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.     

Ecological and evolutionary drivers of range size in Coenagrion damselflies

Geographic range size is a key ecological and evolutionary characteristic of a species, yet the causal basis of variation in range size among species remains largely unresolved. One major reason for this is that several ecological and evolutionary traits may jointly shape species' differences in range size. We here present an integrated study of the contribution of ecological (dispersal capacity, body size and latitudinal position) and macroevolutionary (species' age) traits in shaping variation in species' range size in Coenagrion damselflies. We reconstructed the phylogenetic tree of this genus to account for evolutionary history when assessing the contribution of the ecological traits and to evaluate the role of the macroevolutionary trait (species' age). The genus invaded the Nearctic twice independently from the Palearctic, yet this was not associated with the evolution of larger range sizes or dispersal capacity. Body size and species' age did not explain variation in range size. There is higher flight ability (as measured by wing aspect ratio) at higher latitudes. Species with a larger wing aspect ratio had a larger range size, also after correcting for phylogeny, suggesting a role for dispersal capacity in shaping the species' ranges. More northern species had a larger species' range, consistent with Rapoport's rule, possibly related to niche width. Our results underscore the importance of integrating macroecology and macroevolution when explaining range size variation among species.     

Experimental evidence for extreme dispersal limitation in tropical forest birds

Movements of organisms between habitat remnants can affect metapopulation structure, community assembly dynamics, gene flow and conservation strategy. In the tropical landscapes that support the majority of global biodiversity and where forest fragmentation is accelerating, there is particular urgency to understand how dispersal across habitats mediates the demography, distribution and differentiation of organisms. By employing unique dispersal challenge experiments coupled with exhaustive inventories of birds in a Panamanian lacustrine archipelago, we show that the ability to fly even short distances (< 100 m) between habitat fragments varies dramatically and consistently among species of forest birds, and that this variation correlates strongly with species' extinction histories and current distributions across the archipelago. This extreme variation in flight capability indicates that species' persistence in isolated forest remnants will be differentially mediated by their respective dispersal abilities, and that corridors connecting such fragments will be essential for the maintenance of avian diversity in fragmented tropical landscapes.     

Species traits predict island occupancy in noctuid moths

Knowing how species’ traits relate to processes that underlie occupancy patterns such as colonisation and population persistence, is important for our understanding of how species survive in fragmented and changing landscapes. We used automatic UV light-traps to sample noctuid moths on two remote islands, and compared traits of island occupants with those of a species pool from mainland southeast Sweden. Widely distributed species, generalist species, species with a long adult activity period and species active late in the summer had higher probability of occupancy on the remote islands. The results were consistent between islands. The traits of host plant specificity and species with an adult activity period during late summer remained robust and were statistically significant after controlling for any possible phylogenetic bias. This indicates that species exhibiting those traits survive better when habitat and climate changes. It is crucial to include our results in; (1) conservation planning, e.g. when devising conservation measures in fragmented landscapes; (2) for predictions of future occupancy patterns; and (3) ecosystem impact assessments, e.g. the importance of moths as pollinators, herbivores and being the functional link between parasitoids, plants, consumers and predators.     

Seed dispersal in both space and time is necessary for plant diversity maintenance in fragmented landscapes

Metacommunity theory emphasizes that seed dispersal not only limits but equally maintains plant diversity, though the latter receives little empirical attention. Discerning the temporal and spatial components of seed dispersal and understanding how their interaction shapes fragmented communities and maintains their diversity may be pivotal to further our ecological understanding of spatial and temporal seed dispersal and its implications for landscape‐scale conservation management. To investigate the relative importance of spatial and temporal seed dispersal and their roles in maintaining plant diversity, the herb layer and seed bank of grassland communities were inventoried in 77 sites across abandoned and intact rotational grazing networks in a 100 km² fragmented grassland landscape in the Stockholm archipelago (Baltic Sea, Sweden). Besides analysing alpha‐ and beta‐diversity patterns, nestedness analyses connect deterministic community changes and diversity losses with dispersal‐related life‐history traits and habitat specialization to identify the mechanism driving community changes and maintaining local diversity. The loss of rotational grazing networks caused community diversity declines via non‐random extinctions of spatially and temporally poor dispersers, particularly among grassland specialists. Temporal seed dispersal halted further community disassembly, maintaining diversity in the abandoned grazing networks. Spatial dispersal within the intact grazing networks was found to be an overriding, homogenizing agent conserving diversity in both the herb layer and seed bank. This empirical evidence establishes how spatial and temporal seed dispersal interact to maintain diversity in fragmented landscapes. Poorly connected grasslands appear limited by spatial dispersal, yet are maintained by temporal seed dispersal. In fragmented landscapes where grazing networks are rarely present, temporal rather than spatial seed dispersal may be more important in maintaining species diversity, since effective spatial dispersal may be significantly diminished. The grazing network's efficacy at boosting spatial dispersal and upholding community diversity presents a powerful management tool to conserve local and regional species diversity.     

Species vulnerability to climate change: impacts on spatial conservation priorities and species representation

Climate change may shrink and/or shift plant species ranges thereby increasing their vulnerability and requiring targeted conservation to facilitate adaptation. We quantified the vulnerability to climate change of plant species based on exposure, sensitivity and adaptive capacity and assessed the effects of including these components in complementarity‐based spatial conservation prioritisation. We modelled the vulnerability of 584 native plant species under three climate change scenarios in an 11.9 million hectare fragmented agricultural region in southern Australia. We represented exposure as species' geographical range under each climate change scenario as quantified using species distribution models. We calculated sensitivity as a function of the impact of climate change on species' geographical ranges. Using a dispersal kernel, we quantified adaptive capacity as species' ability to migrate to new geographical ranges under each climate change scenario. Using Zonation, we assessed the impact of individual components of vulnerability (exposure, sensitivity and adaptive capacity) on spatial conservation priorities and levels of species representation in priority areas under each climate change scenario. The full vulnerability framework proved an effective basis for identifying spatial conservation priorities under climate change. Including different dimensions of vulnerability had significant implications for spatial conservation priorities. Incorporating adaptive capacity increased the level of representation of most species. However, prioritising sensitive species reduced the representation of other species. We conclude that whilst taking an integrated approach to mitigating species vulnerability to climate change can ensure sensitive species are well‐represented in a conservation network, this can come at the cost of reduced representation of other species. Conservation planning decisions aimed at reducing species vulnerability to climate change need to be made in full cognisance of the sensitivity of spatial conservation priorities to individual components of vulnerability, and the trade‐offs associated with focussing on sensitive species.     

Non‐equilibrium in plant distribution models – only an issue for introduced or dispersal limited species?

Species distribution models rely on the assumption that species' distributions are at equilibrium with environmental conditions within a region – i.e. they occur in all suitable habitats. If this assumption holds, species occurrence should be predictable from measures of the environment. Introduced species may be poor candidates for distribution models due to their presumed lack of equilibrium within the landscapes they occupy, although predicting their potential distributions is often of critical importance to natural resource managers. We determined if the accuracy of species distribution models differed between 17 native and 17 introduced riparian plant species in the western United States. We also assessed if model accuracy was associated with both environmental and biological factors that can influence dispersal. We used Random Forests to model species distributions and linear regression to determine if model accuracy was associated with dispersal‐related traits. Model accuracy for introduced species was higher than that for native species. Dispersal‐related traits did not affect model accuracy or improvement, though two other traits, family affiliation and rarity on the landscape, did have an effect. Distance‐based measures of dispersal potential improved model fit equally for both native and introduced species and for species with a variety of dispersal traits, suggesting that the importance of regional propagule pressure is relatively constant across species with different dispersal opportunities. Several lines of future questioning are suggested by our results, including why introduced species may in some cases produce more accurate distribution models than native species and how species dispersal traits relate to distribution model accuracy at different spatial scales.     

Spatial synchrony in stream fish populations: influence of species traits

Spatial synchrony in population dynamics has been identified in most taxonomic groups. Numerous studies have reported varying levels of spatial synchrony among closely‐related species, suggesting that species' characteristics may play a role in determining the level of synchrony. However, few studies have attempted to relate this synchrony to the ecological characteristics and/or life‐history traits of species. Yet, as to some extent the extinction risk may be related to synchrony patterns, identifying a link between species' characteristics and spatial synchrony is crucial, and would help us to define effective conservation planning. Here, we investigated whether species attributes and temperature synchrony (i.e. a proxy of the Moran effect) account for the differences in spatial population synchrony observed in 27 stream fish species in France. After measuring and testing the level of synchrony for each species, we performed a comparative analysis to detect the phylogenetic signal of these levels, and to construct various multi‐predictor models with species traits and temperature synchrony as covariates, while taking phylogenetic relatedness into account. We then performed model averaging on selected models to take model uncertainty into account in our parameter estimates. Fifteen of the 27 species displayed a significant level of synchrony. Synchrony was weak, but highly variable between species, and was not conserved across the phylogeny. We found that some species' characteristics significantly influenced synchrony levels. Indeed, the average model indicated that species associated with greater dispersal abilities, lower thermal tolerance, and opportunistic strategy displayed a higher degree of synchrony. These findings indicate that phylogeny and spatial temperature synchrony do not provide information pertinent for explaining the variations in species' synchrony levels, whereas the dispersal abilities, the life‐history strategies and the upper thermal tolerance limits of species do appear to be quite reliable predictors of synchrony levels.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

Species traits and climate velocity explain geographic range shifts in an ocean‐warming hotspot

Species' ranges are shifting globally in response to climate warming, with substantial variability among taxa, even within regions. Relationships between range dynamics and intrinsic species traits may be particularly apparent in the ocean, where temperature more directly shapes species' distributions. Here, we test for a role of species traits and climate velocity in driving range extensions in the ocean‐warming hotspot of southeast Australia. Climate velocity explained some variation in range shifts, however, including species traits more than doubled the variation explained. Swimming ability, omnivory and latitudinal range size all had positive relationships with range extension rate, supporting hypotheses that increased dispersal capacity and ecological generalism promote extensions. We find independent support for the hypothesis that species with narrow latitudinal ranges are limited by factors other than climate. Our findings suggest that small‐ranging species are in double jeopardy, with limited ability to escape warming and greater intrinsic vulnerability to stochastic disturbances.     

Dispersal patterns of Squamata in the Mediterranean: An evolutionary perspective

Mediterranean islands have a high diversity of squamates, although they are unevenly distributed. This variability in the composition of the reptile assemblages across islands may have been influenced by differences in the colonization abilities of these species. To evaluate the dispersal capacities of squamate species, we modeled their sea routes using cost surface models. We estimated the effects of some life‐history traits and the phylogenetic signal in the characteristics of the modeled dispersal paths. We hypothesized that a significant phylogenetic signal should be present if the dispersal ability is enhanced by traits shared among evolutionarily related species. The results showed that no phylogenetic signal was present in the characteristics of the dispersal paths (i.e., in the distance traveled/bypassed sea depth). Thus, no superior island‐colonizer lineages were detected in Mediterranean Squamata. However, our analyses also revealed that small‐sized lizards were superior to other groups of squamates at dispersing over long distances on the sea.     

Climate and habitat availability determine 20th century changes in a butterfly's range margin

Evidence of anthropogenic global climate change is accumulating, but its potential consequences for insect distributions have received little attention. We use a ''climate response surface'' model to investigate distribution changes at the northern margin of the speckled wood butterfly, Pararge aegeria. We relate its current European distribution to a combination of three bioclimatic variables. We document that P. aegeria has expanded its northern margin substantially since 1940, that changes in this species'' distribution over the past 100 years are likely to have been due to climate change, and that P. aegeria will have the potential to shift its range margin substantially northwards under predicted future climate change. At current rates of expansion, this species could potentially colonize all newly available climatically suitable habitat in the UK over the next 50 years or more. However, fragmentation of habitats can affect colonization, and we show that availability of habitat may be constraining range expansion of this species at its northern margin in the UK. These lag effects may be even more pronounced in less-mobile species inhabiting more fragmented landscapes, and highlight how habitat distribution will be crucial in predicting species'' responses to future climate change.     

Pushing the Pace of Tree Species Migration

Plants and animals have responded to past climate changes by migrating with habitable environments, sometimes shifting the boundaries of their geographic ranges by tens of kilometers per year or more. Species migrating in response to present climate conditions, however, must contend with landscapes fragmented by anthropogenic disturbance. We consider this problem in the context of wind-dispersed tree species. Mechanisms of long-distance seed dispersal make these species capable of rapid migration rates. Models of species-front migration suggest that even tree species with the capacity for long-distance dispersal will be unable to keep pace with future spatial changes in temperature gradients, exclusive of habitat fragmentation effects. Here we present a numerical model that captures the salient dynamics of migration by long-distance dispersal for a generic tree species. We then use the model to explore the possible effects of assisted colonization within a fragmented landscape under a simulated tree-planting scheme. Our results suggest that an assisted-colonization program could accelerate species-front migration rates enough to match the speed of climate change, but such a program would involve an environmental-sustainability intervention at a massive scale.     

Illuminating geographical patterns in species' range shifts

Species' range shifts in response to ongoing climate change have been widely documented, but although complex spatial patterns in species' responses are expected to be common, comprehensive comparisons of species' ranges over time have undergone little investigation. Here, we outline a modeling framework based on historical and current species distribution records for disentangling different drivers (i.e. climatic vs. nonclimatic) and assessing distinct facets (i.e. colonization, extirpation, persistence, and lags) of species' range shifts. We used extensive monitoring data for stream fish assemblages throughout France to assess range shifts for 32 fish species between an initial period (1980–1992) and a contemporary one (2003–2009). Our results provide strong evidence that the responses of individual species varied considerably and exhibited complex mosaics of spatial rearrangements. By dissociating range shifts in climatically suitable and unsuitable habitats, we demonstrated that patterns in climate‐driven colonization and extirpation were less marked than those attributed to nonclimatic drivers, although this situation could rapidly shift in the near future. We also found evidence that range shifts could be related to some species' traits and that the traits involved varied depending on the facet of range shift considered. The persistence of populations in climatically unsuitable areas was greater for short‐lived species, whereas the extent of the lag behind climate change was greater for long‐lived, restricted‐range, and low‐elevation species. We further demonstrated that nonclimatic extirpations were primarily related to the size of the species' range, whereas climate‐driven extirpations were better explained by thermal tolerance. Thus, the proposed framework demonstrated its potential for markedly improving our understanding of the key processes involved in range shifting and also offers a template for informing management decisions. Conservation strategies would greatly benefit from identifying both the geographical patterns and the species' traits associated with complex modifications of species' distributions in response to global changes.     

Species interactions during diversification and community assembly in an island radiation of shrews

Background

Closely related, ecologically similar species often have adjacent distributions, suggesting competitive exclusion may contribute to the structure of some natural communities. In systems such as island archipelagos, where speciation is often tightly associated with dispersal over oceanic barriers, competitive exclusion may prevent population establishment following inter-island dispersal and subsequent cladogenesis.

Methodology/Principal Findings

Using a combination of tools, we test the hypothesis that the distributions of shrew (Crocidura) species in the Philippines are the result of competitive exclusion preventing secondary invasion of occupied islands. We first compare ecological niche models between two widespread, allopatric species and find statistical support for their ecological similarity, implying that competition for habitat between these species is possible. We then examine dispersion patterns among sympatric species and find some signal for overdispersion of body size, but not for phylogenetic branch length. Finally, we simulate the process of inter-island colonization under a stochastic model of dispersal lacking ecological forces. Results are dependent on the geographic scope and colonization probability employed. However, some combinations suggest that the number of inter-island dispersal events necessary to populate the archipelago may be much higher than the minimum number of colonization events necessary to explain current estimates of species richness and phylogenetic relationships. If our model is appropriate, these results imply that alternative factors, such as competitive exclusion, may have influenced the process of inter-island colonization and subsequent cladogenesis.

Conclusions/Significance

We interpret the combined results as providing tenuous evidence that similarity in body size may prevent co-occurrence in Philippine shrews and that competitive exclusion among ecologically similar species, rather than an inability to disperse among islands, may have limited diversification in this group, and, possibly other clades endemic to island archipelagos.     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Abundance, distribution and life histories of grassland plants: a comparative study of 81 species

1 This study examines the abundance and distribution of grassland plant species in particular relation to features affecting colonization. Seed production (inversely related to seed size) and recruitment success (positively related) affect colonization ability, suggesting that seed size can be used as a key trait.
2 Data on seed size, dispersal mode, life form, geographical range size and abundance were gathered for 81 grassland plant species in a field study area in Sweden. Seed production and plant size were estimated for 69 of these species. Analyses were performed both across species, with species treated as independent data points, and for 43 'phylogenetically independent contrasts'.
3 The cross-species analyses suggested that local abundance was related to life forms but not dispersal or plant size. Perennials were generally most abundant, as were clonal species. If abundance reflects colonization we predicted that species with intermediately sized seeds (or intermediate seed production) would be most abundant, and this was supported by the phylogenetic contrast but not by cross-species analyses. In the former analysis, a high abundance of species was significantly associated with a small seed size deviation (and seed number deviation) from the median values of these traits in the community.
4 Local abundance, seed production and seed size deviation from the community median value were positively related to geographical range size in the cross-species analysis, but no relationships were seen in the phylogenetic contrast analysis.
5 We conclude that colonization processes do have a significant influence on abundance patterns in grasslands. Seed size is a key trait for colonizing ability, and the effects of the trade-off of seed size vs. seed number must be considered. No single mechanism can be identified that influences both abundance and geographical distribution range.     

Matrix quality and disturbance frequency drive evolution of species behavior at habitat boundaries

Previous theoretical studies suggest that a species' landscape should influence the evolution of its dispersal characteristics, because landscape structure affects the costs and benefits of dispersal. However, these studies have not considered the evolution of boundary crossing, that is, the tendency of animals to cross from habitat to nonhabitat (“matrix”). It is important to understand this dispersal behavior, because of its effects on the probability of population persistence. Boundary‐crossing behavior drives the rate of interaction with matrix, and thus, it influences the rate of movement among populations and the risk of dispersal mortality. We used an individual‐based, spatially explicit model to simulate the evolution of boundary crossing in response to landscape structure. Our simulations predict higher evolved probabilities of boundary crossing in landscapes with more habitat, less fragmented habitat, higher‐quality matrix, and more frequent disturbances (i.e., fewer generations between local population extinction events). Unexpectedly, our simulations also suggest that matrix quality and disturbance frequency have much stronger effects on the evolution of boundary crossing than either habitat amount or habitat fragmentation. Our results suggest that boundary‐crossing responses are most affected by the costs of dispersal through matrix and the benefits of escaping local extinction events. Evolution of optimal behavior at habitat boundaries in response to the landscape may have implications for species in human‐altered landscapes, because this behavior may become suboptimal if the landscape changes faster than the species' evolutionary response to that change. Understanding how matrix quality and habitat disturbance drive evolution of behavior at boundaries, and how this in turn influences the extinction risk of species in human‐altered landscapes should help us identify species of conservation concern and target them for management.     

Colonization of the Galápagos Islands by plants with no specific syndromes for long‐distance dispersal: a new perspective

Since nobody has witnessed the arrival of early plant colonists on isolated islands, the actual long‐distance dispersal (hereafter LDD) has historically been a matter of speculation. In the present study, we offer a new approach that evaluates whether particular syndromes for LDD (i.e. the set of traits related to diaspore dispersal by animals, wind and sea currents) have been favourable in the natural colonization of the Galápagos Islands by plants. Dispersal syndromes of the 251 native genera (509 angiosperm species) presently acknowledged as native were carefully studied, combining data from floristic lists of the Galápagos Islands, diaspore traits, characteristics of continental relatives and our own observations. We used these genera (and occasionally infrageneric groups) as the working units to infer the number of introductions and colonists. A final number of native plants was inferred and analysed after correcting by pollen records of six species from six genera previously considered exotic (palaeobotanical correction). The number of early colonists was also corrected by incorporating information from the few (n= 12) phylogenetic studies of genera from both the Galápagos Islands and the Americas (phylogenetic correction). A total of 372 colonization events were inferred for the native flora using the latest check‐list. The proportions of native colonists grouped into five categories were: endozoochory 16.4%, epizoochory 15.7%, hydrochory 18.6%, anemochory 13.3%, and unassisted diaspores 36.0%. These figures did not vary significantly on analysing only the 99 genera that include endemic species in order to rule out any human‐mediated introductions. Irrespective of the roles of the different agents involved in LDD, diaspores with no special syndrome for LDD (unassisted diapores), such as many dry fruits, have been successful in reaching and colonizing the Galápagos archipelago. This finding leads us to suggest that unpredictable and so far unknown LDD mechanisms should be further considered in the theory of island biogeography.     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.