Geographic variation of wing morphology of great fruit-
eating bats (Artibeus lituratus): environmental, genetic
and spatial correlates of phenotypic differences
RICHARD D. STEVENS
1
*, MARY E. JOHNSON
2
and EVE S. MCCULLOCH
3
1
Department of Natural Resources Management and the Museum of Texas Tech University, Lubbock,
TX, 79409, USA
2
The Great Basin Institute, 16750 Mt Rose Highway, Reno, NV, 89511, USA
3
Division of Biology, 103 Ackert Hall, Kansas State University, Manhattan, KS, 66506, USA
Received 15 December 2015; revised 15 January 2016; accepted for publication 15 January 2016
Responses of species to environmental gradients are important and frequent determinants of geographic
phenotypic variation that can drive adaptive processes. Nonetheless, random genetic processes such as drift can
also result in geographic variation in phenotypes, and should be evaluated before implicating selection as the
process driving phenotypic change. We examined geographic variation in wing morphology of Artibeus lituratus
among 18 different sites distributed across interior Atlantic Forest of Paraguay and Argentina. Moreover, we
contrasted geographic variation with environmental, spatial, and genetic variation to test hypotheses related to
selection and drift and their impacts on wing morphology. For A. lituratus distributed across interior Atlantic
Forest, significant differences among sites characterized variation in wing morphology. Geographic variation was
significantly related to climatic variables but not spatial or genetic distances. Such a pattern suggests that
phenotypic variation is related to selection for particular environmental regimes, and not genetic drift. Four
significant dimensions of phenotypic variation were determined. Three dimensions were related to variation
among individuals in terms of wing tips, whereas one was related to overall body size. Wing tips are important
for manoeuverability during flight and differences among sites likely reflect differences in forest and vegetation
structure that must be managed during foraging. Although climate provides good surrogates for environmental
variation, it is probably only an indirect cue of selection regimes that determine variation in wing morphology.
Future studies should evaluate more direct environmental measures such as vegetation structure when
attempting to interpret geographical variation in wing morphology. © 2016 The Linnean Society of London,
Biological Journal of the Linnean Society, 2016, 118, 734–744.
KEYWORDS: geographical variation – morphometrics – selection – wing morphology.
INTRODUCTION
For many taxa of plants and animals, especially
those with large geographic ranges, there is a com-
mon pattern of geographic phenotypic variation
related to environmental variables (e.g. latitude) or
climatic characteristics (e.g. temperature or precipi-
tation) (Masaki, 1967; Barclay, Fullard & Jacobs,
1999; Roff & Mousseau, 2005; Yom-Tov & Geffen,
2006; Jiang et al., 2013; Sun et al., 2013). Environ-
mental correlations such as these can provide valu-
able insights into the microevolutionary processes
responsible for biotic diversification, with natural
selection arguably being the most important. For
example, when environmental regimes cause differ-
ential reproductive success across different pheno-
types, and there is geographical variation among
sties regarding those environmental regimes, selec-
tion can produce phenotype–environment rela-
tionships. Nonetheless, a number of different
microevolutionary processes can also result in pheno-
type–environment relationships (Armbruster & Sch-
waegerle, 1996).
Other genetic processes unrelated to the environ-
ment, such as mutation or genetic drift, can produce
menacingly similar geographical patterns (Arm-
bruster & Schwaegerle, 1996). Genetic drift can cause
a spatial pattern of isolation-by-distance (IBD) *Corresponding author. E-mail: richard.stevens@ttu.edu
734 © 2016 The Linnean Society of London, Biological Journal of the Linnean Society, 2016, 118, 734–744
Biological Journal of the Linnean Society, 2016, 118, 734–744. With 3 figures.
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