The effect of coachwhip presence on body size of North
American racers suggests competition between these
sympatric snakes
D. A. Steen
1,2
*, C. J. W. McClure
2
, L. L. Smith
1
, B. J. Halstead
3
, C. K. Dodd Jr
4
, W. B. Sutton
5
,
J. R. Lee
6
, D. L. Baxley
7
, W. J. Humphries
8
& C. Guyer
2
1 Joseph W. Jones Ecological Research Center, Newton, GA, USA
2 Department of Biological Sciences, Auburn University, Auburn, AL, USA
3 US Geological Survey, Western Ecological Research Center, Dixon, CA, USA
4 Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL, USA
5 Department of Forestry, Wildlife, and Fisheries, University of Tennessee, Knoxville, TN, USA
6 The Nature Conservancy, Camp Shelby, MI, USA
7 Kentucky Department of Fish and Wildlife Resources, Frankfort, KY, USA
8 North Carolina Wildlife Resources Commission, Chapel Hill, NC, USA
Keywords
character displacement; competition;
morphology; reptile; squamate; snake.
Correspondence
*Current address: Virginia Polytechnic
Institute and State University, 100
Cheatham Hall, Blacksburg, VA 24061, USA.
Email: davidasteen@gmail.com
Editor: Mark-Oliver Rödel
Received 9 April 2012; revised 19 August
2012; accepted 21 August 2012
doi:10.1111/j.1469-7998.2012.00965.x
Abstract
When sympatric species compete, character divergence may help maintain coex-
istence. Snakes are often found in species-rich assemblages while exploiting similar
resources; because snake body size is a relatively plastic trait that determines the
range of prey sizes an individual may consume, divergence in body size between
sympatric species may arise as a result of interspecific interactions. The North
American racer, Coluber constrictor, and the larger coachwhip, Coluber flagellum,
have a close taxonomic relationship and similar foraging strategies. Therefore, we
hypothesized that C. constrictor would be smaller where they co-occur with
C. flagellum, as compared to where C. flagellum is absent, throughout the south-
eastern extent of their range. To evaluate this hypothesis, we obtained data on
body size for 2321 adult C. constrictor and 526 adult C. flagellum, along with
habitat data and other potentially important factors influencing body size.
Coluber constrictor was smaller than elsewhere when in peninsular Florida, in pine
forests, on hydric soils and in the presence of the larger and potentially competing
C. flagellum. Body size of C. flagellum did not vary by any measured habitat
variables. The trends we documented are consistent with the hypothesis that
C. constrictor body size is influenced by several variables, including co-occurrence
with C. flagellum.
Introduction
Given a limited resource, interspecific competition may occur
between closely related species. Morphological divergence is
one potential outcome of this competition (e.g. Grant &
Grant, 2006), a process referred to as character displacement
(Brown & Wilson, 1956). By influencing resource use, charac-
ter displacement may facilitate coexistence over evolutionary
time (Bowers & Brown, 1982). However, it is difficult to dif-
ferentiate the past effects of competition from ongoing
processes. For example, studies of resource partitioning or
morphological variation may demonstrate that sympatric
species use different resources or exhibit differences in mor-
phology. However, it is often unknown whether these differ-
ences result from past evolutionary divergence to reduce
competition, random processes or from plastic adaptations
resulting from current interactions (e.g. Connell, 1980).
Studies that quantify both the diet and morphology of sym-
patric species may provide powerful evidence for character
displacement (e.g. Huey, 1974). However, it is logistically
challenging to measure the availability and consumption of
prey on a large scale. As a result, morphological variation may
be used as a proxy for resource use. If the morphological traits
related to catching and consuming prey differ in relation to the
presence or abundance of potential competitors, this is con-
sistent with the hypothesis that interspecific interactions influ-
ence resource use (e.g. Simberloff et al., 2000; Davies et al.,
2007).
Body sizes may differ among members of species-rich snake
assemblages (Guyer & Donnelly, 1990). Although body size
may not always be an appropriate means to infer differences in
resource use among wildlife species (Wilson, 1975), snakes
swallow prey whole; thus, body size has an important influ-
ence on the maximum size of potential prey (King, 2002). In
Journal of Zoology
Journal of Zoology. Print ISSN 0952-8369
86 Journal of Zoology 289 (2013) 86–93 © 2012 The Authors. Journal of Zoology © 2012 The Zoological Society of London