Journal of

Plant Ecology
VOLUME 3, NUMBER 4, PAGES 243250 DECEMBER 2010 doi: 10.1093/jpe/rtq012 Advanced Access published on 28 July 2010 available online at www.jpe.oxfordjournals.org

Generalized food-deceptive orchid species ower earlier and occur at lower altitudes than rewarding ones
Loc Pellissier1,*, Pascal Vittoz1,2, Antonina Ingrid Internicola1,4 and Luc Daniel Bienvenu Gigord3
Department of Ecology and Evolution, University of Lausanne, Biophore, CH-1015 Lausanne, Switzerland Faculty of Geosciences and Environment, University of Lausanne, Biophore, CH-1015 Lausanne, Switzerland `re Georges, Les Colimac union, France Conservatoire Botanique National de Mascarin, 2 rue du Pe xons, 97436 Saint-Leu, La Re 4 Present address: Department of Biological Sciences, 2500 University Drive N.W., University of Calgary, Calgary AB, Alberta, Canada T2N 1N4 *Correspondence address. Department of Ecology and Evolution, University of Lausanne, Biophore, CH-1015 Lausanne, Switzerland. Tel: +41-21-692-42-79; Fax: +41-21-692-42-65; E-mail: loic.pellissier@unil.ch
2 3 1

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Abstract
Aims Food-deceptive pollination, in which plants do not offer any food reward to their pollinators, is common within the Orchidaceae. As food-deceptive orchids are poorer competitors for pollinator visitation than rewarding orchids, their occurrence in a given habitat may be more constrained than that of rewarding orchids. In particular, the success of deceptive orchids strongly relies on several biotic factors such as interactions with co-owering rewarding species and pollinators, which may vary with altitude and over time. Our study compares generalized food-deceptive (i.e. excluding sexually deceptive) and rewarding orchids to test whether (i) deceptive orchids ower earlier compared to their rewarding counterparts and whether (ii) the relative occurrence of deceptive orchids decreases with increasing altitude. Methods To compare the owering phenology of rewarding and deceptive orchids, we analysed data compiled from the literature at the species level over the occidental Palaearctic area. Since owering phenology can be constrained by the latitudinal distribution of the species and by their phylogenetic relationships, we accounted for these factors in our analysis. To compare the altitudinal distribution of rewarding and deceptive orchids, we used eld observations made over the entire Swiss territory and over two Swiss mountain ranges. Important Findings We found that deceptive orchid species start owering earlier than rewarding orchids do, which is in accordance with the hypotheses of exploitation of naive pollinators and/or avoidance of competition with rewarding co-occurring species. Also, the relative frequency of deceptive orchids decreases with altitude, suggesting that deception may be less protable at high compared to low altitude. Keywords: altitude d timing of owering d food-deception d European orchids d Orchidaceae d biogeography Received: 31 July 2009 Revised: 8 April 2010 Accepted: 16 April 2010

INTRODUCTION
Most animal-pollinated angiosperms offer rewards to their pollinators in the form of nectar or pollen in order to establish a stable mutualistic interaction (Simpson and Neff 1983). However, some plants do not produce such rewards, which have raised curiosity for over two centuries (Darwin 1877; Sprengel 1793). This strategy has evolved in approximately one-third of the species in the Orchidaceae (Dafni 1984;

kova et al. 2006; Little 1983). Food-deceptive orchids Jersa do not offer any nectar and their pollen, clumped into pollinia, is most of the time inaccessible as reward (Johnson and Edwards 2000). Because pollinators learn to avoid deceptive owers (Ackerman 1986; Gigord et al. 2002; Smithson and Macnair 1997), food-deceptive orchids usually show a lower reproductive success than their rewarding counterparts due to pollination limitation (Gill 1989; Neiland and Wilcock 1998; Tremblay et al. 2005). To enhance their chance to be

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pollinated, food-deceptive orchids often rely on mimicry. While cases of strict Batesian mimicry, when owers specically imitate several oral signals of a particular rewarding model, are scarce (Anderson and Johnson 2006; Anderson kova et al. 2006), most food-deceptive orchids et al. 2005; Jersa rely on generalized food-deception, in which the plant exhibits kova general rewarding oral signals (Gigord et al. 2002; Jersa et al. 2006). Generalized food-deceptive orchids exploit the innate food-foraging behaviour of pollinators. Their pollination is assured either by naive insects that have not yet learned to avoid these plants in an associative learning context (Heinrich 1975), or by experienced, but exploratory, pollinators whose food resources become depleted (Internicola et al. kova et al. 2006). 2009; Jersa The reproductive success of generalized food-deceptive orchids can vary according to the characteristics of rewarding co-owering species, such as similarity in oral traits (Internicola et al. 2007; Pellegrino et al. 2008), plant spatial distribution (Internicola et al. 2006) or plant abundance (Sabat and Ackerman 1996), through their effect on pollinator visitation behaviour and learning (Internicola et al. 2007). As the characteristics and composition of plant communities usually change within and among seasons (Elzinga et al. 2007), the reproductive success of deceptive orchids, and hence the selective pressures on their reproductive traits, may vary as well. Flowering at the beginning of the season may be particularly critical for deceptive orchids because their reproductive success may be independent of the characteristics of later-owering rewarding species and thus less variable from year to year (Internicola et al. 2008). Earlyowering deceptive orchids may additionally benet from a higher average density of naive pollinators (Heinrich 1975) and from lower competition with rewarding co-owering species (Internicola et al. 2008). Since deceptive orchids are poorer competitors for pollinator visitation, owering early may be more crucial for them than for rewarding orchids. As a result, generalized food-deceptive orchids may evolve to ower earlier than rewarding orchids. In ac kova (2006) recordance with this, Kindlmann and Jersa cently showed that the peak of owering of deceptive orchids may occur earlier compared to that of rewarding orchids. These authors used the owering period of the species published by Delforge (2005) as an estimate of the timing of their owering peak. However, these data correspond to the whole owering period of the species over their entire European range. According to this denition, species distributed throughout Europe occur in different locations where populations may ower asynchronously, leading to a prolonged estimation of the owering period of these species. This may have biased the estimation of the timing of ower kova (2006), in paring peak used by Kindlmann and Jersa ticular for species with a wide geographical distribution. In addition, although plant owering phenology might be constrained by the phylogeny and the latitudinal distribution of the species, these authors did not account for these effects in

their analysis. Therefore, further studies are needed to clearly demonstrate that early owering may be an important component of the success of generalized food-deception. Interestingly, the starting date of owering may be more relevant than the owering peak to test whether deceptive orchids evolved to ower earlier than rewarding orchids. Indeed, the reproductive success of generalized food-deceptive orchids is usually higher for individuals blooming outside the owering peak of their population than those blooming at peak owering. (Fritz 1990; Parra-Tabla and Vargas 2004, 2007; Sabat and Ackerman 1996; Sun et al. 2009), in particular for early-owering individuals (Ackerman 1981; Nilsson 1980, 1983, 1984). Thus, the most successful individuals usually ower before their populations owering peak, leading selection to favour individuals with an early ower opening date (Sun et al. 2009). Therefore, comparing the oweropening date of rewarding and generalized food-deceptive orchids may be a more suitable estimate of the phenological constraints on the success of generalized food-deception than the owering peak. In addition, the starting date of owering may be more reliable than the peak of owering to compare the owering phenology of rewarding and generalized fooddeceptive orchids. Contrarily to the peak of owering, an unbiased starting date of owering can be easily obtained for many European orchids (Delforge 2005). Also, as a species usually starts owering in its most southern locations, it is possible to accurately account for the effect of latitude on orchid owering phenology. The success of important life-history traits can also be constrained by altitude. The altitudinal gradient offers ideal conditions for exploring evolutionary adaptations over short spatial distances (Ko rner 2007). Any observed variation in the frequency of a trait is related to environmental factors correlated with altitude such as temperature, seasonality (Ko rner 2007) or pollinator abundance (Arroyo et al. 1982). For instance, Jacquemyn et al. (2005b) documented a change in orchids breeding system with altitude on La Reunion Island. At lower altitude, animal-pollinated species, e.g. by hawk moth or ies, were more abundant, while at high altitude, species were more often auto-pollinating and cleistogamous (Jacquemyn et al. 2005b). This can be explained by an adaptive evolution to the unpredictable (Catling 1990) and poorer (Arroyo et al. 1982) pollinator environment at high altitudes. Similarly, the success of generalized food-deception may decrease with increasing altitude. Indeed, in a poor pollinator environment, the reproductive success of both rewarding and generalized food-deceptive orchids may similarly decrease. However, the reproductive success of deceptive orchids is usually lower than that of rewarding orchids (Gill 1989; Neiland and Wilcock 1998; Tremblay et al. 2005). As a result, the reproductive success of rewarding orchids may decrease with increasing altitude, but that of deceptive orchids may become very low or even null. Thus, the maintenance of deceptive orchids may be more compromised at high altitude than that of rewarding orchids.

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We investigated whether the success of generalized fooddeception depends on the owering phenology of the species and on their altitudinal distribution. First, we compared the owering phenology of rewarding and generalized fooddeceptive orchids based on European published data. We accounted for the latitudinal distribution of each species and the phylogenetic relationships among them. Generalized food-deceptive orchids should start owering earlier in the season than rewarding orchids, but this difference should decrease with increasing latitude. Second, we investigated the effect of altitude on the relative frequency of generalized food-deceptive orchids on two data sets, one vegetation survey at the Swiss scale and one at a smaller regional scale. We expected that the frequency of generalized food-deceptive orchids would decrease with increasing altitude.

an equivalent status of each species within a genus. We performed the analyses with all branches scaled to unit length, as branch lengths were not available for the whole tree.

Effect of altitude
We investigated the variation of frequency of generalized food-deception with altitude using 87 865 observations of 60 orchid species recorded in the Swiss Floristic Network (CRSF/ZDSF) database. This database records all the observations of plant species made over the Swiss territory. We calculated the proportion of occurrences of food-deceptive orchids relative to the total number of occurrences of orchids in altitudinal intervals of 100 m ranging from 400 to 2800m. If a trait is signicantly more frequent in some part of an altitudinal gradient, we may assume that individuals exhibiting this trait are better adapted to the conditions found in this part of the gradient and hence have a higher probability of establishment than individuals that do not exhibit it. For instance, Jacquemyn et al. (2005b) discussed the increasing occurrence of selfpollination with altitude in orchids of La Reunion as probably due to better recruitment of self-pollinating siblings compared to cross-pollinating ones. Conversely, if a trait has no effect on the probability of establishment of individuals that exhibit it, no trend regarding its frequency should be found along the gradient. In order to obtain more conservative results, when observations of individuals in a altitudinal class were <10, at the extreme parts of the gradient, we did not take the class into account, because the proportion would be based on an insufcient sample size that might be misleading. We analysed these data with a general linear model (GLM; McCullagh and Nelder 1989) with the altitudinal factor in quadratic form. Such models provide a good estimation of potential trends over an entire gradient, even for non-linear correlations. We also studied the frequency of generalized food-deception along altitude by using eld observations at the regional scale. We used data from 734 non-forest vegetation plots located in the external calcareous Alps and Jura mountain of Canton de Vaud in Switzerland (hereafter Vaud Mountains), investigated in the context of the MODIPLANT (Guisan and Thuiller 2005; Randin et al. 2006) and PERMANENT.PLOT.CH (http:// www.unil.ch/ecospat/page48113_en.html) projects. The sampling points were either randomly stratied by classes of elevation, slope and aspect or by elevation only. Each class of altitude, ranging from 1000 to 2500 m, received identical sampling efforts. The plants observed in each of the sampling points were exhaustively inventoried. From this data set, the occurrences of orchids with their associated altitude were assembled. We analysed 493 observations from 23 species. We calculated the proportion of occurrences of deceptive orchids relative to the total number of occurrences of orchids in altitudinal intervals of 100 m. As above, when observations of individuals in an interval were <10, we did not take the class into account. As previously, we applied a GLM with the altitudinal factor in quadratic form.

MATERIALS AND METHODS

Starting date of owering
We compiled the starting month of owering of 233 rewarding and generalized food-deceptive orchid species from 22 genera es dEurope published in the book Guide des Orchide (Delforge 2005). Our data set included species occurring in the occidental Palaearctic area, which extends from Northern Europe to Northern Africa and from Western Europe to Eastern Europe (49 E). The starting month of owering corresponds to the earliest month when a species owers over its entire range. As the starting date of owering denitely depends on the latitudinal distribution of the species, we also extracted information from the same source on the most southern country where each species is recorded. We transformed the categorical variable country into a continuous variable by linking each country to its mean latitude. Information regarding the reward strategy was provided by the literature (Davies et al. 2005; Delforge 2005, 2007; Jacquemyn et al. kova et al. 2006; Neiland and Wilcock 1998; 2005a; Jersa Smithson 2006). We analysed the effect of the reward strategy on the starting month of owering in an analysis of covariance, using the southern latitudinal limit of the species (latitude) as covariate. We used generalized estimating equations (GEEs) to take phylogenetic dependence among orchid species into account (Paradis and Claude 2002), by using the Compar gee function as implemented in the Analyses of Phylogenetics and Evolution package (APE) (Paradis and Claude 2002). GEEs quantify the phylogenetic dependences of the observations by a correlation matrix based on the phylogenetic relationships among the observed taxa. Using GEE to account for the phylogeny of the species ensures that any other signicant effect obtained is not due to the phylogeny. We used the published phylogenetic trees of European orchids (Bateman et al. 2003; Freudenstein et al. 2004; Salazar et al. 2009; Tyteca and Klein 2008) to build a phylogenetic tree comprising the 233 analysed species. As the phylogeny within genera is not fully resolved, particularly in the Epidendroideae subfamily, we assumed

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All statistics were performed with R 2.6.1 software (R development Core Team 2007).

RESULTS
Starting date of owering
Based on the onset of owering of 233 European orchid species, we found that food-deceptive orchids start owering signicantly earlier than rewarding orchids (Table 1, Fig. 1). This result is unlikely due to phylogenetic patterns among the species as we accounted for such effects in our analysis. We also found that orchids tend to ower later in the season at increasing latitude (Table 1, Fig. 1). Finally, the interaction between starting month of owering and latitude was signicant, indicating that the difference in the starting month of owering between rewarding and deceptive orchids tends to decrease with increased latitude(Table 1, Fig. 1).
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Effect of altitude
Relationship between altitude and frequency of orchids of different reward strategies in Switzerland showed a signicant decrease in the occurrence of generalized food-deceptive orchid species with increasing altitude on both data sets from Switzerland, for the whole Swiss territory (explained deviance of the GLM = 0.74, P < 0.0001, Fig. 2a) and for the Vaud Mountains (explained deviance of the GLM = 0.64, P < 0.0001, Fig. 2b). We examined whether this result may have been due to different levels of rarity and/or geographical distribution between the rewarding and generalized fooddeceptive orchids. To ensure that the frequency of rarity did not differ between species of the two reward types, we counted the number of rewarding and deceptive orchids that occur on the Swiss red list of threatened species for both data sets (at the Swiss territory and the Vaud Mountains scales). The proportion of rare or threatened species did not differ among nectar strategies (chi-square testSwiss territory: chi = 0.8, degree of freedom (df) = 1, P = 0.3711; Vaud Mountains: chi = 0.33, df = 1, P = 0.5637). Similarly, the geographical
Table 1: results of the analysis of covariance testing effect of the nectar strategy (rewarding vs. generalized food-deceptive) on the starting month of owering, with latitude as a co-variable and implemented with the GEEs to account for the phylogeny of the 233 species
Estimation of coefcients Intercept Latitude Nectar strategy Latitude 3 nectar strategy Estimate Standard error t 3.49 0.054 1.84 0.02 0.61 0.007 0.39 0.008 P 5.7 <0.001 8 2.1 <0.001 0.043 dfPa = 35.9 4.7 <0.001

Figure 1: the relationship between the starting month of owering and the latitude of the southernmost country occupied by 233 rewarding (+, solid line) and generalized food-deceptive (3, dashed line) orchid species considered in the analysis.

Boldface indicates signicant effects. a Phylogenetic degrees of freedom (dfP) are residual number of degrees of freedom corrected to account for branch length distances (Paradis 2006).

Figure 2: relationship between altitude and frequency of occurrences of generalized food-deceptive orchid species in Switzerland with the quadratic GLMs based on two data sets, from (a) 87 865 observations of 60 orchid species in the CRSF database over the Swiss territory and (b) 493 observations of 23 orchid species in the Vaud Mountains.

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distributions of only four deceptive and three rewarding orchids were limited to a small part of the Swiss territory, while most species used in these analyses have a large geographical distribution over the Swiss territory (Lauber and Wagner 2007) so that their altitudinal range reects their ecological limits. For these reasons, we conclude that the nding of decreasing frequency of generalized food-deception with increasing altitude is not due to differences in rarity or actual range of the distribution.

DISCUSSION
Although we accounted for the latitudinal distribution of and the phylogenetic relationships among species, generalized food-deceptive orchids start owering signicantly earlier than rewarding orchids throughout the occidental Palaearctic area. This result is in agreement with Heinrichs hypothesis (1975) that owering early may represent an adaptive strategy to benet from the higher abundances of naive pollinators and to avoid competition with later-owering rewarding species (Internicola et al. 2008). This interpretation may seem counterintuitive with regards to the magnet-species hypothesis (Johnson et al. 2003), which stipulates that deceptive plants may benet from higher pollinator visitation rates when they co-ower with rewarding plants (Johnson et al. 2003; Laverty 1992; Pellegrino et al. 2008). However, different food-deceptive species might resort to different mechanisms to increase their chance to be pollinated, probably according to the ecological conditions of their local environment. For instance, some deceptive orchids such as Dactylorhiza sambucina may simply evolve to ower earlier than rewarding co-occurring plants, while others, such as Traunsteinera globosa, may rely on the magnet species effect (Juillet et al. 2007). Thus, our result that generalized food-deceptive orchids globally start owering earlier than rewarding orchids suggests that early owering may be a common strategy among generalized food-deceptive orchids and hence a key factor in the success of generalized food deception. Alternatively, Dafni and Bernhardt (1990) proposed that food-deceptive orchids, which usually experience lower pollinator visitation rates than rewarding orchids, may evolve long-lived owers to maximize opportunities for mating. Based on this hypothesis, Ruxton and Schaefer (2009) argued that the evolution of long-lasting owers may at the same time cause deceptive orchids to ower earlier than rewarding orchids. Accordingly, the early owering of deceptive orchids would only be a consequence of selection for longer ower life span. However, this hypothesis seems unlikely, since several studies showed that food-deceptive orchids are under selection for early owering (Parra-Tabla and Vargas 2004; Sabat and Ackerman 1996; Sun et al. 2009). We also found a signicant interaction between the effects of nectar and latitude on the starting month of owering of rewarding and generalized food-deceptive orchids over the occidental Palaearctic area. As latitude increases, vegetation period usually shortens, which may prevent generalized food-

deceptive orchids to ower far earlier than their rewarding counterparts, as they do in southern areas. This latitudinal effect may reduce the benets of early owering in deceptive orchids, potentially decreasing the occurrence of generalized food-deceptive orchids with increasing latitude. Our analysis based on the starting date of owering did not allow to test whether latitude indeed constrains the success of generalized food-deception, but suggests that such a constraint may exist. The signicant decrease in the frequency of generalized food-deceptive orchid species with altitude over the Swiss territory and the Vaud Mountains indicates that generalized fooddeception may be disadvantageous at high altitude compared to lowlands. While generally decreasing proportion of orchid species with increasing altitude could be caused by environmental factors correlated with altitude, such as temperature or seasonality (Ko rner 2007), this probably does not explain the difference between nectar strategies. In contrast, some studies have documented a decrease in pollinator diversity with increasing altitude (Arroyo et al. 1982), more variable pollinator environments at high altitudes (Catling 1990) and lower pollinator visitation rates in alpine populations (Bingham and Orthner 1998; Warren et al. 1988). A decrease in temperature explains this pattern (Dillon et al. 2006), which could lead to reduced fruit initiation (Malo and Baonza 2002) because such changes in insect community composition and pollinator visitation may decrease the reproductive success of both rewarding and deceptive orchids. However, as the reproductive success of deceptive orchids is usually lower than the one of rewarding orchids (Gill 1989; Neiland and Wilcock 1998; Tremblay et al. 2005), their reproductive success may become very low or even null at high altitudes, which may compromise their maintenance. In addition, vegetation period usually shortens with increasing altitude, so that plant species rather tend to ower simultaneously at high altitude (Ko rner 2003). This may lead to increased competition for access to pollinators and decreased access to naive pollinators, which may explain why deceptive orchids occur less frequently at higher elevations. Moreover, visiting food-deceptive owers may be less disadvantageous in lowlands than at high altitudes, where the time to forage is diminished and lower air temperature imposes increased energetic costs of ight (Dillon et al. 2006). Hence, alpine pollinators may be more careful to avoid food-deceptive plants, e.g., by increasing their learning efciency to visit the most protable owers or by using diverse probing techniques to check whether nectar is present (Heinrich 1979; Howell and Alarcon 2007; Wetherwax 1986). Finally, other factors related to recruitment may also account for differences in occurrences at high altitude between nectar strategies over the Swiss territory and the Vaud Mountains. Indeed, rewarding orchids generally have a higher reproductive success in terms of fruit set than deceptive orchids do (Gill 1989; Neiland and Wilcock 1998; Tremblay et al. 2005). In the context of source-sink dynamics, the realized niche of a species can be extended by dispersal from source into sink populations (Pulliam 2000). If their source

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populations produce more seeds, rewarding orchids may have an increased probability of establishment in constraining habitats located higher in altitude compared to deceptive orchids. The methods we used have a number of limitations. First, the data set from the Swiss territory, although containing a large number of observations, is not strictly rigorous. It might contain several sources of sampling bias due to the nonrandomness of the observations. Second, our classication of each species into rewarding or generalized food-deceptive might be too strict in the light of the more subtle differences between nectar strategies in nature. While the species we classied as deceptive produce no nectar at all, rewarding species might produce a wide range of nectar amounts. Also, some deceptive orchid species might provide a food reward in a different form than nectar. For instance, some insects collect a secretion, located on the stigma surface, composed of sugar and amino acids, meant to retain the dry pollen coming from the pollinia (Delforge 2005). Moreover, some species that provide nectar might depend more on autogamy than insect pollination to ensure their reproduction (Delforge 2005). Third, literature data regarding owering are rather crude. Thus, eld observations might be more informative than the starting date of owering standardized for each species as used here. Fourth, the results presented here are mainly correlative, which is not sufcient to prove the causality of the relationship between pollinator availability or diversity and the temporal and spatial distribution of generalized food-deception in orchids. In the future, similar analyses should be conducted on other data sets to conrm the trends found in this study. For instance, records of the phenology of several populations of rewarding and generalized food-deceptive species at a more local scale, complemented with measures of the ower visitation rate and reproductive success throughout the owering season and for different altitudes, may offer deeper insight into the reproductive strategies of generalized food-deceptive orchids. Additionally, manipulative in situ experiments could be undertaken to investigate which factors are the most constraining on generalized food-deceptive orchids and best account for their decreasing abundance with increasing altitude. In conclusion, we found that generalized food-deceptive orchids start owering earlier compared to rewarding orchids throughout the occidental Palaearctic area. This result suggests that early owering of deceptive orchids may represent an adaptive strategy to exploit the high abundance of naive pollinators and/or the decrease in competition with rewarding coowering species for access to pollinators. We also found that orchids start owering later in the season at increasing latitude and that the difference in the starting date of owering of rewarding and deceptive orchids decreases with increasing latitude. The shortening of vegetation period and stronger environmental constraints with increasing latitude may prevent orchids to ower too early and generalized fooddeceptive orchids to ower far earlier compared to rewarding orchids. Furthermore, we found a clear decrease in the proportion of generalized food-deceptive orchid species with increas-

ing altitude both over the Swiss territory and the Vaud Mountains. This pattern may be due to several environmental factors, in particular including reduced pollinator visitation rates or pollinator-mediated interactions with co-occurring species. To our knowledge, this is the rst clear evidence that a generalized food-deceptive strategy may be constrained by altitude and that generalized food-deceptive orchids ower earlier compared to rewarding orchids.

FUNDING
Centre de Conservation de la Nature du Canton de Vaud; Federal Ofce of the Environment; Swiss National Science Foundation (3100A0-100754/1 to L.D.B.G. and PBLAA122727 to I.A.I.).

ACKNOWLEDGEMENTS
and J. Dwyer for helpful comments. We thank C. Wedekind, J. Durose We are grateful to A. Guisan and C. Randin for providing regional eld sampling of vegetation, to the Centre de Conservation de la Nature du Canton de Vaud and the Federal Ofce of the Environment for nancially supporting their collection and to the CRSF (www.crsf.ch) and AGEO research group (www.ageo.ch) for providing orchid observation data over the Swiss territory. Conict of interest statement. None declared.

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