Lizards Podarcis Muralis and Podarcis Hispanica in a Mountain Region of Central Spain
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HERPETOLOGICAL JOURNAL, Vol. 5, pp. 181-188 (1995) HABITAT SELECTION AND THERMAL ECOLOGY OF THE SYMPATRIC LIZARDS PODARCIS MURALIS AND PODARCIS HISPANICA IN A MOUNTAIN REGION OF CENTRAL SPAIN JA VIER MARTiN-VALLEJ0 1, JA VIER GARCiA-FERNANDEZ2, V ALENTiN PEREZ-MELLAD02•3 & JOSE LUIS VICENTE-VILLARDON 1 1 Departamento de Estadistica y Matematica Aplicadas, Universidad de Salamanca, Cl Espejo 2, E-37007 Salamanca, Sp ain 2 Departamento de Biologia Animal, Universidad de Salamanca, E-37071 Salamanca, Sp ain 3 Present address: Departamento de Ciencias Ambientales y Recursos Naturales, Universidad de Alicante, Sp ain We studied habitat selection, field body temperatures (TB) and activity cycles in sympatric populations of the lacertid lizards Podarcis muralis and Podarcis hispanica in a mountain area in Central Spain. Different patterns of habitat selection for the two species were found: P. muralis occupies N and NW facingtalus sites, while P. hispanica shows a less habitat-specific distribution at the study site. Inter- and intraspecific differences in TBs were found, while thermoregulatory precision analyses showed differences between adult male and female P. muralis. Hypotheses based on activity cycles, habitat selection and basking behaviour are discussed to explain such results in two closely related species. INTRODUCTION (3) inspect whether differences in thermal ecology could be foundbetween the two species. Thermoregulatory behaviour and habitat selection are intimately related and strongly influence life his MATERIAL AND METHODS tory traits and fitness of lizards (Ouboter, 1981; Huey, ANIMALS 1982; van Berkum, Huey & Adams, 1986; Adolph, 1990). Many authors have approached the study of in Lizards of the lacertid genus Podarcis from the Ibe teractions between the conditions of the habitat rian Peninsula seem to be very similar in basic occupied by a given species and its thermal behaviour ecological characteristics such as diet composition, (Hillman, 1969; Moermond, 1979; Roughgarden, Por foraging behaviour, activity cycles and thermoregula ter & Heckel, 1981; Hertz & Huey, 1981; Tracy & tory behaviour (Arnold & Burton, 1978; Christian, 1986; van Damme, Bauwens, Castilla & Perez-Mellado, l 983a,b; Arnold, 1987; Gosa, 1987). Verheyen, 1989; Marquet, Ortiz, Bozinovic & Jaksic, These similarities are especially strong between P. 1989), as well as focusing on the connection between muralis and P. hispanica, two species so closely related thermoregulation and activity time (Porter & Tracy, that there are some problems for their systematic dis 1983; Grant & Dunham, 1988). tinction over a wide part of their distribution range We here report a study of body temperatures, activ (e.g. Martinez-Rica & Laplaza, 1989). These two ity cycles and features of habitat selection in the lizards small lizards are similar in size (adult snout-vent Podarcis muralis and Podarcis hispanica, that are length 49.7-67.6 mm and 46.7-66.8 mm respectively, sympatric in some areas of the Iberian Peninsula, in personal data), morphology (Perez-Mellado & cluding the Sierra de Guadarrama (Central System, Galindo, 1986) and habitat requirements (Arnold, Spain), where this study was undertaken. The analysis 1987). Podarcis muralis occupies a wide range of lati of thermoregulatory precision, activity cycles and tudes in Europe -the southwestern extreme of which is basking behaviour have been developed through field the Sierra de Guadarrama-, while P. hispanica is lim observations, and the conclusions extracted are there ited to the Iberian Peninsula, southern France and fore mainly directed towards suggesting working northern Africa. hypotheses forfuture studies. STUDY AREA Our main objectives were to: (!) quantify habitat occupation by the two species in the area, and identify The study area is located in the westernmostpart of any intra- and interspecific differences between them; the Sierra de Guadarrama, Spain, a mountain range (2) determine some aspects of their thermal ecology, reaching a maximum altitude of 1902 m. The and relate these to the habitat selection features; and altitudinal range surveyed extended from 1450 to 1902 182 J. MARTIN-VALLEJO ET AL. m. The dominant vegetation consists of Pinus we estimated the av ailability of microhabitats in the sy lvestris forest, widely spread throughout the whole environment by measuring the same variables in mountain range. In the high mountain areas, the forest squares forwhich the centres were the points occupied was gradually replaced by small shrubs, with by us afterstoppin g every two minutes during random Juniperus communis subsp. nana being the dominant directed walks. species. Below 1550 m, the forestbecomes very open, STATISTICAL ANALYSIS and Quercus ilex subsp. ballota is the predominant tree species. Important habitat differencesocc urred re We used one-way analysis of variance (ANOVA) sulting from the topography: the southernslopes of the and Bonferroni's multiple comparison test (see mountains present a greater number of open habitats Keppel, 1991) on arcs in x·1 transformeddata to exam with Mediterranean shrubs such as Cistus laurifolius, ine differences between samples (P. mura/is, P. Halymium viscosum, Erica arborea and Thymus hispanica and av ailability) foreach microhabitat vari bracteatus. The northern slopes exhibit closed Pinus able. When variances were heterogeneous across forests with a few accompanying scrubs (mainly samples (Bartlett's test) we used the non-parametric Arctostaphyllos uva-ursi) and have much higher hu Kruskal-Wallis test, followed by Dunn's multiple com midity. parison test (Dunn, 1964; see Hollander & Wolfe, 1973). METHODS Thermoregulatory precision was estimated by the Observations were made at monthly intervals from slope of the least square regressions of TB on TA or May 1988 to September 1989. We captured lizards by TS. Differences between lizard species and age/sex hand or noose, and recorded the following data upon groups were assessed by analyses of covariance each capture: species, date, time, age, sex, reproductive (ANCOV A). As another indicator of thermoregulatory condition of females, behaviour (basking or active), precision we employed the interquartile range. To ana orientation of slope (N, NW, W, SW, S, SE, E, NE), lyse differences between two means we used the altitude, body temperature (cloaca! = TB), air tempera two-tailed Student t-test, with the Welch approxima ture (shaded bulb, 10 cm above substrate = TA), and tion (t) in case of heterogeneity of variances, or a substrate temperature (shaded bulb = TS). Tempera Z-test for n>50. Frequency distributions of distinct be tures were measured with a Schultheis thermometer to havioural categories were compared with x2 tests. The the nearest 0.1°C. significancele vel used was a= 0.05. Three ag e classes were considered: juveniles, the RESULTS individuals bornin the year of capture; subadults, one year-old individuals, which have not yet reached HABITAT SELECTION sexual maturity; and adults, sexually mature individu Podarcis muralis occupied a narrower altitudinal als. range in the study area (1490-1 725 m; Fig. 1), while P. Activity rhythms were determined from the frequen hispanica was present along the whole surveyed range cies of lizard observations made in the diffe rent (1450-1900 m). months and time intervals (data from 1988 and 1989 were pooled since no differences were detected). Cor rection indices (Telleria, 1986) were used to av oid possible biases caused by an irregular survey intensity. 1900 1900 The microhabitat selection field study was devel 1850 1850 oped between April and August 1989, at monthly 1800 intervals. A 5 x 5 m square was used as the survey unit, 1800 the centre of which was a lizard previously observed 1750 175 (these observations were made randomly, using two 1700 1700 hour intervals and considering the first lizard seen af 1650 1650 ter each of these periods of time). Within each square we estimated 28 variables: (1-5) percentage cover at 1600 1600 ground level of gravel, grass, fallen leaves, rock and 1550 1550 soil; (6- 10) percentage cover of vegetation layers with 1500 1500 heights <25 cm, 25-50 cm, 50-100 cm, 1-2 m, >2 m; (1 1-15) percentage cover of walls, and rocks with 1450 1450 heights <25 cm, 25-50 cm, 50- 100 cm, >100 cm; (16) percentage cover of talus structures; (17-18) percent 60 50 40 30 20 10 0 10 20 30 40 50 60 ag e cover of dead tree trunks and branches; ( 19-20) percentage cover of 0-20° and 20-40° slopes; (2 1 -28) FIG. I. Absolute frequencies (on horizontal axis) of SW, S, SE, E, NE, N, NW or W dominant orientations. Podarcis muralis (left hand) and Podarcis hispanica (right Percentages were calculated as means of estimates hand) lizards seen at the diffe rent altitude intervals (on made by eye by two independent observers. Similarly vertical axis). HABITAT SELECTION IN PODARCIS 183 TABLE I. Average percentages and standard deviations for TABLE 2. Results obtained in the habitat selection study. (* the three groups considered: Availability, P. muralis and P. P<0.05; ** P<0.0 1. The last three variables were analysed hispanica. using non-parametric tests). Availability P. hispanica P. muralis Variables Availability/ Availability/ P. muralis/ (n=45) (n=24) (n 22) = P. hispanica P. muralis P. hispanica mean SD mean SD mean SD Grass * * Soil * * Gravel 2.2 14.7 16.7 37.3 15.5 34.6 Rocks < 25cm * * Grass 72.6 41.4 41.7 49.3 13.2 30.3 Talus * * Withered Slope 0-20° * * leaves 8.8 28.5 0.0 0.0 0.0 0.0 Slope >20° * * Slab 4. 1 16.8 9.4 18.1 3.4 11.4 Trees >2m * * Soil 12.2 30.3 24.0 38.1 67.9 43.2 Shrubs < 25cm * * Trees: Rocks 25-50cm * * >2m 20.0 27.1 3.1 13.5 6.2 18.4 <2m 1.2 5.3 6.7 10. I 8.6 13.9 Shrubs: < 25cm 8.0 15.2 2.5 5.9 2.8 9.8 25-50cm 15.3 20.6 6.8 9.