Journal of Thermal Biology 48 (2015) 1–10

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Journal of Thermal Biology

journal homepage: www.elsevier.com/locate/jtherbio

Thermoregulation of two sympatric species of horned lizards in the

Chihuahuan Desert and their local extinction risk
Rafael A. Lara-Reséndiz a,b,n, Héctor Gadsden c, Philip C. Rosen d, Barry Sinervo b,
Fausto R. Méndez-De la Cruz a
a
Laboratorio de Herpetología, Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México, A. P. 70515, C.P. 04510 México,
D.F., Mexico
b
Department of Ecology and Evolutionary Biology, Earth and Marine Sciences Building A316, University of California, Santa Cruz, CA 95064, USA
c
Instituto de Ecología, A. C., Miguel de Cervantes No. 120, Complejo Industrial Chihuahua, C.P. 31109 Chihuahua, Chihuahua, Mexico
d
School of Natural Resources and the Environment, University of Arizona, Tucson, AZ 85721, USA

art ic l e i nf o a b s t r a c t

Article history: Thermoregulatory studies of ectothermic organisms are an important tool for ecological physiology,
Received 2 October 2013 evolutionary ecology and behavior, and recently have become central for evaluating and predicting
Received in revised form global climate change impacts. Here, we present a novel combination of field, laboratory, and modeling
21 November 2014
approaches to examine body temperature regulation, habitat thermal quality, and hours of thermal re-
Accepted 24 November 2014
Available online 27 November 2014
striction on the activity of two sympatric, aridlands horned lizards (Phrynosoma cornutum and Phryno-
soma modestum) at three contrasting Chihuahuan Desert sites in Mexico. Using these physiological data,
Keywords: we estimate local extinction risk under predicted climate change within their current geographical
Climate change distribution. We followed the Hertz et al. (1993, Am. Nat., 142, 796–818) protocol for evaluating ther-
Ecophysiological model
moregulation and the Sinervo et al. (2010, Science, 328, 894–899) eco-physiological model of extinction
Habitat thermal quality
under climatic warming. Thermoregulatory indices suggest that both species thermoregulate effectively
Phrynosoma
Thermal hours of restriction despite living in habitats of low thermal quality, although high environmental temperatures restrict the
Thermoregulatory efficiency activity period of both species. Based on our measurements, if air temperature rises as predicted by
climate models, the extinction model projects that P. cornutum will become locally extinct at 6% of sites
by 2050 and 18% by 2080 and P. modestum will become extinct at 32% of sites by 2050 and 60% by 2080.
The method we apply, using widely available or readily acquired thermal data, along with the modeling,
appeared to identify several unique ecological traits that seemingly exacerbate climate sensitivity of P.
modestum.
& 2014 Elsevier Ltd. All rights reserved.

1. Introduction
Global climate change and habitat alterations due to human
activities exert significant selective pressures and negative effects
on natural populations of many taxa (Cahill et al., 2013; Chen et al.,
2011; Huey et al., 2012; Kareiva et al., 1993; Pereira et al., 2010).
Recent research shows 12% of 200 Mexican populations of Scelo-
porus lizards have become locally extinct since 1975 as result of
rising environmental temperatures (Sinervo et al., 2010). This oc-
curs because when lizards are exposed to operative environmental
temperatures (Te) greater than their preferred body temperatures
n
Corresponding author at: Department of Ecology and Evolutionary Biology,
Earth and Marine Sciences Building A316, University of California, Santa Cruz, CA
95064, USA.
E-mail addresses: rafas.lara@gmail.com (R.A. Lara-Reséndiz),
hgadsden@gmail.com (H. Gadsden), pcrosen@email.arizona.edu (P.C. Rosen),
lizardrps@gmail.com (B. Sinervo), faustor@ib.unam.mx (F.R. Méndez-De la Cruz).
(Tp), activity time is restricted, and thus net energy gain becomes
insufficient for females in their reproductive season (Dunham,
1993; Huey et al., 2010; Sinervo et al., 2010, 2011). Reduction in
activity time is expressed as hours of thermal restriction (hr), or
hours of forced inactivity, which are the cumulative hours each
day during the reproductive season when Te exceeds Tp.
Most lizards must thermoregulate to maintain their field body
temperature (Tb) within an optimal range, but environments with
high temperatures increase the risk of overheating, shortening
activity time (Avery, 1982; Cowles and Bogert, 1944; Dunham,
1993; Huey, 1982), and thus elevating extinction risk. Therefore,
thermoregulation studies of ectothermic organisms highly con-
strained by their environment are essential to evaluate the beha-
vioral capacity of different species to minimize high Tb in ther-
mally severe environments (Kearney et al., 2009; Vickers et al.,
2011). Currently, evaluating temperature regulation in field-active
reptiles can be approached with the protocol proposed by Hertz
et al. (1993), which has been used widely for desert lizards and the

http://dx.doi.org/10.1016/j.jtherbio.2014.11.010
0306-4565/& 2014 Elsevier Ltd. All rights reserved.
2 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10
quantitative indices give more detailed information than previous
methods in thermal biology (Blouin-Demers and Nadeau, 2005;
Vickers et al., 2011). In addition, it is possible to predict extinction
risk using the physiological model proposed by Sinervo et al.
(2010) based on hr during peak reproductive seasons, and to apply
these predictions to future warming scenarios across a species'
distribution. These two methodologies synthesize thermoregula-
tion, habitat thermal quality, and the hours of restriction, and thus
can be used to evaluate how thermal conditions affect lizard po-
pulations in desert areas and to evaluate local population extinc-
tion risks.
In this study, we focused on two sympatric desert species of
horned lizards, Phrynosoma cornutum and Phrynosoma modestum,
that inhabit the Chihuahuan Desert in northern Mexico, and which
may offer a model for studying diurnal lizard thermal biology and
climate-change sensitivity. First, the Chihuahuan Desert represents
an important biogeographic region with wide temperature ranges,
great variation in the values of climatic variables, ecologically
contrasting environments, and a high proportion of reptiles
adapted to unique edaphic, climatic, and hydrologic conditions
(Lemos-Espinal and Smith, 2009; Macmahon and Wagner, 1985).
Second, both species of horned lizards are diurnal, similar in ac-
tivity patterns, microhabitat types, and natural history (ant-spe-
cialist diet, oviparity, breeding seasonality and body form). They
co-exist in varied habitats in Chihuahuan Desert (Pianka and
Parker, 1975; Sherbrooke, 2003), and both species may be espe-
cially vulnerable in distributions confined to hot, variable desert
environments. Third, thermal preference data are available for the
genus Phrynosoma (Brattstrom, 1965a; Cowles and Bogert, 1944;
Heath, 1962, 1964; Lemos-Espinal et al., 1997; Woolrich-Piña et al.,
2012; and others) as are previous general descriptions of aspects of
Phrynosoma thermal ecology (Brattstrom, 1965b; Guyer and Lin-
der, 1985; Heath, 1965; Pianka and Parker, 1975; Prieto and
Whitford, 1971; and others). Thermoregulatory efficiency and ha-
bitat thermal quality of Phrynosoma have been studied in cold
climate conditions (Arias-Balderas, 2011; Christian, 1998; Powell
and Russell, 1985; Vázquez-Urzúa, 2008) but studies necessary for
assessing these parameters have only recently become available in
the most vulnerable, hot desert environments (Lara-Reséndiz
et al., 2014). Fourth, some recent researchers have explored the
limitations of applying correlative models of species niche dis-
tribution to predict future species' range and vulnerability to
global changing (Buckley et al., 2010; Ceia-Hasse et al., 2014;
Clusella-Trullas and Chown, 2014; Huey et al., 2012; Kearney,
2013; and references therein); knowledge of species' thermo-
regulation will be increasingly necessary for integrative, mechan-
istic models that incorporate thermal requirements and environ-
mental constraints for climate-niche modeling. Thus, novel field
and laboratory data are needed to explore horned lizard extinction
risk in order to evaluate conservation status and, therefore assist
critical management decision-making in the face of climate-
change adaptation.
Here we synthesize methods for thermoregulation character-
ization of Hertz et al. (1993) with those of Sinervo et al. (2010) for
extinction modeling to present a new method for applying field,
laboratory, and modeling approaches to evaluate climate sensi-
tivity of ectothermic animals. Our objectives in this paper are to
examine habitat thermal quality, accuracy, efficiency of thermo-
regulation, and hours of thermal restriction of activity for P. cor-
nutum and P. modestum at three contrasting sites in the Mexican
Chihuahuan Desert, and then to evaluate local extinction risks
resulting from climate warming across their geographical dis-
tribution based on a mechanistic physiological niche model.
2. Materials and methods
2.1. Study organisms and sites
The species studied, which are unusual within the genus in that
they occur in sympatry over large portions of their geographic
distribution, are highly divergent in size: P. cornutum are much
larger (adult snout-vent length [SVL] 69–114 mm) than P. mod-
estum (adult SVL 37–68 mm) (Sherbrooke, 2003). Although from
different clades (Leaché and McGuire, 2006; but see Wiens et al.,
2010) the two species share many aspects of their general biology.
They share foraging strategies, are ant specialists, and they coexist
in the same macro-habitat, except for some differences in sub-
strate along the elevational gradient. P. cornutum is most abundant
on fine soils, while P. modestum tends to be further up the gradient
(closer to steep sides of the mountain) on pebbly to rocky soils
(García-De la Peña et al., 2012; Pianka and Parker, 1975; Shaffer
and Whitford, 1981), and it has been suggested as a rock mimic
(Sherbrooke and Montanucci, 1988). Their reproductive seasons
(breeding and egg-laying) are also similar: April to mid-July or
early August in P. cornutum and March–July or August in P. mod-
estum (Ballinger, 1974; Hernández-Ibarra and Ramírez-Bautista,
2002; Howard, 1974).
Three contrasting sites were chosen to represent varied soil
types, temperature regimes, and vegetation of the Chihuahuan
Desert (Fig. 1) and include: A) typical Chihuahuan Desert scrub in
Bolsón de Mapimí on the border between the states of Durango
and Chihuahua (26°52′16.03″N, 103°53′8.59″W; elevation 1114 m;
mean annual temperature and precipitation of 20.8 °C, and
230 mm, respectively), characterized by sandy soils with relatively
high vegetation cover dominated by Acacia greggii, Acacia con-
stricta, Larrea tridentata, Yucca elata, and Lycium berlandieri as well
as grasses and Prosopis juliflora in lower-lying areas (Barbault and
Halffter, 1981; Breimer, 1985); B) Plains Grassland and Semi-desert
Grassland in Janos, Chihuahua state (30°49′31.6″N, 108°28′26.6″W;
elevation 1435 m; mean annual temperature and precipitation of
15.7 °C, and 307 mm, respectively), characterized by high propor-
tion of vegetation dominated by grasses (Bouteloua gracilis, B.
eripoda, Aristida hamulosa, Festuca spp.), Prosopis spp., and Cylin-
dropuntia imbricata (Santos-Barrera et al., 2008); and C) Sama-
layuca Sand Dunes in Chihuahua (31°28′33.1″N, 106°28′33.6″W;
elevation 1260 m; mean annual temperature and precipitation of
17.7 °C, and 257 mm, respectively), characterized by semi-stabi-
lized dunes with low cover of relatively halophytic Chihuahuan
Desert scrub vegetation; the most representative plants are Pro-
sopis glandulosa, Atriplex canescens, L. berlandieri, Gutierrezia sar-
othrae, and Salsola tragus (García-De la Peña et al., 2012).
2.2. Fieldwork
Fieldwork was performed during the reproductive season of
the lizards (spring and summer) from 2006–2010. Activity Tb of
each lizard captured was measured using a digital thermometer
(Fluke™ model 51-II) with the sensor inserted one centimeter into
the cloaca and recorded within the first 30 s after capture. We also
recorded time of capture, gender, SVL (mm), and location with a
Garmin Etrex™ GPS receiver. All lizards were transferred to the
laboratory for studies of thermal physiology.
Simultaneously, during May–July 2009 and 2010, we recorded
field Te at all sites. (Mapimí: 34 days, Janos: 72 days, and Sama-
layuca: 65 days). Operative temperatures represent the equili-
brium of an inanimate object with its habitat and allow explora-
tion of the thermal environment at the spatial scale experienced
by the lizards; additionally, they can be easily replicated in mul-
tiple spatial–temporal conditions (Bakken, 1992), even at sites of
recorded extinction (Sinervo et al., 2010). Recently, Te using
 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10
Fig. 1. Photographs of the study sites and main landscape elements in northern
Mexico showing contrasting vegetation (details in text). Mapimí (A), Janos (B), and
Samalayuca (C).
polyvinylchloride (PVC) pipe models has been validated against
body temperatures for small ectotherms (Adolph, 1990; Lara-Re-
séndiz et al., 2014; Sinervo et al., 2010). Operative temperatures at
each site were recorded using six data-logger sensors connected to
previously field-calibrated PVC models similar in size and shape to
Phrynosoma (8
5
1.2 cm3), painted with 33% reflective gray
spray paint (Rustoleum™ gray primer) to approximate the re-
flectivity of phrynosomatids (Adolph, 1990), and placed in three
microsites occupied by the lizards (two at sunny sites, two in
shade, and two in sun/shade mosaic) (Sinervo et al., 2011). Tem-
perature was recorded every 30 min from 07:30 to 21:30 h to es-
timate the distribution of Te throughout the potential horned li-
zards' activity period (Dzialowski, 2005; Pianka and Parker, 1975).
3
The activity period was defined as including the first lizard ob-
served in morning to the end of potential crepuscular activity
based on our previous observations and the literature on similar
desert lizards (Harris, 1958; Lara-Reséndiz et al., 2013; Williams,
1959).
2.3. Laboratory work and thermoregulation indices
The Tp interquartile range was recorded in a laboratory thermal
gradient consisting of a wooden shuttle box with
150
100
80 cm3 (length, width, and height) dimensions di-
vided into 10 tracks with insulation barriers to prevent behavioral
influence of adjacent lizards, and filled with 2–3 cm of sandy soil.
The shuttle box was housed in a room at a constant temperature of
20 °C. At one end, and at the center of the box, lamps were placed
at different heights to generate a thermal gradient from 20-50 °C.
Lizards remained in the gradient from 07:00 to 19:00 h; Tb was
registered every two hours starting at 9:00 h with the same digital
thermometer used in the field. The 25% and 75% quartiles of each
specie' thermal gradient Tp interquartile range (Tp25 and Tp75) were
used as the lower and upper Tp (Hertz et al., 1993). Laboratory
experiments were conducted two days after capture. After these,
all lizards were released at their capture sites.
Data on field Tb, laboratory Tp, and Te were used to calculate
indices (Hertz et al., 1993) for accuracy of thermoregulation (db)
and habitat thermal quality (de), individually, as follows. If Tb or
Te o Tp interquartile range, then db ¼│Tp25–Tb│ and de ¼│Tp25–
Te│, respectively, and if Tb or Te 4Tp interquartile range then
db ¼ │Tb–Tp75│ and de ¼│Te–Tp75│, respectively. Note that we use
absolute value of Tb–Tp and Te–Tp deviations. When Tb or Te values
were within Tp interquartile range, db and de were considered
equal to zero. High values of db and de indicate a low accuracy of Tb
regulation and a low habitat thermal quality from the organism's
perspective, while values equal to or near to zero indicate high
accuracy of Tb and thermally ideal environments. Subsequently,
thermoregulation efficiency (E) was calculated using averages of db
and de ( ¼ db and de ) using the following equation: E = 1 − ( db/ de)
for each species and population. An E index near to one is in-
dicative of an organism that actively thermoregulates while en-
vironmental temperature availability is far from its physiological
requirements. Such an organism is subject to thermal stress and
must act to increase or decrease Tb with respect to Te. In contrast,
an E value equal or near to zero is indicative of a thermoconformer
that is not regulating temperatures actively (see details in Hertz
et al. (1993)). Following Blouin-Demers and Weatherhead (2001)
and Blouin-Demers and Nadeau (2005) we measured the index of
effectiveness of thermoregulation (de–db) to consider the magni-
tude of db and de for interpreting E and to quantify the extent of
departure from perfect thermoconformity. We calculated standard
errors of E for each species by bootstrapping, using db average for
each species and 1000 iterations of de. We compared E values
between species and populations using bootstrap estimates, that
is, to test differences among species or populations at 95% con-
fidence level (Hertz et al., 1993). Finally, field Tb observations that
fell above or below the Tp interquartile range were used to define
the direction of the Tb deviation with respect to Tp interquartile
range, as well as the percentage of Tb maintained within the Tp
interquartile range (Hertz et al., 1993).
2.4. Physiological model for local extinction risk
Here, we outline the application of Sinervo et al. (2010) method
for physiological modeling of lizard population extinction risk; a
detailed description of this protocol is in Sinervo et al. (2010, 2011)
and corresponding supplementary materials (also see Ceia-Hasse
et al. (2014)).
4 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10

Using Tp interquartile range for each species and Te for the 3. Results
three field sites during the reproductive season, we estimated the
number of hours per day potentially available for activity (ha) as 3.1. Body temperature in field (Tb)
the time when Te was within the Tp interquartile range, and then
hr was estimated as time when Te exceeded Tp, following Sinervo Eighty-three P. cornutum were captured (13 in Mapimí, 2 in
et al. (2010). Samalayuca, and 68 in Janos) while they were active, from 07:16–
Environmental data to estimate time restriction throughout the 21:43 h. Five of these were found in the open after 21:10 h (1–2 h
geographical range of P. cornutum and P. modestum were from the post-sunset) in July at Janos, with suboptimal Tb ( X̄ ¼28.7, range
three weather stations closest in elevation and proximity (airline 27.2–30.8 °C) when Te was 32 °C (Fig. 2A and D). Twenty-four P.
distance) to our Te study populations (Torreón, Coahuila; Ciudad modestum (5 from Mapimí and 19 from Samalayuca) were col-
Juárez International Airport, Chihuahua; and Deming Municipal lected from 09:00–19:45 h; the highest observation frequency was
Airport, New Mexico). Maximum air temperature (Tmax) records from 10:00–12:35 h (Fig. 2B).
during May–July in 2009 and 2010 were used to compute the
functional relationship between hr and activity time. We related hr
on a daily basis to the Tmax observed at each weather station and
fitted the following highly significant linear regression
equation (hr [Te 4 Tp] ¼intercept þslope (Tmax–Tp), following Si-
nervo et al. (2010)). This equation relating hr to Tmax and Tp, cali-
brated specifically for these horned lizards, was then used in
conjunction with predictions of Tmax from global climate circula-
tion models for 2050 and 2080 at a resolution of 10 arcmin (IPCC
IVth Climate Assessment, Hijmans et al., 2005) to compute the
likelihood of extinction due to increasing hr during the species'
reproductive seasons.
Data for geographical distribution of P. cornutum (n ¼516) and
P. modestum (n ¼631) were obtained from a variety of sources:
Global Biodiversity Information Facility (GBIF) when available,
Comisión Nacional para el Conocimiento y Uso de la Biodiversidad
(CONABIO), Colección Nacional de Anfibios y Reptiles (CNAR), our
prior field sampling, and the literature. Localities were georefer-
enced to the nearest minute of latitude and longitude via direct
consultation of various map series, following removal from the
dataset of points with uncertain localities, identification, or
otherwise unusable locality descriptors.

2.5. Statistical analysis

Only Tb and Tp from adult lizards were analyzed: P.

cornutum 4 69 mm SVL; for P. modestum 437 mm SVL (Goldberg,
2012; Lemos-Espinal and Smith, 2009). Data were excluded if
temperature measurements were taken more than 40 s after first
sighting the lizard in the field.
Assumptions of normality and homogeneity of variances were
analyzed using the Kolmogorov–Smirnov and Levene's tests, re-
spectively. Data satisfying requirements for parametric tests were
compared via the Student's t-test or Analysis of Variance (ANOVA);
otherwise the non-parametric Mann–Whitney or Kruskal–Wallis
tests were used and when differences existed at Po 0.05, post hoc
pair-wise comparisons were tested for significance in which the P
value was adjusted by Dunn's method (nonparametric). We used a
Two Way Repeated Measures ANOVA (Two Way RMANOVA) with
within-subjects factor with six levels (every two hours measure of
Tp), between subjects factor with two levels (male and female) for
each species. We used JMP version 10.0.0 (SAS, 2007) and Sig-
maPlot version 11 (Systat Software, San Jose, CA) for statistical
analyses and graphing. The significance value in all statistical tests
was P o0.05 (Sokal and Rohlf, 2000). Comparing Te among sites
was done using only the average of the data per day to avoid
pseudoreplication. Simulations for the extinction model and the E Fig. 2. Activity period of (A) Phrynosoma cornutum (B) Phrynosoma modestum in the
bootstrap resampling were conducted in R software (R Develop- daytime at the Chihuahuan Desert. Tb observations below the preferred tempera-
tures range is depicted in black, whilst observations in range are shown in white
ment Core Team, 2012). Figures and tables in the text show
and those above preferred temperatures range are in gray. (C) shows Te distribution
mean 7standard error ( X̄ 7 SE), (n) sample size, and temperature in the daytime at Mapimí, (D) at Janos, and (E) in Samalayuca. Points indicate
range. Mean 7Standard Error, and the dotted line indicates the average °C.
 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10
Fig. 3. Distribution of field body temperatures (Tb) of Phrynosoma cornutum and
Phrynosoma modestum and operative temperatures (Te) at Mapimí in Durango as
well as Samalayuca and Janos in Chihuahua. The arrow points to the average of the
field body temperatures (A, B) and operative temperatures (C–E.). The preferred
temperatures range (Tp) in thermal gradient for P. cornutum (32.5–36 °C) is re-
presented by diagonal shading and for P. modestum (31.1–36.5 °C) is represented by
crosshatching. Maximum critical temperature (CTMax) of P. cornutum (47.9 °C ac-
cording to Prieto and Whitford (1971)) is depicted by the dashed line. Notice the
difference of scales.
No significant differences in Tb were found among study sites in
P. cornutum (U¼346.0, P¼0.219; Samalayuca excluded due to small
sample size) or P. modestum (U¼ 27.5, P¼0.065). We therefore used
the overall Tb of 33.470.45 °C (n¼83, range 22.1–40.8 °C, Fig. 3A)
for P. cornutum and 31.970.93 °C (n¼24, range 21–38 °C, Fig. 3B)
for P. modestum, even though the difference between the species in
Tb was not significant (t¼1.460, d.f.¼105, P¼ 0.147).
5
3.2. Preferred body temperature in the laboratory (Tp)
Preferred body temperature in the thermal gradient for P. cor-
nutum was 34.27 0.1 °C (n ¼97, range 24–43.5 °C, Fig. 3A), and
33.6 70.51 °C (n ¼8, range 25.6–39 °C, Fig. 3B) for P. modestum,
with the species again not significantly different (U ¼17440.5,
P¼ 0.844). The interquartile range of Tp was 32.5–36 °C for P. cor-
nutum, and 31.1–36.5 °C for P. modestum (Fig. 3). Laboratory Tp for
P. cornutum was not significantly different for the entire activity
period (every measure of Tp) (RMANOVA; Wilk's λ ¼0.975;
F5,568 ¼ 0.463; P ¼0.803) and there was no significant effect found
in Tp between sexes (RMANOVA; Wilk's λ ¼0.919; F1,568 ¼ 1.57;
P¼ 0.177). Similarly, Tp for P. modestum was not significantly dif-
ferent among measures (RMANOVA; Wilk's λ ¼0.029;
F5,46 ¼ 13.449; P ¼0.071) and there was no significant effect found
in Tp between sexes (RMANOVA; Wilk's λ ¼0.103; F1,46 ¼3.475;
P¼ 0.238).
3.3. Operative temperatures (Te)
Calibration of lizard and PVC model (every 3 min over 3 h), in
both sunny and shaded microhabitats (see Lara-Reséndiz et al.,
2014), gave a highly significant linear correlation between field Tb
and Te (r ¼0.98, n ¼51, P o0.001; Tb ¼ 7.4582 þ0.8207 Te). Thus, we
assumed the lizard models accurately measured the available Te
for the horned lizard in this study, especially during the hours of
activity (Tb 420 °C).
Operative temperatures registered at Mapimí in late spring and
early summer averaged 32.8 70.61 °C (n ¼34, range 15.0–54.5 °C;
Fig. 3C); at Janos it was 31.1 7 0.42 °C (n ¼ 72, range 12.9–62.7 °C;
Fig. 3D); and at Samalayuca it was 39.9 71.19 °C (n ¼65, range
22.9–54.1 °C; Fig. 3E). Mean daytime Te for each of the three sites
is shown in Fig. 2C–E. Based on the PVC models, Te differed sta-
tistically among sites (H ¼31.898, d.f. ¼2, P o0.001). There were no
differences between Mapimí and Janos, but both differed from
Samalayuca (Dunn's pairwise comparisons, P o0.05; Figs. 2C–E
and 3C and D). Due to this variation in thermal opportunities,
thermoregulation indices and hours of restriction in activity were
calculated separately for each site, using the Tp and activity time of
each species.
3.4. Thermoregulation indices
The thermoregulation accuracy index (db) for P. cornutum was
2.9 70.30 (n ¼83) and for P. modestum it was 1.470.63 (n ¼24),
with the two species significantly different (t¼2.258, d.f. ¼105,
P¼ 0.026), and thus P. modestum was more accurately maintaining
Tb within its Tp interquartile range. For the Tp interquartile range,
37.3% of P. cornutum Tb records were below the Tp interquartile
range, 33.7% within, and 28.9% above (Fig. 3A). For P. modestum,
25% were below Tp interquartile range, 66.7% within and only
8.33% above the preferred interquartile range (Fig. 3B).
The thermal quality index (de) was estimated separately for
each study site due to significant among-site differences in Te. The
de's for P. cornutum at Mapimí, Janos and Samalayuca are pre-
sented in Table 1; these results differed significantly among sites
(H ¼156.705, d.f. ¼ 2, Po0.001). Thermal quality was highest at
Janos, where the greatest number of individuals of this species was
collected. This site was followed by Mapimí, with moderate
quality. Samalayuca had the lowest thermal quality, where only
two individuals of P. cornutum were collected; however, 19 P.
modestum were found there. The de of the sympatric species, P.
modestum, at Mapimí and Samalayuca were low and sites did not
differ significantly (U¼101244.5, P ¼0.094; Table 1).
For P. cornutum at Mapimí, 57.1% of Te observations were below
the Tp interquartile range, 11.3% within the range and 31.6% above
6 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10

Table 1
Thermoregulatory data and variables in Phrynosoma cornutum and Phrynosoma modestum at three study sites in the Chihuahuan Desert. Mean 7Standard Error are shown.

Species Phrynosoma cornutum Phrynosoma modestum

Variables/Sites Mapimí Janos Samalayuca Mapimí Samalayuca

Sample size (n) 13 68 2 5 19

Field body temperature (Tb; °C) 33.4 7 0.45a 31.9 7 0.93a
Preferred temperature, laboratory (Tp; °C) 34.2 7 0.1a 33.670.51 a

Tp interquartile range (Tp25–Tp75; °C) 32.5  36a 31.1  36.5a

Hours of activity (ha) 5.4 7 0.55 8.0 7 0.32 3.2 7 0.54 3.3 7 0.42 2.4 7 0.53
Hours of restriction (hr) 5.7 7 0.53 3.8 7 0.26 9.0 70.61 5.9 7 0.52 7.8 7 0.60
Thermoregulatory accuracy (db) 2.9 70.30a 1.4 7 0.63a
Habitat thermal quality (de) 6.3 7 0.12 4.8 70.07 7.9 7 0.47 6.3 7 0.11 7.3 7 0.48
Thermoregulatory efficiency (de–db) 3.4 1.9 5.0 4.9 5.9
Thermoregulatory efficiency (E) 0.53 70.10 0.39 7 0.01 0.63 7 0.21 0.737 0.11 0.80 7 0.21

a
Calculated by species.

(Fig. 3C). At Janos, 65.1% of Te observations were below the Tp in-
terquartile range, 15.2% within the range and 19.7% above (Fig. 3D).
In contrast, at Samalayuca, only 27.7% of observations were below,
7.7% were within, and 64.6% were above the Tp interquartile range,
and a substantial fraction of Te exceeded P. cornutum's critical
thermal maximum (CTMax) of 47.9 °C (Prieto and Whitford, 1971;
Fig. 3E). For P. modestum at Mapimí, 53.2% of Te observations were
below its Tp interquartile range, 16.6% within and 30.2% above the
range (Fig. 3C). At Samalayuca, 22.3% of Te observations for P.
modestum were below its Tp interquartile range, 13.1% within and
64.6% above the range (Fig. 3E).
There were significant differences in thermoregulation effi-
ciency index between both species, suggesting that P. modestum
(E ¼0.72 70.001) was more efficient than P. cornutum
(E ¼0.53 70.001) (H ¼2634.1, d.f. ¼2, P o0.001). In addition, there
were differences between localities, with P. modestum more effi-
cient at Samalayuca than Mapimí (U¼ 33146.0, Po0.001; Table 1),
and P. cornutum more efficient at Samalayuca than Mapimí and
Janos (H¼ 2634.1, d.f. ¼2, P o0.001; Table 1) despite the low
thermal quality of the environment at Samalayuca (Fig. 2E). A si-
milar pattern was found in the de – db index (Table 1), where P.
modestum was more efficient than P. cornutum and both species
were more efficient at Samalayuca.
3.5. Physiological model for local population extinction risk
Hours of activity for foraging and hours of restriction for fora-
ging for both species are shown in Table 1. For P. cornutum, Janos
was the most suitable site with eight hours for activity (see
Figs. 2D and 3D); Samalayuca was the least suitable due to the
high Te, with nine hours per day of thermal restriction (Figs. 2E
and 3E); whereas Mapimí was intermediate, with ha and hr nearly
equal (Figs. 2C and 3C). P. modestum at Mapimí and Samalayuca
had hr between six and eight hours, with low ha at both study
sites. Samalayuca had Te 4 Tp for nearly nine hours of activity re-
striction per day and a high percentage of temperatures (33.8%)
above the CTMax of P. cornutum (Fig. 3E).
Formulas for predicting extinctions according to the relationship
between hr and Tmax relative to Tp in P. cornutum was hr ¼0.35
(Tmax–Tp)þ 4.23 and in P. modestum it was hr ¼ 0.54 (Tmax–Tp)þ 6.08.
The linear fit differed between species (F¼ 3.456, d.f.¼2, Po0.037),
and we therefore applied a separate formula for each species.
If climate change (increase in Tmax) continues in México and
the United States as projected by the IPCC III Climate Assessment
(Hijmans et al., 2005), our computation predicts that P. cornutum
will be driven locally extinct at 5.6% of sites by 2050 and 17.6% by
2080 (Fig. 4A) while P. modestum will be driven locally extinct at
32.4% of sites by 2050 and 60% by 2080.

Fig. 4. Contour plots of year of predicted extinction for Phrynosoma cornutum and Phrynosoma modestum, assuming hr ¼ 3.9 and the Tp's calculated in this study.
 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10
4. Discussion
There are relatively few species of horned lizards living sym-
patrically in large areas (Pianka and Parker, 1975). However,
Shaffer and Whitford (1981) and García-De la Peña et al. (2012)
suggested P. cornutum and P. modestum may coexist avoid com-
petition throughout their distribution in Mexico and the United
States by selecting different species of ants, employing different
foraging behaviors, and occupying different microhabitats, as well
as employing different strategies for predator avoidance (Sher-
brooke, 2003). In this paper, no difference was found between the
Tp of the two species, which is consistent with data available for
closely related, sympatric lizard species (Díaz de la Vega-Pérez
et al., 2013; Grigg and Buckley, 2013). Thus, our results should
highlight the effects of thermal microenvironmental opportunities
on two lizard species with similar thermal physiology but different
morphology and behavior.
At Samalayuca mean Te was 40.2 °C, and our sample size of P.
modestum exceeded that of P. cornutum by 8-fold. P. modestum
showed better ability to exploit high environmental temperatures,
exhibiting a slightly wider range of Tp than P. cornutum (Fig. 3E).
Nonetheless, Samalayuca had the lowest habitat thermal quality
due to high Te, with many models exceeding the CTMax of horned
lizards. Though this is based on only one known value for P. cor-
nutum (47.9 °C; Prieto and Whitford, 1971), which according to
Grigg and Buckley (2013) CTMax could differ in P. modestum. We
found fewer horned lizards than did García-De la Peña et al. (2012)
in a previous study at this site, although we cannot draw conclu-
sions from this given potential differences in search effort. How-
ever, we collected the greatest number of P. modestum individuals
in the study from Samalayuca, suggesting that habitat type, scar-
city of P. cornutum as a potential competitor, or other factors may
be of high importance for the species. The Janos site had the
lowest mean Te (30.9 °C) among our sites, resulting in high ther-
mal quality for P. cornutum: 82% of the P. cornutum captured in the
study were at this site, whereas P. modestum was not found there,
although it was recently recorded nearby (Santos-Barrera et al.,
2008). At our Mapimí site, mean Te was 31.5 °C, and thermal
quality (de ¼ 6.3) was identical for P. cornutum and P. modestum,
since thermal opportunities were distributed similarly around the
Tp interquartile range of both species. The sites in the Chihuahuan
Desert with the highest thermal quality were Janos for P. cornu-
tum, and Mapimí for P. modestum, whereas Samalayuca had a high
percentage of extremely high temperatures (Kour and Hutchison,
1970; Prieto and Whitford, 1971).
Results indicated that Tb maintained in field were similar to the
Tp interquartile range for P. cornutum (db ¼2.9) and even more so
for P. modestum (db ¼1.4), reflecting the abilities of these species to
thermoregulate successfully in these desert locations. Further-
more, P. cornutum was moderately efficient and P. modestum highly
efficient at thermoregulation (E ¼0.62 and 0.76, respectively). Both
species reached Tb near their optimal activity thermal ranges,
despite the variability in the thermal quality of their habitats.
Christian (1998) found that for the viviparous lizard P. hernandesi
(formerly P. douglasii) in a montane habitat (elevation 2290 m),
maximum thermal efficiency occurred during April, the coldest
month, with mean Te ¼ 6.1 °C, with the lizards maintaining Tb be-
tween 25.4 and 36.4 °C (mean 32.5 °C), near the species' preferred
temperature. In contrast, during the sunniest months, the species
reached Tb ¼35.5 °C when Te ¼22.6 °C. Thermoregulation accuracy
for P. hernandesi was 0.25, thermal quality was 1.6, and thermal
efficiency was 0.85 (see Blouin-Demers and Nadeau, 2005, Ap-
pendix A). Thus, despite major differences in habitat, latitude, and
elevation, P. hernandesi was more accurate and efficient in ther-
moregulation than the two species studied here. Accuracy of
thermoregulation may be more readily achieved by a species that
7
basks in environments with low Te than by desert species that may
be forced to tolerate high temperatures or retreat to shelter where
they are inactive.
Recent research suggests ectothermic organisms from desert
environments have a high extinction probability under current
rates of global warming, because sufficiently rapid adaptive
change in Tb is unlikely (Ballesteros-Barrera et al., 2007; Barrows,
2011; Deutsch et al., 2008; Dunham, 1993; Gadsden et al., 2012;
Güizado-Rodríguez et al., 2012; Paranjpe et al., 2013; Sinervo et al.,
2010). Nevertheless, population impacts of rising environmental
temperatures depend on an ectotherm's capacity to compensate
through acclimation, thermoregulatory behaviors such as habitat
selection, change in daily activity patterns, and shifts in re-
productive and activity phenology (Deutsch et al., 2008; Huey
et al., 2009; Kearney et al., 2009), and perhaps, ultimately by
dispersal to track its shifting climate-niche. This paper demon-
strates that oviparous species of horned lizards inhabiting deserts
are vulnerable to changes in thermal habitat quality. Our data
suggest that P. cornutum might have a slightly more constrained
range of Tp than P. modestum, but these two sympatric species
appear to be physiologically similar. Both exhibited high field Tb,
reflecting high microclimate temperatures that increase risks of
overheating and dehydration and thus shorten the time available
for feeding and breeding activities, while also increasing tem-
perature-related maintenance cost, potentially leading to local
extinction. The three sites studied in the Chihuahuan Desert show
significant differences in environmental temperature spectra at
the microclimate level, and also in the strategies used by the two
species of horned lizards to regulate Tb in these thermal niches.
According to the lizard extinction model developed by Sinervo
et al. (2010), species are negatively effected when Te rises above
Tp, or when hr 43.85 h/day during the two peak months of the
reproductive period. Thermal restrictions of Sceloporus lizards
are higher in pregnant females, in which exposure to
temperatures 4 35 °C result in developmental abnormalities in
progeny (Beuchat, 1986). At our study sites, both species had a
wide range of times, overall, during which potential activity could
occur (P. cornutum ¼ 14.5 and P. modestum ¼10.7 h), but hours of
activity on any given day were much lower, and hours of restric-
tion (hr) varied from four to nine hr/d (Table 1) with an average of
6.4 h. This hr was higher than values previously calculated for
phrynosomatid lizards based largely upon members of the genus
Sceloporus (Sinervo et al., 2010). However, additional data on hr
earlier in the spring reproductive period (i.e., during a larger
sample of days) may be required to determine how close these
horned lizards species are to the extinction limit defined using the
Sceloporus model.
We suspect that the typical foraging behavior of horned lizards,
which can rapidly pack their guts (see Munger, 1984) during early
morning or late in the afternoon (pre-darkness) foraging (personal
observations, RALR, PCR), and their large gut volume and ability to
eat abundant, highly concentrated prey (usually ants), may allow
them to tolerate greater hr than Sceloporus. Ant-eating horned li-
zards can efficiently consume a large volume of prey in a very
short period of time, and then spend the rest of the day in retreat
sites when Te 4Tp. In contrast, Sceloporus lizards have much
smaller gut volumes and hours of restriction as low as 3.85 during
the two months of reproduction are sufficient to drive populations
to extinction (Sinervo et al., 2010).
Our results indicated a pattern of crepuscular activity by P.
cornutum because 7.4% of the individuals collected were found
active after 21:10 h. In addition, two of us (RALR and HG) ob-
served a copulation at dusk, far from the refuges of both in-
dividuals, during May at Mapimí (Lara-Reséndiz et al., 2013). Our
observation is in general agreement with previous observations of
nocturnal activity patterns (Harris, 1958; Mays and Nickerson,
8 R.A. Lara-Reséndiz et al. / Journal of Thermal Biology 48 (2015) 1–10
1968; Williams, 1959), nocturnal nest-digging (Sherbrooke, 2002)
and oviposition (Sherbrooke, 2000) in Phrynosoma. According to
Huey (1982) some reptile species voluntarily spend the night
outside of their burrows to prolong potential foraging time and
actively select relatively low body temperatures (i.e., suboptimal
body temperature). Furthermore, this condition produces a lower
Tb that reduces metabolic loss throughout the night. This may be a
response of individuals that fail to accomplish necessary activities
during the day, even if suboptimal voluntary temperature risks
increase exposure to nocturnal predators. In addition, according to
Huey et al. (2010), lizards select low temperature shelters during
hot seasons and days, such as burrows with optimal temperatures,
reducing time available for foraging, social interactions, and re-
production, often with deleterious effects on populations. For
diurnal lizards capable of activity under low-light conditions, as
observed here, activity times are extended into dusk hours, when
Te is close to Tp. It is also noteworthy that many prey species of
ants resume their activity after dusk, so crepuscular activity could
allow these lizards the opportunity to fill their guts and digest the
meal during the night, if they can find retreat sites with high Te
(Clarke and Zani, 2012). One of us (PCR) has regularly observed
regal horned lizards (Phrynosoma solare) in Tucson actively fora-
ging between sunset and true darkness, and even peering into ant
colony (Pogonomyrmex rugosus) openings for mass emergence of
ants, which, on hot summer days, often occurs after sunset and at
or just before dark. In this regard, crepuscular activity may be a
novel adaptive mechanism for escaping the effects of climate
warming, although as Williams (1959) noted, poor nocturnal vi-
sion in diurnal lizards may be a constraint. In conclusion, it is
important to model climate change effects on this group of lizards
in desert and temperate environments throughout their distribu-
tion, to learn about the traits that might exacerbate or ameliorate
risk of extinction through climate change scenarios. Therefore the
extinction risk may be ameliorated for Phrynosoma if an adequate
transition to crepuscular activity is achieved.
Field-active Tb’s of P. cornutum and P. modestum (33.4 and
31.9 °C, respectively) reported in our study are similar to, though
slightly lower than those reported for both species elsewhere in
their distribution (Cowles and Bogert, 1944; Kour and Hutchison,
1970; Lemos-Espinal et al., 1997; Pianka and Parker, 1975; Prieto
and Whitford, 1971). In addition, Tb registered for both species in
the Chihuahuan Desert (latitudes 26 to 31°N) were similar to,
though slightly lower than, those for other species of horned li-
zards from arid zones (Ballinger et al., 1998; Ballinger and Schrank,
1970; Brattstrom, 1965a; Cunningham, 1966; Heath, 1962, 1964,
1965; Hodges, 2002; Lemos-Espinal et al., 1997; Pianka and Parker,
1975). Our results may reflect the availability of low Te at two of
our study sites, and the relatively high desert elevations of the
Mexican Altiplano, where we worked. However, viviparous species
of horned lizards at high elevations and latitudes such as P. orbi-
culare, P. hernandesi, and P. douglasii maintain Tb  4 °C lower than
reported for most oviparous species distributed from sea level up
to 1500 m (Arias-Balderas, 2011; Christian, 1998; Powell and
Russell, 1985; Prieto and Whitford, 1971; Vázquez-Urzúa, 2008).
Relatively low field Tb and ha, and high hr of Phrynosoma at our
study sites may also reflect behavioral responses to thermal con-
straints of high Te. In our study, P. modestum achieved high ther-
moregulatory efficiency, by reduced exposure to activity with
potentially high costs associated with hr. This small horned lizard
has low body mass that increases heating rate toward CTMax and
may require more cautious thermoregulatory behavior to reduce
risk of overheating mortality. In southeastern Arizona, P. cornutum
was able to eat more ants, in shorter foraging bouts, than P.
modestum (Munger, 1984), posing another potential cost of fora-
ging as the environment heats up. Further, P. modestum occurs in
more desert-like, less grass-shaded habitat than P. cornutum, and
uses a unique anti-predator mechanism of rock-mimicry by re-
maining immobile with a hunched body among small rocks on the
hot microenvironment of open ground (Sherbrooke and Mon-
tanucci, 1988), and their small size makes them vulnerable to
predators such as grasshopper mice (Onychomys) and snakes that
are active in the vegetation (Sherbrooke, 2003). These unique as-
pects of morphology, coloration, habitat, thermal activity, and
behavior may expose P. modestum to strong climate warming risks,
reflected in the greater computed population extinction prob-
abilities. Both species that we studied are geographically posi-
tioned to extend their ranges northward into the current Plains
Grasslands of the United States as climate warms, but there are
potential problems with this scenario. Distributions of plants in
the arid southwest of North America contracted at their lower, hot
and arid margins but failed to compensate by equivalent upslope
movement (Brusca et al., 2013). During times of high temperature
and drought stress, propagule production and dispersal may be
weak, and tracking of climatic niches may be slow. For P. mod-
estum, habitat specialization on rocky piedmonts and foothills may
present barriers to northward dispersal across fine soils of inter-
vening valleys, and this unique species may thus face additional
threats for climate-change adaptation. The method we present
here appears able to incorporate many of the unique thermo-
regulatory adaptations that can determine high climate sensitivity
of individual species, and these details may prove essential for
management plans to facilitate climate adaptation for the pre-
servation of biodiversity.
Acknowledgments
This paper is part of the thesis research of the first author
(RALR) at the Programa de Posgrado en Ciencias Biológicas, Uni-
versidad Nacional Autónoma de México (UNAM) funded by a
Doctoral grant from CONACyT (165066) and a posdoctoral fel-
lowship from UC MEXUS-CONACyT (FE-14-2). We thank the Na-
tional Science Foundation, United States for the Grants (IOS-
1022031; EF 1241848) awarded to BS, and UNAM PAPIIT/DGAPA
(IN224208 and IN215011) for providing financial support to FRMD.
The collection permit numbers 03699/09 and 05570/10 were given
by the Dirección General de Vida Silvestre of México. Additional
field and laboratory work was carried out with the assistance of V.
Jiménez, A. Díaz de la Vega, R. Huitrón, and G. Castañeda. We also
thank the Reserve “El Uno” of The Nature Conservancy for facilities
granted during our stay and permits to collect on communal land.
We thank Robert Cooper, Elizabeth Bastiaans, Dhanashree Para-
njpe, and two anonymous reviewers for providing helpful com-
ments and suggestions on the manuscript. Finally, we thank GBIF,
CONABIO, and CNAR for providing databases for analysis.
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associated with the prairie dog grasslands ecosystem and surrounding areas at Philip C. Rosen was born in New York City, USA, on
the Janos Casas Grandes complex, northwestern Chihuahua, Mexico. Acta Zool. February 25, 1955. He completed degrees at University
Mex. Nueva Ser. 24, 125–136. of Michigan (BSc, 1980), Arizona State University (MSc,
SAS, 2007. JMP v.10.0.0, Pacific Grove, SAS Institute Inc. 1987), and University of Arizona (PhD, 2000) where he
Shaffer, D.T., Whitford, W.G., 1981. Behavioral-responses of a predator, the round- is currently a research scientist. His research and con-
tailed horned lizard, Phrynosoma modestum and its prey, honey pot ants, servation work has focused on amphibians and reptiles
Myrmecocystus spp. Am. Midl. Nat. 105, 209–216. in desert, grassland, aquatic, and urban environments
Sherbrooke, W.C., 2000. Phrynosoma modestum (Roundtailed Horned Lizard). Noc- in southwestern North America, and he has over 100
turnal nest-digging and oviposition. Herpetol. Rev. 31, 242–243. publications and formal research reports. His current
Sherbrooke, W.C., 2002. Phrynosoma cornutum (Texas Horned Lizard). Nocturnal work involves conservation biology and trends mon-
nesting, eggs, nest predation, hatchlings. Herpetol. Rev. 33, 206–208. itoring, working with ranid frogs, chelonians, and de-
Sherbrooke, W.C., 2003. Introduction to horned lizards of North America. University sert lizards.
of California Press, Berkeley.
Sherbrooke, W.C., Montanucci, R.R., 1988. Stone mimicry in the round-tailed
horned lizard, Phrynosoma modestum (Sauria, Iguanidae). J. Arid Environ. 14,
275–284.
Barry Sinervo was born in Ontario, Canada, on No-
Sinervo, B., Méndez-de la Cruz, F., Miles, D.B., Heulin, B., Bastiaans, E., Villagrán-
vember 24, 1960. He obtained his BSc in Biology/
Santa Cruz, M., Lara-Reséndiz, R., Martínez-Méndez, N., Calderón-Espinosa, M.
Mathematics with honors (1982) at Dalhousie Uni-
L., Meza-Lázaro, R.N., Gadsden, H., Avila, L.J., Morando, M., De la Riva, I.J., Se-
versity, Nova Scotia, Canada, and his PhD (1988) at
pulveda, P.V., Rocha, C.F.D., Ibargüengoytía, N., Puntriano, C.A., Massot, M., Le-
University of Washington, Seattle. He conducted two
petz, V., Oksanen, T.A., Chapple, D.G., Bauer, A.M., Branch, W.R., Clobert, J., Sites,
postdocs, at University of Berkeley (1990) and at Uni-
J.W., 2010. Erosion of lizard diversity by climate change and altered thermal
versity of Washington (1992). Currently, he is a full
niches. Science 328, 894–899.
professor at University of California Santa Cruz; his
Sinervo, B., Miles, D.B., Martínez-Méndez, N., Lara-Reséndiz, R., Méndez-de la Cruz,
research is focused on behavioral ecology and evolu-
F.R., 2011. Response to comment on “Erosion of lizard diversity by climate
tionary biology. He teaches game theory, herpetology,
change and altered thermal niches”. Science 332, 537.
and behavioral ecology. He has more than 100 pub-
Sokal, R.R., Rohlf, F.S., 2000. Biometry. Freeman and Company, New York.
lications. He is working with an international inter-
Vázquez-Urzúa, E., 2008. Termorregulación de la lagartija vivípara Phrynosoma orbicu-
disciplinary group to study the effects of global
lare (Phrynosomatidae) en zonas de alta montaña. Instituto de Biología. Universidad
warming on the extinction of lizards.
Nacional Autónoma de México, Distrito Federal, México (M.Sc. thesis).
Vickers, M., Manicom, C., Schwarzkopf, L., 2011. Extending the cost-benefit model
of thermoregulation: high-temperature environments. Am. Nat. 177, 452–461.
Wiens, J.J., Kuczynski, C.A., Arif, S., Reeder, T.W., 2010. Phylogenetic relationships of
phrynosomatid lizards based on nuclear and mitochondrial data, and a revised Fausto R. Méndez-de la Cruz was born in Mexico City,
phylogeny for Sceloporus. Mol. Phylogenet. Evol. 54, 150–161. Mexico, on September 6, 1957. He obtained his BSc
Williams, K.L., 1959. Nocturnal activity of some species of horned lizards, genus (1983), MSc (1986), and PhD (1989) at UNAM. He
Phrynosoma. Herpetologica 15, 43. conducted a postdoctoral year at the University of
Woolrich-Piña, G.A., Smith, G.R., Lemos-Espinal, J.A., 2012. Body temperatures of Florida, USA, and a sabbatical year at the Virginia
three species of Phrynosoma from Puebla, Mexico. Herpetol. Notes 5, 361–364. Polytechnic Institute and State University. He is a full
professor at the Instituto de Biología, UNAM. His main
research topics are the evolution of viviparity, parthe-
nogenesis, and the vulnerability of amphibians and
reptiles to global warming. He was president of the
Rafael A. Lara-Reséndiz was born in México City,
Sociedad Herpetológica Mexicana and Sociedad Lati-
Mexico, on October 22, 1981. He obtained his Bachelor's
noamericana de Herpetología. Méndez-de la Cruz has
degree at the Universidad Nacional Autónoma de
more than 100 publications.
México (UNAM). Later in 2008 Lara-Reséndiz received
his master's degree at UNAM after studying the thermal
ecology of lizards. He completed his PhD at the In-
stituto de Biología at UNAM, Mexico City, and is cur-
rently a post-doctoral fellow at University of California,
Santa Cruz. He has published scientific papers and book
chapters, and has given presentations at international
scientific meetings. His research is focused on the
thermophysiology of lizards, especially Phrynosoma,
Sceloporus, and Phyllodactylus. He has collaborated with
researchers from U.S., India, Chile, and Argentina.

Héctor Gadsden was born in Mexico City, Mexico, on July

10, 1952. He obtained his BSc (1978), MSc (1982), and PhD
(1988) at UNAM and belongs to the National Researcher
System (SNI). He works as a senior researcher at the In-
stituto de Ecología, A. C., in Chihuahua, Mexico. Gadsden
studies population and community ecology on lizards. He
has coordinated projects funded by CONACyT and CON-
ABIO. He has directed 35 undergraduate, master, and
doctorate students. He has published more than 100
publications. Currently, he is working with an interna-
tional interdisciplinary group to study the effects of global
warming on the extinction of lizards.