The relationship between geographic range size and life history traits: is biogeographic history uncovered? A test using the Iberian butterflies

The geographic range of a species is influenced by past phylogenetic and biogeographic patterns. However, other historical interactions, including the interplay between life history and geography, are also likely involved. Therefore, the range size of a species can be explained on the basis of niche‐breadth or dispersal related hypotheses, and previous work on European butterflies suggests that both, under the respective guise of ecological specialisation and colonising ability may apply. In the present study, data from 205 species of butterflies from the Iberian peninsula were processed through multiple regression analyses to test for correlations between geographic range size, life history traits and geographic features of the species distribution types. In addition, the percentage of variance explained by the subsets of variables analyzed in the study, with and without control for phylogenetic effects was tested. Despite a complex pattern of bivariate correlations, we found that larval polyphagy was the single best correlate of range size, followed by dispersal. Models that combined both life history traits and geographic characteristics performed better than models generated independently. The combined variables explained at least 39% of the variance. Bivariate correlations between range size and body size, migratory habits or egg size primarily reflected taxonomic patterning and reciprocal correlations with larval diet breadth and adult phenology. Therefore, aspects of niche breadth i.e. potential larval diet breadth emerged as the most influential determinants of range size. However, the relationships between these types of ecological traits and biogeographic history must still be considered when associations between life history and range size are of interest.     

A meta-analysis of the traits affecting dispersal ability in butterflies: can wingspan be used as a proxy?

1. Dispersal ability of a species is a key ecological characteristic, affecting a range of processes from adaptation, community dynamics and genetic structure, to distribution and range size. It is determined by both intrinsic species traits and extrinsic landscape-related properties. 2. Using butterflies as a model system, the following questions were addressed: (i) given similar extrinsic factors, which intrinsic species trait(s) explain dispersal ability? (ii) can one of these traits be used as a proxy for dispersal ability? (iii) the effect of interactions between the traits, and phylogenetic relatedness, on dispersal ability. 3. Four data sets, using different measures of dispersal, were compiled from published literature. The first data set uses mean dispersal distances from capture-mark-recapture studies, and the other three use mobility indices. Data for six traits that can potentially affect dispersal ability were collected: wingspan, larval host plant specificity, adult habitat specificity, mate location strategy, voltinism and flight period duration. Each data set was subjected to both unifactorial, and multifactorial, phylogenetically controlled analyses. 4. Among the factors considered, wingspan was the most important determinant of dispersal ability, although the predictive powers of regression models were low. Voltinism and flight period duration also affect dispersal ability, especially in case of temperate species. Interactions between the factors did not affect dispersal ability, and phylogenetic relatedness was significant in one data set. 5. While using wingspan as the only proxy for dispersal ability maybe problematic, it is usually the only easily accessible species-specific trait for a large number of species. It can thus be a satisfactory proxy when carefully interpreted, especially for analyses involving many species from all across the world.     

A macroevolutionary perspective on species range limits

Understanding the factors that determine the geographic range limits of species is important for many questions in ecology, evolution and conservation biology. These limits arise from complex interactions among ecology and dispersal ability of species and the physical environment, but many of the underlying traits can be conserved among related species and clades. Thus, the range limits of species are likely to be influenced by their macroevolutionary history. Using palaeontological and biogeographic data for marine bivalves, we find that the range limits of genera are significantly related to their constituent species richness, but the effects of age are weak and inconsistent. In addition, we find a significant phylogenetic signal in the range limits at both genus and family levels, although the strength of this effect shows interoceanic variation. This phylogenetic conservatism of range limits gives rise to an evolutionary pattern where wide-ranging lineages have clusters of species within the biogeographic provinces, with a few extending across major boundaries.     

Human pressures predict species’ geographic range size better than biological traits

Geographic range size is the manifestation of complex interactions between intrinsic species traits and extrinsic environmental conditions. It is also a fundamental ecological attribute of species and a key extinction risk correlate. Past research has primarily focused on the role of biological and environmental predictors of range size, but macroecological patterns can also be distorted by human activities. Here, we analyse the role of extrinsic (biogeography, habitat state, climate, human pressure) and intrinsic (biology) variables in predicting range size of the world's terrestrial mammals. In particular, our aim is to compare the predictive ability of human pressure vs. species biology. We evaluated the ability of 19 intrinsic and extrinsic variables in predicting range size for 4867 terrestrial mammals. We repeated the analyses after excluding restricted‐range species and performed separate analyses for species in different biogeographic realms and taxonomic groups. Our model had high predictive ability and showed that climatic variables and human pressures are the most influential predictors of range size. Interestingly, human pressures predict current geographic range size better than biological traits. These findings were confirmed when repeating the analyses on large‐ranged species, individual biogeographic regions and individual taxonomic groups. Climatic and human impacts have determined the extinction of mammal species in the past and are the main factors shaping the present distribution of mammals. These factors also affect other vertebrate groups globally, and their influence on range size may be similar as well. Measuring climatic and human variables can allow to obtain approximate range size estimations for data‐deficient and newly discovered species (e.g. hundreds of mammal species worldwide). Our results support the need for a more careful consideration of the role of climate change and human impact – as opposed to species biological characteristics – in shaping species distribution ranges.     

Correlates of diversification in the plant clade Dipsacales: geographic movement and evolutionary innovations

We explore patterns of diversification in the plant clades Adoxaceae and Valerianaceae (within Dipsacales), evaluating correlations between biogeographic change (i.e., movements into new areas), morphological change (e.g., the origin of putative key innovations associated with vegetative and reproductive characters), and shifts in rates of diversification. Our findings indicate that rates of diversification in these plants tend to be less tightly correlated with the evolution of morphological innovations but instead exhibit a pronounced correlation with movement into new geographic areas, particularly the dispersal of lineages into new mountainous regions. The interdependence among apparent novelties (arising from their nested phylogenetic distribution) and the correlation between morphological and biogeographic change suggests a complex history of diversification in Dipsacales. Overall, these findings highlight the importance of incorporating biogeographic history in studies of diversification rates and in the study of geographic gradients in species richness. Furthermore, these results argue against a simple deterministic relationship between dispersal and diversification: like other factors that may influence the probability of speciation and/or extinction, the impact of dispersal on diversification rates depends on being in the right place at the right time.     

The global invasion success of Central European plants is related to distribution characteristics in their native range and species traits

Aim Determining which traits predispose a species to become invasive is a fundamental question of invasion ecology, but traits affect invasiveness in concert with other factors that need to be controlled for. Here, we explore the relative effects of biological traits of plant species and their distributional characteristics in the native range on invasion success at two stages of invasion. Location Czech Republic (for native species); and the world (for alien species). Methods The source pool of 1218 species of seed plants native to Central Europe was derived from the flora of the Czech Republic, and their occurrence in 706 alien floras all over the world was recorded, distinguishing whether they were listed as an ‘alien’ or a ‘weed’ in the latest version of Randall’s ‘Global compendium of weeds’ database. The latter type of occurrence was considered to indicate species ability to invade and cause economic impact, i.e. a more advanced stage of invasion. Using the statistical technique of regression trees, we tested whether 19 biological traits and five distributional characteristics of the species in their native range can be used to predict species success in two stages of invasion. Results The probability of a species becoming alien outside its native distribution range is determined by the size of its native range, and its tolerance of a wide range of climates acquired in the region of origin. Biological traits play only an indirect role at this stage of invasion via determining the size of the native range. However, the ability of species to become a weed is determined not only by the above characteristics of native distribution, but also directly by biological traits (life form and strategy, early flowering, tall stature, generative reproduction, number of ploidy levels and opportunistic dispersal by a number of vectors). Species phylogenetic relatedness plays only a minor role; it is more important at the lowest taxonomic levels and at the later stage of invasion. Main conclusion The global success of Central European species as ‘weeds’ is determined by their distributional characteristics in the native ranges and by biological traits, but the relative importance of these determinants depends on the stage of invasion. Species which have large native ranges and are common within these ranges should be paid increased attention upon introductions, and the above biological traits should be taken into account in screening systems applied to evaluate deliberate introductions of alien plants to new regions.     

Long‐distance dispersal syndromes matter: diaspore–trait effect on shaping plant distribution across the Canary Islands

Oceanic islands emerge lifeless from the seafloor and are separated from continents by long stretches of sea. Consequently, all their species had to overcome this stringent dispersal filter, making these islands ideal systems to study the biogeographic implications of long‐distance dispersal (LDD). It has long been established that the capacity of plants to reach new islands is determined by specific traits of their diaspores, historically called dispersal syndromes. However, recent work has questioned to what extent such dispersal‐related traits effectively influence plant distribution between islands. Here we evaluated whether plants bearing dispersal syndromes related to LDD – i.e. anemochorous (structures that favour wind dispersal), thalassochorous (sea dispersal), endozoochorous (internal animal dispersal) and epizoochorous (external animal dispersal) syndromes – occupy a greater number of islands than those with unspecialized diaspores by virtue of their increased dispersal ability. We focused on the native flora of the lowland xeric communities of the Canary Islands (531 species) and on the archipelago distribution of the species. We controlled for several key factors likely to affect the role of LDD syndromes in inter‐island colonization, namely: island geodynamic history, colonization time and phylogenetic relationships among species. Our results clearly show that species bearing LDD syndromes have a wider distribution than species with unspecialized diaspores. In particular, species with endozoochorous, epizoochorous and thalassochorous diaspore traits have significantly wider distributions across the Canary archipelago than species with unspecialized and anemochorous diaspores. All these findings offer strong support for a greater importance of LDD syndromes on shaping inter‐island plant distribution in the Canary Islands than in some other archipelagos, such as Galápagos and Azores.     

Evolution of fruit traits in ficus subgenus Sycomorus (Moraceae): to what extent do frugivores determine seed dispersal mode?

Fig trees are a ubiquitous component of tropical rain forests and exhibit an enormous diversity of ecologies. Focusing on Ficus subgenus Sycomorus, a phenotypically diverse and ecologically important Old World lineage, we examined the evolution of fruit traits using a molecular phylogeny constructed using 5 kilobases of DNA sequence data from 63 species (50% of global diversity). In particular, we ask whether patterns of trait correlations are consistent with dispersal agents as the primary selective force shaping morphological diversity or if other ecological factors may provide a better explanation? Fig colour, size and placement (axial, cauliflorous, or geocarpic) were all highly evolutionarily liable, and the same fruit traits have evolved in different biogeographic regions with substantially different dispersal agents. After controlling for phylogenetic autocorrelation, we found that fig colour and size were significantly associated with fig placement and plant-life history traits (maximum plant height and leaf area, respectively). However, contrary to prevailing assumptions, fig placement correlated poorly with known dispersal agents and appears more likely determined by other factors, such as flowering phenology, nutrient economy, and habitat preference. Thus, plant life-history, both directly and through its influence on fig placement, appears to have played a prominent role in determining fruit traits in these figs.     

Can population genetic structure be predicted from life-history traits?

Population genetic structure is a key parameter in evolutionary biology. Earlier comparative studies have shown that genetic structure depends on species ecological attributes and life-history traits, but species phylogenetic relatedness had not been accounted for. Here we reevaluate the relationships between genetic structure and species traits in seed plants. Each species is characterized by a set of life-history and ecological features as well as by its geographic range size, its heterozygote deficit, and its genetic structure at nuclear and organelle markers to distinguish between pollen- and seed-mediated gene flow. We use both a conventional regression approach and a method that controls for phylogenetic relationships. Once phylogenetic conservatism and covariation among traits are taken into account, genetic structure is shown to be related with only a few synthetic traits, such as mating system for nuclear markers and seed dispersal mode or geographic range size for organelle markers. Along with other studies on invasiveness or rarity, our work illustrates the fact that predicting the fate of species across a broad taxonomic assemblage on the basis of simple traits is rarely possible, a testimony of the highly contingent nature of evolution.     

Incomplete datasets obscure associations between traits affecting dispersal ability and geographic range size of reef fishes in the Tropical Eastern Pacific

Dispersal is thought to be an important process determining range size, especially for species in highly spatially structured habitats, such as tropical reef fishes. Despite intensive research efforts, there is conflicting evidence about the role of dispersal in determining range size. We hypothesize that traits related to dispersal drive range sizes, but that complete and comprehensive datasets are essential for detecting relationships between species’ dispersal ability and range size. We investigate the roles of six traits affecting several stages of dispersal (adult mobility, spawning mode, pelagic larval duration (PLD), body size, aggregation behavior, and circadian activity), in explaining range size variation of reef fishes in the Tropical Eastern Pacific (TEP). All traits, except for PLD (148 species), had data for all 497 species in the region. Using a series of statistical models, we investigated which traits were associated with large range sizes, when analyzing all TEP species or only species with PLD data. Furthermore, using null models, we analyzed whether the PLD‐subset is representative of the regional species pool. Several traits affecting dispersal ability were strongly associated with range size, although these relationships could not be detected when using the PLD‐subset. Pelagic spawners (allowing for passive egg dispersal) had on average 56% larger range sizes than nonpelagic spawners. Species with medium or high adult mobility had on average a 25% or 33% larger range, respectively, than species with low mobility. Null models showed that the PLD‐subset was nonrepresentative of the regional species pool, explaining why model outcomes using the PLD‐subset differed from the ones based on the complete dataset. Our results show that in the TEP, traits affecting dispersal ability are important in explaining range size variation. Using a regionally complete dataset was crucial for detecting the theoretically expected, but so far empirically unresolved, relationship between dispersal and range size.     

Do short-lived and long-lived alien plant species differ regarding the traits associated with their success in the introduced range?

In spite of the several studies trying to identify the biological traits that are generally associated with the success of alien plant species, only a few traits are consistently shown to be important. Dividing the species into meaningful sub-categories may improve our ability to distinguish successful alien species. We asked whether there are differences between short-lived and long-lived herbaceous aliens regarding the biological traits associated with their success in their introduced range. We used the source-area approach to answer the question by performing a comparative study with those Central-European herbaceous plant species which are invasive or non-invasive aliens in the United States. Biological traits used in the analysis were extracted from European databases. The significant traits (with one exception) conferred invasiveness for only one of the two life history groups. These results outline a particular combination of competition and colonization in both groups, although achieved in different ways. Short-lived invasive species, which are supposed to be good colonizers with effective reproduction and dispersal, are backed by some kind of competitive ability conferred by height; while in the case of competitive and persistent long-lived species, the successful aliens are equipped with traits that make them better colonizers than other perennial alien species (e.g., tolerance for a wide range of anthropogenic disturbance, dispersal through water).     

Dietary generalism accelerates arrival and persistence of coral‐reef fishes in their novel ranges under climate change

Climate change is redistributing marine and terrestrial species globally. Life‐history traits mediate the ability of species to cope with novel environmental conditions, and can be used to gauge the potential redistribution of taxa facing the challenges of a changing climate. However, it is unclear whether the same traits are important across different stages of range shifts (arrival, population increase, persistence). To test which life‐history traits most mediate the process of range extension, we used a 16‐year dataset of 35 range‐extending coral‐reef fish species and quantified the importance of various traits on the arrival time (earliness) and degree of persistence (prevalence and patchiness) at higher latitudes. We show that traits predisposing species to shift their range more rapidly (large body size, broad latitudinal range, long dispersal duration) did not drive the early stages of redistribution. Instead, we found that as diet breadth increased, the initial arrival and establishment (prevalence and patchiness) of climate migrant species in temperate locations occurred earlier. While the initial incursion of range‐shifting species depends on traits associated with dispersal potential, subsequent establishment hinges more on a species’ ability to exploit novel food resources locally. These results highlight that generalist species that can best adapt to novel food sources might be most successful in a future ocean.     

Behavioral Flexibility and the Evolution of Primate Social States

Comparative approaches to the evolution of primate social behavior have typically involved two distinct lines of inquiry. One has focused on phylogenetic analyses that treat social traits as static, species-specific characteristics; the other has focused on understanding the behavioral flexibility of particular populations or species in response to local ecological or demographic variables. Here, we combine these approaches by distinguishing between constraining traits such as dispersal regimes (male, female, or bi-sexual), which are relatively invariant, and responding traits such as grouping patterns (stable, fission-fusion, sometimes fission-fusion), which can reflect rapid adjustments to current conditions. Using long-term and cross-sectional data from 29 studies of 22 species of wild primates, we confirm that dispersal regime exhibits a strong phylogenetic signal in our sample. We then show that primate species with high variation in group size and adult sex ratios exhibit variability in grouping pattern (i.e., sometimes fission-fusion) with dispersal regime constraining the grouping response. When assessing demographic variation, we found a strong positive relationship between the variability in group size over time and the number of observation years, which further illustrates the importance of long-term demographic data to interpretations of social behavior. Our approach complements other comparative efforts to understand the role of behavioral flexibility by distinguishing between constraining and responding traits, and incorporating these distinctions into analyses of social states over evolutionary and ecological time.     

Evolutionary and ecological success is decoupled in mammals

Aim

To identify which factors distinguish ecologically successful mammalian clades (i.e., clades with a large combined range size) from less successful ones.

Location

Global.

Methods

We estimated the total range sizes for each individual mammalian subfamily and used phylogenetic regressions to identify the relative importance of factors related to colonization ability (body size and niche width) and adaptability (rate of evolution of body size and rate of evolution of climatic preference) in determining these ranges. We then estimated the importance of the same factors on the variation in diversification rate within mammals.

Results

We found strong support for a link between total range size and traits related to colonization ability. In particular, we found larger total range sizes among clades containing larger bodied species and clades with wider climatic niche width, while we did not find support for any predictors related to adaptability being linked to total range size. We also found that traits related to increased range size were associated with reduced diversification rate.

Main Conclusions

Range size for mammalian clades is mainly predicted by colonization ability, suggesting that most clades are limited by dispersal rather than their ability to adapt to new environments. The most ecologically successful (i.e., most widespread) mammalian clades tend to possess traits that reduce geographical isolation among populations, but the same traits tend to decrease diversification rates. Our results unveil a decoupling between evolutionary and ecological success in mammals.

     

The influence of seasonal migration on range size in temperate North American passerines

Seasonal migration has been alternately proposed to promote geographic range size in some contexts and to constrain it in others, but it remains unclear if migratory behavior has a general effect on range size. Because migration involves movement, most hypotheses about the relationship between migration and range size invoke an influence of migration on the process of dispersal-mediated range expansion. Intuitively, a positive relationship between migratory behavior and dispersal ability could bolster range expansion among migratory species, yet some biogeographic patterns suggest that long-distance migration may instead impede range expansion, especially in the temperate zone. We conducted a comparative analysis of the relationship between migratory behavior and range size by testing the effect of migratory status, migration distance and morphological dispersal ability on breeding range size among all temperate North American passerines. Further, we assessed whether these traits affect range expansion into suitable habitat by analyzing their relationship with range filling (the proportion of climatically-suitable area occupied, or ‘filled’ by a species). Contrary to previous studies, we found migration and dispersal ability to be poor predictors of range size and range filling in North America. Rather, most variation in range size is explained by latitude. Our results suggest that migratory behavior does not affect range size within the scale of a continent, and furthermore, that temperate North American passerines’ breeding ranges are not influenced by their dispersal abilities. To better understand why migratory behavior appears to promote range size in some contexts and constrain it in others, future studies should investigate how migratory behavior affects dispersal at the individual level, as well as the relationship between the evolution of migratory behavior and the breadth of species’ climatic niches.     

Wing morphology and migration status,but not body size,habitat or Rapoport's rule predict range size in North‐American dragonflies (Odonata: Libellulidae)

Understanding why species range sizes vary is important for predicting the impact of environmental change on biodiversity. Here we use a multi‐variable approach in a phylogenetic comparative context to understand how four morphological, two ecological, and two eco‐geographical variables are associated with range size, latitudinal range and longitudinal range in 81 species of North‐American libellulid dragonflies. Our results show that: 1) migratory species and species with a more expanded basal hindwing lobe have a larger range size; 2) opposite to Rapoport's rule, latitudinal range is negatively correlated with mid‐range latitude; 3) longitudinal range is predicted by wing morphology and migration; 4) body size and larval habitat are not correlated with range size, latitudinal range or longitudinal range. These results suggest that dispersal‐related traits, such as wing shape and migratory status, are important factors in predicting the range size of libellulid dragonflies. In addition, the reverse Rapoport's rule suggests that more northern‐centred species might be more specialized than more southern‐centred species. We suggest that the variables predicting range size are likely imposed by taxon‐specific morphological, ecological, physiological and behavioural traits. Taxon‐specific knowledge is thus necessary to understand the dynamics of range sizes and is important to implement successful restoration and conservation plans of threatened species.     

A comparative analysis of dispersal syndromes in terrestrial and semi‐terrestrial animals

Dispersal, the behaviour ensuring gene flow, tends to covary with a number of morphological, ecological and behavioural traits. While species‐specific dispersal behaviours are the product of each species’ unique evolutionary history, there may be distinct interspecific patterns of covariation between dispersal and other traits (‘dispersal syndromes’) due to their shared evolutionary history or shared environments. Using dispersal, phylogeny and trait data for 15 terrestrial and semi‐terrestrial animal Orders (> 700 species), we tested for the existence and consistency of dispersal syndromes across species. At this taxonomic scale, dispersal increased linearly with body size in omnivores, but decreased above a critical length in herbivores and carnivores. Species life history and ecology significantly influenced patterns of covariation, with higher phylogenetic signal of dispersal in aerial dispersers compared with ground dwellers and stronger evidence for dispersal syndromes in aerial dispersers and ectotherms, compared with ground dwellers and endotherms. Our results highlight the complex role of dispersal in the evolution of species life‐history strategies: good dispersal ability was consistently associated with high fecundity and survival, and in aerial dispersers it was associated with early maturation. We discuss the consequences of these findings for species evolution and range shifts in response to future climate change.     

Phylogenetic effects on functional traits and life history strategies of Australian freshwater fish

Understanding the biogeographic and phylogenetic basis to interspecific differences in species’ functional traits is a central goal of evolutionary biology and community ecology. We quantify the extent of phylogenetic influence on functional traits and life‐history strategies of Australian freshwater fish to highlight intercontinental differences as a result of Australia's unique biogeographic and evolutionary history. We assembled data on life history, morphological and ecological traits from published sources for 194 Australian freshwater species. Interspecific variation among species could be described by a specialist–generalist gradient of variation in life‐history strategies associated with spawning frequency, fecundity and spawning migration. In general, Australian fish showed an affinity for life‐history strategies that maximise fitness in hydrologically unpredictable environments. We also observed differences in trait lability between and within life history, morphological and ecological traits where in general morphological and ecological traits were more labile. Our results showed that life‐history strategies are relatively evolutionarily labile and species have potentially evolved or colonised in freshwaters frequently and independently allowing them to maximise population performance in a range of environments. In addition, reproductive guild membership showed strong phylogenetic constraint indicating that evolutionary history is an important component influencing the range and distribution of reproductive strategies in extant species assemblages. For Australian freshwater fish, biogeographic and phylogenetic history contribute to broad taxonomic differences in species functional traits, while finer scale ecological processes contribute to interspecific differences in smaller taxonomic units. These results suggest that the lability or phylogenetic relatedness of different functional traits affects their suitability for testing hypothesis surrounding community level responses to environmental change.     

The phylogenetic component of food web structure and intervality

Despite the exceptional complexity formed by species and their interactions in ecological networks, such as food webs, regularities in the network structures are repeatedly demonstrated. The interactions are determined by the characteristics of a species. The characteristics are in turn determined by the species’ phylogenetic relationships, but also by factors not related to evolutionary history. Here, we test whether species’ phylogenetic relationships provides a significant proxy for food web intervality. We thereafter quantify the degree to which different species traits remain valuable predictors of food web structure after the baseline effect of species’ relatedness has been removed. We find that the phylogenetic relationships provide a significant background from which to estimate food web intervality and thereby structure. However, we also find that there is an important, non-negligible part of some traits, e.g., body size, in food webs that is not accounted for by the phylogenetic relationships. Additionally, both these relationships differ depending if a predator or a prey perspective is adopted. Clearly, species’ evolutionary history as well as traits not determined by phylogenetic relationships shapes predator-prey interactions in food webs, and the underlying evolutionary processes take place on slightly different time scales depending on the direction of predator-prey adaptations.     

Speciation in Hawaiian angiosperm lineages: cause, consequence, and mode

The biota of Hawaiian Islands is derived entirely from long distance dispersal, often followed by in situ speciation. Species descended from each colonist constitute monophyletic lineages that have diverged to varying degrees under similar spatial and temporal constraints. We partitioned the Hawaiian angiosperm flora into lineages and assessed morphological, ecological, and biogeographic characteristics to examine their relationships to variation in species number (S). Lineages with external bird dispersal (through adhesion) were significantly more species-rich than those with abiotic dispersal, but only weakly more species-rich than lineages with internal bird dispersal (involving fleshy fruits). Pollination mode and growth form (woody vs. herbaceous) had no significant effect on S, in contrast to studies of angiosperm families. S relates positively to the geographic and ecological range size of whole lineages, but negatively to local abundance and mean range sizes of constituent species. Species-rich lineages represent a large proportion of major adaptive shifts, although this appears to be an artifact of having more species. Examination of 52 sister species pairs in numerous lineages provides evidence for allopatric (including peripheral isolates) and parapatric (ecological) modes, with 15 cases of each. Although postspeciational dispersal may obscure these modes in many of the remaining cases, instances of sympatric and hybrid speciation are also discussed. Because speciation is both a consequence and a cause of ecological and biogeographic traits, speciation mode may be integral to relationships between traits. We discuss the role of speciation in shaping the regional species pool.