232
breeding sites sets limits on their distributions. Historical
factors also play a role, as pointed out in Chapter 12. Past
and present locations of dispersal barriers, including moun-
tain ranges, rivers, and oceans, have excluded many species
from invading areas where climatically they could survive
and flourish. Geographic distributions of species or popula-
tions can be limited by microhabitat distributions, presence
or absence of competitors or predators, or even the avail-
ability of prey. Microhabitat specialists such as Xantusia
henshawi are restricted to areas with exfoliating rock;
flat lizards in the genera Tropidurus and Platysaurus are
restricted to granitic outcrops in South America and South
Africa, respectively. Anolis lizards on Caribbean islands
have evolved microhabitat specialization (see Chapter 12)
in response to competition with other Anolis and thus are
limited to specific microhabitats within the same habitat.
Dietary specialists occur only in microhabitats containing
the prey that they eat. Horned lizards, Phrynosoma, which
are ant specialists, do not occur in habitats lacking edible
ants, and some, like Phrynosoma cornutum, may move very
little while active because they sit along trails of harvester
ants. No single factor explains the geographic distribution
of any species. At a local level, a multitude of factors influ-
ences spatial distributions of individuals.
LOCAL DISTRIBUTION OF INDIVIDUALS
Distribution of individuals occurs at a number of levels.
In the context of community ecology, species tend to be
associated with specific microhabitat patches (the “place”
resource or niche discussed in Chapter 12). A relatively
easy and informative exercise is to walk through a natu-
ral habitat and list the animal species and the microhabi-
tats where they were first observed. A tabulation of these
data reveals that each species tends to be associated with
specific microhabitats. Because microhabitats interdigitate,
any given habitat can contain a large number of species
that spatially overlap, but occupy different microhabitats.
Selection of microhabitats is enforced by competitive inter-
actions and the risk associated with activity in unfamiliar
places or patches. An individual may no longer be cryptic
or may be unable to escape predators in unfamiliar patches
(see Chapter 11). The distribution of amphibians and rep-
tiles that use wetlands, especially those that live on land but
breed in water (most amphibians) or live in the water but
reproduce on land (turtles, most snakes), can vary consider-
ably on a seasonal basis as individuals move to reproduce.
Within species, individuals often move within an area
that they do not defend from conspecifics, called the home
range. Foraging and social activities occur in this area.
Adjacent home ranges can overlap or they can be com-
pletely exclusive. Part or all of the home range might be
defended, usually against conspecifics but occasionally
against other species. This defended area is the territory,
PART | IV  Behavioral Ecology
which is introduced in this chapter and discussed in the
context of social behavior in Chapter 9. Spacing typically
implies the spatial distribution of individuals within a spe-
cies and, more specifically, within a local population. As
a result, spacing usually focuses on home ranges and ter-
ritories.
Home Ranges
Home ranges of amphibians and reptiles usually are asso-
ciated with one or more resources. The resources include
food, shelter, mates, thermoregulation sites, escape routes,
and a host of other things. Home range size can vary
between sexes, is often associated with body size, and is
often influenced by population density. For species living
in two-dimensional habitats, such as fringe-toed lizards
(Uma) on sand dunes in southern California or plethodon-
tid salamanders in the Great Smoky Mountains, the home
range can easily be measured as the area that encompasses
all the outer points within which an individual occurs. This
technique is called the minimum polygon method of home
range determination and does not take into consideration
the amount of time or the relative frequency with which
an individual might use different parts of the home range.
Nevertheless, it is the most widely used method of calculat-
ing home range and has many advantages. In particular, it
can be calculated easily in the field, the measurements are
fairly accurate if samples are adequate, and it is based on
actual observations of animal occurrences. Moreover, the
amount of overlap in home ranges between individuals in
the population can be easily calculated. The variation in
sizes of amphibian and reptile home ranges is impressive
(Table 8.1). An association between body size and home
range size exists across many species, but some exceptions
exist. For arboreal amphibians and reptiles, measuring
home range is much more difficult, and even defining it is
not easy. The Amazonian lizard Anolis transversalis, for
example, spends much of its life in the canopy of a single
or a few trees. The home range is three dimensional and
thus is a volume rather than an area. Moreover, because
the lizard can only move on the branches and leaves within
the canopy, many gaps or unusable areas exist. Neverthe-
less, conceptually, a three-dimensional home range is no
different from a two-dimensional one—they both represent
regular use of space by individuals.
Home ranges can and often do vary through time or
space; they can change radically following single events.
For some species they may not even exist. Home ranges are
not usually defended, and other individuals may use parts
of them. Overlap in home ranges among individuals can be
considerable. During the nonbreeding season, many terres-
trial amphibians (e.g., Ambystoma maculatum, Plethodon
cinereus, Rhinella marina, Rana temporaria) have small to
moderate-sized home ranges away from water. An individual
Chapter | 8  Spacing, Movements, and Orientation 233

TABLE 8.1  Home Range and Resource Defense in Select Amphibians and Reptiles

Defense

Taxon Area (m2) Female area Male size (mm) Terr S-S Habits
Salamanders

Batrachoseps pacificus 3.6 ? 42 ? ? Terrestrial

Desmognathus fuscus 1.4 ? 45 ? ? Semiaquatic

Salamandra salamandra 10 > 82 ? ? Terrestrial

Anurans

Atelopus varius <20 = 25 + Terrestrial

Lithobates clamitans 65 = 60 + Semiaquatic

Eleutherodactylus marnockii 328 = 20 + Terrestrial

Ranitomeya imitator 4.6–13.5 = 22 + +

Ranitomeya variabilis 31.3–38.9 = 17 + +

Bufo bufo spinosis 570 = ≈150 – – Terrestrial

Pseudepidalea viridis 2,456 = ≈120 – – Terrestrial

Turtles

Terrapene c. triungis 52,000 = 115a – – Terrestrial

Trachemys scripta 397,500 < 200a – ± Aquatic

Crocodylians

Crocodylus niloticus 7990 < 2100b + Aquatic

Lizards

Sceloporus merriami 535 < 45 + Terrestrial

Varanus olivaceus 20,500 < 450 ± + Arboreal

Xantusia riversiana 17 = 65 – + Terrestrial

Snakes

Acrochordus arafurae 15,000 ? 900 – ? Aquatic

Carphophis amoenus 253 ? 215 ? ? Semifossorial

Natrix natrix 99,000 > 700 ? ? Terrestrial

Pantherophis obsoletus 5,600 < 758 – – Arboreal

Coluber constrictor 24,200 = 59 – – Terrestrial

Note: Terr = territorial, S-S = site or resource specific.

aPlastron length
bTotal length

Sources: Salamanders—Bp, Cunningham, 1960; Df, Ashton, 1975; Ss, Joly, 1968. Anurans—Av, Crump, 1986; Lc, Martof, 1953; Em, Jameson, 1955; Ri and
Rv, Brown et al., 2009; Bbs and Bv, Indermaur et al., 2009. Turtles—Tct, Schwartz et al., 1984; Ts, Schubauer et al., 1990. Crocodilians—Cn, Hutton, 1989.
Lizards—Sm, Ruby and Dunham, 1987; Vo, Auffenberg, 1988; Xr, Fellers and Drost, 1991a. Snakes—Aa, Shine and Lambreck, 1985; Ca, Barbour et al., 1969;
Nn, Madsen, 1984; Po and Cc, Carfagno and Weatherhead, 2008.

can have one or more resting and feeding sites (activity cen-
ters) within its home range, but it might use a single site
for a day, a week, or longer before shifting to another site.
An amphibian may not visit all sites each day or even each
week, but the periodic occurrence at sites and the persistent
occupancy of the total area adjacent to these sites delimit
the individual’s home range. For species that reproduce
in ponds or streams, the home range breaks down during
breeding events because adults breed in aquatic sites that are
not within the home range. Terrestrial-breeding amphibians
234
(e.g., dendrobatid, mantellid, and brachycephalid frogs,
Plethodon) generally deposit eggs within their home ranges.
Some spend their entire lives in a single home range.
Crawfish frogs, Lithobates areolatus, have one of
the smallest home ranges of any frog species, primarily
because they are restricted to crayfish burrows when not
breeding. Their breeding season extends for as little as 2–6
weeks during early spring, so about 10.5 months are spent
in their burrows. The home range, based on the concept
that it is the place where an animal spends its time and car-
ries out most of its activities, includes a small feeding plat-
form at the entrance of the burrow and the burrow itself;
thus, calculations reveal that the home range is 0.05 m2.
Crawfish frogs tracked for a number of years showed fidel-
ity to the same burrows year after year. Abandoned cray-
fish burrows provide several advantages for crawfish frogs:
burrows extend to the water table in years of normal rain-
fall and below the frostline in winter so frogs do not risk
dehydration or freezing. The crayfish burrows are mostly
in grasslands, which can burn either naturally or anthro-
pogenically, but frogs are not harmed during burns. When
threatened by predators, frogs, which always orient with
heads toward the burrow entrance, back down the burrow,
tilt their heads to plug the burrow, and inflate their bodies
against the sides of the burrows to discourage predators.
Shape of home ranges varies considerably and is often
related to the microhabitat specificity of a species and the
physical structure of the microhabitat. In the case of craw-
fish frogs discussed above, including the area covered dur-
ing the migration to the breeding site and back to the burrow
would increase the size of the home range and change its
shape radically. Semiaquatic or aquatic species (Desmog-
nathus monticola, Limnonectes macrodon, mud turtles) are
linearly distributed along streams and lakeshores. As a con-
sequence, individuals within these populations tend to have
elongate, narrow home ranges along the stream edge or
lakeshore, or within the watercourse. The watersnake Nerodia
taxispilota is linearly distributed along rivers, streams, and
edges of ponds and lakes in the southeastern United States.
Along part of the Savannah River that divides Georgia and
South Carolina, these watersnakes are most common adja-
cent to backwater areas, such as oxbow lakes and areas
along the outside banks of curves in the river where water
velocity is highest. They commonly are found on steep
riverbanks or perched on logs and roots out of the water.
Although N. taxispilota moved an average of 270 m during
a 2-year period, most individuals moved very little. These
snakes are capable of long movements but often remain in
a relatively small area because of the availability of good
perch sites. Additionally, steep banks and overhanging logs
and roots provide some protection from aquatic and terres-
trial predators.
Alligator snapping turtles (Macrochelys temminckii) live
on the bottom of streams, lakes and ponds. In streams, home
PART | IV  Behavioral Ecology
ranges of these turtles are effectively linear. Adult females
have larger home ranges than males (878 m versus 481 m,
respectively), and although immature individuals have even
larger home ranges (1073 m), their home ranges are not
statistically different from home ranges of adult females
because of high variability. Each turtle uses several core
sites in the streams, with most movement occurring between
core sites. Core sites usually are deep pools, contain struc-
tural elements (e.g., sunken logs, beaver dens, overhanging
trees and shrubs), and are located where a tree canopy exists.
The smooth softshell turtle Apalone muticus is linearly
distributed in rivers throughout the southern and central
United States. These rivers experience drastic and unpre-
dictable fluctuations in water level, and flooding can cause
major changes in the physical structure of the river channel.
As a result, the home ranges of soft-shelled turtles are short-
lived. Home ranges are associated with sandbars that change
periodically due to erosion, but some softshells change the
location of their home ranges without any apparent change
of habitat (Fig. 8.1). Some individuals often move long dis-
tances from their home ranges and then return, presumably
River flow
Home range
shift
River flow
0 0.5 km
FIGURE 8.1  Shifts in the home ranges of two female Apalone muticus
in the Kansas River. Two time periods are represented: closed circles
represent early sightings during summer, and open circles represent
sighting approximately 1–2 months later (time periods are not the same
for each turtle). The upper panel is a subadult that shifted its home range
1363 m upstream. The lower panel is an adult female that shifted its
home range 1534 m upstream. Because the turtles are aquatic and live
in rivers and streams, their distribution is linear. Adapted from Plummer
and Shirer, 1975.
Chapter | 8  Spacing, Movements, and Orientation
assessing the quality of other areas in the river. Other indi-
viduals maintain approximately the same home range year
after year, even when the habitat structure changes.
Home range size often varies among sexes and with
reproductive state. Home ranges of male Sceloporus
jarrovi are twice the size of female home ranges and
increase in average size as the fall breeding season com-
mences. In contrast, female home range size remains the
same (Fig. 8.2). The increase in male home range size
is due partially to a 50% reduction in male density from
summer to fall and an increase in the proportion of the
home range defended by reproductive males. By the peak
of the breeding season, males defend the entire home
range; during this time, the home range and territory are
the same.
In most species, home range size generally decreases
as food availability or density increases (Fig. 8.3). In at
least one instance, the local climate places constraints on
lizard activity that feed back on the amount of space used
1000
males
800
Home range size (m2)
600
400
200
females
0
May June July Aug Sept
FIGURE 8.2  Seasonal variation in home range size for male and female
Sceloporus jarrovi. Breeding occurs in fall, at which time male home
ranges increase in size. Adapted from Ruby, 1978.
1000
Density (females per hectare)
100
10
0 Grapevine Hills
0 1 10 100 1000
Female home-range size (m2)
FIGURE 8.3  As female density increases, home range size decreases for
most amphibians and reptiles, as shown here for territorial and nonterrito-
rial female lizards. Adapted from Stamps, 1983.
235
by individuals. Based on long-term capture–recapture
studies, Arthur Dunham and his collaborators were able to
demonstrate that variation in home range size results from
complex interactions between resource availability, micro-
climate, and physical structure of the habitat. As in Scelo-
porus jarrovi, home ranges of male Sceloporus merriami
are larger than those of females, but geographically close
populations vary greatly in home range size (Fig. 8.4). This
lizard occurs across an elevational gradient in the Chisos
Mountains of west Texas. Populations at higher elevations
experience a much more mesic environment than those at
low elevations. Males and females at the lowest elevations
at Boquillas have much smaller home ranges than individu-
als at higher elevations, even though food availability is
lowest and lizard density is highest at Boquillas. Although
it appears paradoxical that lizard density could be high
with low food availability, an interaction between repro-
ductive, microhabitat, and energetic requirements accounts
for the small home ranges. The environment at Boquillas
is the most extreme (high temperatures, low rainfall) along
the elevational gradient, and as a result, the amount of time
available to each lizard for activity is reduced. Feeding
rates of Boquillas lizards are low, suggesting that energy
is more limited compared with higher-elevation popula-
tions. High temperature also limits activity, and with food
already in short supply, the lizards further limit their activ-
ity, which reduces home range size. Reduced activity cou-
pled with low food availability ultimately feeds back on
allocation of energy for reproduction and results in lower
reproductive output. Sceloporus merriami is a sit-and-wait
predator. In contrast, lizards that actively search for prey
would be expected to have large home ranges. Actively
Oct
300
Area in square meters
200
100
all
all
all
all
all
all
82 83 84 82 83 84 82 83 84 82 83 84 82 83 84 82 83 84
males females males females males females
Boquillas Maple Canyon
10000
FIGURE 8.4  Home range size in Sceloporus merriami varies between
sexes, among years, and among three different sites in the Chisos
Mountains of west Texas. Boquillas, the site with the most extreme (hot
and dry) environment, imposes thermal constraints on lizard activity,
resulting in small home ranges. Adapted from Ruby and Dunham, 1987.