Theoretical Population Biology 98 (2014) 1–10
Contents lists available at ScienceDirect
Theoretical Population Biology
journal homepage: www.elsevier.com/locate/tpb
The genetic signature of rapid range expansions: How dispersal,
growth and invasion speed impact heterozygosity and allele surfing
Devin W. Goodsman
a,∗
, Barry Cooke
c
, David W. Coltman
a
, Mark A. Lewis
a,b
a
Department of Biological Sciences, CW 405, Biological Sciences Bldg., University of Alberta, Edmonton, Alberta, Canada T6G 2E9
b
Mathematical and Statistical Sciences, 632 CAB, University of Alberta, Edmonton, Alberta, Canada T6G 2G1
c
Canadian Forest Service, Northern Forestry Centre, 5320 122 Street Northwest, Edmonton, Alberta, Canada T6H 3S5
article info
Article history:
Received 6 December 2013
Available online 6 September 2014
Keywords:
Dispersal
Genetic diversity
Heterozygosity
Invasion
Range expansion
abstract
As researchers collect spatiotemporal population and genetic data in tandem, models that connect
demography and dispersal to genetics are increasingly relevant. The dominant spatiotemporal model
of invasion genetics is the stepping-stone model which represents a gradual range expansion in which
individuals jump to uncolonized locations one step at a time. However, many range expansions occur
quickly as individuals disperse far from currently colonized regions. For these types of expansion,
stepping-stone models are inappropriate. To more accurately reflect wider dispersal in many organisms,
we created kernel-based models of invasion genetics based on integrodifference equations. Classic theory
relating to integrodifference equations suggests that the speed of range expansions is a function of
population growth and dispersal. In our simulations, populations that expanded at the same speed but
with spread rates driven by dispersal retained more heterozygosity along axes of expansion than range
expansions with rates of spread that were driven primarily by population growth. To investigate surfing
we introduced mutant alleles in wave fronts of simulated range expansions. In our models based on
random mating, surfing alleles remained at relatively low frequencies and surfed less often compared
to previous results based on stepping-stone simulations with asexual reproduction.
© 2014 Elsevier Inc. All rights reserved.
1. Introduction
Range expansions explain the wide spatial distribution of many
dominant species. Unfortunately however, researchers often have
only a snapshot of the extent of a recently expanded range rather
than a complete spatiotemporal dataset. Genetic data have been
used to elucidate processes underlying range expansions based on
these snapshots, from our own planetary conquest (Ramachandran
et al., 2005) to the post-glacial expansion of grasshoppers (Hewitt,
1999). Such insights, based on snapshots of genetic patterns on the
landscape, are predicated on models that connect the dynamics,
movement and genetics of populations. Thus, spatiotemporal
genetic models are increasingly relevant as we accumulate large
genetic databases. In this research we introduce integrodifference
models as an alternative modeling framework in invasion genetics
∗
Corresponding author.
E-mail addresses: goodsman@ualberta.ca (D.W. Goodsman),
Barry.Cooke@NRCan-RNCan.gc.ca (B. Cooke), dcoltman@ualberta.ca
(D.W. Coltman), mark.lewis@ualberta.ca (M.A. Lewis).
with a sound mathematical and ecological basis. Integrodifference
equations are discrete-time, continuous-space models that apply
to range expansions in which populations have synchronized
growth and dispersal stages (Neubert et al., 1995). Thus, they are
useful for many herbaceous, invertebrate, and vertebrate species
prone to invasion (Kot et al., 1996).
Currently, invasion models with analytical solutions for the
patterns of genetic diversity that they produce are limited to
the island model (Wright, 1951; Buerger and Akerman, 2011)
and the stepping-stone model (Kimura and Weiss, 1964; Thibault
et al., 2009; DeGiorgio et al., 2011; Slatkin and Excoffier, 2012). In
the island model, subpopulations receive migrants at a constant
rate from a single unchanging source population, whereas in the
stepping-stone model, unoccupied demes are colonized sequen-
tially one after another, and only receive migrants from adjacent
subpopulations (Kimura and Weiss, 1964; DeGiorgio et al., 2009,
2011). Many dispersing organisms however, can move to locations
beyond adjacent unoccupied areas (Levin et al., 2003) and dispersal
is an important determinant of the speed of population expansion
in space (Kot et al., 1996). For these reasons, neither the island nor
the stepping-model in their original form is realistic in terms of
population processes or dispersal (Le Corre and Kremer, 1998).
http://dx.doi.org/10.1016/j.tpb.2014.08.005
0040-5809/© 2014 Elsevier Inc. All rights reserved.