Disturbed habitats locally reduce the signal of deep evolutionary history in functional traits of plants.

The functioning of present ecosystems reflects deep evolutionary history of locally cooccurring species if their functional traits show high phylogenetic signal (PS). However, we do not understand what drives local PS. We hypothesize that local PS is high in undisturbed and stressful habitats, either due to ongoing local assembly of species that maintained ancestral traits, or to past evolutionary maintenance of ancestral traits within habitat species-pools, or to both. We quantified PS and diversity of 10 traits within 6704 local plant communities across 38 Dutch habitat types differing in disturbance or stress. Mean local PS varied 50-fold among habitat types, often independently of phylogenetic or trait diversity. Mean local PS decreased with disturbance but showed no consistent relationship to stress. Mean local PS exceeded species-pool PS, reflecting nonrandom subsampling from the pool. Disturbance or stress related more strongly to mean local than to species-pool PS. Disturbed habitats harbour species with evolutionary divergent trait values, probably driven by ongoing, local assembly of species: environmental fluctuations might maintain different trait values within lineages through an evolutionary storage effect. If functional traits do not reflect phylogeny, ecosystem functioning might not be contingent on the presence of particular lineages, and lineages might establish evolutionarily novel interactions.

Root traits explain plant species distributions along climatic gradients yet challenge the nature of ecological trade-offs.

Ecological theory is built on trade-offs, where trait differences among species evolved as adaptations to different environments. Trade-offs are often assumed to be bidirectional, where opposite ends of a gradient in trait values confer advantages in different environments. However, unidirectional benefits could be widespread if extreme trait values confer advantages at one end of an environmental gradient, whereas a wide range of trait values are equally beneficial at the other end. Here, we show that root traits explain species occurrences along broad gradients of temperature and water availability, but model predictions only resembled trade-offs in two out of 24 models. Forest species with low specific root length and high root tissue density (RTD) were more likely to occur in warm climates but species with high specific root length and low RTD were more likely to occur in cold climates. Unidirectional benefits were more prevalent than trade-offs: for example, species with large-diameter roots and high RTD were more commonly associated with dry climates, but species with the opposite trait values were not associated with wet climates. Directional selection for traits consistently occurred in cold or dry climates, whereas a diversity of root trait values were equally viable in warm or wet climates. Explicit integration of unidirectional benefits into ecological theory is needed to advance our understanding of the consequences of trait variation on species responses to environmental change.

Anthropogenic threats to evolutionary heritage of angiosperms in the Netherlands through an increase in high-competition environments.

Present biodiversity comprises the evolutionary heritage of Earth's epochs. Lineages from particular epochs are often found in particular habitats, but whether current habitat decline threatens the heritage from particular epochs is unknown. We hypothesized that within a given region, humans threaten specifically habitats that harbor lineages from a particular geological epoch. We expect so because humans threaten environments that dominated and lineages that diversified during these epochs. We devised a new approach to quantify, per habitat type, diversification of lineages from different epochs. For Netherlands, one of the floristically and ecologically best-studied regions, we quantified the decline of habitat types and species in the past century. We defined habitat types based on vegetation classification and used existing ranking of decline of vegetation classes and species. Currently, most declining habitat types and the group of red-listed species are characterized by increased diversification of lineages dating back to Paleogene, specifically to Paleocene-Eocene and Oligocene. Among vulnerable habitat types with large representation of lineages from these epochs were sublittoral and eulittoral zones of temperate seas and 2 types of nutrient-poor, open habitats. These losses of evolutionary heritage would go unnoticed with classical measures of evolutionary diversity. Loss of heritage from Paleocene-Eocene became unrelated to decline once low competition, shade tolerance, and low proportion of non-Apiaceae were accounted for, suggesting that these variables explain the loss of heritage from Paleocene-Eocene. Losses of heritage from Oligocene were partly explained by decline of habitat types occupied by weak competitors and shade-tolerant species. Our results suggest a so-far unappreciated human threat to evolutionary heritage: habitat decline threatens descendants from particular epochs. If the trends persist into the future uncontrolled, there may be no habitats within the region for many descendants of evolutionary ancient epochs, such as Paleogene.

Amenazas Antropogénicas para la Herencia Evolutiva de las Angiospermas en los Países Bajos a partir del Incremento en los Ambientes de Competencia Elevada Resumen La biodiversidad actual abarca la herencia evolutiva de las épocas de la Tierra. Los linajes de épocas particulares se encuentran con frecuencia en hábitats particulares pero desconocemos si la declinación contemporánea de los hábitats amenaza a la herencia de una época en particular. Nuestra hipótesis supone que dentro de una región determinada, los humanos son una amenaza específica para los hábitats que albergan linajes de una época geológica particular. Suponemos esto pues los humanos amenazan a los ambientes y a los linajes que se diversificaron durante estas épocas. Diseñamos una nueva estrategia para cuantificar, por tipo de hábitat, la diversificación de los linajes de épocas distintas. Cuantificamos para los Países Bajos, una de las regiones mejor estudiada florística y ecológicamente, la declinación de los tipos de hábitat y de especies durante el siglo pasado. Definimos los tipos de hábitat con base en la clasificación de la vegetación y usamos las jerarquías existentes de la declinación de clases y especies de vegetación. Hoy en día, la mayoría de los tipos de hábitat en declinación y el grupo de especies en lista roja se caracterizan por la diversificación incrementada de los linajes que datan del Paleógeno, específicamente el Paleoceno-Eoceno y el Oligoceno. Entre los tipos de hábitat vulnerables con una gran representación de los linajes de estas épocas encontramos a la zona sublitoral e intermareal de los mares templados y dos tipos de hábitats abiertos con deficiencia de nutrientes. Estas pérdidas de linaje evolutivo pasarían desapercibidas con las medidas clásicas de la diversidad evolutiva. La pérdida de la herencia del Paleoceno-Eoceno dejó de estar relacionada con la declinación una vez que contabilizamos la baja competencia, la tolerancia a la sombra y la baja proporción de especies no pertenecientes a la familia Apiaceae, lo que sugiere que estas variables explican la pérdida de herencia del Paleoceno-Eoceno. La pérdida de herencia del Oligoceno estuvo explicada en parte por la declinación de los tipos de hábitat ocupados por competidores débiles y especies tolerantes a la sombra. Nuestros resultados sugieren una amenaza humana para la herencia evolutiva que todavía no ha sido apreciada: la declinación del hábitat amenaza a los descendientes de épocas particulares. Si en el futuro las tendencias siguen sin ser controladas, puede que no haya hábitats en la región para muchos de los descendientes evolutivos de épocas antiguas, como el Paleógeno.

Plant species occurrence patterns in Eurasian grasslands reflect adaptation to nutrient ratios.

Previous studies of Eurasian grasslands have suggested that nutrient ratios, rather than absolute nutrient availabilities and associated productivity, may be driving plant species richness patterns. However, the underlying assumption that species occupy distinct niches along nutrient ratio gradients remains to be tested. We analysed plant community composition and nutrient status of 644 Eurasian wet grassland plots. The importance of nutrient ratios driving variation in species composition was analysed using ordination methods (DCA and CCA). Subsequently, we assessed the niche position and width along the most important nutrient ratio gradient [N:P] for each species. We found that the N:P ratio explained part of the variation in species composition independent from conventional explanatory variables. The N:P ratio explained less variation than soil moisture or pH, but more than productivity or the availability of N and P separately, highlighting its importance for grassland species composition. Species occupied distinct niches along the N:P gradient, and species' niche widths decreased toward extreme nutrient limitation. After correcting for niche position, there was no overall difference in niche width between endangered and non-endangered species. Surprisingly, endangered species with niche optima at the extreme P-limited end of the gradient had broader niches than their non-endangered counterparts. As species occupied distinct niches along a nutrient ratio gradient, future grassland conservation efforts may benefit from targeting changes in nutrient ratios, i.e. the balance between N and P, rather than only focussing on a general reduction in nutrient availability. However, what management interventions can be used for this purpose remains unclear.

Benefits from living together? Clades whose species use similar habitats may persist as a result of eco-evolutionary feedbacks.

Contents 66 I. 67 II. 68 III. 69 IV. 70 V. 73 VI. 75 VII. 77 78 References 78 SUMMARY: Recent decades have seen declines of entire plant clades while other clades persist despite changing environments. We suggest that one reason why some clades persist is that species within these clades use similar habitats, because such similarity may increase the degree of co-occurrence of species within clades. Traditionally, co-occurrence among clade members has been suggested to be disadvantageous because of increased competition and enemy pressure. Here, we hypothesize that increased co-occurrence among clade members promotes mutualist exchange, niche expansion or hybridization, thereby helping species avoid population decline from environmental change. We review the literature and analyse published data for hundreds of plant clades (genera) within a well-studied region and find major differences in the degree to which species within clades occupy similar habitats. We tentatively show that, in clades for which species occupy similar habitats, species tend to exhibit increased co-occurrence, mutualism, niche expansion, and hybridization - and rarely decline. Consistently, throughout the geological past, clades whose species occupied similar habitats often persisted through long time-spans. Overall, for many plant species, the occupation of similar habitats among fellow clade members apparently reduced their vulnerability to environmental change. Future research should identify when and how this previously unrecognized eco-evolutionary feedback operates.

Local dominance of exotic plants declines with residence time: a role for plant-soil feedback?

Recent studies have shown that introduced exotic plant species may be released from their native soil-borne pathogens, but that they become exposed to increased soil pathogen activity in the new range when time since introduction increases. Other studies have shown that introduced exotic plant species become less dominant when time since introduction increases, and that plant abundance may be controlled by soil-borne pathogens; however, no study yet has tested whether these soil effects might explain the decline in dominance of exotic plant species following their initial invasiveness. Here we determine plant-soil feedback of 20 plant species that have been introduced into The Netherlands. We tested the hypotheses that (i) exotic plant species with a longer residence time have a more negative soil feedback and (ii) greater local dominance of the introduced exotic plant species correlates with less negative, or more positive, plant-soil feedback. Although the local dominance of exotic plant species decreased with time since introduction, there was no relationship of local dominance with plant-soil feedback. Plant-soil feedback also did not become more negative with increasing time since introduction. We discuss why our results may deviate from some earlier published studies and why plant-soil feedback may not in all cases, or not in all comparisons, explain patterns of local dominance of introduced exotic plant species.

Museum specimens reveal loss of pollen host plants as key factor driving wild bee decline in The Netherlands.

Evidence for declining populations of both wild and managed bees has raised concern about a potential global pollination crisis. Strategies to mitigate bee loss generally aim to enhance floral resources. However, we do not really know whether loss of preferred floral resources is the key driver of bee decline because accurate assessment of host plant preferences is difficult, particularly for species that have become rare. Here we examine whether population trends of wild bees in The Netherlands can be explained by trends in host plants, and how this relates to other factors such as climate change. We determined host plant preference of bee species using pollen loads on specimens in entomological collections that were collected before the onset of their decline, and used atlas data to quantify population trends of bee species and their host plants. We show that decline of preferred host plant species was one of two main factors associated with bee decline. Bee body size, the other main factor, was negatively related to population trend, which, because larger bee species have larger pollen requirements than smaller species, may also point toward food limitation as a key factor driving wild bee loss. Diet breadth and other potential factors such as length of flight period or climate change sensitivity were not important in explaining twentieth century bee population trends. These results highlight the species-specific nature of wild bee decline and indicate that mitigation strategies will only be effective if they target the specific host plants of declining species.

Phylogenetically poor plant communities receive more alien species, which more easily coexist with natives.

Alien species can be a major threat to ecological communities, but we do not know why some community types allow the entry of many more alien species than do others. Here, for the first time, we suggest that evolutionary diversity inherent to the constituent species of a community may determine its present receptiveness to alien species. Using recent large databases from observational studies, we find robust evidence that assemblage of plant community types from few phylogenetic lineages (in plots without aliens) corresponds to higher receptiveness to aliens. Establishment of aliens in phylogenetically poor communities corresponds to increased phylogenetic dispersion of recipient communities and to coexistence with rather than replacement of natives. This coexistence between natives and distantly related aliens in recipient communities of low phylogenetic dispersion may reflect patterns of trait assembly. In communities without aliens, low phylogenetic dispersion corresponds to increased dispersion of most traits, and establishment of aliens corresponds to increased trait concentration. We conclude that if quantified across the tree of life, high biodiversity correlates with decreasing receptiveness to aliens. Low phylogenetic biodiversity, in contrast, facilitates coexistence between natives and aliens even if they share similar trait states.

Dispersal failure contributes to plant losses in NW Europe.

The ongoing decline of many plant species in Northwest Europe indicates that traditional conservation measures to improve the habitat quality, although useful, are not enough to halt diversity losses. Using recent databases, we show for the first time that differences between species in adaptations to various dispersal vectors, in combination with changes in the availability of these vectors, contribute significantly to explaining losses in plant diversity in Northwest Europe in the 20th century. Species with water- or fur-assisted dispersal are over-represented among declining species, while others (wind- or bird-assisted dispersal) are under-represented. Our analysis indicates that the 'colonization deficit' due to a degraded dispersal infrastructure is no less important in explaining plant diversity losses than the more commonly accepted effect of eutrophication and associated niche-based processes. Our findings call for measures that aim to restore the dispersal infrastructure across entire regions and that go beyond current conservation practices.

Less lineages - more trait variation: phylogenetically clustered plant communities are functionally more diverse.

Functional diversity within communities may influence ecosystem functioning, but which factors drive functional diversity? We hypothesize that communities assembled from many phylogenetic lineages show large functional diversity if assembly is random, but low functional diversity if assembly is controlled by interactions between species within lineages. We combined > 9000 descriptions of Dutch plant communities, a species-level phylogeny, and information on 16 functional traits (including eight dispersal traits). We found that all traits were conserved within lineages, but nevertheless communities assembled from many lineages showed a smaller variation in trait-states of most traits (including dispersal traits) than communities assembled from few lineages. Hence, within lineages, species are not randomly assembled into communities, contradicting Neutral Theory. In fact, we find evidence for evolutionary divergence in trait-states as well as present-day mutual exclusion among related, similar species, suggesting that functional diversity of communities increased due to past and present interactions between species within lineages.

The relationship between global and regional distribution diminishes among phylogenetically basal species.

Phylogenetic legacy and phylogenetic trends affect the ecology of species-except, apparently, for the width of their distribution. As a result, "macroecological" patterns of species distributions emerge constantly in phylogenetically very distinct species assemblages. The width of the global distribution of species, for instance, constantly correlates positively to the width of their regional distribution. However, such patterns primarily reflect the phylogenetically derived species that dominate most assemblages. Basal species, in contrast, might show different macroecological patterns. We tested the hypothesis that the correlation between global and regional distributions of species diminishes among the phylogenetically basal species. We considered central European higher plants and defined global distribution as the occupancy of global floristic zones, regional distribution as the grid occupancy in Eastern Germany, and phylogenetic position as the rank distance to tree base. We also took into account a number of confounding variables. We found that, across all lineages, the global/regional correlation diminished among basal species. We then reanalyzed 19 lineages separately and always found the same pattern. The pattern reflected both increases in global distributions and decreases in regional distributions among basal species. The results indicate that many basal species face a risk of global or at least regional extinction, but have escaped the downward spiral of mutually reinforcing extinction risks at multiple scales. We suggest that many basal species had much time to expand their global ranges but are presently displaced locally by more derived species. Overall, the study shows that macroecological patterns may not be static and universal, but may undergo macroevolutionary trends. Analyses of macroecological patterns across a phylogeny may thus provide insights into macroevolutionary processes.