15 (4): 2-xxx (2008) Preprint web release / Parution Internet préimpression

15 (4): 2-xxx
2008-10-31
(2008)

Correlations of biotic and abiotic variables

with ground surface temperature:
An ectothermic perspective1
Glenn M. CUNNINGTON2, James SCHAEFER, Joseph E. CEBEK & Dennis MURRAY,
Department of Biology, Trent University, 1600 West Bank Drive, Peterborough, Ontario K9J 7B8, Canada.

Abstract: Temperature is often an important determinant of species presence and activity patterns, but observations of
temperature at fine resolution and broad extents—scales relevant to individual organisms—are rare. We analyzed how biotic
and abiotic variables influence ground surface temperature (GST) using high-resolution thermal data for a 1900-ha area of
forest and field near Wasaga Beach, Ontario, Canada. Temperature was recorded at 2-h intervals from May to October, 2004
and 2005, from data loggers approximately 300 m apart. At the same sites, we recorded overstory and understory vegetation,
elevation, and distances to the nearest road and urban infrastructure. Correspondence Analysis (CA) indicated that thermal
conditions varied most strongly seasonally and diurnally. Canonical Correspondence Analysis (CCA) pinpointed which
biotic and abiotic variables were related to surface temperature. Canopy cover and leaf litter cover were positively related to
cool sites during both cool and warm periods. Sites close to urban areas and roads were warmer than those further away from
these features. As thermally variable environments affect the physiology, behaviour, and ecology of ectothermic species,
determination of variables that affect GST provides a more comprehensive depiction of habitat for these species.
Keywords: abiotic, biotic, ectotherm, ground surface temperature, habitat, urban heat island, hognose snake.

Résumé : La température est souvent un déterminant important de la présence d’espèces et de leurs patrons d’activité, mais
les observations de température à résolution fine sur de grandes étendues — les échelles appropriées aux organismes
individuels — sont rares. Nous avons analysé comment les variables biotiques et abiotiques influencent la température de
surface du sol en utilisant des données thermiques à haute résolution pour une aire de 1900 ha de champs et de forêts près
Wasaga Beach, Ontario, Canada. Des enregistreurs électroniques de données espacés d’environ 300 m ont mesuré la température
à chaque 2 heures entre mai et octobre en 2004 et 2005. Aux mêmes sites, nous avons noté la végétation de la canopée et du
sous-étage, l’élévation et la distance à la route et à l’infrastructure urbaine la plus proche. L’analyse des correspondances a
indiqué que les conditions thermiques ont varié de façon la plus importante au sein de la saison et de la journée. L’analyse
canonique des correspondances a identifié précisément quelles variables biotiques et abiotiques étaient reliées à la température
de surface. Le couvert forestier et la couverture de détritus de feuilles étaient reliés de façon positive aux sites frais durant les
périodes tant fraîches que chaudes. Les sites près des zones urbaines et des routes étaient plus chauds que ceux plus éloignés de
ces éléments. Puisque des environnements thermiques variables ont un effet sur la physiologie, le comportement et l’écologie
des espèces ectothermes, la détermination des variables influençant la température de surface du sol permet une description
plus complète de l’habitat de ces espèces.
Mots-clés : abiotique, biotique, couleuvre à nez retroussé, ectotherme, habitat, îlot de chaleur urbain, température de surface
du sol.

Introduction
Ecosystems are influenced by numerous biotic and
abiotic variables (Saunders et al., 1998). The realization of
conservation goals requires an understanding of the manner
in which these environmental variables influence the abil-
ity of species to exist (Corney et al., 2004). Temperature
is often included in studies of environmental variables
that affect species presence and activity patterns; how-
ever, Jarvis and Stuart (2001) indicated that there is a
surprising lack of studies that focus on the determination
of temperature at both high resolution and large extents.
Climatic modeling and the determination of ground surface
temperature (GST) from satellite imagery historically have
1Rec. 2007-09-20; acc. 2008-04-14.
Associate Editor: Christian Messier.
2 Author for correspondence. Ottawa-Carleton Institute of Biology, Carleton
driven interest in broad-scale GST (Nelson, Siegel & Yoder,
2004). Expensive temperature data logger equipment in
the form of national weather stations has been utilized for
decades (Changnon, 1999; Jarvis & Stuart, 2001; Arnfield,
2003; Englehart & Douglas, 2003; Corney et al., 2004), but
technological advancements in the form of mobile thermal
data loggers have allowed biologists to ask questions at fine
scales, pertinent to habitat use of some species. Such inves-
tigations were logistically impossible in the past (Angilletta
& Krochmal, 2003).
As GST varies both spatially and temporally, the deri-
vation of spatially distributed models can be problematic
because of both theoretical complexity and data availability
(Jarvis & Stuart, 2001). Lewis (1998) indicated that differ-
ences in GST in a single year stemmed from both biotic and

University, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6, Canada, e-mail: glenn.
abiotic features. Abiotic variables are often drivers of further
cunnington@gmail.com ecosystem and landscape-level processes, such as species
DOI 10.2980/15-4-3140 distributions, plant growth, and nutrient cycling (Saunders

et al., 1999). Alteration of the landscape through natural or
anthropogenic events may also influence the thermal condi-
tions at a site (Saunders et al., 1998). Determination of the
variables that affect GST would thus provide a basis to infer
potential ecological effects of both natural and anthropo-
genic processes.
There have been few fine-scale studies that have uti-
lized small electronic devices to gather extensive thermal
data (Halley, Eriksson & Nunez, 2003). Newmark (2005)
utilized a limited number of electronic data loggers to deter-
mine differences in the diurnal thermal regime between
forest edge and interior. These devices are often expensive
(Angilletta & Krochmal, 2003) and utilized only in small
numbers (Newmark, 2005). When such devices are not
available, fine-scale studies rely on point sampling in lim-
ited quantities (Stabler, Martin & Brazel, 2004).
Given that temperature plays a significant ecological
role in the life history of many species, especially ecto-
therms, it is important to establish what variables affect
surface temperature (Jarvis & Stuart, 2001). We tested the
hypothesis that biotic factors, such as canopy cover and
ground vegetation, lower GST, while abiotic factors, like
urban features and roads, increase GST. Canopy cover may
act to reduce the effects of climatic heating events by shift-
ing the thermally active surface vertically from the ground
surface (Oke, 1997). Thus, we anticipated that temperature
at locations further from anthropogenic influences would be
driven by biotic variables and would be cooler, while areas
closer to anthropogenic features would experience higher
temperatures than those farther away. We then considered
the implications of these results for an ectotherm, the east-
ern hog-nosed snake (Heterodon platirhinos), a threatened
species in our study area (Cunnington & Cebek, 2005).
Methods
Field data were collected over 2 seasons, May to
October 2004 and May to October 2005, in a 1900-ha
site near the town of Wasaga Beach, Ontario (44°  30' n,
80° 01' w). Wasaga Beach is located on Georgian Bay on
Lake Huron. Given its wide variety of vegetative communi-
ties (Brunton, 1989) and nearby anthropogenic influences,
our study area provided an excellent opportunity to study
the effects of biotic and abiotic variables on GST.
Ambient temperatures were collected by a weather sta-
tion equipped with a Stevenson Screen (Davis Instruments
Weather Monitor II, Hayward, California, USA). Ground
surface temperatures were collected by Thermochron iBut-
ton temperature data loggers (Maxim Dallas Semiconductor
DS1921G, Sunnyvale, California, USA, accuracy ± 1 °C)
placed throughout the park. To facilitate their retrieval,
iButtons were taped to 0.75-m wooden stakes. The stakes
were driven into the ground such that the iButton faced
south and was located at the ground surface. Temperatures
were collected every 2 h at the weather station and iButton
locations. The locations for data loggers in 2004 (n = 112)
were generated from random numbers. The locations of
data loggers that were successfully retrieved in 2004 were
reused in 2005. The remaining locations for 2005 (n = 325)
were obtained by systematically placing data loggers 300 m
ÉCOSCIENCE, vol. 15 (4), 2008
apart in a grid. The data loggers were deployed with a GPS
unit (Garmin 72, Olathe, Kansas, USA; accuracy ± 4 m),
and the iButton locations were mapped electronically using
ArcView 3.2 (Environmental Systems Research Institute,
1999a). To determine if the iButton temperatures accu-
rately reflected ambient conditions a single iButton was
placed adjacent to the weather station. Data collected by the
weather station and adjacent iButton were analyzed with a
correlation analysis.
Vegetation surveys were conducted in plots (of 1 and
5 m radius) centred on each data logger location. These data
included elements assumed to affect the thermal conditions
on the ground surface (Table I; Nielson, 1985; Lewis, 1998;
Halley, Eriksson & Nunez, 2003). Vegetation surveys were
conducted in June and July of 2004 and 2005.
Additional variables were determined with GIS (Table I).
ASPECT was calculated on a scale of 1 to 180, with 1
representing north and 180 representing south; east and
west were given equal values (90). ELEVATION, ROAD,
and URBAN were calculated with the ArcView 3.2
Geoprocessing extension. SLOPE was produced with Spatial
Analyst (Environmental Systems Research Institute, 1999b).
Multivariate statistical analyses were carried out using MVSP
(Kovach Computing Services, 2006).
Temperature data collected by iButton data loggers were
analyzed with Correspondence Analysis (CA) to determine
how season and time of day affected site-specific tempera-
tures in 2004 and 2005. To illustrate how CA extracted tem-
poral variation in temperature, extreme sites were selected
confirm the axes interpretation. Data were also analyzed
with Canonical Correspondence Analysis (CCA) to deter-
mine which biotic and abiotic variables were related to
Table I. Environmental variables used in the analysis of ground
surface temperature.
Source Variable Description
Plot1 CANOPY Canopy closure (%)
Plot1 LEAF Coverage (%) of leaf litter
Plot1 LOG Coverage (%) of downed woody debris
(> 5 cm diameter)
Plot1 VEG Coverage (%) of herbaceous vegetation
Plot1 SOIL Coverage (%) of bare soil
Plot1 STUMP Coverage (%) of stumps
Plot1 VEGDENS Number of herbaceous stems
Plot2 OVER D Presence or absence of deciduous
tree > 15 cm DBH3
Plot2 UNDER D Presence or absence of deciduous tree
< 15 cm DBH
Plot2 OVER C Presence or absence of coniferous tree
> 15 cm DBH
Plot2 UNDER C Presence or absence of coniferous tree
< 15 cm DBH
GIS4 ELEVATION Elevation at 5 m resolution
GIS4 ROAD Reciprocal of Euclidean distance (m) to
nearest paved road
GIS4 URBAN Reciprocal of Euclidean distance (m) to
nearest urban area
GIS4 SLOPE Slope (%)
GIS4 ASPECT Aspect
1 1-m-radius plots.
2 5-m-radius plots.
3 DBH = diameter at breast height.
4 Calculated with Geographic Information System software.
3
Cunnington et al.: Variation in ground surface temperatures
surface temperature. CCA seeks to determine coefficients of
environmental variables to obtain a site score to maximize
the variance of the average position of species (Schaefer,
Stevens & Messier, 1996) or, in this case, temperature.
Twenty eastern hog-nosed snakes were opportunisti-
cally captured by hand and radio-tagged. Observations
of behaviour and body temperature (T b) were obtained
by radio telemetry between May 2002 and October 2005.
Only snakes with a body mass > 200 g were implanted
(R.  Willson, unpubl. data) with a temperature-sensitive
radio transmitter (Holohill SI-2T, Carp, Ontario, Canada;
9 g) with 0.5 °C resolution and 1.0 °C accuracy. To allow
snakes to return to their normal movement patterns, no
behavioural or Tb observations were included within 10
d of release. Snakes were classified as active (on ground
surface) or inactive (below ground surface). Each time an
individual was located by telemetry, Tb, behaviour (active
or inactive), and ground temperature (Tg) were recorded. Tg
was collected with a digital hygrometer (VWR International
35519-050, Mississauga, Ontario, Canada) by placing the
temperature probe on the ground surface within 5 m of the
snake. Linear regression was used to test if Tg and Tb were
related. We also determined the percentage of snakes active
versus Tb. To maintain precision, only temperatures with >
20 observations were included.
Results
Temperatures were recorded every 2 h from May 30 at
0000 to October 2 at 2200 in 2004 and 2005, resulting in
Figure 1. Correspondence Analysis ordination of 2004 temperatures at data logger sites. Black symbols represent sites selected for illustrations of diurnal
temperature profiles (mean temperature versus time of day for May 31; inset).
4
1512 temperature readings per data logger each year. One
hundred and twelve data loggers were placed in the field in
2004, but only 79 were recovered at the end of the season.
Similarly, only 259 of 325 data loggers were recovered
in 2005. Thermal data collected by the weather station in
2004 and 2005 indicated that the mean spring temperature
in 2005 (18.6 °C) was higher than in 2004 (10.7 °C); how-
ever, the mean summer temperature was warmer in 2004
(17.3 °C) than in 2005 (15.7 °C). Thermal data collected
by the weather station and the adjacent iButton were cor-
related (r2 = 0.471).
Results from the 2004 CA indicated that 50.3% of the
temporal variation in temperature was explained by the first
axis and 14% was explained by the second axis (Figure 1).
To interpret Axis 1, CA scores were plotted against time of
day and month (Figure 2). This indicated that the first axis
ordered the sites based on diurnal variation and the second
axis on seasonal variation (Figure 1). Data from 4 selected
locations (Figure 1 inset) showed that high CA1 values were
indicative of warmer conditions during daytime, especially
from 1000 to 2000, but not during nighttime.
Similarly, for the 2005 CA, 48.5% of the variation was
explained by the first axis and 12.7% was explained by the
second axis (Figure 3). Diurnal variation, the first axis, was
represented by a gradient of cool days/warm nights to warm
days/cool nights (Figure 3). The second axis revealed a gra-
dient of cool spring/summer and warm fall to warm spring/
summer and cool fall (Figure 3).
 ÉCOSCIENCE, vol. 15 (4), 2008

The 2004 CCA indicated that distances to urban areas
and roads had the greatest effect on surface temperature
(Figure 4). Deciduous overstory and understory cover, soil,
and elevation were related to warm sites both seasonally and
diurnally (Figure 4). Coniferous overstory and understory,
leaf, and canopy cover were related to diurnally cool sites.
The 2005 CCA produced somewhat different results.
While roads and soil were still related to diurnally warm
sites, proximity to urban areas was no longer a factor
(Figure 5). Canopy and leaf cover continued to be related
to diurnally cool sites; however, overstory and understory
deciduous were related to diurnally cool sites as well
(Figure 5). Urban area was not a driver of either seasonal or
diurnal thermal variation in 2005.
A total of 1964 observations of snake behaviour, Tb,
and Tg were obtained from 20 adults between 2002 and
2005. Linear regression with T b as the dependent vari-
able and Tg as the independent variable generated a slope

Figure 2. Interpretation of CA axes for 2004. a) CA axis 1 scores plotted against time of day and b) mean CA axis 2 scores plotted against month.

Figure 3. Correspondence Analysis ordination of 2005 temperatures at data logger sites. Black symbols represent sites selected for illustrations of diurnal
temperature profiles (mean temperature versus time of day for May 31; inset).

5
 Cunnington et al.: Variation in ground surface temperatures

Figure 4. Axis 1 and 2 of canonical correspondence analysis (CCA)
associating site temperature conditions and environmental variables for
2004. See Table I for definitions of variables.
Warm Spring/summer
size and limited spatial extent studies and understand the fine
scale thermal dynamics of the landscape.
The 2 y of study differed markedly. This provided us
with an opportunity to compare not only which variables
affect GST, but also how those variables vary in years with
contrasting ambient conditions. In 2004 (cooler spring,
warmer summer), anthropogenic variables such as proxim-
ity to urban areas and roads had the greatest influence on
diurnal variation in temperature (Figure 4). Given the urban
heat island effect (Oke, 1979), it is not surprising that these
variables contributed to increases in GST. It is of interest
then that in 2005 (warmer spring, cooler summer) distance
from urban areas did not greatly influence GST (Figure 5).
The vector representing URBAN does not appear in Figure 5
as it is too short to be visible in conjunction with the other
environmental variables. We propose that the effects of some
variables, like urban areas, interact with prevailing ambi-
ent conditions in complex ways that need further inquiry.
Previous studies have indicated that natural processes do
not have as great an impact on GST as anthropogenic distur-
bances (Lewis, 1998; Saunders et al., 1998).
Biotic variables such as percent canopy cover and leaf
litter were related to cool sites (both seasonally and diur-

nally) in 2004 (Figure 4). Plant canopies have been shown

Axis 2 to provide a climatic buffer for the ground surface, provid-
ELEVATION ing shading from extreme temperatures during the day and
cool fall

a downward flux of heat at night (Oke, 1997). The effects

of canopy and leaf litter cover were accentuated in 2005
OVER C (Figure 5); it is likely that the shading provided by these
SLOPE variables decreased GST. While this result is intuitive, it
ROAD D echoes Lewis (1998), who found that deforestation resulted
LEAF
A VEG in an immediate and long-term increase in GST.
Axis 1
Seasonal
variation

CANOPY
UNDER D SOIL The presence of deciduous understory and overstory
C
OVER D trees was positively related to warm sites in the cool year;
LOG B however, these same variables were positively related to
Cool Spring/summer

cool sites in the warm year. This result is somewhat con-

fusing. Nielson (1985) found similar temperatures existed
warm fall

below both deciduous and coniferous canopy; however,

Cool days Diurnal Warm days
warm nights variation cool nights
Figure 5. Axis 1 and 2 of canonical correspondence analysis (CCA)
associating site temperature conditions and environmental variables for
2005. See Table I for definitions of variables.
(95% CI: 0.85 < b1 < 0.91) that was significantly less than
unity and an intercept (2.02 < b0 < 3.59) greater than zero
(y = 0.877x + 2.805, r2 = 0.477, F1,1014 = 924). This indicat-
ed that while Tg successfully predicted Tb, there was much
variation not explained by Tg. Snakes were not active when
Tb was below 14 °C or above 37 °C. Snakes were always
observed active when Tb was above 21 °C.
Discussion
The results of our study supported our hypothesis that
biotic factors, such as canopy cover and ground vegetation, are
associated with lower GST and that abiotic factors, like urban
features and roads, are linked to higher GST. Furthermore, our
study methodology allowed us to move beyond small sample
light penetration was less when coniferous trees were pres-
ent. Our result was likely caused by seasonal variation in
deciduous canopy cover, while coniferous canopy cover is
essentially static. It would appear that the amount of bare
soil at a site affects GST in a similar manner; shorter dis-
tance to nearest road in both 2004 and 2005 was associated
with warm sites (Figures 4 and 5).
Our regression analysis of Tg and Tb indicates that
ground temperature is related to a snake’s body tempera-
ture; however, it is not a simple relationship. The lack of
strong fit (r2 = 0.477) implies that snakes may have mod-
est scope to modulate body temperature (Brattstrom, 1965;
Huey et al., 1989). The eastern hog-nosed snake Tb data
allow for an interesting assessment of the landscape thermal
data. The upper lethal limit for eastern hog-nosed snakes is
41.4 ± 1.8 °C, and these snakes seek cooler body tempera-
tures when they reach 36.1 ± 1.2 °C (Figure 5; Kroll, 1973).
The thermal limits of the species demonstrate that under-
standing the variables that affect GST can provide insight
into the habitat requirements for an ectotherm. Given the
strong effects of urbanization and roadways on GST (Oke,
1973), it is apparent that the impacts of these anthropogenic

6

features extend beyond reduction in habitat in a purely
structural sense.
Since temperature fluctuates both temporally and spa-
tially, our analysis lends itself to the formation of a broader
definition of habitat. The dramatic seasonal and diurnal
fluctuations of temperature require that habitat be redefined
as a dynamic variable. Thermally suitable habitat varies not
only on a seasonal basis, as conventionally treated, but even
more markedly on a diurnal basis (Figures 1 and 2). The
definition of an ectothermic species’ habitat may need to
account for such thermal dynamics of the landscape at bio-
logically meaningful spatial and temporal scales.
Acknowledgements
We gratefully acknowledge the assistance of M.  Shoreman,
M.  Beecroft, K.  Janke, C.  VanNess, and P.  Wilson. Research was
funded by the Ontario Ministry of Natural Resources, the Friends
of Nancy Island Historic Site, Ontario Parks, and a Natural Sciences
and Engineering Research Council Discovery Grant to J. A.
Schaefer. This study was conducted following protocols approved
by the Ontario Ministry of Natural Resources (MHFW1004).
Literature Cited
Angilletta, M. J. & A. R. Krochmal, 2003. The Thermochron:
A truly miniature and inexpensive temperature-logger.
Herpetological Review, 34: 31–32.
Arnfield, J. A., 2003. Two decades of urban climate research: A
review of turbulence, exchanges of energy and water, and the
urban heat island. International Journal of Climatology, 23: 1–26.
Brattstrom, B. H., 1965. Body temperature of reptiles. American
Midland Naturalist, 73: 376–422.
Brunton, D. F., 1989. Biological Inventory and Evaluation of Wasaga
Beach Provincial Park and Adjacent Natural Areas, Simcoe
County, Ontario. Parks and Recreational Areas Section, Ontario
Ministry of Natural Resources, Richmond Hill, Ontario.
Changnon, S. A., 1999. A rare long record of deep soil tempera-
tures defines temporal temperature changes and an urban heat
island. Climatic Change, 42: 531–538.
Corney, P. M., M. G. LeDuc, S. M. Smart, K. J. Kirby, R. G. H.
Bunce & R. H. Marrs, 2004. The effect of landscape-scale
environmental drivers on the vegetation composition of British
woodlands. Biological Conservation, 120: 491–505.
Cunnington, G. M. & J. E. Cebek, 2005. Mating and nesting
behavior of the eastern hognose snake (Heterodon platirhi-
nos) in the northern portion of its range. American Midland
Naturalist, 154: 474–478.
Englehart, P. J. & A. V. Douglas, 2003. Urbanization and seasonal
temperature trends: Observational evidence from a data-sparse
part of North America. International Journal of Climatology,
23: 1253–1263.
Environmental Systems Research Institute, 1999a. ArcView GIS.
Version 3.2. Redlands, California.
ÉCOSCIENCE, vol. 15 (4), 2008
Environmental Systems Research Institute, 1999b. Spatial Analyst.
Version 1.1. Redlands, California.
Halley, V., M. Eriksson & M. Nunez, 2003. Frost prevention
and prediction of temperatures and cooling rates using GIS.
Australian Geographical Studies, 41: 287–302.
Huey, R. B., C. R. Peterson, S. J. Arnold & W. P. Porter, 1989.
Hot rocks and not-so-hot rocks: Retreat-site selection by garter
snakes and its thermal consequences. Ecology, 70: 931–944.
Jarvis, C. H. & N. Stuart, 2001. A comparison among strategies for
interpolating maximum and minimum daily air temperatures,
Part I: The selection of “guiding” topographic and land cover
variables. Journal of Applied Meteorology, 40: 1060–1074.
Kovach Computing Services, 2006. Multivariate Statistical
Package (MVSP), Version 3.13n. Anglesey, Wales.
Kroll, J. C., 1973. Comparative Physiological Ecology of Eastern
and Western Hognose Snakes (Heterodon platirhinos and
Heterodon nasicus). Ph.D. thesis. Texas A&M University,
College Station, Texas.
Lewis, T., 1998. The effect of deforestation on ground surface tem-
peratures. Global and Planetary Change, 18: 1–13.
Nelson, N. B., D. A. Siegel & J. A. Yoder, 2004. The spring bloom
in the northwestern Sargasso Sea: Spatial extent and relation-
ship with winter mixing. Deep Sea Research, 51: 987–1000.
Newmark, W., 2005. Diel variation in the difference in air tem-
perature between the forest edge and interior in the Usambara
Mountains, Tanzania. African Journal of Ecology, 43: 177–180.
Nielson, E. T., 1985. Seasonal and diurnal leaf movements of
Rhododendron maximum L. in contrasting irradiating environ-
ments. Oecologia, 65: 296–302.
Oke, T. R., 1973. City size and the urban heat island. Atmospheric
Environment, 7: 769–779.
Oke, T. R., 1979. Review of Urban Climatology, 1973–1976.
World Meteorological Organization Technical Note No. 169.
Secretariat of the World Meteorological Organization, Geneva.
Oke, T. R., 1997. Surface climate processes. Pages 21–43 in W. G.
Bailey, T. R. Oke & W. R. Rouse (eds.). The Surface Climates
of Canada. McGill-Queens’ University Press, Montreal, Quebec
and Kingston, Ontario.
Saunders, S. C., J. Chen, T. R. Crow & K. D. Brosofske, 1998.
Hierarchical relationships between landscape structure and
temperature in a managed forest landscape. Landscape Ecology,
13: 381–395.
Saunders, S. C., J. Chen, T. D. Drummer & T. R. Crow, 1999.
Modeling temperature gradients across edges over time in a
managed landscape. Forest Ecology and Management, 117:
17–31.
Schaefer, J. A., S. D. Stevens & F. Messier, 1996. Comparative
winter habitat use and associations among herbivores in the
High Arctic. Arctic, 49: 387–391.
Stabler, L. B., C. A. Martin & A. J. Brazel, 2004. Microclimates
in a desert city were related to land use and vegetation index.
Urban Forestry and Urban Greening, 3: 137–147.