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REVIEW AND
SYNTHESIS Niche conservatism as an emerging principle
in ecology and conservation biology
Abstract
John J. Wiens,1* David D. The diversity of life is ultimately generated by evolution, and much attention has focused
Ackerly,2 Andrew P. Allen,3 on the rapid evolution of ecological traits. Yet, the tendency for many ecological traits to
Brian L. Anacker,4 Lauren B. instead remain similar over time [niche conservatism (NC)] has many consequences for
Buckley,5 Howard V. Cornell,4 the fundamental patterns and processes studied in ecology and conservation biology.
Ellen I. Damschen,6 T. Jonathan Here, we describe the mounting evidence for the importance of NC to major topics in
Davies,7,8 John-Arvid Grytnes,9
ecology (e.g. species richness, ecosystem function) and conservation (e.g. climate change,
Susan P. Harrison,4 Bradford A.
invasive species). We also review other areas where it may be important but has generally
Hawkins,10 Robert D. Holt,11
Christy M. McCain12 and
been overlooked, in both ecology (e.g. food webs, disease ecology, mutualistic
Patrick R. Stephens13 interactions) and conservation (e.g. habitat modification). We summarize methods for
testing for NC, and suggest that a commonly used and advocated method (involving a
test for phylogenetic signal) is potentially problematic, and describe alternative
approaches. We suggest that considering NC: (1) focuses attention on the within-
species processes that cause traits to be conserved over time, (2) emphasizes connections
between questions and research areas that are not obviously related (e.g. invasives, global
warming, tropical richness), and (3) suggests new areas for research (e.g. why are some
clades largely nocturnal? why do related species share diseases?).
Keywords
Climate change, community assembly, conservation, disease ecology, food webs, habitat
destruction, invasive species, niche conservatism, phylogeny, species richness.
cases where observed evolutionary change is slow or absent, Below we provide our working definitions of the niche
and species seem to retain similar traits over long periods of and NC. We then describe potential tests of NC, the
time. These instances of slow evolution may have many increasing evidence for NC in many areas of ecology and
fundamental consequences for ecology. Furthermore, the conservation, new areas where NC has not been widely
current biodiversity crisis may reflect the consequences of applied, and areas for future research.
slow evolution, particularly with regards to niches of
species.
What is the niche?
The niche (sensu Hutchinson; see Holt 2009) describes the
set of abiotic and biotic conditions where a species is able to The niche has been defined in terms of the distribution of
persist. Outside the niche, individuals are not expected species, the functions they perform and the resources they
to leave descendants, nor populations to persist, nor clades consume (reviewed in Soberon 2007). These definitions can
to endure and proliferate. This abstract trait, the niche, is a be separated into two classes (Soberon 2007). The Grin-
function of many organismal traits (e.g. body size, tolerance nellian niche or non-interactive niche, is important for
to pH, feeding adaptations). Sometimes, these niche-related understanding the large-scale geographic distribution of
traits evolve rapidly (e.g. Schluter 2000). But quite often, species. The Eltonian niche focuses on biotic interactions
these traits seem to change very slowly (e.g. Peterson et al. and resourceconsumer dynamics, often at more local
1999; Wiens & Graham 2005). scales. NC is relevant to both facets of the Hutchinsonian
The tendency of species and clades to retain their niches niche.
and related ecological traits over time is called niche Following Hutchinson (1957), we consider the niche as
conservatism (NC hereafter). This term was first coined by describing the set of biotic and abiotic conditions where a
Harvey & Pagel (1991; although the concept has many species can persist (Holt 2009). This includes both the
antecedents) and was subsequently popularized by Holt & distribution of a species and its interactions with other
Gaines (1992), Peterson et al. (1999), Prinzing et al. (2001) species. Much literature on NC has focused on climate and
and many others. NC is relevant to a variety of traits, from geographic distribution. However, the niche is also relevant
those determining the abiotic niche axes of a species (e.g. to the fine-scale distribution of species (e.g. microhabitats),
tolerance to cold and drought) to those determining the resources they consume, and biotic interactions. For
resource utilization (e.g. microhabitat, diet) and other many parasitic or symbiotic organisms, their hosts may
aspects of interspecific interactions. Furthermore, NC can determine the abiotic environment they experience (e.g.
occur at a variety of different spatial, temporal and temperature, moisture, pH) and the Grinnellian and
phylogenetic scales. The fact that NC can occur at different Eltonian concepts converge. Similarly, Grinnellian niche
scales is part of what makes it relevant to so many different dimensions (e.g. temperature) may influence key aspects of
topics, from intraspecific patterns and conservation biology the Eltonian niche (e.g. food availability, competition),
over decades (e.g. responses of species to anthropogenic which in turn influence large-scale distributions of species.
climate change and spread of invasive species), to
speciation and community ecology, to large-scale patterns
What is niche conservatism?
of biogeography and species richness generated over tens
or hundreds of millions of years (e.g. Wiens & Graham We define NC as the retention of niche-related ecological
2005). traits over time. This definition is intentionally broad,
Here, we review the importance of NC to ecology and because NC can involve many traits and time scales, and can
conservation. Ours is not the first review of NC. Wiens & be detected using many tests (see below). NC is more
Graham (2005) discussed the relevance of NC to many inclusive than phylogenetic niche conservatism (PNC),
ecological and evolutionary questions, but focused on a defined here as retention of ecological traits over time
single (general) trait, the tolerance of species to factors that among related species. Importantly, NC can occur within
limit their geographic ranges. Similarly, Pearman et al. (2008) species (e.g. constraining responses to global warming and
reviewed some applications of phylogenies and environ- spread of invasives), a level at which phylogeny may be
mental niche modelling to the study of NC. Here, we irrelevant. Thus, defining NC based solely on a phylogenetic
address NC more generally, and address several topics not test may be inappropriate. Furthermore, there is a distinc-
previously considered in an NC framework. Losos (2008) tion between the definition of a concept and the specific test
reviewed tests for NC and whether NC is prevalent based used to measure its effects empirically (e.g. competition is
on his preferred method [i.e. a test for phylogenetic signal not defined based on a particular test).
using a Brownian motion (BM) model of trait evolution on a Niche conservatism in a species or clade may be most
tree]. We argue that this method may sometimes be apparent when contrasted with an alternative set of
misleading and suggest a variety of alternative methods. ecological conditions or resources that they fail to occupy
or utilize, and which are instead occupied by other species climatic NC leading to local extinction as climate changes).
or clades (Fig. 1). Depending upon the organism and Many other terms share this property of being a pattern at
question, these alternative conditions may include temperate one level (and requiring a causal explanation) and a process
environments for tropical clades, different host or prey at another (and providing a causal explanation for patterns
types, high vs. low pH soils, or the before and after climate at that level). For example, speciation is a pattern of one
of a locality that has become 3 C warmer. species splitting into two (a pattern explained by various
Some authors (e.g. Losos 2008) have expressed concern evolutionary processes), and a process that creates more
over whether NC is a pattern or process. We argue that at species (and thus part of a causal explanation for many
one level, NC is a pattern of ecological similarity over time. species richness patterns). At the population-level, several
However, NC can also be viewed as a process, if this pattern different processes may give rise to NC, and distinguishing
of ecological similarity helps create other patterns (e.g. these processes is a major area for future research
(see below). Importantly, the pattern of NC is created by
these finer-scale processes, but NC can also be viewed as a
process that helps create other patterns (e.g. diversity
gradients).
Resource or environment 1 Resource or environment 2
phylogeny (Revell et al. 2008), whereas the pattern expected identified and tested using SDMs. For example, a hypothesis
from NC is no change. Therefore, a pattern of no signal or of climatic NC predicts that invasive species will spread
weak signal could either mean that the trait varies randomly primarily in regions that are climatically similar to their
across the phylogeny, or shows stasis. As this result could native range. SDMs can be used to test the spatial limits of
mean either no NC or strong NC, this test is potentially this predicted range, and whether species diverge from these
problematic. expectations (e.g. Peterson 2003), offering both an applica-
One alternative approach is to compare the relative fit of tion and test of NC. SDMs and related approaches can also
different evolutionary models to the data, including a BM be used to help determine which climatic factors (if any) set
model, a model of stasis or stabilizing selection (as in an the range limits of clades and species. Such analyses have
Ornstein-Uhlenbeck, OU, model, with one or more optima), been used to help explain patterns of species richness (e.g.
and a model of white noise (e.g. Kozak & Wiens 2010). climate prevents tropical clades from invading temperate
Finding that a character fits a model of stasis would potentially regions; Wiens et al. 2006) and community structure (e.g.
support NC, whereas a model of white noise would not. climate restricts clades with different microhabitat preferences
Importantly, significant fit to the BM model (phylogenetic to different regions; Stephens & Wiens 2009). Once poten-
signal) could also be consistent with NC (and previous studies tially limiting climatic variables are identified with SDMs, their
claiming to find NC by this criterion have not necessarily been fit to the phylogeny (or rate of change) can then be tested
misled). We find no compelling argument for claiming that as described above (e.g. Wiens et al. 2006; Stephens &
NC is present only when phylogenetic signal is stronger than Wiens 2009).
expected under BM. Peterson et al. (1999) proposed a test of NC based on
This type of model-fitting approach seems promising, but whether the SDM for one species predicts the geographic
may also have limitations. Testing its efficacy under different range of its sister species, and Warren et al. (2008) proposed
simulated evolutionary scenarios should be a priority for several variations on this test. Such tests are complementary
future research. to those based on entire phylogenies, but may be more
Another approach is to use a time-calibrated phylogeny relevant to smaller phylogenetic scales. An important
and estimate rates of trait evolution (e.g. Ackerly 2009b). direction for future work is to take NC tests based on
This approach can then be used to compare rates of change SDMs and combine them with mechanistic modelling of
in different traits and clades (e.g. OMeara et al. 2006). species ranges, which incorporates physiological parameters
However, two caveats should be made. First, estimates of and other factors in addition to climatic data (Kearney &
trait disparity (i.e. variance) within clades are not necessarily Porter 2009).
equivalent to rates of change, particularly if the phylogeny
within clades is ignored (OMeara et al. 2006). Second,
Other tests
comparisons of rates alone do not address whether niches
are significantly conserved or not, only whether they are Again, a plethora of tests may be applied to a diversity of
higher or lower than in another trait or clade. However, NC-related questions. Some tests are similar to those
these rate comparisons could be combined with compari- described above. For example, Cattin et al. (2004) found
sons of alternative models (e.g. BM, OU) to provide a more phylogenetic structure in food webs (a potential manifesta-
complete assessment of NC than gained from either tion of NC) by showing a negative relationship between the
approach alone. phylogenetic distance between species and the similarity in
Similar tests can be applied to both continuous variables the prey taxa consumed, using a Mantel test. This approach
and those treated as categorical or discrete (e.g. arboreality, is related to a test of phylogenetic signal, but focuses on
herbivory). For example, given a phylogeny and a categorical species in a given community, rather than a clade.
variable, one can measure the fit of the character to the tree, Rangel et al. (2007) used an innovative approach to test
randomize states among taxa, and compare the observed fit whether climatic NC drives large-scale patterns of species
to that in the randomizations (e.g. Crisp et al. 2009). richness in South American birds. They simulated the
evolution of species distributions under different rates of
niche evolution among species, and evaluated which rate
Tests based on species distribution modelling
generated richness patterns most closely matching empirical
Many tests of the relationship between NC, climate and patterns. They found that low rates of change (strong NC)
geographic distribution involve species distribution models offered the best fit. This general approach might be applied
(SDMs, also called environmental niche models). For to many other NC-related questions (e.g. community
biogeographic hypotheses, a key idea is that climatically assembly).
unsuitable conditions can limit geographic ranges when Other tests may apply to finer phylogenetic and temporal
there is NC, and such conditions can potentially be scales. For example, range shifts and local extinctions in
response to climate change offer strong evidence for NC Ricklefs 2004). These processes can be viewed as evolu-
over short time scales (e.g. Parmesan & Yohe 2003), as do tionary or biogeographic. Yet, large-scale patterns of species
analyses of climatic niches over time from modern and diversity are often strongly associated with climate, and with
historical locality data (Tingley et al. 2009). other ecological factors at local scales (e.g. soil pH). These
Paleontological studies offer additional evidence. For two perspectives are sometimes seen as being in conflict
example, studies reviewed by Eldredge et al. (2005) (i.e. evolution vs. ecology; Algar et al. 2009). The concept of
showed that many fossil species did not go extinct or NC offers a bridge between them (e.g. Wiens & Donoghue
change morphologically in response to climate change, 2004). NC may explain why species fail to disperse between
but instead shifted their geographic ranges to remain different climates and habitats (e.g. a tropical species cannot
within their ancestral environment (i.e. habitat tracking), rapidly adapt to cold winter temperatures, and fails to
a pattern we attribute to NC. In fact, NC is sufficiently colonize temperate regions). The tendency for a group to
prevalent in the fossil record that certain plant clades remain in its ancestral environment as it diversifies could
are used as indicators of past climates (e.g. palms = trop- lead to higher richness in some climates (e.g. tropical vs.
ical). Stasis in morphological traits over time, a major temperate) or habitats (e.g. mesic vs. arid) than in others,
topic in paleobiology (see Eldredge et al. 2005), may even without differences in rates of speciation and
also offer evidence of NC, provided that the traits are extinction in different environments (Fig. 1). The idea that
niche-related. regions or habitats occupied longer will have more species is
called the time-for-speciation effect (TSE; review in
Stephens & Wiens 2003). There is growing evidence that
Summary
the combination of NC and TSE may help explain many
All tests have limitations, and there is no universal test for richness patterns, including latitudinal, elevational, and local
NC. But the diversity of available tests allows one to diversity.
confirm results with many approaches. Also, different tests Several studies have supported the role of NC and TSE in
may be relevant to NC in different contexts. generating high tropical species richness. Wiens et al. (2006)
It is also important to remember that the niche is multi- showed that high tropical richness in treefrogs (Hylidae) is
dimensional, and in any clade, some aspects may be related to their origin and longer time in tropical regions
conserved while others diversify. For example, Galapagos (TSE), and that expansion of tropical clades into temperate
finches and Rift-lake cichlids are classic examples of North America is limited by a climatic variable (temperature
adaptive radiation, particularly in trophic niches (Schluter seasonality), which shows significant phylogenetic signal
2000). Yet, within these radiations, all species occur under (consistent with NC). Algar et al. (2009) claimed to refute
similar climatic conditions, and exhibit NC from that this, but ignored both time and temperature seasonality, did
perspective. In addition, NC may be evident at some not explicitly compare tropical and temperate regions, and
temporal and phylogenetic scales but not others. Any claim offered no comparable alternative hypothesis. A similar
about conserved or labile niches may only apply to the pattern of ancient tropical origin and recent temperate
trait(s) and time scale under study. dispersal occurs in ranid frogs (Wiens et al. 2009), which
dominate the Old World tropics (unlike the predominately
Neotropical hylids). Local diversity in New World bats also
EMERGING EVIDENCE FOR NICHE CONSERVATISM shows a strong latitudinal TSE (Stevens 2006). In birds,
IN ECOLOGY AND CONSERVATION BIOLOGY tropical regions globally are dominated by more basal clades,
a pattern consistent with NC and TSE (Hawkins et al. 2007).
Overview
A simulation study of South American birds showed that
In this section, we review the growing evidence for the their richness patterns are best explained by strong climatic
importance of NC to many research areas in ecology and NC (Rangel et al. 2007). Recent analyses of global distribu-
conservation. We also point out important areas in need of tion patterns in plants showed evidence for NC constraining
further research. dispersal between major biomes (Crisp et al. 2009) and
for TSE in explaining lowland tropical species richness
(Jansson & Davies 2008). Analyses of climate and richness
Species richness patterns
across mammal clades suggest that the mammalian latitu-
A major goal of ecology is to explain patterns of species dinal diversity gradient may be related to NC and the TSE
richness, from global to local scales. Ultimately, explanations (Buckley et al. 2010).
for diversity patterns must include the processes that Other studies have suggested that tropical richness may
directly change species numbers in a region or community: be explained instead by higher rates of tropical speciation or
speciation, extinction (local or global) and dispersal (e.g. temperate extinction (many reviewed in Mittelbach et al.
2007). However, most did not test for a biogeographic to affect the richness and composition of local communities
TSE at all, making it difficult to evaluate which hypothesis (see also Ackerly 2009a).
(rates vs. time) is more important in explaining diversity In addition to abiotic factors (e.g. climate, pH), biotic
patterns. Furthermore, even if higher rates of tropical factors might also be involved in the interplay of NC and
diversification (speciation extinction) prove to be more TSE in explaining patterns of local diversity. For example,
important than the TSE, NC might still be important in Brown et al. (2000) argued that local species richness
generating latitudinal diversity patterns, for example, by of Enallagma damselfly larvae in lakes with fish as top
limiting dispersal of tropical species into temperate regions predators (fish lakes) is higher than in lakes where dragonfly
(e.g. Allen & Gillooly 2006). Reconciling the relative larvae are top predators (fishless), because use of fish-lake
importance of diversification rates, TSE, and NC in habitat has been conserved in Enallagma for tens of millions
generating the latitudinal diversity gradient is a major of years. In contrast, fishless lakes (which require special
challenge for future research, and future studies should adaptations to cope with predation by dragonflies) represent
consider all of these processes, not just diversification rates. a habitat that has been colonized much more recently by
Niche conservatism-based hypotheses can potentially Enallagma, leaving less time for speciation to build up
explain many other diversity patterns beyond high tropical diversity in these lakes.
richness. For example, some groups actually have higher In summary, there is now evidence that NC may be
richness in temperate regions than in tropical regions. relevant to many richness patterns at many scales. Yet, most
Analyses of predominately temperate clades of frogs and patterns to date have been addressed with only a handful
snakes (Smith et al. 2005; Pyron & Burbrink 2009) suggest of studies, and few have explicitly tested for both NC and
that TSE and NC (i.e. temperate origins and climatic the TSE.
constraints on dispersal, respectively) explain their unusual
diversity patterns. Richness varies elevationally as well as
Community assembly
latitudinally, and in many clades and areas, regional richness
is highest at mid-elevations (e.g. McCain 2005; Oommen & Many recent studies have addressed the conservatism of
Shanker 2005; Smith et al. 2007; Li et al. 2009; Kozak & niche-related traits among species in local communities (e.g.
Wiens 2010). This mid-elevation hump also appears to be microhabitat preference), often with the goal of under-
caused by the TSE (based on studies in frogs, salamanders standing community assembly (e.g. Prinzing et al. 2001;
and fish; Smith et al. 2007; Wiens et al. 2007; Li et al. 2009; Cavender-Bares et al. 2004; Swenson & Enquist 2009).
Kozak & Wiens 2010), with major clades seemingly These studies offer many examples both for and against NC,
originating in environments presently situated at mid- depending on the clade and trait. Many studies involve
elevations, followed by dispersal to lower and higher community phylogenetics (reviews in Cavender-Bares et al.
elevations. NC is hypothesized to limit dispersal between 2009; Vamosi et al. 2009), where NC is often key to
elevational climatic zones, although rigorously demonstrat- interpreting patterns. Assuming strong NC, communities of
ing this remains a major challenge (but see Kozak & Wiens closely related species may represent the effects of habitat
2010). NC may help explain other elevational diversity filtering (close relatives with similar traits are filtered in to a
patterns as well (e.g. decreasing richness at higher eleva- community from the regional species pool) and communi-
tions). ties of distantly related species may represent the effects of
Perhaps the least explored interface of NC and species competition (limiting coexistence of close relatives sharing
richness relates to local-scale diversity. Local and regional similar traits and resource requirements). Tests of NC for
species richness patterns are often strongly correlated each trait are essential for interpreting these patterns, as
(review in Harrison & Cornell 2008), and recent analyses rapid trait evolution may lead to very different conclusions
demonstrate that effects of NC on regional diversity can (e.g. Webb et al. 2002; Losos 2008). However, even given
trickle down to local communities. For example, Partel NC in the relevant traits, these expectations and interpreta-
(2002) showed that local plant richness increased with tions (e.g. ecologically similar species competitively exclude
increasing soil pH in regions of generally high pH but each other) are surprisingly controversial (e.g. Mayfield &
decreased in regions of low pH, and attributed this Levine 2010).
difference to the larger pool of species adapted to the
prevailing pH level in each region. Harrison & Grace (2007)
Ecosystem function
showed that the positive productivity-richness relationship
in the California flora is driven by the large proportion of Recent studies suggest that NC may have important
species regionally with evolutionary affinities to high- consequences for ecosystem function. Maherali & Kliron-
productivity conditions (moist, north-temperate environ- omos (2007) used experimental communities of mycorrhizal
ments) and that the consequences of this NC filtered down fungi to show that plant productivity (a common index of
Loeuille & Loreau 2005; Rossberg et al. 2006; Ingram et al. (a) h1
2009). The importance of trait conservatism in food webs p
p1
suggests that NC could be important for related topics as
well, such as energy flow and nutrient cycling. h2
p2
millions of years. For figs and their pollinating fig-wasps, a et al. 2010; for an older example, see Warwick & Clarke
tight mutualism has been maintained for > 60 million years, 1995). Specifically, anthropogenic disturbances tend to
and is conserved among c. 800 fig species distributed around decrease phylogenetic diversity, suggesting that only some
the world (Rnsted et al. 2005). More generally, phylogenetic clades can tolerate a given modification, leaving a pool of
studies of interaction networks between plants and animals more closely related species afterwards (Fig. 4). For
(i.e. pollinators, seed dispersers) show that related animal example, a detailed study of zooplankton in north-temperate
species tend to interact with sets of related plant species, and lakes (Helmus et al. 2010) demonstrated this pattern, and
vice versa (Rezende et al. 2007), suggesting NC. Other also suggested that sensitivity to each disturbance regime
mutualistic interactions may also be conserved and ancient. (e.g. decreased pH, increased nitrogen) was phylogenetically
For example, fungal lineages associated with lichen symbi- conserved and differed between clades. These latter analyses
oses appear to be very old, suggesting an ancient symbiosis pave the way for understanding how these sensitivities
(Lutzoni et al. 2001). Bacterial endosymbionts are important evolved across the phylogeny.
to many organismal functions, such as herbivory. A recent Given information on the ecological requirements nec-
summary (Moran et al. 2008; their Table 1) of the estimated essary for persistence of a species in intact vs. modified
ages of mutualistic interactions between insects and their habitats (e.g. microclimates, food resources), studying
bacterial endosymbionts reveals seven systems each con- habitat modification from a NC perspective can help
served for > 100 million years. NC in endosymbionts may elucidate the potential for those requirements to evolve,
also drive NC in their hosts, as aphid thermal tolerances are
determined (at least in part) by their endosymbiotic bacteria
(Dunbar et al. 2007). Conversely, analyses of facilitation in
plants (Valiente-Banuet et al. 2006) show that mesic-adapted
clades can expand their niches into arid regions if they grow
under the canopy of xeric-adapted lineages (i.e. nurse
plants). These are just a few examples of how NC and
positive interspecific interactions may be intertwined.
However, as with food webs and disease ecology, the
ecological mechanisms that underlie NC in these cases
remain understudied.
Finally, a recent study (Gomez et al. 2010) found evidence
for phylogenetic conservatism in all types of interspecific
interactions (hostparasite, predatorprey, mutualism) in
116 clades (genera) across the Tree of Life. These authors
including generalists and specialists and viruses, bacteria,
fungi, plants and animals.
Figure 4 Niche conservatism, phylogeny and anthropogenic
Conservation and habitat modification change. Six species belong to two clades (black vs. white circles)
and occur in three communities (squares; where circles represent
Habitat destruction is often considered the most important species from each clade). In the top row of communities, the
current threat to biodiversity (e.g. Dirzo & Raven 2003), and distribution of species is associated with phylogenetically conserved
can also be seen from the framework of NC: habitat is tolerances to a range of conditions along a natural environmental
modified faster than a species can adapt to these changes gradient. In the bottom set of communities, species composition
(Holt & Gomulkiewicz 2004). To some, this may seem has now changed due to conserved tolerances to anthropogenic
trivial; little evolutionary perspective seems necessary to changes, with the loss of one of the clades from these communities
understand why cutting down a forest leads to extinction of and a reduction in phylogenetic diversity. Empirical studies have
endemic, forest-dwelling species. But threats from habitat now demonstrated changes in the phylogenetic composition of
modification will depend on the interaction between the communities in response to climate change, pollution, invasive
species, agriculture, urbanization, and other human modifications
type of modification (e.g. clearcutting, selective logging,
(Knapp et al. 2008; Willis et al. 2008; Dinnage 2009; Helmus et al.
agriculture, pollution), tolerances of species to that modi- 2010), and in some cases have shown the specific biological traits
fication (e.g. ability to withstand heat, low pH), and whether that are conserved and seemingly underlie these responses (e.g.
those tolerances will evolve rapidly or be conserved. flowering time and climate change; Willis et al. 2008). Anthropo-
Recent studies have shown that habitat modification can genic changes have also been shown to lead to a loss of
have non-random phylogenetic effects on impacted com- phylogenetic diversity (e.g. Helmus et al. 2010), seemingly through
munities (e.g. Knapp et al. 2008; Dinnage 2009; Helmus conservatism of niche-related traits.
based on their ability (or inability) to evolve in the past. Pleiotropy and trade-offs between traits may also lead to
At this point, we know little about whether the relevant selection against niche evolution (e.g. Jenkins & Hoffman
traits might evolve rapidly or be conserved, and if processes 1999; Etterson & Shaw 2001). NC may also be enhanced by
that maintain NC might be circumvented (excepting the very sharp contrasts in conditions in space and time (e.g. the
obvious interest in maintaining diversity within species). edge between marine and terrestrial environments, serpen-
A deeper understanding of what habitat modifications a tine and non-serpentine soils), whereas niche evolution may
species or clade can (or cannot) tolerate and adapt to may be facilitated when environmental gradients are more
also suggest how to modify human-altered landscapes to gradual (Holt & Gomulkiewicz 2004).
allow their persistence. Third, niche evolution may be impeded by gene flow. For
example, where species ranges are limited by unfavourable
environmental conditions, adaptation to those extralimital
THE FINAL FRONTIER: CAUSES OF NICHE
conditions may be reduced by gene flow from the centre of
CONSERVATISM
the range (e.g. Kirkpatrick & Barton 1997). Analogous
We argue that the major area for future research in NC is processes should limit evolution of novel niche-related traits
to obtain a better mechanistic understanding of why it within populations, leading to conservatism (assuming no
occurs. In many cases, researchers have shown phylogenetic trait-related assortative mating).
patterns (e.g. association between host and parasite The processes described above have generally been
phylogenies), but the specific ecological traits that underlie demonstrated in both theoretical and empirical studies.
these patterns remain unclear. In some cases, correlative However, the empirical studies have not done so with the
studies may identify the specific traits that underlie these intention of addressing NC per se, and the relative
ecological patterns. In other cases, extensive experimental importance of these processes remains largely unknown.
work may be needed. Such studies may be trait and taxon It is also possible that the causes of NC in a trait might
specific. But once the specific trait is identified, the next change across the history of a clade or range of a species,
question is: what limits changes in this trait over time? even as the trait itself remains constant.
The role of competition and other biotic interactions in In summary, we argue that the major area for future
constraining niche evolution remains poorly studied and is a research in NC is to understand its ecological and
major question for future research in NC. For example, have evolutionary underpinnings, rather than simply document-
most species in clade A remained in habitat X because clade ing it. Some important questions include: what is the relative
B already occupies habitat Y? This could potentially be importance of competition relative to tolerance to abiotic
supported if, within clade A, there are repeated shifts to factors in NC? What microevolutionary processes underlie
habitat Y in regions where clade B is absent. Other NC? For example, is lack of genetic variation typically a
interspecific interactions could also limit or enhance niche limiting factor, or is selection more important? How do the
evolution in a species or clade. As mentioned above, the answers to these questions vary with the topic (large-scale
abiotic tolerance of an endosymbiont may limit the environ- species richness vs. hostparasite relationships), environ-
mental niche of its host (in aphids), and positive interspecific ment, traits and organisms involved?
interactions may expand the environmental niche (nurse
plants provide shade for mesic lineages in deserts).
CONCLUSIONS
Several population-level processes may underlie NC.
First, the evolution of niche-related traits may be con- In this article, we have argued that NC may be relevant to
strained by limited genetic variation in those traits (e.g. many different patterns and questions in ecology and
Bradshaw 1991). conservation biology, and have suggested many areas for
Second, even if there is genetic variation, natural selection future research. But why think about these questions in
may still lead to NC. For example, behavioural habitat terms of NC?
choice should cause animals to consistently avoid habitats in For many questions, the importance of NC is related to
which their fitness will be lower (e.g. deserts for forest the importance of phylogeny. Our review highlights the
dwellers), leading to NC (e.g. Holt & Barfield 2008). need for ecologists and conservation biologists to be aware
Similarly, selection should favour individuals that choose the that many of the traits and patterns they study may have
dietary or microhabitat resources they are best adapted to ancient roots that go far deeper than the species and
utilize (e.g. small seeds for small finches), and these choices ecological conditions seen today (and may not be fully
may increase specialization and reduce opportunities to understood by examining those species and conditions
adapt to alternate resources. In sessile organisms, traits are alone). Thinking about NC encourages thinking about
expected to evolve towards those conditions where most phylogeny. However, this is hardly new (e.g. Brooks &
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Editor, Arne Mooers
Wiens, J.J., Parra-Olea, G., Garcia-Paris, M. & Wake, D.B. (2007).
Phylogenetic history underlies elevational patterns of biodiver- Manuscript received 4 February 2010
sity in tropical salamanders. Proc. R. Soc. Lond. B, 274, 919928. First decision made 17 March 2010
Wiens, J.J., Sukumaran, J., Pyron, R.A. & Brown, R.M. (2009). Second decision made 10 June 2010
Evolutionary and biogeographic origins of high tropical Manuscript accepted 25 June 2010