base de estudos do Pantanal of the Federal university of mato Grosso do sul, Corumbá, mato Grosso do sul, Brazil. abb. 2: absolute häufigkeit der 29 beobachteten aktiven adulten von Ameiva ameiva (L innaeus , 1758) und die mittlere substrattemperatur (°C) an deren jeweiligem aufenthaltsort (■) im tagesverlauf auf dem untersuchungsgelände (Base de estudos do Pantanal der universität von mato Grosso do sul, Corumbá, mato Grosso do sul, Brasilien).

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Fig. 3: substrate temperatures (°C) of the 29 sites where active Ameiva ameiva (L innaeus , 1758), were observed, sorted by the time of observation (local time). study site: Base de estudos do Pantanal of the Federal university of mato Grosso do sul, Corumbá, mato Grosso do sul, Brazil. abb. 3: substrattemperaturen (°C) an jenen stellen, an denen Ameiva ameiva (L innaeus , 1758) beobachtet wurde, geordnet nach der Beobachtungszeit. untersuchungsgelände: Base de estudos do Pantanal der universität von mato Grosso do sul, Corumbá, mato Grosso do sul, Brasilien. Benício_etal_Microhabitat_selection_Ameiva_ameiva:HERPETOZOA.qxd 12.02.2019 14:53 Seite 5

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216 R. a. BeníCio & z. oRteGa & a. menCía & d. C. P assos

table 2: Results of the Generalized Linear models analysis of substrate temperature (°C), type of substrate (categories: grass, leaflitter and exposed soil) and exposure to the sun (categories: full sun, shade and filtered sun) in 145 points on the probability of representing a habitation of an Ameiva ameiva (L innaeus , 1758), individual. tab. 2: ergebnisse der analyse des verallgemeinerten linearen modells. untersucht wurden substrattemperatur (°C), substrattyp (Kategorien: Gras, Laubstreu und blanker Boden) und sonnenexposition (Kategorien: volle Besonnung, schatten, halbschatten) an 145 meßpunkten hinsichtlich der Wahrscheinlichkeit, ein aufenthaltsort eines individuums von Ameiva ameiva (L innaeus , 1758) zu sein .

estimate std. error Z value P value schätzwert standardfehler Z-Wert P-Wert substrate temperature / substrattemperatur 0.04892 0.03130 1.563 0.11 type of substrate / substrattyp 0.28925 0.73933 0.391 0.69 exposure to sun / sonnenexposition -0.10989 0.68181 -0.161 0.87

ResuLts and disCussion

twenty-nine adult individuals of A. 17:00 h; saRtoRi us et al. 1999; zaLuaR & ameiva (116 unused points, see the com - RoCha 2000; s aLes et al. 2011a). plete dataset in table 1) were observed. the the selected study design revealed average temperature at the used points was that neither the sun exposure condition (full 36 °C (ranging from 27 °C to 51 °C; sd = sun, shade or filtered sun), the type of sub - 6.8 °C). the first individual was observed strate (grass, leaflitter or exposed soil) nor around 09:00 h, foraging, and the last one substrate temperature, affected the probabil - ended its activity at approximately 16:00 h ity of occurrence at a particular point of A. (Fig. 2). the pattern of the temperature of ameiva in the Pantanal. the present results the substrate (t s) on which A. ameiva was showed that the studied environmental vari - observed showed peaks, occurring alterna - ables did not affect the microhabitat selec - tively either at very high but also milder tion of A. ameiva , at least during in the dry temperatures along its daily activity period season (september). (Fig. 3). the CLR model revealed that none Ameiva ameiva lizards show a rela - of the environmental variables (t s, type of tively high plasticity in microhabitat use substrate or sun exposure) significantly depending on the habitat in which they affected the probability of occurrence of occur. For instance, while a population of Ameiva lizards ( P = 0.11, P = 0.69, P = 0.87, A. ameiva from the amazonian savanna respectively; table 2). showed the narrowest niche breadth within Previous studies found that A. ameiva its lizard community regarding microhabitat maintained high and stable body tempera - use ( mesQuita et al. 2006), another popula - tures in thermally and spatially heteroge - tion from a sensu stricto Caatinga habitat neous habitats, and that they stopped activi - showed one of the broadest microhabitat ty when environmental temperatures were niche breadths within its lizard community low ( saRtoRius et al. 1999). the studied (Vitt 1995). in fact, even among open lizards started their activity later in the habitats, A. ameiva is reported to use differ - morning and finished it sooner in the ent types of microhabitats. in seasonally evening than other saurian species living in dry forests from northeastern Brazil, this the same habitat (e.g., Tropidurus lagun - species was found virtually on all types of ablanca CaRVaLho , 2016, was active be- substrate at the ground level, except for tween 07:30 h and 17:00 h during the same rocky ones ( saLes et al. 2011a). none- days and in the same habitat; the authors’ theless, in a sand dune habitat form south - personal observations). thus, the studied eastern Brazil, A. ameiva only used three population seemed to follow the same strat - kinds of microhabitats, in a similar pattern egy reported for several other populations of as found for a Pantanal population in the A. ameiva : remaining inactive under unsuit - present study. however, it is important to able environmental temperatures (e.g., tem - remember that A. ameiva lizards are active peratures prevailing before 09:00 h and after foragers that move constantly through space Benício_etal_Microhabitat_selection_Ameiva_ameiva:HERPETOZOA.qxd 12.02.2019 14:53 Seite 7

microhabitat selection of Ameiva ameiva (Linnaeus , 1758) in the Brasilian Pantanal 217

(RoCha et al. 2009), and which maintain predation risk can map the microhabitat their preferential body temperature through - choice of these lizards. in addition, it is out their wide geographic distribution, possible that substrate temperature, type of despite the thermal differences between substrate or sun exposure are important in ecosystems ( Vitt & C oLLi 1994 ). determining microhabitat selection of A. in this sense, the present results and ameiva in other periods of its annual activi - the existing literature show that A. ameiva is ty. the results of the selected approach show not a microhabitat specialist, as other coex - that neither the in situ type or temperature of isting neotropical lizards are (see, e.g., Vitt the substrate nor the sun exposure influenced 1991). nonetheless, it is possible that other the microhabitat use of A. ameiva during the variables that were not considered in the dry season (september) in the Pantanal of present study, such as food availability or Brazil.

aCKnoWLedGments

this article is the result of a research of the 20th Coordenação de aperfeiçoamento de Pessoal de nível Curso de Campo do Pantanal (ecoPan2017) of the superior (CaPes) for financial support through a post - universidade Federal of mato Grosso do sul. Ronildo doctoral fellowship (Programa nacional de Pós- a. BeníCio thanks Fundação de amparo à Pesquisa do doutorado - PnPd/CaPes). the authors also thank estado de são Paulo for financial support (Proc. FaPe - Luíz Gustavo oLiVeiRa -s antos (Federal university of sP 2015/11821-0), z. oRteGa and a. menCía thank mato Grosso do sul ) for help with the data analyses.

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date oF suBmission: may 25, 2018 Corresponding editor: heinz Grillitsch

authoRs: Ronildo a. BeníCio (Corresponding author, < [email protected] >) 1) , zaida oRteGa 2) , abraham menCía 3) & daniel C. Passos 4) 1) Programa de Pós-Graduação em ecologia e Recursos naturais, universidade Federal de são Carlos, Rodovia Washington Luiz, Km 235, 13565-905, são Carlos, sP, Brazil. 2) Programa de Pós-Graduação em ecologia e Conservação, universidade Federal do mato Grosso do sul, av. Costa e silva, s/nº, 79070-900, Campo Grande, ms, Brazil. 3) Programa de Pós-Graduação em Biologia animal, universidade Federal do mato Grosso do sul, av. Costa e silva, s/nº, 79070-900, Campo Grande, ms, Brazil. 4) Programa de Pós-Graduação em ecologia e Conservação, Laboratório de ecologia e Comportamento animal, universidade Federal Rural do semi-Árido, Rua Francisco mota, 572, 59625-900, mossoró, Rn, Brazil.