1377
Limits to the niche and range margins of alien species
Jake M. Alexander and Peter J. Edwards
J. M. Alexander (jake.alexander@env.ethz.ch) and P. J. Edwards, Inst. of Integrative Biology, ETH Zürich,
Universitätstrasse 16, ETH Zentrum CHN, CH–8092 Zürich, Switzerland.
We discuss the apparent paradox that while introduced populations often adapt rapidly to conditions in the new range,
it is normally assumed that the species’ niche remains unchanged. Focusing on plants, we argue that studies of the niche
dynamics of alien species are useful for understanding the constraints acting on species in their native ranges, and vice versa.
Most hypotheses about species ecological range margins are more consistent with there being a niche shift than niche stasis
in the new range. After reviewing the evidence for niche shifts in alien species, we suggest that the probability of a shift
occurring depends primarily upon the ecological and genetic processes limiting the species in its native range. For example,
a fundamental niche shift might occur if introduced individuals are released from maladaptive gene flow from central
populations, or if genetic diversity is increased by the mixing of individuals from different sources. In addition, other
factors such as species characteristics, introduction history and conditions in the new range may also influence whether
a niche shift occurs. Based on these considerations, we propose conditions under which niche shifts are most likely. Such
understanding is important for predicting and mitigating current and future anthropogenic impacts on species ranges.
he spatial distribution of a species is determined by all
those factors that constrain positive population growth. One
such factor – reflected in the species’ fundamental niche – is
the inability of populations to adapt to abiotic conditions
beyond the species’ boundary. A second – reflected in the
realized niche – is the species’ exclusion from some parts
of its fundamental niche through negative interactions with
other organisms. A third is the inability of the species to
disperse to all potentially suitable areas (the intersection of
regions A, B and M, Fig. 1B) (Pulliam 2000, Soberón 2007,
Svenning et al. 2008). In seeking to understand the relative
importance of these factors, we can learn much from the
many natural experiments produced by the reshuffling of
the Earth’s biota by human activities (Holt and Keitt 2005,
Parmesan et al. 2005). For example, if a species spreads
rapidly when introduced to a new region, it is clear that
it was previously limited by its capacity to disperse; and if
the species occupies the same niche in the introduced and
native ranges (Peterson 2003, huiller et al. 2005), we can
conclude that similar factors limit its distribution in both
regions. hus, alien species provide excellent model systems
for understanding both niche dynamics and the factors
influencing species ranges (Levin 2003b, Holt et al. 2005,
Sax et al. 2007, Sexton et al. 2009).
In this paper, we explore how theories about the range
margins of species, especially plants, can help in understand-
ing niche dynamics of alien species, and vice versa. We begin
by attempting to reconcile evidence for rapid evolution in
introduced populations with the observation that many spe-
cies occupy the same niche in their native and introduced
regions. We then consider whether the various hypotheses
that have been proposed to explain range margins (Table 1,
Fig. 1) (Hoffmann and Blows 1994, Gaston 2003) are more
consistent with niche conservatism (stasis) or shifts. We
review evidence for niche shifts in introduced populations,
and based upon this analysis propose circumstances under
which niche shifts are most likely to occur. Finally, we sug-
gest potentially fruitful avenues for future research.
Reconciling rapid evolution and niche dynamics
of alien species
At a geographic scale, the niche of a species is defined by
the range of environmental conditions within which it can
maintain positive population growth rates (Grinnell 1917,
Hutchinson 1957). However, populations within species
may be genetically adapted to local conditions, implying that
the niches of these populations are different (Mau-Crimmins
et al. 2006). Indeed, individuals of one ecotype recipro-
cally transplanted to the habitat of another may be unable
to reproduce, in which case local adaptation is not merely
a “fine-tuning” of the genotype to its environment but is
crucial for its persistence (Potvin 1986). Alien populations
provide many examples of how rapidly such genetic differen-
tiation can arise. For example, genetically determined clines
in plant traits along latitudinal gradients have been observed
in introduced populations of Echinochloa crus-galli (Roy et al.
2000), Hypericum perforatum (Maron et al. 2004), Impatiens
glandulifera (Kollmann and Bañuelos 2004), Lythrum
salicaria (Montague et al. 2008), Solidago species (Weber
Oikos 119: 1377–1386, 2010
doi: 10.1111/j.1600-0706.2009.17977.x
© 2010 he Authors. Journal compilation © 2010 Oikos
Subject Editor: Eric Seablom. Accepted 20 April 2010