A process‐based model supports an association between dispersal and the prevalence of species traits in tropical reef fish assemblages

Habitat dynamics interacting with species dispersal abilities could generate gradients in species diversity and prevalence of species traits when the latter are associated with species dispersal potential. Using a process‐based model of diversification constrained by a dispersal parameter, we simulated the interplay between reef habitat dynamics during the past 140 million years and dispersal, shaping lineage diversification history and assemblage composition globally. The emerging patterns from the simulations were compared to current prevalence of species traits related to dispersal for 6315 tropical reef fish species. We found a significant spatial congruence between the prevalence of simulated low dispersal values and areas with a large proportion of species characterized by small adult body size, narrow home range mobility behaviour, pelagic larval duration shorter than 21 days and diurnal activity. Species characterized by such traits were found predominantly in the Indo‐Australian Archipelago and the Caribbean Sea. Furthermore, the frequency distribution of the dispersal parameter was found to match empirical distributions for body size, PLD and home range mobility behaviour. Also, the dispersal parameter in the simulations was associated to diversification rates and resulted in trait frequency matching empirical distributions. Overall, our findings suggest that past habitat dynamics, in conjunction with dispersal processes, influenced diversification in tropical reef fishes, which may explain the present‐day geography of species traits.     

High connectivity among habitats precludes the relationship between dispersal and range size in tropical reef fishes

The hypothesis that pelagic larval duration (PLD) influences range size in marine species with a benthic adult stage and a pelagic larval period is intuitively attractive; yet, studies conducted to date have failed to support it. A possibility for the lack of a relationship between PLD and range size may stem from the failure of past studies to account for the effect of species evolutionary ages, which may add to the dispersal capabilities of species. However, if dispersal over ecological (i.e. PLD) and across evolutionary (i.e. species evolutionary age) time scales continues to show no effect on range size then an outstanding question is why? Here we collected data on PLD, evolutionary ages and range sizes of seven tropical fish families (five families were reef‐associated and two have dwell demersal habitats) to explore the independent and interactive effects of PLD and evolutionary age on range size. Separate analyses on each family showed that even after controlling for evolutionary age, PLD has an insignificant or a very small effect on range size. To shed light on why dispersal has such a limited effect on range size, we developed a global ocean circulation model to quantify the connectivity among tropical reefs relative to the potential dispersal conferred by PLD. We found that although there are several areas of great isolation in the tropical oceans, most reef habitats are within the reach of most species given their PLDs. These results suggest that the lack of habitat isolation can potentially render the constraining effect of dispersal on range size insignificant and explain why dispersal does not relate to range size in reef fishes.     

Factors shaping the range-size frequency distribution of the endemic fish fauna of the Tropical Eastern Pacific

Aim To assess the effect of habitat fragmentation and isolation in determining the range‐size frequency distribution (RFD) of the shorefish fauna endemic to a discrete biogeographical region. Location The Tropical Eastern Pacific (TEP). Methods Habitat isolation represents the separation between oceanic islands and the continental shore of the TEP and habitat fragmentation the degree of spatial continuity of habitats (i.e. reefs, soft bottom, nearshore waters) along the continental coast of the TEP. The effects of habitat isolation and fragmentation were quantified by comparing the RFDs of (1) the species found on oceanic islands vs. the continental shore, and (2) species on the continental shore that use different habitat types. Results The RFD of the entire TEP fauna was bimodal, with peaks at both small‐ and large‐range ends of the spectrum. The small‐range peak was due almost entirely to island species and the large‐range peak due mainly to species found in both the continental shore and oceanic islands. RFDs varied among species using different habitats on the continental shore: reef‐fishes had a right‐skewed RFD, soft‐bottom species a flat RFD, and coastal‐pelagic fishes a left‐skewed RFD. Main conclusions Variation in dispersal capabilities associated with habitat isolation and fragmentation in the TEP appears to be the main mechanism contributing to differences among RFD structure, although variation in tolerances arising from the dynamic regional environment may contribute to some patterns. Because diversity patterns are strongly affected by RFD structure, it is now evident that the insular and continental components of a fauna should be treated separately when analysing such patterns. Furthermore, contrasts in RFD structure among species using different habitats demonstrate that a full understanding of the causes of diversity patterns requires analyses of complete regional faunas in relation to regional geography.     

Relationship between pelagic larval duration and abundance of tropical fishes on temperate coasts of Japan

The influence of pelagic larval duration (PLD) and egg type dispersal capabilities of 35 demersal and pelagic-spawning tropical fish species is examined in relation to their abundance on the temperate coasts of Japan. The PLDs of pelagic spawners were significantly longer than those of demersal spawners, and a high occurrence of pelagic spawners on the temperate coasts suggests that these fishes are more easily transported to temperate coasts than demersal spawners. For demersal spawners, the common species on the temperate coasts had significantly longer PLDs than the rare species; this suggests that PLD is a major factor influencing the distribution patterns of tropical demersal spawners on temperate coasts. Moreover, a negative correlation between PLD and the abundance of some species of pelagic and demersal spawners suggests the presence of reproductively active fishes in northern subtropical and even in temperate waters.     

The relationship between pelagic larval duration and range size in tropical reef fishes: a synthetic analysis

We address the conflict in earlier results regarding the relationship between dispersal potential and range size. We examine all published pelagic larval duration data for tropical reef fishes. Larval duration is a convenient surrogate for dispersal potential in marine species that are sedentary as adults and that therefore only experience significant dispersal during their larval phase. Such extensive quantitative dispersal data are only available for fishes and thus we use a unique dataset to examine the relationship between dispersal potential and range size. We find that dispersal potential and range size are positively correlated only in the largest ocean basin, the Indo-Pacific, and that this pattern is driven primarily by the spatial distribution of habitat and dispersal barriers. Furthermore, the relationship strengthens at higher taxonomic levels, suggesting an evolutionary mechanism. We document a negative correlation between species richness and larval duration at the family level in the Indo-Pacific, implying that speciation rate may be negatively related to dispersal potential. If increased speciation rate within a taxonomic group results in smaller range sizes within that group, speciation rate could regulate the association between range size and dispersal potential.     

Phylogenetic perspectives on reef fish functional traits

Functional traits have been fundamental to the evolution and diversification of entire fish lineages on coral reefs. Yet their relationship with the processes promoting speciation, extinction and the filtering of local species pools remains unclear. We review the current literature exploring the evolution of diet, body size, water column use and geographic range size in reef‐associated fishes. Using published and new data, we mapped functional traits on to published phylogenetic trees to uncover evolutionary patterns that have led to the current functional diversity of fishes on coral reefs. When examining reconstructed patterns for diet and feeding mode, we found examples of independent transitions to planktivory across different reef fish families. Such transitions and associated morphological alterations may represent cases in which ecological opportunity for the exploitation of different resources drives speciation and adaptation. In terms of body size, reconstructions showed that both large and small sizes appear multiple times within clades of mid‐sized fishes and that extreme body sizes have arisen mostly in the last 10 million years (Myr). The reconstruction of range size revealed many cases of disparate range sizes among sister species. Such range size disparity highlights potential vicariant processes through isolation in peripheral locations. When accounting for peripheral speciation processes in sister pairs, we found a significant relationship between labrid range size and lineage age. The diversity and evolution of traits within lineages is influenced by trait–environment interactions as well as by species and trait–trait interactions, where the presence of a given trait may trigger the development of related traits or behaviours. Our effort to assess the evolution of functional diversity across reef fish clades adds to the burgeoning research focusing on the evolutionary and ecological roles of functional traits. We argue that the combination of a phylogenetic and a functional approach will improve the understanding of the mechanisms of species assembly in extraordinarily rich coral reef communities.     

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.     

A phylogenetic analysis of egg size,clutch size,spawning mode,adult body size,and latitude in reef fishes

Theoretical treatments of egg size in fishes suggest that constraints on reproductive output should create trade-offs between the size and number of eggs produced per spawn. For marine reef fishes, the observation of distinct reproductive care strategies (demersal guarding, egg scattering, and pelagic spawning) has additionally prompted speculation that these strategies reflect alternative fitness optima with selection on egg size differing by reproductive mode and perhaps latitude. Here, we aggregate data from 278 reef fish species and test whether clutch size, reproductive care, adult body size, and latitudinal bands (i.e., tropical, subtropical, and temperate) predict egg size, using a statistically unified framework that accounts for phylogenetic correlations among traits. We find no inverse relationship between species egg size and clutch size, but rather that egg size differs by reproductive mode (mean volume for demersal eggs = 1.22 mm³, scattered eggs = 0.18 mm³, pelagic eggs = 0.52 mm³) and that clutch size is strongly correlated with adult body size. Larger eggs were found in temperate species compared with tropical species in both demersal guarders and pelagic spawners, but this difference was not strong when accounting for phylogenetic correlations, suggesting that differences in species composition underlies regional differences in egg size. In summary, demersal guarders are generally small fishes with small clutch sizes that produce large eggs. Pelagic spawners and egg scatterers are variable in adult and clutch size. Although pelagic spawned eggs are variable in size, those of scatterers are consistently small.     

‘Stick with your own kind,or hang with the locals?’ Implications of shoaling strategy for tropical reef fish on a range‐expansion frontline

Range shifts of tropical marine species to temperate latitudes are predicted to increase as a consequence of climate change. To date, the research focus on climate‐mediated range shifts has been predominately dealt with the physiological capacity of tropical species to cope with the thermal challenges imposed by temperate latitudes. Behavioural traits of individuals in the novel temperate environment have not previously been investigated, however, they are also likely to play a key role in determining the establishment success of individual species at the range‐expansion forefront. The aim of this study was to investigate the effect of shoaling strategy on the performance of juvenile tropical reef fishes that recruit annually to temperate waters off the south east coast of Australia. Specifically, we compared body‐size distributions and the seasonal decline in abundance through time of juvenile tropical fishes that shoaled with native temperate species (‘mixed’ shoals) to those that shoaled only with conspecifics (as would be the case in their tropical range). We found that shoaling with temperate native species benefitted juvenile tropical reef fishes, with individuals in ‘mixed’ shoals attaining larger body‐sizes over the season than those in ‘tropical‐only’ shoals. This benefit in terms of population body‐size distributions was accompanied by greater social cohesion of ‘mixed’ shoals across the season. Our results highlight the impact that sociality and behavioural plasticity are likely to play in determining the impact on native fish communities of climate‐induced range expansion of coral reef fishes.     

Higher investment in flight morphology does not trade off with fecundity estimates in a poleward range‐expanding damselfly

1. Evolutionary increases in dispersal‐related traits are frequently documented during range expansions. Investment in flight‐related traits is energetically costly and a trade‐off with fecundity may be expected during range expansion. 2. However, in contrast to wing‐dimorphic species, this trade‐off is not general in wing‐monomorphic species. In the absence of a dispersal‐‐fecundity trade‐off, an increased investment in clutch size at the expansion front is expected possibly at a cost of reduced offspring size. 3. The study evaluated investment in female flight morphology and fecundity‐related traits (clutch size, hatchling size) and potential trade‐offs among these traits in replicated populations of the poleward range‐expanding damselfly Coenagrion scitulum. 4. Females at the expansion front had a higher relative thorax length, indicating an increased investment in flight; this can be explained by spatial sorting of dispersal ability or in situ natural selection at the expansion front. Edge females produced larger hatchlings, however, this pattern was totally driven by the population‐specific thermal larval regimes and could not be attributed to the range expansion per se. By contrast, clutch sizes did not differ between core and edge populations. There was no signal of a dispersal–fecundity trade‐off either for a trade‐off between clutch size and hatchling size. 5. These results indicate that evolution of a higher dispersal ability at the expansion front of C. scitulum does not trade off with investment in fecundity, hence a dispersal–fecundity trade‐off is unlikely to slow down range expansion of this species.     

Do early life history characteristics determine species ranges and dispersal of coral reef fishes in the Indo‐Pacific?

The Indo‐Pacific consists of extensive continuous coastlines and many island groups of varying sizes and isolation. The species ranges of coral reef fishes vary enormously from Indo‐Pacific wide to highly endemic. There is also great variation in the early life history characteristics of coral reef fishes ( e.g . pelagic larval durations, spawning strategies and swimming abilities). We use individual‐based models (IBMs) to simulate the dispersal of coral reef fishes in the Indo‐Pacific. The development of dispersal strategies is explored based on ecological and geographical constraints. Simulations are presented for climatic and anthropogenically‐induced events.     

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.     

Isolation drives taxonomic and functional nestedness in tropical reef fish faunas

Taxonomic nestedness, the degree to which the taxonomic composition of species‐poor assemblages represents a subset of richer sites, commonly occurs in habitat fragments and islands differing in size and isolation from a source pool. However, species are not ecologically equivalent and the extent to which nestedness is observed in terms of functional trait composition of assemblages still remains poorly known. Here, using an extensive database on the functional traits and the distributions of 6316 tropical reef fish species across 169 sites, we assessed the levels of taxonomical vs functional nestedness of reef fish assemblages at the global scale. Functional nestedness was considerably more common than taxonomic nestedness, and generally associated with geographical isolation, where nested subsets are gradually more isolated from surrounding reef areas and from the center of biodiversity. Because a nested pattern in functional composition implies that certain combinations of traits may be represented by few species, we identified these groups of low redundancy that include large herbivore‐detritivores and omnivores, small piscivores, and macro‐algal herbivores. The identified patterns of nestedness may be an outcome of the interaction between species dispersal capabilities, resource requirements, and gradients of isolation among habitats. The importance of isolation in generating the observed pattern of functional nestedness within biogeographic regions may indicate that disturbance in depauperate and isolated sites can have disproportionate effects on the functional structure of their reef fish assemblages.     

Global patterns in marine dispersal estimates: the influence of geography, taxonomic category and life history

We examine estimates of dispersal in a broad range of marine species through an analysis of published values, and evaluate how well these values represent global patterns through a comparison with correlates of dispersal. Our analysis indicates a historical focus in dispersal studies on low-dispersal/low-latitude species, and we hypothesize that these studies are not generally applicable and representative of global patterns. Large-scale patterns in dispersal were examined using a database of correlates of dispersal such as planktonic larval duration (PLD, 318 species) and genetic differentiation (FST, 246 species). We observed significant differences in FST (p<0.001) and PLD (p<0.001) between taxonomic groups (e.g. fishes, cnidarians, etc.). Within marine fishes (more than 50% of datasets), the prevalence of demersal eggs was negatively associated with PLD (R2=0.80, p<0.001) and positively associated with genetic structure (R2=0.74, p<0.001). Furthermore, dispersal within marine fishes (i.e. PLD and FST) increased with latitude, adult body size and water depth. Of these variables, multiple regression identified latitude and body size as persistent predictors across taxonomic levels. These global patterns of dispersal represent a first step towards understanding and predicting species-level and regional differences in dispersal, and will be improved as more comprehensive data become available.     

Body size and spatial scales in avian response to landscapes: a meta‐analysis

Response of species to their environment is dependent on the scales that individuals interact with landscapes. Several life‐history traits have been suggested to influence scales of response to landscapes, but little empirical work has addressed this issue, limiting our ability to predict the relevant scale(s) at which species respond to their environment. Body size is frequently hypothesized to be relevant for predicting the effects of landscape structure on species given correlations between body size and various measures of mobility or perceptual range, yet evidence of this relationship remains equivocal. We conducted a meta‐analysis of 22 studies to address the question of whether body size is related to the characteristic scale at which bird species respond to landscape structure. On the basis of correlation coefficients and linear mixed models, we show that body size is positively related to the characteristic scale of response to landscapes. The strength of this relationship is less than body size – dispersal allometries, suggesting that response to landscapes may be influenced, in part, by factors other than dispersal (e.g. territory size). However, the strength of the observed body size – scale relationship may have been affected by several factors, including limitations in the range of body sizes and landscape extents tested. Our findings suggest that life‐history traits mediate aspects of the scales at which species respond to the landscape and contribute to developing a more predictive framework for investigating scalar relationships in nature.     

Range size: disentangling current traits and phylogenetic and biogeographic factors

The range size of a species can be determined by its current traits and by phylogenetic and biogeographic factors. However, only rarely have these factors been studied in combination. We use data on the geographic range sizes of all 26 Sylvia warblers to explicitly test whether range size was determined by current species-specific traits (e.g., body size, dispersal ability), phylogenetic factors (e.g., age of the lineage), or environmental, biogeographic factors (e.g., latitudinal position of the range). The results demonstrated that current traits and phylogenetic and biogeographic factors were interrelated. While a number of factors were significant in simple regression analyses, only one factor determined range size in the multiple regression analyses--dispersal ability. Species with better dispersal ability had larger ranges than species with poorer dispersal ability. Apparent increases of range size with latitude or with the age of the species resulted from correlations with dispersal ability. While the most significant factor that influences the range size of a group of species might differ from one group to the next, these results demonstrate that studies that focus only on a single, for example, phylogenetic, factor might yield misleading results.     

Correlates of range size and dispersal ability: a comparative analysis of sphingid moths from the Indo-Australian tropics

Aim  We tested various species-level traits for their potential to explain species' range sizes and dispersal abilities.
Location  Southeast Asia and Malay Archipelago.
Methods  We used published maps of geographical distribution estimates for sphingid moths to calculate range areas and classify species according to their dispersion across (present or historical) water straits in the Malay Archipelago. We tested forewing length (FWL), wing load (thorax width/FWL), presence or absence of a functional proboscis (i.e. adult feeding), larval diet breadth and larval diet composition for univariate correlations with range size and inter-island dispersion. We used multivariate, phylogenetically controlled models to test for independent effects of parameters.
Results  Range size correlated strongly with larval diet breadth, a result that was also confirmed in the multivariate model. Adult feeding had a significant impact on range sizes only within the multivariate model, but not in the univariate correlation. Dispersal class also correlated with larval diet breadth, but was additionally influenced by forewing length, wing load and larval diet composition. A univariate effect of adult feeding became non-significant in the multivariate, phylogenetically controlled model.
Main conclusions  Larval diet breadth is the best predictor of range size as well as inter-island dispersion, confirming the importance of niche breadth on the geographic ranges of species. A number of other factors are shown to have additional impact on predictions of range size or inter-island dispersal ability. Our analyses cannot determine the causal mechanisms of these correlations, but may stimulate further research on the adaptive significance of traits affecting range size and dispersal in this system.     

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.