2007 SOUTHEASTERN NATURALIST 6(4):715–726
Use of Altered Habitats by the Endemic
Sand Skink (Plestiodon reynoldsi Stejneger)
David A. Pike1,2,*, Kelley S. Peterman1, and Jay H. Exum1
Abstract - Endemic species with limited geographic ranges are particularly sensitive
to habitat loss and degradation. In order to conserve such species, we tend to
focus on optimal habitats (e.g., for land acquisition), but human-altered habitats
may also be of conservation value if species can persist in these environments.
We evaluated the occurrence of a lizard endemic to scrub and sandhill habitats in
Florida, the threatened Plestiodon reynoldsi (Sand Skink), to determine whether
it occurs in human-altered vegetation types. From 2003–2004, we quantitatively
sampled 46.5 ha (composed of 7 vegetation and 10 soil types) for the distinctive
trails this fossorial lizard leaves in sandy areas. The Sand Skink was present in
equal relative densities across all vegetation types, including disturbed areas such
as pastures and abandoned citrus groves. Further, we detected Sand Skinks on all
well-drained sandy soils we sampled. Thus, Sand Skinks can persist in humanaltered
habitats, at least when underlying soils are suitable for their presence
and have not been modified. In general, more Sand Skinks were found in larger
habitat patches, regardless of whether patches were characterized by vegetation or
soil. Our results suggest that anthropogenically altered habitats have conservation
value for Sand Skinks (and possibly other ecologically similar species) and that
future studies should focus on the effects of restoring these habitats on resident
populations of this species.
Many species are declining due to habitat loss and fragmentation
(Schneider 2001), and current research and protection efforts often focus
on remaining high-quality habitats (e.g., McCoy et al. 1999). However,
studying species in optimal habitats alone may not provide sufficient
information to determine which specific environmental characteristics affect
species presence. Historically, natural disturbance events maintained
vegetative structure in many habitats. However, increasing habitat fragmentation
has suppressed disturbances, resulting in overgrown habitats
that may be unsuitable for many species (e.g., Pringle et al. 2003, Webb
et al. 2005). Understanding which habitat changes are occurring and
how species react to these changes provides important information about
ways to protect biodiversity (e.g., Jäggi and Bauer 1999, Webb and Shine
1998, Webb et al. 2005). For example, past research proposed that habitat-
specific endemic lizards (e.g., Plestiodon reynoldsi Stejneger [Sand
1Environmental Services Group, Glatting Jackson Kercher Anglin Inc., 120 North
Orange Avenue, Orlando, fl32801. 2Current address - School of Biological Sciences
A08, University of Sydney, NSW 2006, Australia. *Corresponding author - dpike@
716 Southeastern Naturalist Vol. 6, No. 4
Skink], Scincidae) might not require habitats containing high-quality
natural vegetation to persist (Campbell and Christman 1982). Instead,
the abundance and distribution of this fossorial, sand-swimming lizard
may largely be determined by soil characteristics because locomotion and
movement of this species is impeded by compacted soils (Andrews 1994,
If human-altered habitats harbor important populations of habitat-specific
endemic species (e.g., Panzer et al. 1995), protection and restoration of these
areas may aid in species persistence (Ries et al. 2001). Conversely, if such
habitats are ignored, naturally occurring and important populations may be
destroyed. Without a thorough understanding of all factors restricting a species’
distribution, important populations may go unnoticed. For example,
habitats are often characterized by vegetation type, even though other factors
such as elevation, rainfall, soil type, or thermal environments may be largely
responsible for species presence (Campbell and Christman 1982, Webb and
Shine 1998, Webb et al. 2005). In fact, a combination of habitat characteristics
(aside from simple vegetation types) may be what shapes species
distributions, especially on small spatial scales and in severely fragmented
habitats (Webb and Shine 1998).
We studied the habitat associations of Plestiodon (formerly Neoseps
[Schmitz et al. 2004]) reynoldsi, a lizard restricted to central Florida that
is locally abundant in scrub and sandhill habitats, which are habitats frequently
subject to some degree of human disturbance. Despite this, little
is understood about local-scale distribution patterns. To help understand
the local-scale distribution of the Sand Skink, we surveyed altered habitat
types and focused on the influence of vegetation type on relative density
of Sand Skinks. As ectotherms, lizards rely on many aspects of the habitat
for thermoregulation, which can affect foraging efficiency, digestion,
growth, and egg development. For example, structural aspects of the
vegetation heavily influence light penetration, resulting in direct effects
on thermal profiles (Pringle et al. 2003, Webb et al. 2005). Altered habitats
may contain the habitat characteristics (e.g., structure) that provide
adequate thermal regimes for Sand Skinks. Additionally, because Sand
Skinks are fossorial, they may require soil types that allow subsurface
movements, particularly loose, dry soils (Campbell and Christman 1982).
We also studied the relationship between soil type and Sand Skink presence,
irrespective of the vegetation type present.
Materials and Methods
Sand Skinks are small fossorial lizards (≈57 mm snout–vent length;
Ashton 2005) that “swim” just beneath the surface of the soil. This swimming
motion leaves distinct sinusoidal trails on the surface of the sand.
2007 D.A. Pike, K.S. Peterman, and J.H. Exum 717
Because Sand Skinks spend much of their time belowground, these trails
are used as a surrogate for animal presence (Andrews 1994, Sutton et al.
1999). Several other declining or habitat-restricted reptile species also
occur sympatrically (Branch et al. 2003, Campbell and Christman 1982,
Means and Simberloff 1987), including P. [Eumeces] egregius lividus
Mount (Blue-tailed Mole Skink), another elusive species of semi-fossorial
skink that is of conservation concern (United States Fish and Wildlife
Service 1993). The conservation of Sand Skinks and other sympatric,
imperiled species is critical to maintaining an important segment of the
highly diverse reptile fauna of the southeastern United States (Gibbons
and Stangel 1999).
Habitat associations of Sand Skinks
Several broad-scale habitat characteristics have been emphasized as
supporting Sand Skink populations. The overall geographic distribution is
largely defined by relatively high elevations (>30 m above sea level) occurring
along relict sand dunes remaining from periods of elevated sea levels
during the Pleistocene (Telford 1959). Associated habitats include rosemary
scrub, overgrown scrub, high pine, and longleaf pine-turkey oak sandhills
(Campbell and Christman 1982; Cooper 1953; Telford 1959, 1962). Ecotonal
areas are also important, particularly those between rosemary scrub
and palmetto-pine flatwoods habitats, and between longleaf pine and sand
pine scrub habitats (Cooper 1953; McCoy et al. 1999; Telford 1959, 1962).
Within these habitats, Sand Skinks are often associated with open, sandy
patches that allow subsurface movement (e.g., free of roots and grasses that
may impede fossorial locomotion), but are most often found underneath
cover (Cooper 1953, Telford 1959). Campbell and Christman (1982) also
hypothesized that reptiles in scrub and sandhill habitats are distributed in
relation to physical characteristics of the habitat itself, rather than to specific
vegetation types. Therefore, areas that contain well-drained, sandy soils
and open patches free of root mats may be suitable for skinks (Campbell
and Christman 1982), even though surface vegetation types are disturbed or
converted to agricultural production, particularly citrus.
We studied skinks on a 158-ha tract of private land in Lake County,
FL. The site is composed of several upland habitat types growing on
well-drained, sandy soils (Table 1, Fig. 1). Historic cattle ranching and
the production of citrus fruit have altered the natural vegetation, which is
now mostly characterized as early stage successional systems (Table 1).
Little natural vegetation of the kind normally associated with natural
Sand Skink habitat remains due to these historical habitat alterations, and
the habitat surrounding our site is extensively modified, and much of it is
currently being developed. Thus, little (if any) habitat typically thought
of as appropriate for Sand Skink exists in the area.
718 Southeastern Naturalist Vol. 6, No. 4
We established 9 sampling plots of varying sizes (mean = 5.18 ± 1.84
ha; range = 0.43–13.58 ha) in vegetation and soil types representative of our
site (totaling 46.5 ha; Fig. 1). Plots were based on representative vegetation
types and avoided wetland areas, resulting in irregular sizes and shapes
Table 1. A description of the vegetation types at our study site in Florida that are considered
uplands, and thus potentially contain Plestiodon reynoldsi (Sand Skinks). Vegetation types were
classified using the Florida Land Use, Cover, and Forms Classification System (flDOT1999),
and then categorized below to distinguish the major characteristics of each. Open (e.g., sandy)
patches are categorized by size (absent, small, large), while all other variables are categorized
by relative abundance (absent, present, dominant). All vegetation types were sampled according
to their availability at our study site (see text).
Vegetation type Canopy Shrub layer Groundcover Open patches
Improved pasture Absent Absent Dominant Small
Unimproved pasture Absent Absent Dominant Small
Palmetto prairie Absent Dominant Dominant Absent/small
Shrub and brush Absent Dominant Dominant Absent
Pine flatwoods Present Present Present Absent
Xeric scrub Absent Dominant Present Large
Sand live oak Dominant Present Present Large
Figure 1. Outline of our study site in Florida showing the location and shape of our 9
sampling plots and their associated vegetation (A) or soil (B) types. The large white
(open) areas represent wetlands, which are not suitable as habitat for Plestiodon reynoldsi
(Sand Skink). Note that sampling plots may each contain more than one patch of each
vegetation or soil type (e.g., vegetation type was independent of the soil underneath).
2007 D.A. Pike, K.S. Peterman, and J.H. Exum 719
(Fig. 1). Each plot contained plywood coverboards (60 x 60 x 1.25 cm)
spread evenly in rows at a density of 64 boards per ha (i.e., 28 m apart). We
cleared all groundcover underneath each board so that only a sandy substrate
remained, allowing greater visibility of Sand Skink trails. Since vegetation
can mask trails, this was especially important in areas with dense herbaceous
vegetation where there are few areas of open sand.
We sampled coverboards once weekly for four consecutive weeks
during the mating season (March–May), when activity is thought to be
greatest (Andrews 1994, Sutton et al. 1999, but see Ashton and Telford
2006 for high levels of activity during other months). When sampling,
we looked underneath each coverboard to determine whether Sand Skink
trails were present. All fieldworkers were trained in identifying trails, and
in the rare case that a question arose about a particular trail, two or more
researchers reached consensus as to whether the trail was made by a Sand
Skink. At each trail, we recorded the exact location of the coverboard by
using a Global Positioning System (Trimble XL GPS; accurate to less than 1 m).
We assumed only one skink was responsible for trails underneath each
coverboard, resulting in conservative estimates of relative density (Sutton
et al. 1999; D.A. Pike, unpubl. data). Additionally, each coverboard was
counted only once when calculating measures of relative density to help
avoid pseudosampling. Relative density is defined herein as the proportion
of unique coverboards with Sand Skink trails over the 4-week sample
period. Although we sampled different portions of the site in successive
years (2004 or 2005), we were interested in presence and relative density
only, so we combined years for analysis.
We spatially mapped all sampling plots and skink locations in
ArcView 3.2 using GPS locations. We layered these points onto grounddetermined
vegetation types (see Table 1 for vegetation types consistent
with the Florida Land Use, Cover, and Forms Classification System; FL
DOT1999) and soil types mapped by the Natural Resources Conservation
Service (NRCS; see Fig. 2 for a list of all soil types sampled). Each of our
9 sampling plots contained multiple vegetation and soil types (i.e., multiple
patches). For statistical purposes, we calculated presence or relative
density separately for each vegetation or soil patch within each sampling
plot, and treated patches of the same habitat type as replicates (Fig. 1).
Since vegetation types were not necessarily related to the underlying soil,
numbers of patches and their sizes varied between these two variables.
To test the hypothesis that relative density differed among habitat types,
taking into account the size of each habitat patch, we used analysis of
covariance (ANCOVA). Habitat type (for vegetation and soil) was the
factor, with the size of each habitat patch as the covariate and relative
density of Sand Skinks as the dependent variable. We also determined the
720 Southeastern Naturalist Vol. 6, No. 4
proportion of each habitat type sampled and the availability of each using
ArcView, and tested whether we surveyed habitats according to their
availability at our site using chi-squre goodness of fit tests. We also used
chi-squre tests to determine whether the presence of Sand Skinks differed
among habitat types (for vegetation and soil). Alpha was set at 0.05 for
all tests, and assumptions of statistical tests were checked and met unless
Overall survey results
We checked 2975 coverboards once weekly for 4 consecutive weeks,
resulting in a total of 11,900 possible encounters with Sand Skink trails.
During this time we encountered trails underneath 726 individual coverboards
(24.5% of possible boards; Table 2). We sampled all vegetation
types present on site (n = 7 types) that were designated as uplands using
the Florida Land Use, Forms, and Classification System (Table 1; FL
DOT 1999). Additionally, we sampled all soil types present at our study
site (n = 10; Table 2, Fig. 1).
We found Sand Skinks in 7 out of the 8 sampling plots (87.5%), and
presence did not depend upon the vegetation type contained within the sampling
plots (χ2 = 8.15, df = 10, P = 0.61; Table 2). Sand Skinks were present
in each vegetation type sampled, and those that were not sampled completely
were sampled in equal proportion to their availability (χ2 = 6.75, df
Figure 2. Soil
for Sand Skinks
in Lake County,
soil refers to a
taking the expected
of plots occupied
by skinks (as calculated by a chi-square goodness-of-fit test) and subtracting
them from the observed number of sampling plots occupied (after Kuhnz et al. 2005).
In soil types with an [observed - expected] value of 0, skinks were found in similar
proportions to the calculated expected values. No soil types contained fewer skinks
than expected. Note that these values are relative to other soils at our site, and may
not represent absolute soil suitability.
2007 D.A. Pike, K.S. Peterman, and J.H. Exum 721
= 3, P = 0.08). Thus, our sampling design controlled for area by sampling
representative areas of each available vegetation type. Additionally, Sand
Skink relative density did not differ by vegetation type (ANCOVA: F10,30 =
0.74, P = 0.68), but depended upon the amount of area sampled (F1,30 = 5.26,
P = 0.03; habitat * area interaction: F9,23 = 0.11, P = 0.99). The larger the
area sampled, the higher the relative density, regardless of vegetation type
(Fig. 3). A mean relative density of 48.8 ± 9.2 skinks/ha was found within
each vegetation type.
The proportion of habitat occupied was dependent upon soil type (χ2 =
28.1, df = 10, P = 0.002); however, all soil types contained Sand Skinks
(Fig. 2) except one plot containing hydric soils (a mixture of Pompano,
Felda, and Oklawaha depressional soils, and Myakka sands; Table 2).
All of the occupied soil types were xeric (dry). Most soil types contained
more-occupied habitat than would be expected based on values calculated
from chi-square tests (Fig. 2). Because a chi-squre test compares the actual
proportion of occupied habitat to that expected based on the amount
and type surveyed, this indicates that each soil type containing Sand
Skinks (n = 8; Table 2, Fig. 2) contained greater proportions of occupied
habitat than would be expected by random chance. Sand Skink relative
density did not differ by soil type (ANCOVA: F8,13 = 1.00, P = 0.48), but
Table 2. Results of our surveys for Plestiodon reynoldsi (Sand Skink) in Florida showing the
amount of area sampled in each vegetation and soil type, the number of coverboards sampled in
each habitat, and the number and proportion of coverboards under which we found P. reynoldsi
or signs of their presence. See text for details regarding the size and density of coverboards and
sizes of the habitat patches that were sampled.
Coverboards Coverboards showing
Habitat Ha sampled (%) sampled P. reynoldsi (%)
Improved pasture 21.65 (73.1) 1386 523 (37.7)
Shrub and brush 0.94 (8.7) 60 4 (6.6)
Palmetto prairie 2.13 (40.0) 137 19 (13.9)
Pine flatwoods 5.25 (54.8) 336 10 (3.0)
Sand live oak 9.07 (99.6) 580 117 (20.2)
Unimproved pasture 2.78 (27.9) 298 40 (22.4)
Xeric scrub 4.66 (99.8) 178 18 (6.0)
Anclote 2.82 (5.8) 180 55 (30.5)
Candler 12.26 (79.9) 784 263 (33.5)
Immokalee 7.14 (22.0) 457 16 (3.5)
Lake 2.50 (99.1) 160 99 (61.9)
Myakka 0.70 (17.1) 49 0 (0.0)
Placid and Myakka 0.16 (3.1) 10 3 (29.7)
Pompano 4.16 (88.3) 266 84 (31.6)
Pompano, Felda, and Oklawaha 0.14 (2.3) 9 0 (0.0)
Tavares 13.87 (48.2) 888 188 (21.2)
722 Southeastern Naturalist Vol. 6, No. 4
depended upon the amount of area sampled (F1,13 = 19.54, P < 0.001; soil
* area interaction: F3,11 = 0.54, P = 0.66). The larger the area sampled, the
higher the relative abundance of Sand Skinks, regardless of the specific
type of xeric soil (Fig. 3).
Our results support previous studies demonstrating that habitatspecific
endemic species can persist in human-altered habitats (Franken
and Hik 2004, Scott et al. 2006). Specifically, the Sand Skink was found
in all of the human-altered vegetation types we surveyed, which are representative
of disturbed lands within central Florida. This result provides
evidence that over small spatial scales the distribution of Sand Skinks
may not be limited solely to specific vegetation types or ecotonal boundaries,
and may be more general than previously thought. This confirms
and strengthens the hypothesis of Campbell and Christman (1982), who
suspected that soil type may be more important than vegetation in determining
Sand Skink distribution, but limited their investigations to natural
habitats in different stages of disturbance-induced succession. Although
previous studies found high abundances in natural habitats (e.g., McCoy
et al. 1999), relative densities did not differ among any of our altered vegetation
or soil types, suggesting that there is no difference in perceived
quality among these habitats.
The behavioral ecology of fossorial reptiles may partially explain our
observed patterns. For example, the space use of fossorial reptiles may
Figure 3. Relationship between the size of the area sampled and Sand Skink relative
density for habitats characterized by vegetation (triangles, solid line) and soil types
(squares, dotted line). See text for statistical results.
2007 D.A. Pike, K.S. Peterman, and J.H. Exum 723
be extremely limited; their lives are spent underneath the soil surface
in relatively small habitat patches (Andrews 1994, Branch et al. 2003,
Campbell and Christman 1982). Therefore, soil composition may be the
most important factor limiting fossorial lizard distributions, rather than
the composition of the plant communities growing on the sandy soils
(Campbell and Christman 1982, Lee 1969). This ecological characteristic
may allow persistence of Sand Skink populations in human-altered vegetation
types well after alteration occurs, especially when underlain by xeric
soils. Since Sand Skinks were found in not only all of the xeric soil types
sampled, but in all vegetation types as well, we cannot elucidate whether
one of these factors is more important in shaping local-scale distribution.
Environmental impact surveys searching for protected species prior
to extensive habitat manipulation (e.g., land clearing, development) must
take into consideration the fact that endemic species may also exist within
severely modified habitat types, even if they are not considered optimal.
Environmental assessments have the obligation to fairly determine the
presence and distribution of focal species, usually species protected by
law. However, when protected species have the reputation of being habitat-
specific (often determined by vegetation type), researchers may focus
on intact natural habitats, and ignore adjacent areas. Our data revealed
that this approach to environmental impact surveys would underestimate
the presence of Sand Skinks, and potentially other sympatric endemics,
within the landscape.
Rapid destruction of habitat is leaving few areas of intact natural habitat
available for conservation purposes (Myers and White 1987, Shine et
al. 1998). However, the finding that an endemic, and supposedly habitatspecific,
lizard species persists in human-altered landscapes suggests that
habitat protection and restoration of areas including cattle ranchland and
citrus groves is potentially of considerable conservation value. This may
be critical in extreme cases where limited tracts of intact natural lands are
available for purchase by conservation organizations or public entities for
conservation. In fact, our recent efforts to find natural habitat appropriate
for conserving populations of Sand Skink indicate that this is the case in
Florida, and thus, habitat restoration in altered lands containing populations,
if successful, may be the most pressing conservation measure for
this and ecologically similar species. In addition to protected species,
many other declining or even common species composing portions of
communities are present in altered habitats (e.g., Ballinger and Watts
1995, Fitch 2006). Therefore, protection and restoration efforts may also
benefit the composition of rare ecological communities, in addition to
single focal species.
With a thorough understanding of rare species distributions, habitat
associations at multiple scales, and ecological tolerances, the probability
724 Southeastern Naturalist Vol. 6, No. 4
that conservation practices will succeed increases (Pringle et al. 2003,
Webb et al. 2005). Additionally, the more habitat types (especially
human-altered habitats) that contain rare species, the higher the chances
are that habitat can be obtained, protected, and restored in such a way as
to ensure persistence. We emphasize that natural habitats should be the
focus of conservation efforts whenever possible; however, human-altered
lands should not be written off as unsuitable for rare species before
thorough research indicates otherwise.
Access to land and funding were provided by Ms. Virginia Dio and Ms. Liz
Tapado. K. Nelson, R. Mejeur, and K. Muddle were responsible for much of the
fieldwork; H.R. Mushinsky provided guidance and insight during our research.
We are especially grateful to K.G. Ashton and E.A. Roznik for greatly improving
the clarity and quality of a previous draft of this manuscript. Monkey Bar Orlando
provided a work environment conducive to the generation of novel ideas, along
with an excellent writing atmosphere.
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