A roadmap to plant functional island biogeography

Island biogeography is the study of the spatio-temporal distribution of species, communities, assemblages or ecosystems on islands and other isolated habitats. Island diversity is structured by five classes of process: dispersal, establishment, biotic interactions, extinction and evolution. Classical approaches in island biogeography focused on species richness as the deterministic outcome of these processes. This has proved fruitful, but species traits can potentially offer new biological insights into the processes by which island life assembles and why some species perform better at colonising and persisting on islands. Functional traits refer to morphological and phenological characteristics of an organism or species that can be linked to its ecological strategy and that scale up from individual plants to properties of communities and ecosystems. A baseline hypothesis is for traits and ecological strategies of island species to show similar patterns as a matched mainland environment. However, strong dispersal, environmental and biotic-interaction filters as well as stochasticity associated with insularity modify this baseline. Clades that do colonise often embark on distinct ecological and evolutionary pathways, some because of distinctive evolutionary forces on islands, and some because of the opportunities offered by freedom from competitors or herbivores or the absence of mutualists. Functional traits are expected to be shaped by these processes. Here, we review and discuss the potential for integrating functional traits into island biogeography. While we focus on plants, the general considerations and concepts may be extended to other groups of organisms. We evaluate how functional traits on islands relate to core principles of species dispersal, establishment, extinction, reproduction, biotic interactions, evolution and conservation. We formulate existing knowledge as 33 working hypotheses. Some of these are grounded on firm empirical evidence, others provide opportunities for future research. We organise our hypotheses under five overarching sections. Section A focuses on plant functional traits enabling species dispersal to islands. Section B discusses how traits help to predict species establishment, successional trajectories and natural extinctions on islands. Section C reviews how traits indicate species biotic interactions and reproduction strategies and which traits promote intra-island dispersal. Section D discusses how evolution on islands leads to predictable changes in trait values and which traits are most susceptible to change. Section E debates how functional ecology can be used to study multiple drivers of global change on islands and to formulate effective conservation measures. Islands have a justified reputation as research models. They illuminate the forces operating within mainland communities by showing what happens when those forces are released or changed. We believe that the lens of functional ecology can shed more light on these forces than research approaches that do not consider functional differences among species.     

What explains patterns of species richness? The relative importance of climatic‐niche evolution,morphological evolution,and ecological limits in salamanders

A major goal of evolutionary biology and ecology is to understand why species richness varies among clades. Previous studies have suggested that variation in richness among clades might be related to variation in rates of morphological evolution among clades (e.g., body size and shape). Other studies have suggested that richness patterns might be related to variation in rates of climatic‐niche evolution. However, few studies, if any, have tested the relative importance of these variables in explaining patterns of richness among clades. Here, we test their relative importance among major clades of Plethodontidae, the most species‐rich family of salamanders. Earlier studies have suggested that climatic‐niche evolution explains patterns of diversification among plethodontid clades, whereas rates of morphological evolution do not. A subsequent study stated that rates of morphological evolution instead explained patterns of species richness among plethodontid clades (along with “ecological limits” on richness of clades, leading to saturation of clades with species, given limited resources). However, they did not consider climatic‐niche evolution. Using phylogenetic multiple regression, we show that rates of climatic‐niche evolution explain most variation in richness among plethodontid clades, whereas rates of morphological evolution do not. We find little evidence that ecological limits explain patterns of richness among plethodontid clades. We also test whether rates of morphological and climatic‐niche evolution are correlated, and find that they are not. Overall, our results help explain richness patterns in a major amphibian group and provide possibly the first test of the relative importance of climatic niches and morphological evolution in explaining diversity patterns.     

Impacts of climate warming on hybrid zone movement: Geographically diffuse and biologically porous “species borders”

Abstract The ecology and evolutionary biology of insect–plant associations has realized extensive attention, especially during the past 60 years. The classifications (categorical designations) of continuous variation in biodiversity, ranging from global patterns (e.g., latitudinal gradients in species richness/diversity and degree of herbivore feeding specialization) to localized insect–plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious. Semantic and biosystematic (taxonomical) disagreements sometimes detract from more important ecological and evolutionary processes that drive diversification, the dynamics of gene flow and local extinctions. This review addresses several aspects of insect specialization, host‐associated divergence and ecological (including “hybrid”) speciation, with special reference to the climate warming impacts on species borders of hybridizing swallowtail butterflies (Papilionidae). Interspecific hybrid introgression may result in collapse of multi‐species communities or increase species numbers via homoploid hybrid speciation. We may see diverging, merging, or emerging genotypes across hybrid zones, all part of the ongoing processes of evolution. Molecular analyses of genetic mosaics and genomic dynamics with “divergence hitchhiking”, combined with ecological, ethological and physiological studies of “species porosity”, have already begun to unveil some answers for some important ecological/evolutionary questions. (i) How rapidly can host‐associated divergence lead to new species (and why doesn't it always do so, e.g., resulting in “incomplete” speciation)? (ii) How might “speciation genes” function, and how/where would we find them? (iii) Can oscillations from specialists to generalists and back to specialists help explain global diversity in herbivorous insects? (iv) How could recombinant interspecific hybridization lead to divergence and speciation? From ancient phytochemically defined angiosperm affiliations to recent and very local geographical mosaics, the Papilionidae (swallowtail butterflies) have provided a model for enhanced understanding of ecological patterns and evolutionary processes, including host‐associated genetic divergence, genomic mosaics, genetic hitchhiking and sex‐linked speciation genes. Apparent homoploid hybrid speciation in Papilio appears to have been catalyzed by climate warming‐induced interspecific introgression of some, but not all, species diagnostic traits, reflecting strong divergent selection (discordant), especially on the Z (= X) chromosome. Reproductive isolation of these novel recombinant hybrid genotypes appears to be accomplished via a delayed post‐diapause emergence or temporal isolation, and is perhaps aided by the thermal landscape. Changing thermal landscapes appear to have created (and may destroy) novel recombinant hybrid genotypes and hybrid species.     

Trait-based species richness: ecology and macroevolution

Understanding the origins of species richness patterns is a fundamental goal in ecology and evolutionary biology. Much research has focused on explaining two kinds of species richness patterns: (i) spatial species richness patterns (e.g. the latitudinal diversity gradient), and (ii) clade-based species richness patterns (e.g. the predominance of angiosperm species among plants). Here, I highlight a third kind of richness pattern: trait-based species richness (e.g. the number of species with each state of a character, such as diet or body size). Trait-based richness patterns are relevant to many topics in ecology and evolution, from ecosystem function to adaptive radiation to the paradox of sex. Although many studies have described particular trait-based richness patterns, the origins of these patterns remain far less understood, and trait-based richness has not been emphasised as a general category of richness patterns. Here, I describe a conceptual framework for how trait-based richness patterns arise compared to other richness patterns. A systematic review suggests that trait-based richness patterns are most often explained by when each state originates within a group (i.e. older states generally have higher richness), and not by differences in transition rates among states or faster diversification of species with certain states. This latter result contrasts with the widespread emphasis on diversification rates in species-richness research. I show that many recent studies of spatial richness patterns are actually studies of trait-based richness patterns, potentially confounding the causes of these patterns. Finally, I describe a plethora of unanswered questions related to trait-based richness patterns.     

Replicate patterns of species richness, historical biogeography, and phylogeny in Holarctic treefrogs

In recent decades, the field of historical biogeography has become increasingly divorced from evolutionary biology, ecology, and studies of species richness. In this paper, we explore the evolutionary causes of patterns of biogeography and species richness in Northern Hemisphere treefrogs, combining phylogenetics, ancestral area reconstruction, molecular dating methods, and ecological niche modeling. We reconstructed phylogenetic relationships among 58 hylid taxa using data from two mitochondrial genes (12S, ND1) and two nuclear genes (POMC, c-myc). We find that parallel patterns of species richness have developed in Europe, Asia, and in two separate clades of North American hylids, with the highest richness at midtemperate latitudes (30-35 degrees) on each continent. This pattern is surprising given that hylids overall show higher species richness in the New World tropics and given many standard ecological explanations for the latitudinal diversity gradient (e.g., energy, productivity, mid-domain effect). The replicate pattern in Holarctic hylids seems to reflect specialized tolerance for temperate climate regimes or possibly the effects of competition. The results also suggest that long-range dispersal between continental regions with similar climatic regimes may be easier than dispersal between geographically adjacent regions with different climatic regimes. Our results show the importance of ecology and evolution to large-scale biogeography and the importance of large-scale biogeography to understanding patterns of species richness.     

The causes of species richness patterns across space, time, and clades and the role of "ecological limits"

A major goal of research in ecology and evolution is to explain why species richness varies across habitats, regions, and clades. Recent reviews have argued that species richness patterns among regions and clades may be explained by "ecological limits" on diversity over time, which are said to offer an alternative explanation to those invoking speciation and extinction (diversification) and time. Further, it has been proposed that this hypothesis is best supported by failure to find a positive relationship between time (e.g., clade age) and species richness. Here, I critically review the evidence for these claims, and propose how we might better study the ecological and evolutionary origins of species richness patterns. In fact, ecological limits can only influence species richness in clades by influencing speciation and extinction, and so this new "alternative paradigm" is simply one facet of the traditional idea that ecology influences diversification. The only direct evidence for strict ecological limits on richness (i.e., constant diversity over time) is from the fossil record, but many studies cited as supporting this pattern do not, and there is evidence for increasing richness over time. Negative evidence for a relationship between clade age and richness among extant clades is not positive evidence for constant diversity over time, and many recent analyses finding no age-diversity relationship were biased to reach this conclusion. More comprehensive analyses strongly support a positive age-richness relationship. There is abundant evidence that both time and ecological influences on diversification rates are important drivers of both large-scale and small-scale species richness patterns. The major challenge for future studies is to understand the ecological and evolutionary mechanisms underpinning the relationships between time, dispersal, diversification, and species richness patterns.     

Out of the dark: 350 million years of conservatism and evolution in diel activity patterns in vertebrates

Many animals are active only during a particular time (e.g., day vs. night), a partitioning that may have important consequences for species coexistence. An open question is the extent to which this diel activity niche is evolutionarily conserved or labile. Here, we analyze diel activity data across a phylogeny of 1914 tetrapod species. We find strong phylogenetic signal, showing that closely related species tend to share similar activity patterns. Ancestral reconstructions show that nocturnality was the most likely ancestral diel activity pattern for tetrapods and many major clades within it (e.g., amphibians, mammals). Remarkably, nocturnal activity appears to have been maintained continuously in some lineages for ～350 million years. Thus, we show that traits involved in local‐scale resource partitioning can be conserved over strikingly deep evolutionary time scales. We also demonstrate a potentially important (but often overlooked) metric of niche conservatism. Finally, we show that diurnal lineages appear to have faster speciation and diversification rates than nocturnal lineages, which may explain why there are presently more diurnal tetrapod species even though diurnality appears to have evolved more recently. Overall, our results may have implications for studies of community ecology, species richness, and the evolution of diet and communication systems.     

Individuality in seaweeds and why we need to care

Documenting the causes and consequences of intraspecific variation forms the foundation of much of evolutionary ecology. In this Perspectives piece, we review the importance of individual variation in ecology and evolution, argue that contemporary phycology often overlooks this foundational biological unit, and highlight how this lack of attention has potentially constrained our understanding of seaweeds. We then provide some suggestions of promising but underrepresented approaches, for instance: conducting more studies and analyses at the level of the individual; designing studies to evaluate heritability and genetic regulation of traits; and measuring associations between individual variation in functional traits and ecological outcomes. We close by highlighting areas of phycological research (e.g., population biology, ecology, aquaculture, climate change management) that could benefit immediately from including a focus on individual variation. Algae, for their part, provide us with a powerful and diverse set of ecological and evolutionary traits to explore these topics. There is much to be discovered.     

Endemics and their common congener plant species on an East Mediterranean island: a comparative functional trait approach

Understanding evolution and ecology of endemic plants is of great importance for conservation of those rare and endangered species. Pairwise comparisons of plant functional traits could be an adequate method to get insights in evolutionary and ecological processes. We examined whether morphological traits representing competitive ability and habitat specificity differ between endemics and common plants. Therefore, we performed pairwise comparison analyses of 9 plant functional traits in 36 congeneric pairs of endemics and their common congeners on the East Mediterranean island of Cyprus, i.e., the first such study conducted on a Mediterranean island. We found that endemic species prefer higher elevations and more extreme habitats. Endemics were smaller and they had smaller flowers than their common congeners. Common species had higher chromosome numbers than endemic ones. Endemic and common species showed no significant differences in canopy height, inflorescence height, leaf length and width, and flowering period. Our study showed that the situation on a large oceanic island does not differ from results in mainland research areas.     

Integrating animal temperament within ecology and evolution

Temperament describes the idea that individual behavioural differences are repeatable over time and across situations. This common phenomenon covers numerous traits, such as aggressiveness, avoidance of novelty, willingness to take risks, exploration, and sociality. The study of temperament is central to animal psychology, behavioural genetics, pharmacology, and animal husbandry, but relatively few studies have examined the ecology and evolution of temperament traits. This situation is surprising, given that temperament is likely to exert an important influence on many aspects of animal ecology and evolution, and that individual variation in temperament appears to be pervasive amongst animal species. Possible explanations for this neglect of temperament include a perceived irrelevance, an insufficient understanding of the link between temperament traits and fitness, and a lack of coherence in terminology with similar traits often given different names, or different traits given the same name. We propose that temperament can and should be studied within an evolutionary ecology framework and provide a terminology that could be used as a working tool for ecological studies of temperament. Our terminology includes five major temperament trait categories: shyness-boldness, exploration-avoidance, activity, sociability and aggressiveness. This terminology does not make inferences regarding underlying dispositions or psychological processes, which may have restrained ecologists and evolutionary biologists from working on these traits. We present extensive literature reviews that demonstrate that temperament traits are heritable, and linked to fitness and to several other traits of importance to ecology and evolution. Furthermore, we describe ecologically relevant measurement methods and point to several ecological and evolutionary topics that would benefit from considering temperament, such as phenotypic plasticity, conservation biology, population sampling, and invasion biology.     

Trait evolution rates shape continental patterns of species richness in North America's most diverse angiosperm genus (Carex,Cyperaceae)

Ecological opportunity has been associated with increases in diversification rates across the tree of life. Under an ecological diversification model, the emergence of novel environments is hypothesized to promote morpho- and ecospace evolution. Whether this model holds at the clade level within the most species-rich angiosperm genus found in North America (Carex, Cyperaceae) is yet to be tested. Recent works demonstrate a temporal coupling of climate cooling and widespread colonization of Carex in North America, implicating ecological diversification. In addition, research has consistently found asymmetric patterns of lineage-level diversification in the genus. Why does variation in clade sizes exist in the genus? Is ecological diversification involved? In this study, we tested whether rates of morphological and ecological trait evolution are correlated with clade-level species richness in Carex of North America north of Mexico. We constructed a phylogeny of 477 species—an almost complete regional sample. We estimated rates of evolution of morphological traits, habitat, and climatic niche and assessed whether differences in rates of evolution correlate with species richness differences in replicate non-nested sister clades. Our work demonstrates significant positive correlations between climatic niche rates, habitat and reproductive morphological evolution, and species richness. This coupling of trait and niche evolution and species richness in a diverse, continental clade sample strongly suggests that the ability of clades to explore niche and functional space has shaped disparities in richness and functional diversity across the North American flora region. Our findings highlight the importance of the evolutionary history of trait and niche evolution in shaping continental and regional floras.     

Patterns of species diversity and phylogenetic structure of vascular plants on the Qinghai‐Tibetan Plateau

Large-scale patterns of species richness and the underlying mechanisms regulating these patterns have long been the central issues in biogeography and macroecology. Phylogenetic community structure is a result of combined effects of contemporary ecological interactions, environmental filtering, and evolutionary history, and it links community ecology with biogeography and trait evolution. The Qinghai-Tibetan Plateau provides a good opportunity to test the influence of contemporary climate on shaping species richness because of its unique geological history, cold climate, and high biodiversity. In this study, based on high-resolution distributions of ˜9000 vascular plant species, we explored how species richness and phylogenetic structure of vascular plants correlate with climates on the highest (and species rich) plateau on the Earth. The results showed that most of the vascular plants were distributed on the eastern part of the plateau; there was a strong association between species richness and climate, even after the effects of habitat heterogeneity were controlled. However, the responses of richness to climate remarkably depended on life-forms. Richness of woody plants showed stronger climatic associations than that of herbaceous plants; energy and water availability together regulated richness pattern of woody plants; whereas water availability predominantly regulated richness pattern of herbaceous plants. The phylogenetic structure of vascular species clustered in most areas of the plateau, suggesting that rapid speciation and environment filtering dominated the assembly of communities on the plateau. We further propose that biodiversity conservation in this area should better take into account ecological features for different life-forms and phylogenetic lineages.     

Protecting rare and endangered species under climate change on the Qinghai Plateau,China

Climate change‐induced species range shift may pose severe challenges to species conservation. The Qinghai‐Tibet Plateau is the highest and biggest plateau, and also one of the most sensitive areas to global warming in the world, which provides important shelters for a unique assemblage of species. Here, ecological niche‐based model was employed to project the potential distributions of 59 key rare and endangered species under three climate change scenarios (RCP2.6, RCP4.5 and RCP8.5) in Qinghai Province. I assessed the potential impacts of climate change on these key species (habitats, species richness and turnover) and effectiveness of nature reserves (NRs) in protecting these species. The results revealed that that climate change would shrink the geographic ranges of about a third studied species and expand the habitats for two thirds of these species, which would thus alter the conservation value of some local areas and conservation effectiveness of some NRs in Qinghai Province. Some regions require special attention as they are expected to experience significant changes in species turnover, species richness or newly colonized species in the future, including Haidong, Haibei and Haixi junctions, the southwestern Yushu, Qinghai Nuomuhong Provincial NR, Qinghai Qaidam and Haloxylon Forest NR. The Haidong and the eastern part of Haibei, are projected to have high species richness and conservation value in both current and future, but they are currently not protected, and thus require extra protection in the future. The results could provide the first basis on the high latitude region to formulate biodiversity conservation strategies on climate change adaptation.     

Conservation of Chinese Theaceae species under future climate and land use changes

Aim

Climate and land use changes are two major pervasive and growing global causes of rapid changes in the distribution patterns of biodiversity, challenging the future effectiveness of protected areas (PAs), which were mainly designed based on a static view of biodiversity. Therefore, evaluating the effectiveness of protected areas for protecting the species threatened by climate and land use change is critical for future biodiversity conservation.

Location

China.

Methods

Here, using distributions of 200 Chinese Theaceae species and ensemble species distribution models, we identified species threatened by future climate and land use change (i.e. species with predicted loss of suitable habitat ≥30%) under scenarios incorporating climate change, land use change and dispersal. We then estimate the richness distribution patterns of threatened species and identify priority conservation areas and conservation gaps of the current PA network.

Results

Our results suggest that 36.30%–51.85% of Theaceae species will be threatened by future climate and land use conditions and that although the threatened species are mainly distributed at low latitudes in China under both current and future periods, the mean richness of the threatened species per grid cell will decline by 0.826–3.188 species by the 2070s. Moreover, we found that these priority conservation areas are highly fragmented and that the current PA network only covers 14.21%–20.87% of the ‘areas worth exploring’ and 6.91%–7.91% of the ‘areas worth attention’.

Main Conclusions

Our findings highlight the necessity of establishing new protected areas and ecological corridors in priority conservation areas to protect the threatened species. Moreover, our findings also highlight the importance of taking into consideration the potential threatened species under future climate and land use conditions when designating priority areas for biodiversity conservation.     

Insights from intralocus tactical conflict: adaptive states,interactions with ecology and population divergence

Alternative reproductive tactics (ARTs) have improved our understanding of the evolution of adaptive variation; for instance, their study has led us to understand that the best phenotype (e.g. large and flashy) for a tactic that uses one mating behavior (e.g. court females) is often not the best phenotype (e.g. small and inconspicuous) for a tactic that uses a different mating behavior (e.g. chase and force‐copulate females). However, genetic correlations of shared traits across ARTs can constrain ARTs from reaching their optimal states, resulting in intralocus tactical conflict (IATC). While constraints on evolution in general have been well‐established and studied, there are some important implications of constraints due to intralocus tactical conflict on ARTs that have not been incorporated into the field of evolutionary ecology. Here we describe how an appreciation of IATC, including how to detect it and when to expect it, can change our perspectives in three areas: 1) adaptive states for traits associated with ARTs (e.g. growth rates, behavioural plasticity); 2) how selection due to ecological variation across populations can produce patterns of divergence between ARTS; 3) and the evolutionary stability of polymorphisms (e.g. how IATC can explain losses of one ART, and why this can lead to rapid speciation).     

Bat fly species richness in Neotropical bats: correlations with host ecology and host brain

Patterns of ectoparasite species richness in mammals have been investigated in various terrestrial mammalian taxa such as primates, ungulates and carnivores. Several ecological or life traits of hosts are expected to explain much of the variability in species richness of parasites. In the present comparative analysis we investigate some determinants of parasite richness in bats, a large and understudied group of flying mammals, and their obligate blood-sucking ectoparasite, streblid bat flies (Diptera). We investigate the effects of host body size, geographical range, group size and roosting ecology on the species richness of bat flies in tropical areas of Venezuela and Peru, where both host and parasite diversities are high. We use the data from a major sampling effort on 138 bat species from nine families. We also investigate potential correlation between bat fly species richness and brain size (corrected for body size) in these tropical bats. We expect a relationship if there is a potential energetic trade-off between costly large brains and parasite-mediated impacts. We show that body size and roosting in cavities are positively correlated with bat fly species richness. No effects of bat range size and group size were observed. Our results also suggest an association between body mass-independent brain size and bat fly species richness. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The evolution of crocodilian nesting ecology and behavior

Crocodilians comprise an ancient and successful lineage of archosaurs that repeatedly raises questions on how they survived a mass extinction and remained relatively unchanged for ~100 million years. Was their success due to the change‐resistant retention of a specific set of traits over time (phylogenetic conservatism) or due to flexible, generalist capabilities (e.g., catholic diets, phenotypic plasticity in behavior), or some combination of these? We examined the evolution of reproductive ecology and behavior of crocodilians within a phylogenetic perspective, using 14 traits for all 24 species to determine whether these traits were phylogenetically constrained versus (ecologically) convergent. Our analysis revealed that the ancestral crocodilian was a mound nester that exhibited both nest attendance and defense. Nesting mode exhibited 4–5 transformations from mound to hole nesting, a convergence of which habitat may have been a driving factor. Hole nesters were more likely to nest communally, but this association may be biased by scale. Although there were exceptions, mound nesters typically nested during the wet season and hole nesters during the dry season; this trait was relatively conserved, however. About two‐thirds of species timed their nesting with the wet season, while the other third timed their hatching with the onset of the wet season. Nest attendance and defense were nearly ubiquitous and thus exhibited phylogenetic conservatism, but attendance lodging was diverse among species, showing multiple reversals between water and burrows. Collectively, our analysis reveals that reproductive trait evolution in crocodilians reflects phylogenetic constraint (nest attendance, nest defense), ecological convergence (seasonal timing of nesting, nest attendance lodging), or both (mode of nesting). Some traits (e.g., communal nesting and mode of nesting) were autocorrelated. Our analysis provides a framework for addressing hypotheses raised for why there has been trait convergence in reproductive ecology and behavior in crocodilians and why some traits remained phylogenetically conserved.     

The relationship between local and regional diversity of indigenous forest fauna in KwaZulu-Natal Province, South Africa

The relationship between local and regional diversity was tested by regressing local community richness against regional species diversity for three taxa, birds, butterflies and mammals, in subtropical forest. The quadratic model best fits the relationship between local and regional diversity for birds. Local bird species richness is theoretically independent of the size of the regional pool of species and may represent saturated communities. A linear model best describes the relationship for mammals and butterflies. For mammals, the slope is shallow (0.264) and regional richness overestimates local species richness, suggesting communities are undersaturated. Extinction filtering may explain this pattern. Past climatic changes have filtered out many mammalian species, these changes have been too recent for autochthanous speciation, and the relatively low vagility of mammals has prevented extensive recolonisation. Differences in the nature of the diversity relationship between taxa are as much due to independent evolutionary histories as to differences in vagility and colonising potential. A pervasive role is suggested for regional biogeographic processes in the development of faunal assemblage structure. Large-scale processes are not considered in current conservation plans. We encourage the shift of conservation emphasis from local ecological processes and species interactions, to whole communities and consideration of regional processes.     

Alien species richness is currently unbounded in all but the most urbanized bird communities

Urban areas suffer high pressure of introductions of alien species compared to other habitats due to intensive human activities. As trading globally continues to rise, more species will likely be introduced into urban areas. To determine whether this increase in introduction pressure will lead to increased alien species richness in urban areas, or whether other processes would act to impose an upper limit on species richness, we examined how the shape of the relationship between alien species richness and the number of introduced species over time (i.e. introduction pressure) varies along gradients of urbanization. We collected species composition data from urban bird surveys worldwide and used a global database of alien bird introductions to quantify how many species have been introduced over time at different sites. We found that urbanization gradually modified the shape of the studied relationship from linear to asymptotic. Only communities in extremely urbanized environments were associated with an asymptotic relationship, suggesting that alien bird richness has likely not reached its ecological limit in most urban areas. Our results show that urbanization can reduce the importance of introduction pressure in determining alien species richness. Additionally, the results predict that alien species richness will increase at finer spatial scales, especially if the introduced species can survive in urban areas outside of their native range.     

Stochastic losses of fire‐dependent endemic herbs revealed by a 65‐year chronosequence of dispersal‐limited woody plant encroachment

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.