Natural ultraviolet-b exposure of the Texas Horned Lizard (Phrynosoma

cornutum) at a North Texas Wildlife Refuge

Author(s): Gary W. Ferguson, William H. Gehrmann, Andrew M. Brinker, Glenn C. Kroh, and Donald C.
Ruthven III
Source: The Southwestern Naturalist, 60(2-3):231-239.
Published By: Southwestern Association of Naturalists
DOI: http://dx.doi.org/10.1894/0038-4909-60.2-3.231
URL: http://www.bioone.org/doi/full/10.1894/0038-4909-60.2-3.231

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THE SOUTHWESTERN NATURALIST 60(2-3): 231–239 JUNE-SEPTEMBER 2015

NATURAL ULTRAVIOLET-B EXPOSURE OF THE TEXAS HORNED

LIZARD (PHRYNOSOMA CORNUTUM) AT A NORTH TEXAS WILDLIFE
REFUGE

GARY W. FERGUSON,* WILLIAM H. GEHRMANN, ANDREW M. BRINKER, GLENN C. KROH, AND DONALD C. RUTHVEN III

Department of Biology, Box 209030, Texas Christian University, Fort Worth, TX 76129 (GWF, WHG, AMB, GCK)
Texas Parks and Wildlife Department, 3036 FM 3256, Paducah, TX 79248 (DCR)
*Correspondent: g.ferguson@tcu.edu

ABSTRACT—We monitored daily patterns of natural ultraviolet-B (UVB) exposure (measured using the
ultraviolet index [UVI]) of the Texas horned lizard (Phrynosoma cornutum) in North Central Texas. The
animals were active on unpaved road surfaces on warm sunny days in May and June in the morning (0800–
1200 h) and in late afternoon and early evening (1800–2100 h). The UVI was high during the morning and
was positively correlated with hour of the day and body temperature. The UVI was lower during the late
afternoon and early evening and was negatively correlated with hour of the day. Body temperatures of the
animals were higher than air temperature during both periods of road activity. On warm sunny days between
the two periods of high road activity, lizards remained active in shaded off-road areas and received variable
amounts of UVB exposure. Two lizards that were followed most hours on warm sunny days had similar UVB
exposure doses (irradiance · time) for the day but showed different patterns of UVI irradiance. The UVI and
dose were lower for an additional individual followed on an overcast day with thermal conditions that
shortened aboveground activity time. On warm sunny days in May and June, P. cornutum received the highest
daily UVB dose of any lizard or snake monitored to date.

RESUMEN—Monitoreamos los patrones diarios de la exposición natural a la radiación ultravioleta-B (UVB)

(medida por el ı́ndice ultravioleta [UVI]) en Phrynosoma cornutum en el centro-norte de Texas. Los animales
estuvieron activos sobre las superficies de caminos no pavimentados en los cálidos y soleados dı́as en mayo y
junio por la mañana (0800–1200 h) y tarde por las tardes y las horas tempranas de la noche (1800–2100 h).
UVI fue alta en las mañanas y estuvo positivamente correlacionada con la hora del dı́a y la temperatura
corporal. UVI estuvo más bajo tarde por la tarde hasta las horas tempranas de la noche, y se correlacionó
negativamente con la hora del dı́a. La temperatura corporal del animal estuvo más alta que la temperatura del
aire durante los dos periodos de actividad en los caminos. En dı́as tibios y soleados entre los dos periodos de
alta actividad en los caminos, las lagartijas siguieron activas en las áreas sombreadas y alejadas de los caminos, y
los animales recibieron cantidades variadas de exposición a UVB. Entre dos lagartijas que fueron observadas
por la mayorı́a de las horas de dı́as cálidos y soleados, hubo una dosis semejante de exposición a la UVB
(radiación · tiempo) durante el dı́a, pero mostraron diferentes patrones de UVI . UVI y la dosis de exposición
fueron más bajos para una lagartija adicional que fue observada durante un dı́a nublado con condiciones
termales que cortaron el tiempo de actividad terrestre. Cuando hacı́a calor y sol en mayo y junio, P. cornutum
recibió la dosis diaria más alta de UVB que cualquier lagartija o culebra que haya sido monitoreada hasta el
presente.

Active basking in the sun by many lizards is a critical
mechanism for thermoregulation (Cowles and Bogert,
1944; Huey, 1982). Some lizard species quickly attain
preferred body temperatures higher than ambient by
basking early in the day. They might maintain this
preferred temperature throughout their activity during
the rest of the day either within narrow limits or with less
precision. Other lizard species are thermal conformers,
and bask little or not at all. When active, they maintain a
variable body temperature that conforms to the range of
their environment (Huey, 1982). Along this spectrum of
thermoregulatory behavior, the Texas horned lizard
(Phrynosoma cornutum: Phrynosomatidae) is a thermoreg-
ulator that maintains its body temperature within a zone
ranging from approximately 35–398C (Heath, 1965;
Prieto and Whitford, 1971). Thermoregulation and
adaptation to environmental temperature changes are
primary factors responsible for daily and seasonal cycles
of activity and basking in this and other horned lizard
species (Burrow et al., 2001; Sherbrooke, 2003).
232 The Southwestern Naturalist
The sun is not only a source of heat but also ultraviolet
(UV) radiation. Exposure to UV radiation from sunlight
can have both negative and positive effects on the health
and well-being of terrestrial vertebrates. The most
detrimental effects of excess UV radiation are tissue and
DNA damage that can lead to sterilization and cancer
(Hays et al., 1995; Blaustein et al., 1998; Miller et al., 2002;
Chang and Zheng, 2003). Inappropriately high exposure
doses of short-wave UV radiation causes skin and eye
damage and reduced reproductive success in captive
reptiles (Ferguson et al., 2002; Gardiner et al., 2009).
The most studied positive effect of ultraviolet-B (UVB;
290–315 nm) is bone health via endogenous vitamin D3
production. In many vertebrates, the exposure of skin to
UVB causes photolysis of provitamin D3 (7-dehydrocho-
lesterol) to previtamin D3 followed by thermal isomeriza-
tion to vitamin D3, which then enters the circulatory
system. Subsequent hydroxylation in the liver and kidneys
produces the biologically active form of vitamin D3, 1,25-
dihydroxyvitamin D3, a hormone critical to calcium and
phosphorus metabolism (Holick, 1996, 1999, 2007).
Vitamin D3 can also enter the circulatory system by
ingestion of food containing vitamin D3. Deficiency of
vitamin D3 in reptiles leads to nutritional metabolic bone
disease, reproductive failure, and death (Ferguson et al.,
1996; Mader, 2006).
Given the costs and benefits of UV exposure, verte-
brates should adaptively regulate their exposure to
optimal levels. A protective adaptation for many lizards
that bask extensively is a dark subdermal or peritoneal
lining that absorbs UV radiation and protects deeper
tissues from UV damage (Porter, 1967). A positive
adaptation would be the ability to detect UV and
voluntarily regulate exposure when there is a need for
vitamin D3 or, alternatively, when potential UV damage is
sensed. One lizard species, the panther chameleon
(Furcifer pardalis), can do the former (Ferguson et al.,
2003; Karsten et al., 2009). The Cuban brown anole
(Anolis sagrei) also appears to possess this ability (Fergu-
son et al., 2013). To our knowledge the ability for
behavioral avoidance of high, potentially detrimental,
UV exposure has not yet been rigorously documented in
reptiles but has been in frogs and fish (Han et al., 2007;
Holtby and Bothwell, 2008).
Researchers have only recently studied UV light
exposure of reptiles in nature (Carman et al., 2000;
Ferguson et al., 2005; Ferguson et al., 2010). Monitoring
of UV index (UVI, a unitless scale of UVB irradiance) of
15 lizard and snake species in the field during their daily
and seasonal peak activity has revealed a range of average
voluntary UVB exposures. Ferguson et al. (2010) have
divided these exposures into four categories or UVB
zones. UVB zone 1 includes crepuscular or shade-dwelling
species with a median daytime UVI of 0.35; zone 2
includes daytime partial sun or occasional basking
species, with a median UVI of 0.9; zone 3 includes full
vol. 60, no. 2-3
or partial sun basking species, with a median UVI of 1.8;
zone 4 includes midday basking species, with a median
UVI of 3.1. Lizards inhabiting a higher UVB zone have
skins less photosensitive to UVB regarding vitamin D
synthesis than those inhabiting a lower UVB zone
(Ferguson et al., 2005). This might also reflect a lower
sensitivity to UVB damage.
The Texas horned lizard occurs in the arid and
semiarid low- and middle-elevation habitats from south-
eastern Arizona south through northern Mexico and
north to central Kansas (Sherbrooke, 2003; Stebbins,
2003; Hodges, 2009). However, populations are declining
across its range and the species is listed as threatened in
Texas. It is insectivorous, mostly myrmecophagous,
subsisting to a great extent on harvester ants (Pogonomyr-
mex).
The objective of the study was to obtain preliminary
baseline information on daily patterns of voluntary UVB
exposure and UVB dose of the Texas horned lizard in the
context of its thermal environment during sunny warm
days at a North Central Texas locality where the species is
abundant and protected. Based on its high thermal
tolerance (Heath, 1965; Prieto and Whitford, 1971) and
the open nature of its desert and grassland habitat, we
hypothesized that it is a UVB zone 4 (highest exposure
category) species.
MATERIALS AND METHODS—We conducted observational studies
on the Texas horned lizard at Matador Wildlife Management
Area (Matador WMA, Cottle Co., Texas; 3487 0 N, 100819.8 0 W;
elevation 570 m) during May and June from 2008–2012. The
management area includes 11,406 ha of rolling plains habitat,
including managed grassland and savannah. The terrain consists
of bluffs and bottom land of the Middle Pease River. A network
of unpaved roads is maintained and allows access to the habitat,
which is managed from an ecosystem standpoint to sustain
native wildlife populations. Texas horned lizards are common
throughout the refuge and lizards use the roads for morning
and evening activity.
We utilized two survey methods: spot-check surveys to
measure UVB irradiance at the locations of individual lizards
encountered in the field, and focal-retrace observations to
measure UVB dose of single individuals observed over periods of
time. Spot-check surveys included driving a vehicle on Matador
WMA roads through the refuge at slow speeds (8 km/h) from
approximately 0800–1300 h or from 1600–2100 h on days
judged to be sufficiently warm for Texas horned lizard activity
(air temperature [Ta] > 208C and little cloud cover). We usually
terminated morning surveys after 1300 h when we had seen no
lizards from 1200–1300 h. If we spotted a lizard during this time,
we continued searching for one additional hour. Each survey
covered approximately 33 km of unpaved Matador WMA roads
with no portion of any of the roads resurveyed during the same
survey. For the UVB study reported here, we conducted 23 spot-
check surveys (12 morning, 11 afternoon–early evening) from
2008–2012. As recommended by Hurlbert (1984) for studies
designed to answer questions such as ‘‘How does UVB exposure
intensity and temperature vary throughout the day?’’, our
replicate sample units (numbers) for each observational
June-September 2015 Ferguson et al.—UVB exposure of the Texas horned lizard
category (hour of the day) were single readings taken at the
precise location of an individual lizard or means of readings of
multiple Texas horned lizards found for an hour of a particular
survey. As a hypothetical example, one particular morning
survey might include three sightings during the 0900 h, two
sightings during the 1000 h, and one during the 1100 h. This
would provide one datum (mean) for the 0900 h, one (mean)
for the 1000 h, and one (single value) for the 1100 h. Another
survey on another day might include four sightings for the 1100
h but none for either the 0900 h or the 1000 h. This would
provide only one datum for the 1100 h. We did not include data
from three individual sightings that represented the only data
for that particular hour for all surveys in subsequent analyses.
We also used these sample units to analyze variation among the
categories and relationships among variables.
During six of the morning spot-check surveys conducted in
2011 and 2012, we took two readings for each individual, one
initial and one after a period of 5–20 min of observation. We
chose the number of minutes between the first and second
readings for each individual using a table of random numbers
between 5 and 20. The goal was to analyze the effect of delayed
processing of lizards discovered on the road on UVB exposure
intensity and to correct ‘‘road bias’’ (see Discussion).
In order to avoid interfering with additional ongoing
research at Matador WMA, we did not mark Texas horned
lizards during our study. Although we do not know the exact
number of lizards recaptured during the UVB study, lizards at
most of the 71 sighting locations were likely to be different
individuals. During the same years an independent mark-and-
recapture study was conducted by author DCR surveying the
same roads by vehicle. All Texas horned lizards encountered by
him and his associates were permanently marked by toe-clipping
or pit-tagging. During 63 surveys of the mark-and-recapture
study only 31 of 154 horned lizards encountered were
recaptured individuals (20.13%). Thus, the probability of
recapturing a given lizard at the refuge during the years of the
two studies was about 0.2. Therefore, we considered that any bias
of error variance due to measuring a few individuals twice
(pseudoreplication; Hurlbert, 1984) would be small. According-
ly, the sample unit for the analysis of initial vs. delayed readings
of individuals using a paired t-test was each individual lizard
encountered during the six surveys of Matador WMA conducted
in 2011 and 2012.
When the goal was to take only a single reading for each
Texas horned lizard encountered, we stopped the vehicle upon
detection of the lizard and noted the initial location of the
lizard. We captured the lizard immediately by hand or by noose,
and measured body temperature (Tb) using a quick-reading
cloacal thermometer (Miller and Weber, Inc., Ridgewood, New
York). We recorded snout–vent length (SVL) in millimeters and
body mass in grams with a plastic ruler and Pesola spring scale
(Forestry Suppliers Inc. Jackson, Mississippi), respectively. We
determined sex for adults by presence or absence of hemi-penal
bulges at the base of the tail. We released each lizard in less than
5 min at the location where it was first seen. After release, we
measured UVB irradiance, Ta, and substrate temperature (Ts) at
the location (see description of techniques below).
When we were to monitor a lizard twice to test for the effect
of delayed processing, the lizard was watched by one investigator
while the UVB and environmental temperatures were measured
by a second investigator. The second investigator took readings
233
at a location close to the lizard and with similar sun exposure
but sufficiently far away to avoid causing the lizard to change
location (approximately 3 m). After the predetermined delay,
we captured and processed the lizard at the second location as
described above.
We employed the focal-retrace method on single individuals
followed for most of a day. The goal of a focal-retrace session was
to estimate the UVB dose (irradiance · time) of an individual
Texas horned lizard for each hour of the day throughout most of
an entire day. The focal involved locating a single lizard on a day
designated as the ‘‘focal day.’’ We located a lizard as close to
0900 h as possible and followed it, if possible, until after 1800 h.
To enable close future monitoring of UVB exposure, we
recorded details of location and time every few minutes (»10
min), and also recorded whenever the lizard moved, or
whenever its sun exposure changed. After the lizard left a
location, we marked that location by inserting a numbered
survey flag into the substrate at the site. We took care to neither
disturb the lizard nor cause it to relocate when marking its
previous locations. Texas horned lizards at the refuge have a
relatively short approach distance before showing a predatory
flight reaction to a human, e.g., alertness, orientation away from
the observer, and flight behavior. We generally remained >3 m
from the lizard and moved slowly during observations.
Throughout the observation period we recorded general notes
on behavior, such as feeding and social interaction, and on
environmental changes, such as temporary cloud cover or
changes in air movement. We followed three adult Texas horned
lizards during most of their activity. We followed each on a
separate focal day, one male and one female in 2010 and one
male in 2012. At the end of the focal day we captured, weighed,
and measured each lizard. We took its body temperature and
released it at the site of capture within 5 min.
We could not directly monitor continuous UVB irradiance of
a lizard at the lizard’s location on a focal day without disturbing
the lizard and altering its activity or habitat use pattern.
Therefore, we assessed UVB irradiance in the field for each of
the three lizards as follows: On the day following the focal day,
designated ‘‘retrace day,’’ we monitored UVB irradiance, Ts, and
Ta at lizard locations marked on the previous day at the same
local time as they were occupied on the focal day. For each
lizard, we also monitored a single control site, located where it
would receive full sun exposure throughout the day, at the
beginning of each hour on the retrace day as well as on the focal
day to assess the climatic similarity of the focal and retrace days.
For all three lizards, values for UVB irradiance at the control
sites were similar on the 2 days (mean UVI difference < 1; Table
1), so no adjustments of the estimates for the focal-day UVI were
considered necessary. However, temperature values on the
retrace day at the control site for the female in 2010 were
warmer (>38C), so for each hour the estimate for the mean
temperature for that hour of her focal day was the retrace-day
value of the lizard site reduced by the mean difference in air
temperature at the control site for that hour of the 2 days. For
the male monitored in 2012 the UVB irradiance was also similar
for both the focal and retrace days (UVI difference < 1; Table
1), so no adjustment was needed. However, the temperatures
were considerably cooler (<68C) on the retrace day, so air
temperature estimates for his locations on the focal day were
adjusted in similar fashion, increasing the values for lizard
locations obtained on the retrace day by the hourly mean
234 The Southwestern Naturalist vol. 60, no. 2-3

TABLE 1—Cloud cover, ultraviolet index (UVI), and temperature data for the control sites monitored on both focal days (Day 1)
and retrace days (Day 2) for three Phrynosoma cornutum followed for most of a focal day at the Matador Wildlife Management Area,
Texas. Because value estimates for the lizard locations on the focal day were obtained on the retrace day, the control sites provided
information on the climatic similarity between the 2 days for each lizard. A mean UVI difference between the control values on the 2
days of >1 and a mean temperature difference >28C were arbitrarily considered grounds for adjusting the estimates for the lizard
locations obtained on the retrace day and applied to the focal day. Abbreviations in the table are as follows: indiv. (individual) , diff.
(Day 1 minus Day 2), Ts (substrate temperature), Ta (air temperature).

% Clouds UVI UVI UVI Ts Ts Ts Ta Ta Ta

h Indiv. Date Day 1 Day 1 Day 2 Diff. Day 1 Day 2 Diff. Day 1 Day 2 Diff.

0900 Female 2010 27 and 28 May 0.0 1.9 2.1 -0.2 30.2 34.0 -3.8 23.7 26.3 -2.6
1000 Female 2010 27 and 28 May 0.0 5.0 4.9 0.1 37.5 40.0 -2.5 27.6 34.2 -6.6
1100 Female 2010 27 and 28 May 5.0 7.7 7.3 0.4 40.2 44.6 -4.4 28.0 33.6 -5.6
1200 Female 2010 27 and 28 May 10.0 9.4 9.3 0.1 45.6 49.0 -3.4 30.7 37.0 -6.3
1300 Female 2010 27 and 28 May 10.0 9.5 9.6 -0.1 44.6 48.8 -4.2 33.1 39.2 -6.1
1400 Female 2010 27 and 28 May 10.0 9.7 7.5 2.2 47.8 48.8 -1.0 34.0 34.2 -0.2
1500 Female 2010 27 and 28 May 10.0 8.9 6.8 2.1 48.8 48.6 0.2 35.8 34.8 1.0
1600 Female 2010 27 and 28 May 10.0 7.2 6.4 0.8 39.2 46.0 -6.8 33.9 37.8 -3.9
1700 Female 2010 27 and 28 May 5.0 4.1 3.5 0.6 37.2 43.6 -6.4 31.7 37.2 -5.5
1800 Female 2010 27 and 28 May 5.0 1.5 1.4 0.1 34.2 40.6 -6.4 32.4 35.6 -3.2
— Mean female 2010 6.0 6.4 5.9 0.6 40.5 44.4 -3.9 31.1 35.0 -3.9
0900 Male 2010 16 and 17 June 0.0 1.7 1.5 0.2 29.2 29.2 0 27.3 26.6 0.7
1000 Male 2010 16 and 17 June 0.0 4.2 4.5 -0.3 30.4 35.4 -5.0 30.3 30.1 0.2
1100 Male 2010 16 and 17 June 5.0 7.5 7.7 -0.2 38.4 37.4 1.0 31.8 30.7 1.1
1200 Male 2010 16 and 17 June 15.0 9.1 10.1 -1.0 41.8 45.0 -3.2 34.5 32.8 1.7
1300 Male 2010 16 and 17 June 36.0 8.5 10.7 -2.2 42.8 45.2 -2.4 34.0 35.0 -1.0
1400 Male 2010 16 and 17 June 40.0 5.3 10.4 -5.1 43.4 47.4 -4.0 34.8 35.1 -0.3
1500 Male 2010 16 and 17 June 0.0 10.3 7.3 3.0 45.0 41.6 3.4 35.5 35.2 0.3
1600 Male 2010 16 and 17 June 0.0 7.4 7.8 -0.4 44.8 42.0 2.8 34.8 37.2 -2.4
1700 Male 2010 16 and 17 June 0.0 5.0 4.5 0.5 42.2 45.8 -3.6 35.1 36.9 -1.8
1800 Male 2010 16 and 17 June 0.0 2.5 2.6 -0.1 40.2 42.8 -2.6 34.0 35.5 -1.5
— Mean male 2010 9.6 6.2 6.7 0.6 39.8 41.2 -1.4 33.2 33.5 -0.3
0900 Male 2012 4 and 5 June 100.0 1.8 0.5 1.3 40.0 23.8 16.2 32.2 22.8 9.4
1000 Male 2012 4 and 5 June 80.00 3.1 5.2 -2.1 45.0 34.0 11.0 39.5 28.0 11.5
1100 Male 2012 4 and 5 June 100.0 4.5 3.2 1.3 44.6 31.2 13.4 36.5 28.2 8.3
1200 Male 2012 4 and 5 June 90.0 6.5 8.3 -1.8 51.6 38.2 13.4 38.0 31.8 6.2
1300 Male 2012 4 and 5 June 50.0 7.8 10.4 -2.6 52.8 38.8 14.0 39.5 34.5 5.0
1400 Male 2012 4 and 5 June 0.0 9.2 7.2 2.0 55.4 39.2 16.2 41.0 35.0 6.0
1500 Male 2012 4 and 5 June 0.0 8.6 8.3 0.3 59.0 43.6 15.4 42.5 36.2 6.3
1600 Male 2012 4 and 5 June 10.0 6.3 6.1 0.2 57.8 43.2 14.6 41.0 35.0 6.0
1700 Male 2012 4 and 5 June 96.0 1.2 3.7 -2.5 43.6 41.2 2.4 38.0 35.5 2.5
1800 Male 2012 4 and 5 June 10.0 0.5 1.0 -0.5 37.4 37.4 0 35.0 34.0 1.0
— Mean male 2012 53.6 5.0 5.4 0.4 48.8 37.1 11.7 38.3 32.1 6.2

difference between the temperatures of control site for the 2
days.
We measured UVB irradiance at lizard locations and control
sites using two handheld broad-band UVB meters, the Solar-
meter 6.2 (displays lW/cm2) and Solarmeter 6.5 (displays UVI)
(Solartech, Inc. Harrison Township, Michigan). These are
sensitive to irradiance within the UV range (280–400 nm) and
respond most strongly to wavelengths in the UVB range below
320 nm. We took readings by placing the base of the unit on the
substrate and orienting the flat sensor disk on the top of the unit
(»10.8 cm from the base) toward the sun or to the adjacent
open sky. If the line to the sun was obscured by features of the
habitat at the lizard location on the retrace day, we did not
relocate the meter but adjusted the orientation of the meter to
generate the maximum reading at each site that the lizard might
have experienced. When measuring, we avoided casting a
shadow from our bodies upon the meter.
We obtained Ts using a digital infrared thermometer (Raytek
PM-30 0 , Santa Cruz, California). We held the unit within 5 cm of
the substrate and activated it with the laser perpendicular to the
substrate. We obtained other temperatures (Ta and Tb) using a
cloacal thermometer. We obtained Tb by inserting the bulb of
the thermometer into the cloaca of the lizard. We obtained Ta by
placing the bulb of the thermometer approximately 5 cm above
the substrate location. If the location was sun-exposed, we
shaded the bulb from direct sunlight with a shadow provided by
obstructing direct sunlight approximately 10 cm above the
thermometer.
At control sites on focal days and retrace days, we estimated
percentage of cloud cover each hour using a spherical crown
June-September 2015 Ferguson et al.—UVB exposure of the Texas horned lizard 235

densitometer (Model A, Forestry Suppliers Inc. Jackson,

Mississippi).
We analyzed data on UVI and temperatures from spot-checks
as survey means or individual values as described above. To
generate UVB dose (irradiance · time) from focal or retrace
data, we averaged the UVB readings of the Solarmeter 6.2 from
two successive observations of an individual obtained on retrace
day and multiplied the average by the elapsed time interval in
seconds between the readings. Then we summed the interval
dose calculations over a longer observation period to obtain
dose value (lJ/cm2). To facilitate the presentation, we convert-
ed values to J/cm2 (lJ/cm2 ‚ 1,000,000). We totaled hourly
doses over 3-h time periods and for the full observation period.
When violations of assumptions of parametric analyses occurred,
we analyzed data with appropriate nonparametric tests. We
reported values as means – 1 SE or SD as indicated. All data were
analyzed and processed using Microsoft Excel 2007 or 2013, and
SigmaPlot 11.0 (Jandel Corp.).

RESULTS—Daily Patterns of the Thermal Environment, Body

Temperature, and Exposure to UVB during Road Activity—On
warm sunny days Texas horned lizards surveyed during
the 5 years were exposed to UVB irradiance throughout
the day when active between 0900–1900 h (Fig. 1a). When
lizards were on roads, Tb exceeded Ta (Fig. 1b). In the
morning (0900–1300 h), both Tb and Ts were significantly
> Ta (Tb = 37.1 – 0.59 SE, n = 23; Ts = 35.6 – 1.13 SE, n
= 25; Ta = 30.6 – 0.52 SE, n = 25; Kruskal-Wallace
ANOVA of ranks, H = 27.6, P < 0.001; Dunn’s method, P
< 0.05). In the late afternoon and early evening (1700–
2100 h), Tb was significantly > Ta (Tb = 34.4 – 0.80 SE, n
=19; Ta = 30.3 – 0.77 SE, n = 19; ANOVA of values F2,47 =
5.3; P = 0.001, P < 0.05, Holm-Sidak t-test) During this
period, Ts was only slightly > Ta (Ts = 31.8 – 1.01 SE, n =
19) and not significantly different (P > 0.05, Holm-Sidak
t-test). We used the nonparametric ANOVA of ranks test
and Dunn’s method for analysis of morning values due to
a violation of the equality of variance assumption of the
parametric ANOVA of values.
UVI exposure was significantly higher for Texas
horned lizards when first encountered in the morning
activity period (5.07 – 0.45 SE) than it was for those first FIG. 1—Mean – SD (vertical capped lines) of a) ultraviolet
encountered in the late-afternoon and early-evening index (UVI) and b) air and body temperatures (Ta and Tb,
activity period (0.66 – 0.46 SE; Mann-Whitney rank sum respectively) taken simultaneously at Phrynosoma cornutum
test t = 215.8, n = 19 [late afternoon and early evening], locations where discovered during sunny periods on unpaved
25 [morning], P < 0.001). There were significant roads at Matador Wildlife Management Area, Texas. Data are
hourly single or mean survey values from readings accumulated
correlations of UVI exposure with time of day, positive
during 12 morning and 11 late-afternoon and early-evening
in the morning (y = -16.1 + 2.1x; R2 = 0.80, P <0.001, n surveys conducted in May and June from 2008–2012. a)
= 25) and negative in the late afternoon and early Numbers shown on graph below the symbols are number of
evening (y = 5.3 - 0.27x; R2 = 0.53, P < 0.001, n = 18). values contributing to the mean and SD values for UVI, Tb, and
There was also a significantly positive correlation Ta at lizard’s location for panels a and b. b) Ta for late-afternoon
between UVI and Tb in the morning (y = -14.3 + and early-evening symbols are offset for clarity.
0.5x; R2 = 0.36; P = 0.002, n = 19) as lizards were
exposed to both solar infrared and UVB irradiation
while on roads (Fig. 1). reading resulted in a significantly lower average UVI
For those individuals monitored twice during road exposure of the second reading due to shade-seeking by
surveys in the mornings of 2011 and 2012, an initial some lizards. Readings were reduced from a mean UVI of
reading of UVI followed by a short delay and a second 5.6 to 4.2 (paired t = 2.33; df = 14; P = 0.035).
236 The Southwestern Naturalist
FIG. 2—Temporal pattern of ultraviolet-index exposure (UVI)
of a) male and b) female Phrynosoma cornutum and that of their
respective full-sun control sites. We followed each for most of a
day in 2010. Total ultraviolet-B doses for the observation period
were similar for the two lizards but the temporal UVI pattern
differed.
Pattern of Daily UVI exposure and UVB Dose for Focal
Individuals in 2010—Road surveys and analysis of spot-
check data provided little insight into the activity and
UVB exposure of Texas horned lizards during most of the
afternoon hours, because they were not visible from the
road. However, focal-retrace data of individuals revealed
patterns of variation throughout the day. These patterns
differed among the individuals (Figs. 2 and 3). A male
(Figs. 2a and 3a) observed during most of a warm (Ta =
34.68C), sunny day on 16 June 2010 moved into
vegetation at the edge of the road after 1200 h and
travelled approximately 140 m during a 5-h period along
the road to the east, shuttling between shaded and
partially shaded locations on the ground. During this
period, he was never more than 1 m from the edge of the
road but unlikely to be visible to people in passing
vehicles. His UVI exposure was relatively low during the
afternoon. At 1700 h he returned to exposed locations on
vol. 60, no. 2-3
FIG. 3—a) Mean – SD (capped vertical line) hourly UVB dose
of the male and female Phrynosoma cornutum from Fig. 2 and that
of their respective full-sun control sites for three 3-h time
periods on two sunny days. b) Mean – SD hourly UVB dose of a
male P. cornutum and that of its respective full-sun control site
for three 3-h time periods on an unusually hot, partially overcast
day in 2012. Total dose for the day was lower (0.84 J) than for
the male or the female on sunny days in 2010 (see Fig. 2).
Temperatures for each lizard are average estimated air
temperature at the lizard site on the focal day for each period
(n = 3) or for the total observation period (n = 9).
the road and received a higher level of UVI exposure
throughout the remainder of the observation period. His
Tb, taken after the period of observation at 1839 h, was
398C. A female (Fig. 2b, 3a) observed on a cooler (Ta =
31.28C) sunny day on 27 May 2010 went deeper into the
vegetation adjacent to the road (1–2 m) at about 1000 h
and remained off of the road surface until about 1800 h.
Her eastward movement along the road was for a much
June-September 2015 Ferguson et al.—UVB exposure of the Texas horned lizard
shorter distance (about 30 m, 19 location changes) than
that of the male (140 m, 28 location changes). The
female experienced a higher afternoon UVI exposure
than the male by climbing into herbaceous vegetation
and shuttling vertically between fully and partially sun-
exposed locations. She returned to the open road, where
she likely would be visible to people in passing vehicles, at
1800 h. Her Tb, taken after her period of observation at
1850 h, was 368C. Both lizards occupied the same general
area along the same road in the morning of their
respective focal days but the male travelled much farther
to the east than the female.
The male and female Texas horned lizards in 2010 had
different patterns of UVB exposure yet received similar
daily UVB doses (Figs. 2 and 3). Thus, on the warmer
sunny day in June the male received most of his daily dose
in the morning and late afternoon (Fig. 3a). He shunned
full sun exposure in the afternoon until 1700 h. By
contrast, on the cooler sunny day in May, the female
received most of her daily dose in the morning and early
afternoon (Fig. 3a). She exposed herself to full sun in the
morning and during the warmest time of day in early
afternoon. During this time she displayed thermoregula-
tory cooling behavior by opening her mouth on four
occasions for periods of up to 2 min. The male never
displayed this behavior during his focal observation
period on the warmer day.
The duration of full sun exposures were generally
short throughout the day (<5 min) for both the male and
female and were interspersed with relocation to and from
shaded locations (Fig. 2). Only the male spent an
extended period in full sun (1 h) between 1730 and
1830 h.
Effect of the Thermal Extremes and Cloud Cover on UVB
Dose—Observation on a day with more extreme weather
conditions in 2012 revealed a different pattern of UVB
exposure of a Texas horned lizard male. That day (4 June
2012) was unusually hot (Ta = 37.48C, peak afternoon Ta
= 408C at the lizard site; >428C at the control site; Table
1, Fig. 3b) and partly overcast (54% cloud cover; Table 1,
Fig. 3b). He avoided all light exposure after 1400 h by
burying into the sandy substrate in the shade and
remained until late afternoon. His daily dose of UVB
was less than one-third that of the male and female
observed on the sunny days in 2010, and who remained
active aboveground all day.
Size and Sex of Monitored Texas Horned Lizards—When
conducting spot-check surveys, we sighted 71 lizards (19
males, 26 females, 19 unsexed juveniles, 7 where age and
sex was undetermined). Of those, we captured, measured,
and weighed 58 lizards. Adult males had an SVL of 83.1 –
7.2 SD mm and a mass of 32.9 –7.2 SD g, n = 15. Adult
females had an SVL of 87.5 – 8.3 SD mm and a mass of
43.0 – 13.5 SD g, n = 26. Juveniles had an SVL of 52.5 –
4.1 SD mm and a mass of 7.3 – 2.3 SD g, n = 17. Sizes for
the male observed during the focal-retrace study in 2010
237
were an SVL of 81 mm and mass of 30 g. Those for the
female observed in 2010 were an SVL of 90 mm and a
mass of 52 g. Those for the male observed in 2012 were an
SVL of 74 mm and a mass of 24 g.
DISCUSSION—Texas horned lizards studied at Matador
WMA in North Central Texas in 2011 and 2012, when
initially encountered on the road in the morning, were
found to be exposing themselves to UVB with a mean UVI
exposure of 5.6. When the same lizards were followed for
a short period and some individuals chose to seek shade,
the mean dropped to 4.2. During road surveys by
automobile, it is less likely for investigators to see Texas
horned lizards when lizards are in shade than when in
fully exposed locations on the road. This leads to a ‘‘road
bias’’ when comparing the UVB exposure of Texas
horned lizards surveyed from a moving vehicle to that
of other lizard species surveyed by walking through the
habitat. When walking and searching, investigators are
more likely to see lizards in partial- and full-shade than
when they are searching from a moving vehicle, even at
slow speeds. Thus, following and repeating the collection
of data can correct the road bias and facilitate compar-
ison. Both the ‘‘uncorrected’’ (UVI » 5) and ‘‘corrected’’
(UVI » 4) exposure estimates for Texas horned lizards
exceed the UVI median exposure of 3.1 for zone-4 lizards
reported to date (Ferguson et al., 2010) and support its
designation as a UVI zone-4 species. The mean UVI value
of 4.2 is similar to that of 4.34 measured for Sceloporus
graciosus gracilis in northern California measured through-
out the day (Ferguson et al., 2014).
The UVB dose measured for two individual Texas
horned lizards followed throughout most of their daily
activity period on sunny days (»3 J/cm2) exceeds that for
any other squamate reptile measured to date. It surpassed
even that of S. g. gracilis in northern California, which was
»2 J/cm2 for the two individuals monitored by Ferguson et
al. (2014). Higher latitude, higher altitude, a shorter
activity period, and more haze due to wildfires near the
study areas of northern California than at those of North
Texas might have reduced the availability of solar UVB and
the opportunity for exposure for S. g. gracilis in northern
California. More research is needed to determine if the
daily UVB doses for the two species are species-specific.
Although we monitored only three individuals for most
of the day using the focal-retrace method, researchers can
test hypotheses of the causes of variation in the daily
pattern of UVB exposure of Texas horned lizards in North
Texas with future focal-retrace research. We hypothesize
that activity and UV exposure patterns vary with the
degree of overcast and extreme thermal conditions. On
warmer sunny days shade-seeking increases in late
morning and midday UVI exposure is reduced. Thus,
the male that was monitored in 2010 was observed on a
warmer day than the female and restricted his afternoon
sun exposure more than did the female. However, on a
238 The Southwestern Naturalist
hot day when air temperature approached and exceeded
upper levels of the optimal thermal range for Texas
horned lizards, a male Texas horned lizard ceased
aboveground activity, which interrupted his UVB expo-
sure, and reduced his overall dose for the day. As hot days
become more frequent after June, Texas horned lizards
often escape heat by burrowing into the substrate for
much of the afternoon (Fair and Henke, 1998, 1999;
Burrow et al., 2001; Sherbrooke, 2003). When thermal
conditions (Ta and Ts) exceed the optimum thermal
range of the species, aboveground activity and UVB
exposure is likely to be constrained.
The effect of high environmental temperatures on
solar exposure may be relevant to conservation of the
Texas horned lizard. Throughout much of the distribu-
tion of this species, overall temperature increase caused
by global warming is likely to alter the availability of
suitable thermal microclimates for many lizard species
(Angilletta, 2009; Sinervo et al., 2010; Huey et al., 2010).
Climate change models that attempt to predict impacts
on lizard species (e.g., Buckley, 2008; Williams et al.,
2012) should incorporate not only reductions of nutri-
tional energy intake as a parameter but also nutritional
quality. The opportunity to endogenously produce
vitamin D by exposure to UVB is a component of
nutritional quality that could be compromised by
increased thermal constraint of aboveground activity.
As shown in previous research, Texas horned lizards
rely on open, often bare, ground in the morning and late
afternoon for basking (Whiting et al., 1993; Fair and
Henke, 1998; Burrow et al., 2001). Accordingly, unpaved
roads at Matador WMA are used by lizards as basking sites
in the morning until body temperatures, and possibly UVI
tolerance, approach the upper limits of the lizard’s
optimal range. Because Texas horned lizards have a
thermal tolerance zone of several degrees Centigrade
(Heath, 1965; Sherbrooke, 2003), they appear to allow
their Tb to gradually increase within this zone as they
periodically expose themselves to increasing solar inten-
sity until late morning. At that time they begin to seek
shade and might remain on the ground but some, such as
the female that we observed in 2010, may continue to
seek solar exposure by climbing vertically into vegetation,
a behavior also reported previously for this species
(Whitford and Bryant, 1979; Sherbrooke, 2003). The
significant positive correlation between body temperature
and UVI in the morning from spot-check data is
consistent with this interpretation of individual behavior.
The stimulus to return to roads in late afternoon might
be to bask for thermoregulation as the availability of other
heat sources wane. However, significant enhancement of
daily UVB dose is possible during the early part of their
late-afternoon and evening activity period until 1900 h.
Knowledge of the daily patterns of thermal exposure
and the UVB zone of reptiles including the Texas horned
lizard provides useful information for captive maintenance
vol. 60, no. 2-3
programs for conservation, education, and research. With
this knowledge, captive-animal managers can establish
appropriate thermal and UVB gradients and establish
natural daily and seasonal patterns for their animals.
Because Texas horned lizards shuttle in and out of sunlight
in nature, a UVI light gradient ranging from 0–8 should be
available to the lizards for up to 9 h per day during spring
and summer in captivity. Thermal gradients that allow
them to maintain body temperatures ranging from 34–
408C should also be provided for up to 12 h per day in
spring and summer. While experimental documentation is
needed to verify whether or not Texas horned lizards are
able to regulate their UVB dose behaviorally, studies
support the hypothesis that other basking lizard species
are capable of adjusting their UVB exposure depending on
their internal vitamin D status (Ferguson et al., 2003;
Karsten et al., 2009; Ferguson et al., 2013).
We thank M. Vaughan, S. Fredlake, S. Hammack, J. Crowley, J.
Chavez, J. Le Van, and S. Bucklin for assistance in the field. K.
Karsten made helpful suggestions during the formulation of
research procedures. M. Nakamoto provided assistance with the
Spanish translation of the abstract. Research was authorized by a
permit from Texas Parks and Wildlife to GWF, SPR 0690-146, and
permission from Matador WMA. We especially thank the staff of
Matador WMA for help and guidance during the field work.
Research was approved by Texas Christian University Institutional
Animal Care and Use Committee protocol 11/04. Partial funding
of the project was by grants from the Faculty Emeritus Research
Fund from Texas Christian University to GWF.
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