Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 1

HERPETOZOA 26 (1/2): 77 - 90 77 Wien, 30. Juni 2013

Thermoregulation, activity and microhabitat selection in the rare and endangered Batuecan Rock , martinezricai (ARRiBAs , 1996) (: sauria: )

Thermoregulation, Aktivität und Mikrohabitatwahl der seltenen und gefährdeten Felseidechse Iberolacerta martinezricai (ARRiBAs , 1996) (squamata: sauria: Lacertidae)

OscAR J. A RRiBAs

KURZFAssUNG

Zur Naturgeschichte von Iberolacerta martinezricai (ARRiBAs , 1996), der seltensten Reptilienart Kontinen- taleuropas, liegen nur wenige Angaben vor. Das zahlenmäßige Geschlechterverhältnis betrug nach informationen aus der Datenbank des Autors 0,74:1 (Männchen:Weibchen), während im deutlich größeren Datensatz der Feld- beobachtungen (dieser schließt allerdings Beobachtungswiederholungen ein) die Männchen überwogen (1,27:1). Alle Altersgruppen und Geschlechter zeigten ähnliche, die Morgenstunden betonende Tagesaktivitätszyklen. Nachmittagsaktivität erfolgte gehäuft an schattigen Orten, wobei sonnenexposition bei hohen Lufttemperaturen vermieden wurde. Zwei Häutungsperioden (eine im April, eine um Ende Juni und Juli) wurden beobachtet. Der Lebensraum war mäßig geneigtes, ausnahmslos von Felstrümmern und großen Blöcken, mit reichlich Moos- und Flechtenbewuchs dominiertes Gelände, was auf erhöhte Feuchte und geologische Hang-stabilität hinwies. Besondere Vorlieben für einzelne Mikrohabitate waren nicht feststellbar. Die Körpertemperatur aktiver individuen war die höchste innerhalb der Gattung Iberolacerta : 33.31±0.12 °c (spannweite 22.8-39.2 °c), ohne signifikanten Geschlechtsunterschied. im Mittel hatten blaufärbige Männchen eine höhere Körpertemperatur als grüne (33.73±0.19 °c gegenüber 32.8±0.28 °c). Die Aufenthaltsorte adulter Weibchen wiesen eine höhere Lufttemperatur auf (23.77±0.20 °c, spannweite 13.8-37.1 °c) als die der Jungtiere. Die substrattemperaturen (30.08±0.31 °c, spannweite 16.5-52.1 °c) waren an den Aufenthaltsorten der Weibchen höher als an denen der Männchen. Iberolacerta martinezricai war mäßig thermokonformistisch (spannweite der Aktivitätstemperatur: 16.4 °c) und ein effektiver und ziemlich präziser Thermoregulator was die substrattemperatur betrifft ( r = 0.36, Anstieg 0.14), aber zeigte kein thermoregulatorisches Verhalten hinsichtlich der Lufttemperatur ( r = 0.15, Anstieg = 0.09). Tigmothermie, Aktivität im schatten bei hoher Lufttemperatur und Aestivation werden diskutiert. ABsTRAcT

The natural history of Iberolacerta martinezricai (ARRiBAs , 1996), the rarest of continental Europe, is practically unknown. sex ratio calculated from the author’s morphological database was 0.74:1 (males:females), whereas from the clearly higher number of field observations (repetitions included) was (1.27:1), favoring males. All sexes and ages revealed similar daily activity cycles, skewed toward the morning hours, with the after - noon activities concentrating at shadowy sites with elevated air temperatures and avoiding sun basking. There were two periods of molting, one around April and another between late June and July. The macro - was strictly rocky (large boulders covered with plentiful mosses and lichens that denote elevated moisture and geological habitat stability) in gentle steeped slopes. specific microhabitat selection was not observed. Body temperatures of active were the highest within the Iberolacerta : 33.31±0.12 °c (range: 22.8-39.2 °c), without significant differences among sexes or ages. On the average, blue males had higher body temperatures than green ones (33.73±0.19 °c versus 32.8±0.28 °c). Ambient air temperatures measured (23.77 ±0.20 °c, range: 13.8-37.1 °c) were higher at sites where adult females than where juveniles were observed. sub- strate temperatures (30.08±0.31 °c, range: 16.5-52.1 °c) were higher in places where females than where males were observed. Iberolacerta martinezricai is a moderate thermoconformist (total temperature range of activity 16.4 °c) and an effective and fairly precise thermoregulator with respect to substrate temperature ( r = 0.36, slope 0.14), but seemed not to thermoregulate relative to the air temperature ( r = 0.15, slope = 0.09). Tigmothermy, activity in the shadow with high ambient air temperatures and aestivation are also discussed. KEY WORDs Reptilia: squamata: sauria: Lacertidae; Iberolacerta , Iberolacerta martinezricai ; microhabitat, body tem - peratures, skin moults, habitat selection, thermoregulation, activity, ecology, behavior, physiology, sistema central, Las Batuecas y Peña de Francia Natural Park, spain Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 2

78 O. J. A RRiBAs

iNTRODUcTiON

Iberolacerta martinezricai (ARRiBAs , (ARRiBAs 1999a). His tentative estimation 1996) was described from the sierra de Peña of the population density in the top region of de Francia as a of cyreni the mountain (the only known locality of the MüLLER & H ELLMicH , 1937 , coincidental ) resulted in the theoretical density of with the raising of this latter to species level 45 individuals/ha or a total number of some (from former L. monticola cyreni ). The thirty individuals. presence of iberian Rock Lizards (at that Fortunately, I. martinezricai was later time collectively named Lacerta monticola localized in another area in the Batuecas BOULENGER , 1905 ) in the Mountain Peña de Valley ( ARRiBAs 2004), in the framework of Francia, Province of salamanca, spain, was genetic, osteological and karyological stud - known from KLEMMER (1957) who studied a ies that showed this lizard was not a sub - male collected by A. EPPLE in 1954. species of I. cyreni , but a valid species, When this German collector visited the more related to I. monticola than to the for - Peña de Francia (2-iX-1954) some days mer (MAYER & A RRiBAs 2003; cROcHET et before the annual pilgrimage to the mona - al. 2004; ARRiBAs & c ARRANZA 2004; ARRi- stery, only a dirt track constructed in 1920 BAs & O DiERNA 2004). existed. in 1961 the street was asphalted Iberolacerta martinezricai , is the facilitating massive access of motor vehi - rarest and probably one of the most threat - cles, and in the 1970’s, the characteristic an - ened reptile species from continental tenna was in stalled, well visible from a dis - Europe. A recent survey on the habitat and tance. These landmarks show the increasing distribution of I. martinezricai confirms that human pressure upon the area over the past its distribution area is very small (12-15 hundred years. km 2) inside three 10 km x 10 km UTM grid in the last decades of 20th century, squares (29TQE48; 29TQE38; 29TQE39). reference was made to the presence of Rock The total occupied area is between 20-25 Lizards in the western spanish sistema km 2 and the estimation of the total popula - central, in a paper about the morphology tion of the species between 1200 to 1500 in - and biology of L. monticola (PEREZ -MEL- dividuals (L iZANA & c ARBONERO 2007; cAR- LADO 1982) as well in the Atlas of amphib - BONERO pers. comm.). Despite the promis - ians and of the Province of sala - ing find of the species at a lower altitude manca ( PEREZ -M ELLADO 1983). There, re - site, the species has not been found in other ferring to the seventies, this author recalled such lowland environment or more to the attention to the rareness of the Peña de West, as previously expected. Francia population: “the situation is more Iberolacerta martinezricai is consid - serious in Peña de Francia where the num - ered critically Endangered (cR, B2ab(v); ber of visitors is growing every year and c2a(ii)), because of its small distribution where a dramatic numerical regression of area (< 10 km 2), population comprising less individuals, which are scarce by them - than 250 adults and its decline from various selves, has been produced in the last threats ( ARRiBAs 2006; PéREZ -M ELLADO et decade”. al. 2009). Despite that the entire population it was not until 1996 when ARRiBAs occurs inside a protected area, namely the described Lacerta cyreni martinezricai in “Parque Natural de las Batuecas y sierra de the framework of a wider revision of ‘ L. ’ Francia”, a specific program and manage - monticola and ‘ L.’ cyreni . After this, the ment for the recovery of this species is still author added new information about the required. Peña de Francia’s population including data However, in spite of this high threat from live specimens and again pointed to category, little is known on this lizard’s biol - the scarcity of the lizards and the increasing ogy and ecology (see ARRiBAs 2006 for a anthropogenic transformation of the area résumé of our knowledge). Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 3

Iberolacerta martinezricai , Batuecan Lizard, activity, microhabitat selection and thermoregulation 79

MATERiALs AND METHODs

study area and quartzite surface in the peaks and crests forms large areas of boulders in the slopes of Data processed in this study come the valleys, where slates are more common. from the whole altitudinal range of the in the area, three bioclimatic stages species; from the Peña de Francia summit to are represented: the Oromediterranean belt the Puerto del Portillo in the Batuecas (>1600 m a.s.l.), only in the summits of the Valley (between 886 and 1735 m a.s.l.). mountain ranges, with broom shrubs (main - The main area of the species lies in the ly Cytisus balansae ); the supramediterrane - north of the Natural Park of Las Batuecas, at an belt (approx. 1000-1600 m a.s.l.) origi - medium and high altitudes in the mountain - nally vegetated by Pyrenean oak ( Quercus ous range of the sierra de Francia. This first pyrenaica ) and a shrub stratum of heath area is centered around the peak “Peña de (Erica arborea and E. australis ), as well as Francia” (1723 m), extending northward to some pine stands ( Pinus nigra and P. the peak “Pico de los Robledos” (1611 m) sylvestris ) interspersed. The low altitudes and surrounding areas, to the northeast as are occupied by the Mesomediterranean belt far as the “sierra del Guindo” (“Hastiala” (400-1000 m a.s.l.) typically constituted by Peak, 1735 m; “Alto del copero”, 1560 m), open forests of Evergreen oaks ( Quercus southward to the “Mesa del Francés” (1640 ilex ) and cork oaks ( Quercus suber ) in the m), and toward the southeast, up to the wetter areas. Azonal vegetation includes northern slopes of the “sierra de la Grajera” saxicolous and riverine species in the (“Rongiero”, 1627 m) ( ARRiBAs 2006; ravines ( ARRiBAs 2004, 2006; LiZANA & LiZANA & c ARBONERO 2007). in the peaks, cARBONERO 2007). records of the species are merely anecdotal, Rainfall is concentrated during fall with densities below 10 individuals/ha in and winter and exceeds 1800 mm in the the few sites where it was found (densities summit areas. summers are sunny, with of 45 individuals/ha estimated in the previ - high insolation and lack of precipitation; ous decade, show the marked regression in drought is restricted to July-september. in parallel to increased human activity). in the lizard , annual and daily insola - other places of the area, the population can tion cycles are relatively short (just a few reach up to 60 individuals/ha ( LiZANA & hours of direct sun per day), due to the relief cARBO NERO 2007). of the mountains and the orientation and in - The second area where the Batuecan clination of the slopes. The sun reaches the Rock Lizard occurs is called “Puerto El habitats late in the morning, especially the Portillo” (“El Portillo Pass”, from 840 m to lowermost portions of the slopes ( MARTiN 1400 m a.s.l.), on the southern slope of a 1998 ; ARRiBAs unpublished data). mountain range named “sierra de la Alberca”. This mountain-wall drops away Field data from the Meseta Norte to the considerably lower Batuecas Valley. its population Localities were visited from 1996 to which was only described in 2004 ( AR - 2012 at irregular intervals: 1997 (April, July, RiBAs 2004) exhibits densities of 25-30 in - and August), 2001 (July and August), and in dividuals/ha (LiZANA & c ARBONERO 2007) . July of 1996, 2002, 2003, 2004, 2005, 2007, The bedrock of the area is mainly of Paleo - 2008, 2011 and 2012. calcula tions were zoic age, with granites, quartzite and slate based on time, habitat and temperature data outcrops. gathered from 619 observ ations of I. mar - The relief is generally smooth but tinezricai . shows active geological phenomena of From each specimen found active, the meteorisation, which produced accumula - following data was taken: tions of large boulders and loose rocks - Date and time of the day (GMT). (called “canchales”, averaging 80-120 cm in - Activity: [inactive, basking, and diameter) and deep valleys with fast flow ing active]. Temperature data (see below) were streams during the winter. The basal granite taken from active specimens only. Activity Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 4

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was quantified by counting the observation Four parameters were used to define events of active individuals, irrespective of the thermoregulatory behavior: the duration of the observation time. Mean Body temperature: The average - sex [coding: 1 - male, 2 - fe male, of temperatures at which the lizards were and 3 - adult of undetermined sex]. Those found active. specimens of undetermined sex were used Thermoconformity : The larger the for global adult comparisons with young range of the body temperature, the more individuals only. pronounced is the thermoconformity. its - Approximate age [adult (coding: standard deviation or standard error of the 1 - males, 2 - females, 3 - specimens of mean are significant measures and resistant undetermined sex), subadults (later also to outliers, but are not available from other called young, juvenile or immature speci - studies for comparison. mens and hatchlings)]. subadults were clas - Efficiency of thermoregulation: De - si fied in calendar-year age classes (coded gree of deviation from the perfect poikilo - “40” if of undetermined age). Hatchlings of thermy (the slope in a diagram of body ver - the year are in their “first calendar year” sus ambient temperatures is equal to 1). This (1cY; coded “41”). After their first hiberna - measures the capacity of the lizards to devi - tion they are in their “second calendar year” ate from the environmental temperatures. (2cY; “42”); after their second hibernation Precision of thermoregulation: Mea - they are in their “third calendar year” (3cY; sured as Pearson’s correlation between body “43”), etc. in total, 40 specimens were stud - and ambient air temperatures or body and ied from: 1st cY, 11 from 2nd cY, 20 from substrate temperatures. 3rd cY, 4 from 4th cY and 1 young of unde - - Habitat availability was calcu - termined age. lated by photointerpretation, overlaying a - Habitat structures (in percent): grid and calculating the percent covers of in a 2-meter radius around the spot of first rocks, stones, bare ground, shrubs and grass localization of the , a visual estimate in the different squares. of the percent cover of rocks, stones, bare Availability and use of resources ground, shrubs and grass was assessed by (Table 1) were compared by testing the null means of visual estimation scales ( PRODON hypothesis of random selectivity, using a & L EBRETON 1981; EMBERGER 1983). Chi -square log-likelihood statistics (G-test; - slope inclination was measured p < 0.05). selection or avoidance of habi - with a clinometer in discrete categories: 0°- tats was quantified by means of an electivi - 10°, 10°-20°, 20°-40°, 40°-50°, 50°-70°, ty index, a normalized forage ratio: w i = o i / 70°-80°, 80°-90°). pi; where w i = forage ratio for the habitat - Temperatures were taken (from category i; o i = proportion or percent of the active specimens only) with schultheis-type habitat category i used; p i = proportion or thermometers (Wesco ®, 0 °c-50 °c, preci - percent of the habitat category i available in sion: ± 0.1 °c) from the cloaca (body tem - the environment ( KREBs 1989, 1999). perature, BT) within 20 seconds of the Electivity indexes are presented as standard - lizards’ first spotting, and ambient air (AT) ized ratios, which sum up to 1.0 for all habi - in the shadow, 20 cm above the ground. tat categories. standardized ratios of 1/ substrate temperatures (sT) were taken with (number of resources) (in our case 1/7 = a TFA® infrared thermometer. Mercury 0.14) point to an indifferent situation (nei - and electronic thermometers were inade - ther preference nor avoidance); values quate instruments to obtain reliable temper - below this indicate relative avoidance, and atures from a surface, as the contact area above relative preference (calculations are with the measured object is not two-dimen - after KREBs 1989, 1999). Only deviations sional but punctual, whereas infrared ther - greater than 5 % ( p < 0.05) were considered mometers measure temperature from a cir - and discussed in the text. cular surface area and were most suitable Total counts differ among variables for measuring substrate temperatures (but and age-classes, as they were not consistent - totally unsuitable to take body tem - ly available for each observation. For in- peratures; see ARRiBAs 2009). stance, data from an adult of unknown sex Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 5

Iberolacerta martinezricai , Batuecan Lizard, activity, microhabitat selection and thermoregulation 81

or juvenile of unknown age were not includ - Ns (not significant) was stated. Diversity ed under males, females, or the definite age measurements and selection indexes were categories, but in the collective comparisons calculated with the program Ecological of adults against juveniles. Methodology ® 6.1.1 ( KREBs 1999). The following comparisons between categories statistical study were done: - separately for all sex and age cate - in general, statistical calculations gories (see under field data above): (“sex were made using the Ncss-2002 ® software and ages”) (Tables 1 and 2). package ( HiNTZE 2001). Univariate com - - Adult males versus adult females parisons among categories were done by versus juveniles. ANOVA with subsequent Tukey-Kramer - Adults versus subadults: Adults cate - post-hoc multiple comparison tests. At p < gory included adult individuals of undeter - 0.05, results were considered significant, at mined sex. p < 0.01 highly significant. concrete p val - - Differences in the temperatures be- ues were given only in the case of signifi - tween the green and blue color morphs cance or near significance, otherwise only (ARRiBAs et al 2008).

REsULTs AND DiscUssiON

sex ratio ARRiBAs & c ARRANZA 2004 and ARRiBAs 2007), based on a smaller number of 50 sex ratio of natural populations was database records, was similar to the current calculated from specimens of the author’s one. in Iberolacerta aranica (A RRiBAs , morphological database, and from speci - 1993), the proportion of sexes in adults was mens observed in the field during transect different from 1:1 (0.55:1), favoring surveys (certainly with specimens repeti - females (however, hatchlings’ sex ratio in tion). Results of both approaches are quite the laboratory was the contrary (1.8:1); this different. shift may be explained by temperature- in specimens from field observations depending sex determination; see ARRiBAs (usually July - August), the sex ratio was 1997). in I. cyreni more females than males 1.27:1 (260 observations of male versus (0.69:1) were observed. The usual situation 204 of female individuals). The difference in Rock Lizard species, however, is that the in the proportion of sexes was significant proportion does not differ significantly from (Multinomial test: Chi 2 = 6.75, P = 0.009). 1:1 as in I. bonnali (L ANTZ , 1927) (0.98:1), The male biased counts may be due to the I. aurelioi (A RRi BAs , 1994) (1.2:1), I. hor - timing of the fieldwork, which was mainly vathi (M éHELY , 1904) (1.01:1) and I. monti - done in the first week of July, and in some cola (1.14:1). in this last species however, years coincided with the gravid period for the females outnumbered the males (0.27:1) females which makes them more stealthy in the highest places of the serra de Estrela, and less detectable. This is probably also Portugal, where densities are extremely high the reason for the generally increased (all above Iberolacerta data from ARRiBAs abundance of male observations in this 1997). study. in laboratory-born/bred I. monticola in the specimens of the morphological and I. cyreni , the sex ratio did not differ sig - database (captured from from April to nificantly from 1:1, although, it happened August), the sex ratio was 0.74:1 (29 males frequently that in lacertids, slightly more and 39 females determined); which does not males than females were born (1.17:1 in I. differ significantly from 1:1 (Multinomial monticola , 1.21:1 in I. aurelioi , 1.8:1 in I. test: Chi 2 = 1.47, Ns). aranica [data from BARBADiLLO 1985; RúA A previously published sex ratio for I. & G ALáN 2003; ARRiBAs & G ALáN 2005] ). martinezricai (males:females = 0.78:1 in The samples of I. martinezricai and I. bon - Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 6

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Table 1: Iberolacerta martinezricai (ARRiBAs , 1996). Habitat structures at the sites of first spotting and selection of these structures with respect to their availability. First line: arithmetic mean ± standard error. second line: in parentheses, minimum and maximum values. Third line: selection index (w, see Materials and Methods). Fourth line: standardized proportions in % . Positive or negative signs indicate selection or avoidance. Only devi - ations exceeding 5 % ( p < 0.05; highlighted in bold) are considered and discussed in the text. Habitat category entries in lines three and four, respectively, sum up to 1.0 in each column. in the lower part of the table, G-test val - ues and their probabilities are indicated. Tab. 1: Iberolacerta martinezricai (ARRiBAs , 1996). Habitatstrukturen, jeweils am Ort der ersten Beobach- tung und deren selektion in Bezug zur Verfügbarkeit. Erste Zeile: arithmetisches Mittel ± standardfehler. Zweite Zeile: in Klammern, Minimum- und Maximumwerte. Dritte Zeile: selektionsindex (w, siehe Material und Metho- den). Vierte Zeile: normalisierte Anteile (%). Positive bzw. negative Vorzeichen bedeuten selektion bzw. Vermei- dung. Nur Abweichungen über 5 % ( p < 0,05; in Fettschrift) sind berücksichtigt und im Text diskutiert. Die summe für alle Habitatkategorien je spalte ergibt für die Einträge in den Zeilen drei und vier jeweils den Wert 1,0. im unteren Tabellenteil werden G-Werte und ihre Wahrscheinlichkeit angegeben.

sex and Ages Males Females Young Availability in the habitat Geschlecht und Alter (n = 49) (n = 28) (n = 85) (%) Männchen Weibchen Junge Verfügbarkeit im Lebensraum inclination of habitat 26.87±2.2 24.57±2.6 23.38±1.5 ) 24.5±1.10 (angular degrees) (0-90) (0-70) (0-80) Hangneigung (Grad) w = 0.2175 w = 0.1825 w = 0.1074 + 7.4 % + 3.9 % - 3.5 % % Rocks 92.79±2.4 92.5±2.5 86.7±2.6 89.7±1.59 % Felsen (30-100) (60-100) (0-100) w = 0.2057 w = 0.1882 w = 0.1091 + 6.2 % + 4.5 % - 3.3 % % stones 0±0 0±0 13.96±2.2 0.93±0.57 % steine (0-0) (0-0) (0-70) w = 0.0001 w = 0.0001 w = 0.2220 - 14.2 % - 14.2 % + 7.9 % % Bare soil 0.34±0.3 0.96±0.9 1.64±0.9 1.09±0.51 % Nackter Boden 0-15 0-25 0-60 w = 0.0620 w = 0.1607 w = 0.1698 - 8.08 % + 1.7 % + 2.7 % % shrubs 1.39±0.8 0.38±0.3 1.96±1.0 1.45±0.60 % Gebüsch (0-30) (0-10) (0-80) w = 0.1906 w = 0.0478 w = 0.1525 + 4.7 % - 9.5 % + 0.9 % % Grass 3.37±1.5 5.76±2.4 5.69±1.3 5.06±0.90 % Gras (0-40) (0-40) (0-40) w = 0.1324 w = 0.2077 w = 0.1269 - 1 % + 6.4 % - 1.6 % % Moss/Lichens 10.89±2 13.18±3.3 11.25±1.6 11.29±1.15 % Moos/Flechten (0-30) (5-30) (0-30) w = 0.1918 w = 0.2130 w = 0.1124 + 4.9 % + 7.02 % - 3.04 % G-test ( df = 6) 2.19 2.160 0.595 p 0.9007 0.9044 0.9953

nali are of insufficient size to derive any and age categories analyzed ( F5,606 = 0.43, conclusion in this respect. Ns; see Table 2 for details of each category), even if they were grouped into only three Activity categories (males, females and juveniles) (F2,600 = 0.14, Ns) [Males: n = 259, mean = The circadian activities (expressed by 1043± 12.93, 742-1705; Females: n = 201, the number of observation events) of the mean = 1056±14.71, 756-1722; Juveniles: n Batuecan Lizard are visualized in Fig. 1. = 143, mean = 1040±15.26, 726-1653]. There were no significant differences in the Equally, juvenile specimens showed identi - daily activity schedule among all the sex cal patterns to adults ( F1,610 = 0.19, Ns Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 7

Iberolacerta martinezricai , Batuecan Lizard, activity, microhabitat selection and thermoregulation 83

Males Females Juveniles

Fig. 1: Bar graph representing the number of active specimens (males, females and juveniles) of Iberolacerta martinezricai (ARRiBAs , 1996) as observed during the hours of the day ( n = 612 observations). Abb. 1: Balkendiagramm der Anzahl aktiver individuen (Männchen, Weibchen und Jungtiere) von Iberolacerta martinezricai (ARRiBAs , 1996) aufgrund von Beobachtungen im Tagesverlauf ( n = 612 Beobachtungen).

[Adults: n = 469; mean = 1048±9.55; 742- only when running from crevice to crevice, 1722; Juve niles: n = 143; mean = which makes the author suppose that there 1040±15.26; 726-1653). exists a certain degree of subterranean and The magnitude of the mean activity is less detectable activity in crevices among similar within the three age and sex classes the large boulders, that retain fresh air dur - studied, and its period extends over 9 or 10 ing the hottest hours of the day ( ARRiBAs hours per day. All three classes exhibited 2004; A RRiBAs & c ARRANZA 2004). activity curves skewed towards the morning hours, whereas the afternoon was usually Molts spent at shadowy places with air tempera - tures warm enough to allow for activity The first molting of the year occurred without sun basking and the risk of over - just shortly after leaving the hibernacula heating. During the morning hours, the ani - (end of March or early April), perhaps mals basked until they reached activity tem - somewhat earlier in males (3 molted and 5 peratures, appropriate to begin to hunt and unmolted seen at this moment) than in fe - display social behavior. During hot weath - males (1 molted versus 3 not molted). These er, the extent of this morning sun basking proportions are indicative but a sound finding could be reduced strongly if the air temper - would require further analyses of a larger ature sufficed for being active. At mid-sum - sample. At the end of April, all individuals mer, after the end of the reproduction period observed were molted. (mid-July), lizard activity declined, in part There is no data from May and June so due to the excessively hot and dry weather that it is not clear whether or not there is conditions. Then, sun basking, was restrict - another molt between the first and the next ed to the first hours of their total daily activ - observed molt seen towards the end of June ity. As the day proceeded, lizards were seen and early July in males, and a little ad - Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 8

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Table 2: Iberolacerta martinezricai (ARRiBAs , 1996). Habitat structures [inclination of slope (°, angular degrees), proportion (%) of rocks, stones, bare soil, shrubs, grass and moss/lichens in the place where the individ - ual was spotted], time of activity (GMT) and temperatures (°c) of body (cloaca, BT), ambient air (AT) and sub - strate (sT) for all the sexand age classes [2cY, 3 cY - juveniles at their second and third calendar year, respec - tively] studied. First line: arithmetic mean ± standard error. second line: minimum and maximum values. The sample named ‘ Σ Adults’ includes males and females plus adults of undetermined sex. Temperatures refer to active specimens only. sample size as indicated in the column headers unless otherwise noted. --- No data. Tab. 2: Iberolacerta martinezricai (ARRiBAs , 1996). Habitatstrukturen [Geländeneigung (°, Winkelgrade), Anteil (%) an Felsen, steinen, nacktem Edboden, Gebüsch, Gras und Moos/Flechten am Aufenthaltsort des individuums], Uhrzeit der Aktivität (GMT) und die Temperaturen (°c) von Körper (Kloakaltemperatur, BT), umgebender Luft (AT) und substrat (sT) für die untersuchten Geschlechter- und Altersklassen [2cY, 3 cY – Jungtiere in ihrem zweiten und dritten Kalenderjahr]. Erste Zeile: arithmetisches Mittel ± standardfehler. Zweite Zeile: Minimum- and Maximumwerte. Die stichprobe ‘ Σ Adults’ umfaßt die Männchen, Weibchen und Erwach- senen unbestimmten Geschlechtes. Temperaturangaben beziehen sich nur auf aktive individuen. Die stich- probenumfänge der spaltenköpfe gelten - falls nicht anders angegeben - für die ganze spalte. --- Keine Daten.

sex and Ages Males Females immatures immatures Σ Adults Σ Young Geschlecht und Alter Männchen Weibchen Junge Junge Σ Adulte Σ Junge (n = 49) (n = 28) (2cY)( n = 36) (3cY)( n = 44) (n = 84) (n = 85) Time of the day 1043±12.93 1056±14.71 1043±29.8 1041±18.17 1048±9.55 1040±15.26 (GMT) 742-1705 756-1722 726-1653 743-1628 742-1722 726-1653 Tageszeit (n = 259) (n = 201) (n = 36) (n = 100) (n = 469) (n = 143) inclination of habitat 26.87±2.26 24.57±2.6 23.14±2.32 24.04±2.21 25.78±1.59 23.38±1.52 Geländeneigung (°) 0-90 0-70 0-80 0-70 0-90 0-80 % Rocks 92.79±2.45 92.5±2.57 82.94±4.8 88.59±3.27 92.82±1.66 86.7±2.64 % Fels 30-100 60-100 0-100 0-100 30-100 0-100 % stones 0±0 0±0 3.67±2.5 0.51±0.51 0±0 1.83±1.11 % steine 0-0 0-0 0-80 0-20 0-0 0-80 % Bare soil 0.34±0.34 0.96±0.96 3.82±2.15 0.52±0.38 0.52±0.38 1.64±0.94 % Erdboden 0-15 0-25 0-60 0-25 0-25 0-60 % shrubs 1.39±0.85 0.38±0.38 1.32±0.77 2.82±2.07 0.92±0.5 1.96±1.07 % Gebüsch 0-30 0-10 0-20 0-80 0-30 0-80 %Grass 3.37±1.5 5.76±2.41 6.17±2.23 5.89±1.87 4.40±1.22 5.69±1.33 % Gras 0-40 0-40 0-40 0-40 0-40 0-40 % Moss/Lichens 10.89±2 13.18±3.32 7.75±1.79 15.62±2.65 11.34±1.62 11.25±1.65 % Moos 0-30 5-30 0-30 5-30 0-30 0-30 BT (°c) 33.52±0.17 33.00±0.20 --- 33.34±0.39 33.31±0.13 33.34±0.39 25-39.2 22.8-36.8 --- 28.3-37.9 22.8-39.2 28.3-37.9 (n = 166) (n = 115) (n = 19) (n = 37) (n = 281) (n = 37) AT (°c) 23.64±0.28 24.7±0.37 20.04±0.67 23.47±0.54 24.05±0.22 22.58±0.47 13.9-34.7 13.8-37.1 16-26.5 13.9-30 13.8-37.1 13.9-30 sT (°c) 29.35±0.43 31.55±0.15 24.96±0.97 31.36±1.01 30.19±0.34 29.63±0.84 18.6-52.1 16.5-50.2 19.9-32.7 16.5-46.1 16.5-52.1 16.5-46.1

vanced towards the second half of June in in other Iberolacerta species, differ - females. Early in July, about half of the ences were found in the molting cycles of males and almost all females appeared most males and females. in male I. aranica there recently molted. in the second half of July, are two defined periods of molting during all males and of course females had finished the activity period (concentrated around molting. in August there were no traces of beginning of July and the second week of molting. For september, information on August, respectively), whereas in females molting activities is lacking. in summary, and juveniles there are three (two similar in only two molts were noted during the activ - date to the males’ periods, the August molt ity period (April to september): one that is perhaps slightly earlier, plus a third one centered around April and another between towards the second week in september) ( AR - late June and July. RiBAs 2007). in I. horvathi there are also Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 9

Iberolacerta martinezricai , Batuecan Lizard, activity, microhabitat selection and thermoregulation 85

Fig. 2: Habitat of Iberolacerta martinezricai (A RRiBAs , 1996) . The species only occurs at these big, geologically stable, rock boulders (or equivalent old stone walls). inside this macrohabitat the lizard does not select particular microhabitat characteristics. Abb. 2: Lebensraum von Iberolacerta martinezricai (A RRiBAs , 1996) . Die Art kommt nur an diesen großen, geologisch stabilen Felsblöcken (oder vergleichbaren steinwänden) vor. innerhalb dieses Lebensraumes bevorzugt die Eidechse bestimmte Mikrohabitatstrukturen nicht deutlich.

Fig. 3: Male blue morph of Iberolacerta martinezricai (ARRiBAs , 1996) basking. Peña de Francia (salamanca, spain). it is shown that males of the blue morph have slightly higher body temperatures than green morph males. Abb. 3: Männliche blaue Morphe von Iberolacerta martinezricai (ARRiBAs , 1996) beim sonnen. Peña de Francia (salamanca, spanien). Es wird gezeigt, daß Männchen der blauen Morphe eine leicht höhere Körpertemperatur haben als die der grünen Morphe. Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 10

86 O. J. A RRiBAs

three molts, one in spring (first half of ty, and would not have developed to such an May), one in summer (July) and one in extent if the rocks were geologically active autumn (towards end of september or early (i.e., moving). The presence of large rock in October) ( LAPiNi et al. 1993). in other accumulations is important for the availabil - lacertids there are four or even more molts ity of deep crevices which represent shelters but during longer activity cycles (for a containing cool and fresh air, allowing for résumé see ARRiBAs 2007). the lizards’ subterranean daytime activities even when external temperatures are too high ( ARRiBAs 2004; A RRiBAs & c ARRANZA Habitat selection 2004). in the center of these rock boulders, Iberolacerta martinezricai is a strictly lizards are less exposed to enemies than in saxicolous species. its presence is narrowly the outer ecotonal areas, although the Weasel linked to large boulders and talus screes and (Mustela nivalis ) and Vipera latastei BOscA , thus, cannot be found in other habitats (Fig. 1878 were seen hunting in sympatry with I. 2) ( ARRiBAs 2004; A RRiBAs & c AR RANZA martinezricai (Coronella austriaca LAU- 2004; A RRiBAs 2006; LiZANA & c ARBONERO RENTi , 1768 was detected but not in the bare 2007). center of big rock boulders.). Also flying Habitat use results, detailed by sex predators such as kestrels may predate upon and age are presented in Table 2, and habi - lizards in these areas. in the bare rock areas, tat availability and selection tests in Table 1. the trophic resources exploited by lizards Regarding habitat preferences, there was a constitute what VALVERDE (1957) called tendency in males towards more stepped “aerial plankton”, i. e., Diptera, flying bee - slopes, rocky surfaces and avoidance of tles and carried to the rocks largely bare ground and shrubby places; in females by air currents, and developed outside the towards the increased use of grassy and lizard’s habitat ( ARRiBAs unpublished). mossy spots, and avoidance of shrubby and, especially, stony areas; and in juveniles Thermoregulation towards these stony habitats. Nonetheless, the null hypothesis (i. e., absence of prefer - Body temperatures (BT).- De - ences for any particular habitat types in the tails of the body, substrate and air tempera - categories males, females and juveniles) tures with regard to sex and age classes are cannot be rejected, as in all three cases, G- shown in Table 2. The mean cloacal body tests were not significant (Table 1). in sum - temperature (BT) in active I. martinez ricai mary, the species is strictly saxicolous with - was 33.31±0.12 °c (range = 22.8-39.2 °c; n out preferences for particular substructures = 318 measurements). Body temperature within this habitat. did not differ significantly among sexes and None of the ANOVA comparisons ages ( F2,315 = 1.83, Ns), or ages alone among the different habitat categories and (adults vs. juveniles; F1,316 = 0.01, Ns). factors (males, females and young separate - However, a significant difference was ly, or adults against juveniles) led to signif - found between the BTs of specimens of the icant results. significance level was barely blue and green morphs ( T248 = 2.68, P = approached solely in the use of rocks when 0.007) [green morph: n = 105, mean = 32.95 adults and juveniles were compared ( F1,153 ±0.20 °c, range = 25.0-36.4 °c; blue morph: = 3.77; P = 0.054), in that rocky habitat was n = 145, mean = 33.73±0.19 °c, range = slightly less preferred in the juveniles. This 22.8-39.2 °c], where the temperature of blue age class occurred more frequently in stony specimens was slightly higher than of green areas (as is typical in lacertids, where juve - ones. strictly speaking, this difference arose niles are the pioneer age class). from the male specimens ( T163 = -3.31, P = The species lives in gently stepped 0.001) [green males: n = 66, mean = slopes where large boulders are dominant, 32.8±0.28 °c, range = 25-36.3 °c; blue with mosses and lichens growing on their males: n = 99, mean = 34.00±0.22 °c, range sur faces. These plants indicate compara - = 29.2-39.2 °c], not from the female temper - tively moist conditions and boulder stabili - atures which did not differ among the morphs Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 11

Iberolacerta martinezricai , Batuecan Lizard, activity, microhabitat selection and thermoregulation 87

(T = 0.03, Ns) [green females: n = 39, 8

t 83 i 8 3 e 9 9 mean = 33.15±0.26 °c, range = 30.3-36.4 b 1 9 r 1 R

A °c; blue females: n = 46, mean = 33.13±0.40

O R 2 9 0 0 e D O 9 s 0 1 1 °c, range = 22.8-36.8 °c]. As a hypothesis, e A D e 9 0 0 0 i c V A 1 e 2 2 2 d L n l V different BTs of blue and green males could

l e A / A L s s s

e r c A A A A s r e u mirror differences in the biological compe - U B B B e f s i i i

Q e p & L R R R a &

R tence and fitness of both color morphs, R R R E O p

L N A A A t D i

L which merits in depth study (Fig. 3). n T E e R U s

A Air temperatures (AT).- Ambi - e G r R M P ent air temperatures when lizards were act- A ive [ n = 394, mean = 23.77±0.20 °c, range g e

i = 13.8-37.1 °c], differed between adults

t s T n 9 1 2 8 6 6 and juveniles ( F = 7.66, P = 0.005). in - A 1,392 0 2 5 2 7 3 - - A ...... - / T 0 0 0 0 0 0

e particular, this difference arose from the B p o .

l comparison of females (active at higher air n s e

t temperatures) and juveniles ( F = 6.51, P

g 2,388 a e i D t = 0.001). For details of air temperatures rel -

s e T n 4 4 0 5 0 1 n - s ative to sex and age classes see Table 2. i 1 1 1 3 5 2 - - A ...... e - / T 0 0 0 0 0 0 e

K summer temperatures in the area can reach B

p

o - l

- high values (maximum measured was 37.1 - s

. °c, but can probably be higher during ther - . a a t t n r

a mal waves of African air masses).

e o 5 3 6 3 7 9 d i c

T t 1 3 5 4 7 4 ...... a o a substrate temperatures (sT).- - A l l 0 0 0 0 0 0 - -

o N e -

T r r ======substrate temperatures at which lizards r e - B o r r r r r r - b - I

were active [ n = 393, mean = 30.08±0.31 c

. g s n °c, range = 16.5-52.1 °c] did not differ e u i t c t between adults and juveniles ( F = 0.47, e a 1,391 p n G s

o 6 7 6 8 4 8 Ns), but among adults, females were found i r t T a 3 3 5 3 6 5 e t ...... a - s d r l 0 0 0 0 0 0 active at higher substrate temperatures than - -

e e - T n r c ======r e B

a males ( F = 4.99, P = 0.007). substrate t 2,388 o r r r r r r l r o c r A temperatures according to sex and age class -

e n b e

I es are summarized in Table 2. )

b n c e i ) ° e Thermoregulative strategies.- s ( v

c i e e °

e t s ( Thermoregulation parameters of the various 4 8 5 8 4 i

. . . . b . 5 5 e

e n 6 6 4 7 4 i 1 1 n g Iberolacerta species are compared in Table 3.

w 1 1 1 1 1 e s n n r ß a n

e Iberolacerta martinezricai showed the ö t a R r e p g highest mean body temperature (33.3 °c) s m n a n r

e within the genus (from 28.13 °c in I. aure -

a K 2 2 2 4 4 n p . . . . .

i

5 r

. lioi to 29.4 °c in I. aranica and I. cyreni ). 9 5 5 5 5 4 y ) e 6 r M 3 3 3 3 3 3 - h ------c 3 o This high BT plausibly evolved during a º t - c x 8 4 8 4 0 6 ( . . . . . a s a 2 2 l i 2 8 0 0 6 2 r period of adaptation to the Mediterranean u 2 1 2 2 1 M o g t e a climate: this did, however, not happen in r l o u

g other congeneric species. Nonetheless, BT m e r r e

o of I. martinezricai is most similar to the h

t

m T r e median temperature calculated from 53 lac - e 6 9 9 3 B v ) 3 4 3

i h . . 9 . 4 6 1 t t . . . . n c

ertid lizard species (33.8 ºc), mainly not 3 9 9 a º a 8 9 8 8 r h ( 3 2 2 e 2 2 2 2 a c AsTiLLA

i high mountain species ( c et al. p M e l m

g 1999). These authors claimed that within a o r e c

given species, lizards, if freed from other V

i

:

a : 3

c constraints, frequently show fairly constant 3 i e

r a l . l z i b b AN

r mean BTs (thermal rigidity sensu V i i o e a h a a l e o t c i n c t i a i T T i i a l t e

n AMME

t D et al. 1989), whereas even among n n v e e r n n n r r m a r a o o o r u o a y closely related species, body temperatures r x m b m c a a h

a a ...... I P I I I T I I I tend to be different and vary related to the Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 12

88 O. J. A RRiBAs

lizards’ biogeographical origins rather than Regarding air temperatures (AT), the current environmental conditions ( BAU- slope of the regression line is even more dif - WENs et al. 1995). The BTs found in I. mar - ferent from 1, but the correlation is so low tinezricai do not comply with this model, in that there is no indication that the lizard being clearly higher than of other congeners thermoregulates in this respect. in fact they and rather in agreement with the ecological seem to do nothing in particular reaction to characteristics of the inhabited localities. increased ambient air temperatures; they The range of observed BTs at which I. hide when ATs rise in parallel to the habi - martinezricai was active (39.2-22.8 = 16.4 tat’s increased exposure to the sun, to cool °c) was similar as in other low mountain in the crevices and later utilize high ATs to Iberolacerta , which all are moderate ther - be active without direct sun-basking. moconformists and able to deploy their ac- To visualize the precision of ther - tivity within a rather wide range of temper - moregulation in I. martinezricai , the body atures (for references see Table 3). The max - temperatures (BT) and their relationships imum body temperature registered (39.2 °c) with both air (AT) and substrate tempera - was exceptional, as already short-termed tures (sT) are shown in Figs. 4A and 4B. exposure to BTs above 40 °c is lethal correlation of BT was higher with sT ( r = (ARRiBAs 2007 and unpublished) if the lizard 0.36; P = 0.0001) than with AT ( r = 0.15; P is not given the opportunity to hide and cool. = 0.0000). This difference was still present Even at substrate temperatures as high as when the effect of the third variable was 47.8 °c, the BT of a lizard that had taken extracted (partial correlation BT-sT, effect refuge under a stone was only 36.5 °c. of AT extracted: r = 0.33; partial correlation The slopes of the BT versus sT and BT-AT, effect of sT extracted: r = 0.07). As AT correlations were significantly different a result, the species is fairly precise when from 1 (if there were no thermoregulatory thermoregulating relative to the substrate behavior, the lizards’ rhythm of warming up temperature (sT) but totally imprecise (or would exactly parallel the heating rhythm of not at all thermoregulating) with air temper - the substrate or air, resulting in a slope of 1). ature (AT). in the relationship between BT and sT, the Once the BT declined below activity slope was 0.14 (BT = 28.9225 + 0.1396 sT; level, warmth gained by tigmothermy (e.g., R2 = 0.1324), whereas it was 0.09 in the when lying under a flat stone on top of a relationship of BT and AT (BT = 31.0305 + warm big rock) may not result in BT rise 0.0929 AT; R2 = 0.0254). sufficient to resume activity but to continue This data shows that I. martinezricai digestion for a longer time. is an effective (and fairly precise, see below in summer, when ambient morning paragraphs) thermoregulator in respect of temperatures rise quickly, activity began al- substrate temperatures (sT) with the slope most without any preceding basking, as the of the regression line being clearly different animals were able to achieve their activity from 1. This thermoregulatory efficiency temperatures without direct sun exposure. relative to the substrate temperatures (sT) is similarly, activity in absence of direct sun equivalent to the corresponding abilities of (due to orographic conditions) was observed the high mountain lizards of the sugbenus during the afternoon. in general, and as is Pyrenesaura (I. bonnali , I. aranica and I. usual with other mountain lizards (A RRiBAs aurelioi ) and exceeds that of other low 2007 and pers. obs. in all the ibero-Pyrenean mountain species such as I. monticola and I. species), a moderate wind increased basking horvathi the slopes of which are closer to 1 time and prolonged the daily activity of the (for references see Table 3). in fact, the cru - animals. The passage of a cloud produced a cial thermal problem that the high mountain drop in air temperature of up to six degrees. Pyre nesaura do face is not the cold, but the And thus in summer, lizards be came in- overheating of the substrate ( ARRiBAs 2007, creasingly active with the lowering of the 2010). similarly, high sTs could be respon - temperatures after a storm. On cloudy days sible for the increased thermoregulation with mild temperatures, the animals left their efficacy of the middle mountain species I. shelters at BTs from 29.2 to 33.8 °c (22.7- martinezricai . 26.4 °c AT and 26.4-33.1 °c sT). Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 13

Iberolacerta martinezricai , Batuecan Lizard, activity, microhabitat selection and thermoregulation 89

Fig. 4: scatter plots and regression lines of body temperatures (BT, °c) versus (A) substrate temperatures (sT, °c - left), and (B) ambient air temperatures (AT, °c - right) of Iberolacerta martinezricai (ARRiBAs , 1996). Regression line equations and determination coefficients (R 2) are given in the text. Abb. 4: streudiagramme und Regressionslinien zur Darstellung der Beziehung zwischen Körpertemperatur (BT, °c) und (A) substrattemperatur (sT, °c - links) sowie (B) Lufttemperatur (AT, °c - rechts) bei Iberolacerta martinezricai (ARRiBAs , 1996). Die Gleichungen der Regressionslinien und die Determinationskoeffizienten (R 2) sind im Text angegeben.

During another meteorological ex- BT measured (39.2 °c) was detected in a treme, very dry and hot weather as can oc- lizard trapped under a culvert without a cur in the middle of August, there were days chance of escape. The animal displayed when no lizards were seen during day as anomalous behavior, including quick move - they did not leave their shelters (aestiva - ments and tremors, and probably was close tion?). The animals sheltered deep in the to a critical temperature. At the accidental crevices when the ambient temperature rose death of a juvenile, the BT was registered to excessively. The BT of a burrowed lizard be 43.6 °c (50 °c sT). Thus, lethality must was e.g., 36.6 °c while the rock surface be expected at body temperatures above 40 temperature was 47.8 °c. Maximum sT °c, similar to other Iberolacerta (40.6 °c measured on a rock was 61 °c. The highest for I. aranica ; ARRiBAs 2007).

AcKNOWLEDGEMENTs

The author acknowledges Junta de castilla y Leon who issued the permissions to survey the study area.

REFERENcEs

ARGUELLO , J. A. & s ALVADOR , A. (1998): ricai .- Boletín de la Asociación Herpetológica Actividad, selección de hábitat y temperaturas corpo - Española, salamanca; 15 (2): 96-97. rales de Lacerta monticola en una localidad de la ARRiBAs , O. J. (2006): Lagartija batueca – Ibero - cordillera cantábrica (sauria, Lacertidae).- Revista lacerta martinezricai (ARRiBAs , 1996). in: sALVADOR , Española de Herpetologia, salamanca; 3 (1): 29-40. A. & MARcO , A. (Eds.): Enciclopedia virtual de los ARRiBAs , O. (1996): Taxonomic revision of Vertebrados Españoles. Madrid (Museo Nacional de the iberian ‘Archaeolacertae’ i: A new interpretation ciencias Naturales). WWW document available at of the geographical variation of ‘ Lacerta ’ monticola < http://www.vertebradosibericos.org/ >. BOULENGER , 1905 and ‘ Lacerta ’ cyreni MüLLER & ARRiBAs , O. J. (2007): istòria naturau e evolucion HELLMicH , 1937 (squamata: sauria: Lacertidae).- dera cernalha Aranesa, Iberolacerta aranica .- Huesca Herpetozoa, Wien; 9 (1/2): 31-56. (Graf icas Alós - conselh Generau d’Aran), pp. 446. ARRiBAs , O. (1999a): New data on the Peña de ARRiBAs , O. J. (2009): Habitat selection, ther - Francia Mountain Lizard ‘ Lacerta ’ cyreni martinezri - moregulation and activity of the Pyrenean Rock Lizard cai ARRiBAs , 1996 (squamata: Lacertidae).- Herpeto - Iberolacerta bonnali (LANTZ , 1927) (squamata: sauria: zoa, Wien; 12 (3/4): 119-128. Lacertidae).-Herpetozoa, Wien; 22 (3/4): 145-166. ARRiBAs , O. (2004): Nuevos datos sobre la dis - ARRiBAs , O. J. (2010): Activity, microhabitat tribución de la lagartija batueca: Iberolacerta martinez - selection and thermal behavior of the Pyrenean Rock Arribas_Iberolacerta_martinezricai_ecology_HERPETOZOA.qxd 25.06.2013 11:35 Seite 14

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DATE OF sUBMissiON: October 29, 2012 corresponding editor: Heinz Grillitsch

AUTHOR: Oscar J. ARRiBAs < [email protected] >, Avda. Fco. cambó 23; 08003 Barcelona, spain