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Perspectives in Plant Ecology, Evolution and Systematics 15 (2013) 328–337

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Perspectives in Plant Ecology, Evolution and Systematics 15 (2013) 328–337 Perspectives in Plant Ecology, Evolution and Systematics 15 (2013) 328–337 Contents lists available at ScienceDirect Perspectives in Plant Ecology, Evolution and Systematics jo urnal homepage: www.elsevier.com/locate/ppees Research article Conservatism of responses to environmental change is rare under natural conditions in a native grassland ∗ Jonathan A. Bennett , James F. Cahill Jr. Department of Biological Sciences, University of Alberta, Edmonton, AB T6G 2E9, Canada a r t i c l e i n f o a b s t r a c t Article history: Whether or not niche conservatism is common is widely debated. Despite this uncertainty, closely related Received 27 April 2013 species are often assumed to be ecologically similar. This principle has led to the proposed use of phyloge- Received in revised form netic information in forecasting species responses to environmental change. Tests of niche conservatism 23 September 2013 often focus on ‘functional traits’ and environmental tolerances, but there have been limited tests for con- Accepted 16 October 2013 servatism in species’ responses to changes in the environment, especially in the field. The prevalence Available online 24 October 2013 of functional convergence and the likelihood of functional trade-offs in a heterogeneous environment suggest that conservatism of the response niche is unlikely to be detectable under natural conditions. To Keywords: test the relevance of evolutionary information in predicting ecological responses, we tested for conser- Phylogenetic community ecology vatism (measured as phylogenetic signal) of grassland plant population responses to 14 treatments (e.g. Phylogenetic signal Phylogenetic niche conservatism light, nutrients, water, enemies, mutualists), each manipulated for 2–3 years, and 4 treatment categories Belowground ecology (aboveground, belowground, resource, and herbivory) at a single site. Individual treatment responses Grazing showed limited evidence of conservatism, with only weak conservatism in plant responses to mycorrhi- Mycorrhizae zae and grazing. Aspects of the response niche were conserved among monocots both aboveground and belowground, although the pattern varied. Conservatism was limited to grazing aboveground, but below- ground responses were conserved as a group, suggesting fundamental differences in how selection has led to niche conservatism in aboveground and belowground environments. Overall, our results suggest that conservatism of the response niche is not common, but is actually rare. As such, evolutionary rela- tionships are likely to be of limited relevance for predicting species responses under field conditions, at least over the short time scales used in this study. © 2013 Elsevier GmbH. All rights reserved. Introduction many ecological factors differentially affect certain lineages within the community, causing phylogenetic clustering (Helmus et al., Plant populations often respond idiosyncratically to changes in 2010; Verdú and Pausas, 2007). This suggests that phylogeny can be their environment (Tilman, 1987; Turkington et al., 2002). Efforts used as a tool to predict species responses to changes in their envi- have been made to identify species characteristics that can be ronment, but for phylogeny to be a useful predictor of ecological used to develop a predictive framework for changes in the relative responses, the niche must be conserved. However, the prevalence abundance of plant populations (e.g. Grime, 1977; Westoby, 1998). of niche conservatism has been questioned (Knouft et al., 2006; Based upon the idea that related species are more ecologically Lavergne et al., 2010; Losos, 2008; Silvertown et al., 2006b). similar (Darwin, 1859), hypothesized patterns of descent (e.g. a Niche conservatism can have multiple definitions. Here, we phylogeny) have been used with some success in determining how define niche conservatism broadly as the tendency of related species respond to both biotic (Burns and Strauss, 2011; Reinhart species to respond similarly to abiotic or biotic environmental et al., 2012) and abiotic (Niinemets and Valladares, 2006; Prinzing, conditions (Wiens et al., 2010; Wiens and Graham, 2005). This def- 2001; Willis et al., 2008) elements of their environments. Further, inition is more liberal than other definitions that consider niche conservatism to require species being more similar than expected under a model of Brownian evolution (Losos, 2008). While phy- logenetic relatedness is often considered an integrative measure ∗ Corresponding author at: B715 Biological Sciences Building, University of of functional similarity (Mouquet et al., 2012; Webb et al., 2002), Alberta, Edmonton, AB T6G 2E9, Canada. Tel.: +1 780 492 1577; for plants, ecologically relevant traits are often labile (Cavender- fax: +1 780 492 9234. Bares et al., 2006; Grime, 2006) or environmentally plastic (Berg E-mail address: [email protected] (J.A. Bennett). 1433-8319/$ – see front matter © 2013 Elsevier GmbH. All rights reserved. http://dx.doi.org/10.1016/j.ppees.2013.10.001 J.A. Bennett, J.F. Cahill Jr. / Perspectives in Plant Ecology, Evolution and Systematics 15 (2013) 328–337 329 and Ellers, 2010; Burns and Strauss, 2012). Further, there are many tested for niche conservatism (as measured by phylogenetic sig- ways to respond to different aspects of the environment. For exam- nal) in responses to each individual treatment and in responses ple, defensive compounds are produced using different pathways, to four categories of ecological treatments representing resource, but all reduce herbivory (Howe and Jander, 2008) and competitive herbivory, aboveground, and belowground treatment groupings. response is associated with many traits representing different ways of coping with reduced resource availability (Wang et al., 2010). Materials and methods Additionally, traits may be associated with multiple functions, yet multiple traits may determine a species’ functional response to a Site description given factor. High volumes of fine roots can increase both nitrogen and water uptake (Craine et al., 2003), but periodic drought tol- All experiments occurred at the approximately 5000 ha Univer- ◦ erance also requires that the plant be able to store water for later sity of Alberta research ranch at Kinsella, Alberta, Canada (53 05 N, ◦ use, which is not an adaptation related to nitrogen uptake (Craine, 111 33 W). Research occurred in three fields located in two sep- 2009). This suggests that conservatism of a trait does not mean arate sections of the ranch totalling 100 ha. Field 1 was located that a plant’s response to one factor related to that trait can predict in the northern part of the ranch, whereas fields 2 and 3 are in its response to other related factors. Many of the traits necessary the southern part of the ranch, with the two sites separated by to respond to environmental conditions also involve functional approximately 6 km. The fields used are unseeded and unbroken trade-offs, such as those between shade and drought tolerance and represent a savannah habitat with mixed grass prairie (pri- (Niinemets and Valladares, 2006). As a consequence, plant species marily Hesperostipa curtiseta (Hitchc.) Barkworth, Poa pratensis L. may be suited to cope with certain environmental conditions, but and Festuca hallii (Vasey) Piper) interspersed with stands of aspen not others. Thus, for many reasons, ecological responses are often (Populus tremuloides Michx.). Though historically lightly grazed by less conserved than morphological or physiological traits (Losos, cattle, grazing was halted for the duration of each experiment. 2008; Prinzing, 2001). This suggests that evolutionary informa- As is true for many grasslands (Foster et al., 2004; Silvertown tion may be of limited use for predicting how species respond to et al., 2006c; Tilman, 1996), plant community structure and func- environmental conditions in nature. tion varies spatially and temporally. Soils at the site have a thin When suites of traits appear to confer specific functioning, they topsoil layer over glacial till (Lamb, 2008), but are spatially vari- have often been grouped into plant functional strategies (Reich able in texture, chemistry, and topography (Bennett et al., 2013). ◦ et al., 2003; Westoby, 1998). Most commonly, plant strategies are The site has an average annual temperature of 2.8 C and receives associated with responses to resource availability and disturbance, approximately 430 mm of precipitation in an average year, but is where some species are adapted to quick growth and rapid resource subject to periodic drought (Cahill, 2003). Fig. 1 shows the vari- acquisition, while others are adapted to disturbances such as her- ability in species richness, productivity, and phylogenetic diversity bivory (Craine, 2009; Grime, 1977; Reich et al., 2003). Responses to over the duration of the experiments. These data were taken from both resources and herbivory are often consistent within broad, un-manipulated plots at the field site, with species richness and 2 phylogenetically distinct functional groups (Coughenour, 1985; phylogenetic diversity data derived from cover estimates (0.25 m ) Lavorel et al., 1997; Niinemets and Valladares, 2006; Turkington and productivity estimates from live biomass clipped in small plots 2 et al., 2002), yet the evidence for conservatism of traits represent- (0.10 m ), dried, and weighed. Phylogenetic diversity was calcu- ing these plant strategies is mixed (Brunbjerg et al., 2012; Diaz lated using the constructed phylogeny (see below) as abundance
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