The experimental evolution of specialists,generalists, and the maintenance of diversity

Environmental heterogeneity may be a general explanation for both the quantity of genetic variation in populations and the ecological niche width of individuals. To evaluate this hypothesis, I review the literature on selection experiments in heterogeneous environments. The niche width usually – but not invariably – evolves to match the amount of environmental variation, specialists evolving in homogeneous environments and generalists evolving in heterogeneous environments. The genetics of niche width are more complex than has previously been recognized, particularly with respect to the magnitude of costs of adaptation and the putative constraints on the evolution of generalists. Genetic variation in fitness is more readily maintained in heterogeneous environments than in homogeneous environments and this diversity is often stably maintained through negative frequency‐dependent selection. Moreover environmental heterogeneity appears to be a plausible mechanism for at least two well‐known patterns of species diversity at the landscape scale. I conclude that environmental heterogeneity is a plausible and possibly very general explanation for diversity across the range of scales from individuals to landscapes.     

An integrated framework to improve the concept of resource specialisation

Resource specialisation, although a fundamental component of ecological theory, is employed in disparate ways. Most definitions derive from simple counts of resource species. We build on recent advances in ecophylogenetics and null model analysis to propose a concept of specialisation that comprises affinities among resources as well as their co‐occurrence with consumers. In the distance‐based specialisation index (DSI), specialisation is measured as relatedness (phylogenetic or otherwise) of resources, scaled by the null expectation of random use of locally available resources. Thus, specialists use significantly clustered sets of resources, whereas generalists use over‐dispersed resources. Intermediate species are classed as indiscriminate consumers. The effectiveness of this approach was assessed with differentially restricted null models, applied to a data set of 168 herbivorous insect species and their hosts. Incorporation of plant relatedness and relative abundance greatly improved specialisation measures compared to taxon counts or simpler null models, which overestimate the fraction of specialists, a problem compounded by insufficient sampling effort. This framework disambiguates the concept of specialisation with an explicit measure applicable to any mode of affinity among resource classes, and is also linked to ecological and evolutionary processes. This will enable a more rigorous deployment of ecological specialisation in empirical and theoretical studies.     

Local adaptation,dispersal evolution,and the spatial eco‐evolutionary dynamics of invasion

Local adaptation and dispersal evolution are key evolutionary processes shaping the invasion dynamics of populations colonizing new environments. Yet their interaction is largely unresolved. Using a single‐species population model along a one‐dimensional environmental gradient, we show how local competition and dispersal jointly shape the eco‐evolutionary dynamics and speed of invasion. From a focal introduction site, the generic pattern predicted by our model features a temporal transition from wave‐like to pulsed invasion. Each regime is driven primarily by local adaptation, while the transition is caused by eco‐evolutionary feedbacks mediated by dispersal. The interaction range and cost of dispersal arise as key factors of the duration and speed of each phase. Our results demonstrate that spatial eco‐evolutionary feedbacks along environmental gradients can drive strong temporal variation in the rate and structure of population spread, and must be considered to better understand and forecast invasion rates and range dynamics.     

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

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

Variation in the degree of specialization can maintain local diversity in model communities

We hypothesize that the continuum between generalist and specialist adaptations is an important general trade-off axis in the maintenance of local diversity, and we explore this idea with a simple model in which there are patch types to which species arrive as propagules and compete. Each patch type is defined by a competitive ranking of all species. A highly specialist species is the top competitor in one patch type but has a relatively low average ranking across different patch types, while a generalist species has a high average rank across patch types but is not the top competitor in any patch type. We use random dispersal and vary the fecundity of all species together to vary total propagule density and therefore recruitment limitation and density-dependent mortality. When fecundity is very high, each patch becomes occupied by its specialist species and generalists go extinct, so the number of species at equilibrium is equal to the number of patch types. If fecundity is very low, generalists dominate and specialists go extinct. There is a range of fecundity levels in which specialists, generalists, and intermediates coexist, and the number of species is substantially greater than the number of patch types. While coexistence of specialists and generalists has been considered a problem in evolutionary ecology, our results suggest to the contrary that this trade-off contributes to the maintenance of local diversity.     

Multitrophic diversity effects depend on consumer specialization and species‐specific growth and grazing rates

Ecosystem functioning is affected by horizontal (within trophic groups) and vertical (across trophic levels) biodiversity. Theory predicts that the effects of vertical biodiversity depend on consumer specialization. In a microcosm experiment, we investigated ciliate consumer diversity and specialization effects on algal prey biovolume, evenness and composition, and on ciliate biovolume production. The experimental data was complemented by a process‐based model further analyzing the ecological mechanisms behind the observed diversity effects. Overall, increasing consumer diversity had no significant effect on prey biovolume or evenness. However, consumer specialization affected the prey community. Specialist consumers showed a stronger negative impact on prey biovolume and evenness than generalists. The model confirmed that this pattern was mainly driven by a single specialist with a high per capita grazing rate, consuming the two most productive prey species. When these were suppressed, the prey assemblage became dominated by a less productive species, consequently decreasing prey biovolume and evenness. Consumer diversity increased consumer biovolume, which was stronger for generalists than for specialists and highest in mixed combinations, indicating that consumer functional diversity, i.e. more diverse feeding strategies, increased resource use efficiency. Overall, our results indicate that consumer diversity effects on prey and consumers strongly depend on species‐specific growth and grazing rates, which may be at least equally important as consumer specialization in driving consumer diversity effects across trophic levels. Synthesis In a microcosm experiment, we investigated multitrophic consumer diversity and specialization effects using ciliate consumers and microalgal prey. Consumer diversity increased consumer biovolume, which was highest in combinations containing both generalists and specialists. Specialist consumers showed a stronger negative effect on prey biovolume and evenness than generalists. These experimental data were supported by a process‐based model, indicating that the large effect of the specialists was based on high per capita grazing rate on the two most productive prey species. Species‐specific traits such as growth and grazing rates were equally important for multitrophic diversity effects than consumer specialization.     

Ecological and evolutionary drivers of the elevational gradient of diversity

Ecological, evolutionary, spatial and neutral theories make distinct predictions and provide distinct explanations for the mechanisms that control the relationship between diversity and the environment. Here, we test predictions of the elevational diversity gradient focusing on Iberian bumblebees, grasshoppers and birds. Processes mediated by local abundance and regional diversity concur in explaining local diversity patterns along elevation. Effects expressed through variation in abundance were similar among taxa and point to the overriding role of a physical factor, temperature. This determines how energy is distributed among individuals and ultimately how the resulting pattern of abundance affects species incidence. Effects expressed through variation in regional species pools depended instead on taxon‐specific evolutionary history, and lead to diverging responses under similar environmental pressures. Local filters and regional variation also explain functional diversity gradients, in line with results from species richness that indicate an (local) ecological and (regional) historical unfolding of diversity–elevation relationships.     

No need for speed: slow development of fungi in extreme environments

Microbial growth under extreme conditions is often slow. This is partly because large amounts of energy are diverted into cellular mechanisms that allow survival under hostile conditions. Because this challenge is universal and diversity in extreme environments is low compared to non-extreme environments, slow-growing microorganisms are not overgrown by other species. In some cases, especially when nutrients are scarce, slow growth was even shown to increase stress tolerance. And in at least some species of extremotolerant and extremophilic fungi, growth rate appears to be coupled with their very unusual morphologies, which in turn may be an adaptation to extreme conditions. However, there is more than one strategy of survival in extreme environments. Fungi that thrive in extremes can be divided into (i) ubiquitous and polyextremotolerant generalists and (ii) rarely isolated specialists with narrow ecological amplitudes. While generalists can compete with mesophilic species, specialists cannot. When adapting to extreme conditions, the risk of an evolutionary trade-off in the form of reduced fitness under mesophilic conditions may limit the maximum stress tolerance achievable by polyextremotolerant generalists. At the same time, specialists are rarely found in mesophilic environments, which allows them to evolve to ever greater extremotolerance, since a reduction of mesophilic fitness is likely to have little impact on their evolutionary success.     

Evolution in a Changing Environment

We propose a simple model for genetic adaptation to a changing environment, describing a fitness landscape characterized by two maxima. One is associated with “specialist” individuals that are adapted to the environment; this maximum moves over time as the environment changes. The other maximum is static, and represents “generalist” individuals not affected by environmental changes. The rest of the landscape is occupied by “maladapted” individuals. Our analysis considers the evolution of these three subpopulations. Our main result is that, in presence of a sufficiently stable environmental feature, as in the case of an unchanging aspect of a physical habitat, specialists can dominate the population. By contrast, rapidly changing environmental features, such as language or cultural habits, are a moving target for the genes; here, generalists dominate, because the best evolutionary strategy is to adopt neutral alleles not specialized for any specific environment. The model we propose is based on simple assumptions about evolutionary dynamics and describes all possible scenarios in a non-trivial phase diagram. The approach provides a general framework to address such fundamental issues as the Baldwin effect, the biological basis for language, or the ecological consequences of a rapid climate change.     

Insights on the functional composition of specialist and generalist birds throughout continuous and fragmented forests

A decline in species number often occurs after forest fragmentation and habitat loss, which usually results in the loss of ecological functions and a reduction in functional diversity in the forest fragments. However, it is uncertain whether these lost ecological functions are consistently maintained throughout continuous forests, and so the importance of these functions in continuous forests remains unknown. Point counts were used to assess both the taxonomic and functional diversity of specialist and generalist birds from sampling in a continuous primary forest compared with forest fragments in order to investigate the responses of these groups to forest fragmentation. We also measured alpha and beta diversity. The responses of specialists and generalists were similar when we assessed all bird species but were different when only passerines were considered. When examining passerines we found lower total taxonomic beta diversity for specialists than for generalists in the continuous forest, while taxonomic beta diversity was higher in the fragmented forest and similar between bird groups. However, total functional beta‐diversity values indicated clearly higher trait regularity in continuous forest for specialists and higher trait regularity in fragments for generalists. Specialists showed significantly higher functional alpha diversity in comparison with generalists in the continuous forest, while both groups showed similar values in fragments. In passerines, species richness and alpha functional diversity of both specialist and generalist were explained by forest connectivity; but, only fragment size explained those parameters for specialist passerines. We suggest that considering subsets of the community with high similarity among species, as passerines, provides a better tool for understanding responses to forest fragmentation. Due to the regularity of specialists in continuous forest, their lost could highly affect functionality in forest fragments.     

Microbial generalist or specialist: Intraspecific variation and dormancy potential matter

Microbial generalists and specialists coexist in the soil environment while having distinctive impacts on microbial community dynamics. In microbial ecology, the underlying mechanisms as to why a species is a generalist or a specialist remain ambiguous. Herein, we collected soils across a national scale and identified bacterial generalists and specialists according to niche breadth at the species level (OTU level), and the single-nucleotide differences in each species were measured to investigate intraspecific variation (at zero-radius OTU level). Compared with that of the specialists, the intraspecific variation of the generalists was much higher, which ensured their wider niche breadth and lower variability. The higher asynchrony and different niche preferences of conspecific individuals and the higher dormancy potential within the generalists further contributed to their stability in varying environments. Besides, generalists were less controlled by environmental filtering, which was indicated by the stronger signature of stochastic processes in their assembly, and had higher diversification and transition rates that allowed them to adapt to environmental changes to a greater extent than specialists. Overall, this study provides a new comprehensive understanding of the rules of assembly and the evolutionary roles of bacterial generalists and specialists. It also highlights the importance of intraspecific variation and the dormancy potential in the stability of species.     

Host and habitat specialization of avian malaria in Africa

Studies of both vertebrates and invertebrates have suggested that specialists, as compared to generalists, are likely to suffer more serious declines in response to environmental change. Less is known about the effects of environmental conditions on specialist versus generalist parasites. Here, we study the evolutionary strategies of malaria parasites (Plasmodium spp.) among different bird host communities. We determined the parasite diversity and prevalence of avian malaria in three bird communities in the lowland forests in Cameroon, highland forests in East Africa and fynbos in South Africa. We calculated the host specificity index of parasites to examine the range of hosts parasitized as a function of the habitat and investigated the phylogenetic relationships of parasites. First, using phylogenetic and ancestral reconstruction analyses, we found an evolutionary tendency for generalist malaria parasites to become specialists. The transition rate at which generalists become specialists was nearly four times as great as the rate at which specialists become generalists. We also found more specialist parasites and greater parasite diversity in African lowland rainforests as compared to the more climatically variable habitats of the fynbos and the highland forests. Thus, with environmental changes, we anticipate a change in the distribution of both specialist and generalist parasites with potential impacts on bird communities.     

Parallelism in adaptive radiations of experimental Escherichia coli populations

Adaptive radiations are major contributors to species diversity. Although the underlying mechanisms of adaptive radiations, specialization and trade‐offs, are relatively well understood, the tempo and repeatability of adaptive radiations remain elusive. Ecological specialization can occur through the expansion into novel niches or through partitioning of an existing niche. To test how the mode of resource specialization affects the tempo and repeatability of adaptive radiations, we selected replicate bacterial populations in environments that promoted the evolution of diversity either through niche expansion or through niche partitioning, and in a third low‐quality single‐resource environment, in which diversity was not expected to evolve. Colony size diversity evolved equally fast in environments that provided ecological opportunities regardless of the mode of resource specialization. In the low‐quality environments, diversity did not consistently evolve. We observed the largest fitness improvement in the low‐quality environment and the smallest the glucose‐limited environment. We did not observe a change in the rate of evolutionary change in either trait or environment, suggesting that the pool of beneficial mutations was not exhausted. Overall, the mode of resource specialization did not affect the tempo or repeatability of adaptive radiations. These results demonstrate the limitations of eco‐evolutionary feedbacks to affect evolutionary outcomes.     

The predictability of phytophagous insect communities: host specialists as habitat specialists

The difficulties specialized phytophagous insects face in finding habitats with an appropriate host should constrain their dispersal. Within the concept of metacommunities, this leads to the prediction that host-plant specialists should sort into local assemblages according to the local environmental conditions, i.e. habitat conditions, whereas assemblages of host-plant generalists should depend also on regional processes. Our study aimed at ranking the importance of local environmental factors and species composition of the vegetation for predicting the species composition of phytophagous moth assemblages with either a narrow or a broad host range. Our database consists of 351,506 specimens representing 820 species of nocturnal Macrolepidoptera sampled between 1980 and 2006 using light traps in 96 strict forest reserves in southern Germany. Species were grouped as specialists or generalists according to the food plants of the larvae; specialists use host plants belonging to one genus. We used predictive canonical correspondence and co-correspondence analyses to rank the importance of local environmental factors, the species composition of the vegetation and the role of host plants for predicting the species composition of host-plant specialists and generalists. The cross-validatory fit for predicting the species composition of phytophagous moths was higher for host-plant specialists than for host-plant generalists using environmental factors as well as the composition of the vegetation. As expected for host-plant specialists, the species composition of the vegetation was a better predictor of the composition of these assemblages than the environmental variables. But surprisingly, this difference for specialized insects was not due to the occurrence of their host plants. Overall, our study supports the idea that owing to evolutionary constraints in finding a host, host-plant specialists and host-plant generalists follow two different models of metacommunities: the species-sorting and the mass-effect model.     

Taming fitness: Organism‐environment interdependencies preclude long‐term fitness forecasting

Fitness is a central but notoriously vexing concept in evolutionary biology. The propensity interpretation of fitness is often regarded as the least problematic account for fitness. It ties an individual's fitness to a probabilistic capacity to produce offspring. Fitness has a clear causal role in evolutionary dynamics under this account. Nevertheless, the propensity interpretation faces its share of problems. We discuss three of these. We first show that a single scalar value is an incomplete summary of a propensity. Second, we argue that the widespread method of “abstracting away” environmental idiosyncrasies by averaging over reproductive output in different environments is not a valid approach when environmental changes are irreversible. Third, we point out that expanding the range of applicability for fitness measures by averaging over more environments or longer time scales (so as to ensure environmental reversibility) reduces one's ability to distinguish selectively relevant differences among individuals because of mutation and eco‐evolutionary feedbacks. This series of problems leads us to conclude that a general value of fitness that is both explanatory and predictive cannot be attained. We advocate for the use of propensity‐compatible methods, such as adaptive dynamics, which can accommodate these difficulties.     

Stasis of functionally versatile specialists

A classic hypothesis posits that lineages exhibiting long-term stasis are broadly adapted generalists that remain well-adapted despite environmental change. However, lacking constraints that steepen adaptive peaks and stabilize the optimum, generalists’ phenotypes might drift around a broad adaptive plateau. We propose that stasis would be likely for morphological specialists that behave as ecological generalists much of the time because specialists’ functional constraints stabilize the optimum, but those with a broad niche, such as generalists, can persist despite environmental change. Tree squirrels (Callosciurinae and Sciurini) exemplify ecologically versatile specialists, being extreme in adaptations for forceful biting that expand rather than limit niche breadth. Here, we examine the structure of disparity and the evolutionary dynamics of their trophic morphology (mandible size and shape) to determine if they exhibit stasis. In both lineages, a few dietary specialists disproportionately account for disparity; excluding them, we find compelling evidence for stasis of jaw shape but not size. The primary optima of these lineages diverge little, if at all over approximately 30 million years. Once their trophic apparatus was assembled, their morphological specialization steepened the slopes of their adaptive peak and constrained the position of the optima without limiting niche breadth.     

Phylogenetic and ecological structure of Mediterranean caddisfly communities at various spatio‐temporal scales

Aim The evolutionary processes structuring the composition of communities remain unclear due to the complexity of factors active at various spatial and temporal scales. Here, we conducted ecological and evolutionary analyses of communities of caddisflies in the genus Hydropsyche (Insecta: Trichoptera) composed of ecomorphologically differentiated species. Location River ecosystems in the Iberian Peninsula and northern Morocco. Methods Nineteen environmental variables were assessed at 180 local study sites and species presence/absence at these sites was used to determine their ecological niche. The evolutionary framework for all 19 species of Hydropsyche encountered was generated by phylogenetic analysis of the mitochondrial cytochrome c oxidase subunit I gene and three nuclear genes: wingless, elongation factor 1‐alpha and 28S RNA. The phylogenetic tree was used: (1) to assess evolutionary niche conservatism by ecological trait correlation with the tree; and (2) to analyse the phylogenetic relatedness of community member species, at three spatial scales (local stream reaches, drainage basins, biogeographical regions). Results Ecological measurements grouped most species into either headwater, mid‐stream or lowland specialists, and traits presumably relevant to river zonation were found to be phylogenetically conservative. Species assemblages at local stream reaches were mostly mono‐ or dispecific. Species diversity increased at larger spatial scales, by adding species with non‐overlapping ecological niches at the level of river basins and by turnover of anciently differentiated lineages at the level of biogeographical regions. This indicates the effects of competition and niche filtering on community structure locally, and ancient ecological diversification and allopatric speciation, respectively, in building up the species pool at basin and biogeographical scales. Main conclusions The study demonstrates the importance of scale (grain size) in studying what determines community composition. Current ecological factors (i.e. competitive exclusion) in Hydropsyche were evident only when studying narrow local sites, while studies of assemblages at larger spatial scales instead demonstrated the roles of ecological niche differentiation, phylogenetic history of trait diversification and allopatric speciation. Increasing the grain size of investigation reveals different portions of correlated spatial and evolutionary processes.     

Eco‐evolutionary feedbacks between private and public goods: evidence from toxic algal blooms

The importance of ‘eco‐evolutionary feedbacks’ in natural systems is currently unclear. Here, we advance a general hypothesis for a particular class of eco‐evolutionary feedbacks with potentially large, long‐lasting impacts in complex ecosystems. These eco‐evolutionary feedbacks involve traits that mediate important interactions with abiotic and biotic features of the environment and a self‐driven reversal of selection as the ecological impact of the trait varies between private (small scale) and public (large scale). Toxic algal blooms may involve such eco‐evolutionary feedbacks due to the emergence of public goods. We review evidence that toxin production by microalgae may yield ‘privatised’ benefits for individual cells or colonies under pre‐ and early‐bloom conditions; however, the large‐scale, ecosystem‐level effects of toxicity associated with bloom states yield benefits that are necessarily ‘public’. Theory predicts that the replacement of private with public goods may reverse selection for toxicity in the absence of higher level selection. Indeed, blooms often harbor significant genetic and functional diversity: bloom populations may undergo genetic differentiation over a scale of days, and even genetically similar lineages may vary widely in toxic potential. Intriguingly, these observations find parallels in terrestrial communities, suggesting that toxic blooms may serve as useful models for eco‐evolutionary dynamics in nature. Eco‐evolutionary feedbacks involving the emergence of a public good may shed new light on the potential for interactions between ecology and evolution to influence the structure and function of entire ecosystems.     

Understanding processes at the origin of species flocks with a focus on the marine Antarctic fauna

Species flocks (SFs) fascinate evolutionary biologists who wonder whether such striking diversification can be driven by normal evolutionary processes. Multiple definitions of SFs have hindered the study of their origins. Previous studies identified a monophyletic taxon as a SF if it displays high speciosity in an area in which it is endemic (criterion 1), high ecological diversity among species (criterion 2), and if it dominates the habitat in terms of biomass (criterion 3); we used these criteria in our analyses. Our starting hypothesis is that normal evolutionary processes may provide a sufficient explanation for most SFs. We thus clearly separate each criterion and identify which biological (intrinsic) and environmental (extrinsic) traits are most favourable to their realization. The first part focuses on evolutionary processes. We highlight that some popular putative causes of SFs, such as key innovations or ecological speciation, are neither necessary nor sufficient to fulfill some or all of the three criteria. Initial differentiation mechanisms are diverse and difficult to identify a posteriori because a primary differentiation of one type (genetic, ecological or geographical) often promotes other types of differentiation. Furthermore, the criteria are not independent: positive feedbacks between speciosity and ecological diversity among species are expected whatever the initial cause of differentiation, and ecological diversity should enhance habitat dominance at the clade level. We then identify intrinsic and extrinsic factors that favour each criterion. Low dispersal emerges as a convincing driver of speciosity. Except for a genomic architecture favouring ecological speciation, for which assessment is difficult, high effective population sizes are the single intrinsic factor that directly enhances speciosity, ecological diversity and habitat dominance. No extrinsic factor appeared to enhance all criteria simultaneously but a combination of factors (insularity, fragmentation and environmental stability) may favour the three criteria, although the effect is indirect for habitat dominance. We then apply this analytical framework to Antarctic marine environments by analysing data from 18 speciose clades belonging to echinoderms (five unrelated clades), notothenioid fishes (five clades) and peracarid crustaceans (eight clades). Antarctic shelf environments and history appear favourable to endemicity and speciosity, but not to ecological specialization. Two main patterns are distinguished among taxa. (i) In echinoderms, many brooding, species‐rich and endemic clades are reported, but without remarkable ecological diversity or habitat dominance. In these taxa, loss of the larval stage is probably a consequence of past Antarctic environmental factors, and brooding is suggested to be responsible for enhanced allopatric speciation (via dispersal limitation). (ii) In notothenioids and peracarids, many clades fulfill all three SF criteria. This could result from unusual features in fish and crustaceans: chromosome instability and key innovations (antifreeze proteins) in notothenioids, ecological opportunity in peracarids, and a genomic architecture favouring ecological speciation in both groups. Therefore, the data do not support our starting point that normal evolutionary factors or processes drive SFs because in these two groups uncommon intrinsic features or ecological opportunity provide the best explanation. The utility of the three‐criterion SF concept is therefore questioned and guidelines are given for future studies.     

Increased competition as a cost of specialization during the evolution of resource polymorphism

Identifying the factors that promote or preclude the evolution of resource polymorphism is essential for understanding the origins of diversity. Although such polymorphisms have long been viewed as an adaptive response to intraspecific competition, they are by no means ubiquitous, even in populations experiencing strong competition. In the present study, we examined a potentially important cost of resource polymorphism. Specifically, resource polymorphism typically entails the evolution of one or more resource‐use specialists, and these specialists may suffer more from competition with other specialists than generalists would with other generalists. Using spadefoot toad tadpoles as a model system, we combined stable isotope analyses with an experiment aiming to characterize dietary differences between alternative carnivore and omnivore morphs and to assess the potential ecological consequences of any such differences. We found that carnivores and omnivores represent alternative trophic specialists and generalists, respectively. We also established that the specialist morph (carnivores) experienced greater intramorph competition than the generalist morph (omnivores). We hypothesize that the greater intramorph competition faced by specialists stems ultimately from functional limitations associated with trophic specialization, which prevent specialists from switching to alternative resources when their resource is depleted. These costs may even preclude the evolution of distinct resource‐use specialists, and hence resource polymorphism, in certain populations. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.