Evaluating life‐history strategies of reef corals from species traits

Classifying the biological traits of organisms can test conceptual frameworks of life‐history strategies and allow for predictions of how different species may respond to environmental disturbances. We apply a trait‐based classification approach to a complex and threatened group of species, scleractinian corals. Using hierarchical clustering and random forests analyses, we identify up to four life‐history strategies that appear globally consistent across 143 species of reef corals: competitive, weedy, stress‐tolerant and generalist taxa, which are primarily separated by colony morphology, growth rate and reproductive mode. Documented shifts towards stress‐tolerant, generalist and weedy species in coral reef communities are consistent with the expected responses of these life‐history strategies. Our quantitative trait‐based approach to classifying life‐history strategies is objective, applicable to any taxa and a powerful tool that can be used to evaluate theories of community ecology and predict the impact of environmental and anthropogenic stressors on species assemblages.     

Density‐dependent survival varies with species life‐history strategy in a tropical forest

Species coexistence in diverse communities likely results from multiple interacting factors. Mechanisms such as conspecific negative density dependence (CNDD) and varying life‐history strategies related to resource partitioning are known to influence plant fitness, and thereby community composition and diversity. However, we have little understanding of how these mechanisms interact and how they vary across life stages. Here, we document the interaction between CNDD and life‐history strategy, based on growth‐mortality trade‐offs, from seedling to adult tree for 47 species in a tropical forest. Species’ life‐history strategies remained consistent across stages: fast‐growing species had higher mortality than slow‐growing species at all stages. In contrast, mean CNDD was strongest at early life stages (i.e. seedling, sapling). Fast‐growing species tended to suffer greater CNDD than slow‐growing species at several, but not all life stages. Overall, our results demonstrate that coexistence mechanisms interact across multiple life stages to shape diverse tree communities.     

Rarity in Chilean forest birds: which ecological and life‐history traits matter?

While it is a truism that species rarity is non-randomly distributed across regions, habitats, and taxa, there is little consensus on which factors are the best predictors of low abundances and restricted geographical ranges. In this study, we evaluate the effects of ecological and life-history traits, as well as phylogeny, on rarity in the abundance and distribution of land birds inhabiting forest habitats in the Mediterranean and temperate regions of Chile. We use data on abundance collected at 16 sites and data on latitudinal distribution obtained from a literature compilation. Statistical analyses were based on multiple regression and multivariate models. We used Signed Mantel test to analyse the relationship between species ecological and life-history traits and rarity, taking into account the effect of phylogenetic relatedness. We found that rarity, in terms of distribution, is associated with a low investment in reproduction, non-migratory status, and degree of habitat specialization. These ecological and life-history traits, in association with forest loss due to climatic changes and human impacts, may explain the narrow distribution of most endemic forest birds species. Rarity in abundance, on the other hand, is more difficult to explain. However, the fact that large species with an insectivorous diet showed low density in the assemblages studied suggests that abundance is mostly regulated by energy (resource) requirements and availability. Finally, our study shows that there is no phylogenetic influence in the observed patterns.     

Population structure,spatial distribution and life‐history traits of blacktip reef sharks Carcharhinus melanopterus

During a survey of the population of blacktip reef shark Carcharhinus melanopterus in Moorea (French Polynesia) between 2007 and 2011, population structural characteristics were estimated from 268 individuals. Total length (L_T) ranged from 48 to 139 cm and 48 to 157 cm for males and females, respectively, demonstrating that the average L_T of females was larger than that of males. The C. melanopterus population at Moorea showed an apparent spatial sexual segregation with females preferentially frequenting lagoons and males the fore‐reefs. Mean growth rate was c. 6 cm year^?1. Males reached sexual maturity at 111 cm L_T. This study reports on the population characteristics of this widespread carcharhinid shark species and makes comparisons with other locations, confirming high geographic variability in the population structure of the species.     

Responses of coral reef fishes to past climate changes are related to life‐history traits

Coral reefs and their associated fauna are largely impacted by ongoing climate change. Unravelling species responses to past climatic variations might provide clues on the consequence of ongoing changes. Here, we tested the relationship between changes in sea surface temperature and sea levels during the Quaternary and present‐day distributions of coral reef fish species. We investigated whether species‐specific responses are associated with life‐history traits. We collected a database of coral reef fish distribution together with life‐history traits for the Indo‐Pacific Ocean. We ran species distribution models (SDMs) on 3,725 tropical reef fish species using contemporary environmental factors together with a variable describing isolation from stable coral reef areas during the Quaternary. We quantified the variance explained independently by isolation from stable areas in the SDMs and related it to a set of species traits including body size and mobility. The variance purely explained by isolation from stable coral reef areas on the distribution of extant coral reef fish species largely varied across species. We observed a triangular relationship between the contribution of isolation from stable areas in the SDMs and body size. Species, whose distribution is more associated with historical changes, occurred predominantly in the Indo‐Australian archipelago, where the mean size of fish assemblages is the lowest. Our results suggest that the legacy of habitat changes of the Quaternary is still detectable in the extant distribution of many fish species, especially those with small body size and the most sedentary. Because they were the least able to colonize distant habitats in the past, fish species with smaller body size might have the most pronounced lags in tracking ongoing climate change.     

Tracking changes in life‐history traits related to unnecessary virulence in a plant‐parasitic nematode

Evaluating trade‐offs in life‐history traits of plant pathogens is essential to understand the evolution and epidemiology of diseases. In particular, virulence costs when the corresponding host resistance gene is lacking play a major role in the adaptive biology of pathogens and contribute to the maintenance of their genetic diversity. Here, we investigated whether life‐history traits directly linked to the establishment of plant–nematode interactions, that is, ability to locate and move toward the roots of the host plant, and to invade roots and develop into mature females, are affected in Meloidogyne incognita lines virulent against the tomato Mi‐1.2 resistance gene. Virulent and avirulent near‐isogenic lines only differing in their capacity to reproduce or not on resistant tomatoes were compared in single inoculation or pairwise competition experiments. Data highlighted (1) a global lack of trade‐off in traits associated with unnecessary virulence with respect to the nematode ability to successfully infest plant roots and (2) variability in these traits when the genetic background of the nematode is considered irrespective of its (a)virulence status. These data suggest that the variation detected here is independent from the adaptation of M. incognita to host resistance, but rather reflects some genetic polymorphism in this asexual organism.     

Interactive life‐history traits predict sensitivity of plants and animals to temporal autocorrelation

Temporal autocorrelation in demographic processes is an important aspect of population dynamics, but a comprehensive examination of its effects on different life‐history strategies is lacking. We use matrix population models from 454 plant and animal populations to simulate stochastic population growth rates (log λ_s) under different temporal autocorrelations in demographic rates , using simulated and observed covariation among rates. We then test for differences in sensitivities, or changes of log λ_s to changes in autocorrelation among two major axes of life‐history strategies, obtained from phylogenetically informed principal component analysis: the fast‐slow and reproductive‐strategy continua. Fast life histories exhibit highest sensitivities to simulated autocorrelation in demographic rates across reproductive strategies. Slow life histories are less sensitive to temporal autocorrelation, but their sensitivities increase among highly iteroparous species. We provide cross‐taxonomic evidence that changes in the autocorrelation of environmental variation may affect a wide range of species, depending on complex interactions of life‐history strategies.     

Macroevolution of life‐history traits in passerine birds: adaptation and phylogenetic inertia

We used a recent passerine phylogeny and comparative method to evaluate the macroevolution of body and egg mass, incubation and fledging periods, time to independence and time with parents of the main passerine lineages. We hypothesised that passerine reproductive traits are affected by adaptation to both past and present environmental factors and phenotypic attributes such as body mass. Our results suggest that the evolution of body and egg mass, time to independence, incubation and fledging times are affected by strong phylogenetic inertia and that these breeding traits are all affected by body mass. Time with parents, where major lineages exhibit their own fixed optima and body mass does not have an effect, and clutch size which is affected by body mass and additionally by climate regimes, do not exhibit any phylogenetic inertia.     

How plant life‐history and ecological traits relate to species rarity and commonness at varying spatial scales

Comparative studies investigating relationships between plant traits and species rarity and commonness were surveyed to establish whether global patterns have emerged that would be of practical use in management strategies aimed at the long‐term conservation of species. Across 54 studies, 94 traits have been examined in relation to abundance, distribution and threatened status at local, regional and geographical spatial scales. Most traits (63) have yet to be the focus of more than one study. Half of the studies involved less than 10 species, and one‐quarter did not replicate rare–common contrasts. Although these features of the literature make it difficult to demonstrate robust generalizations regarding trait relationships with species rarity, some important findings surfaced in relation to traits that have been examined in two or more studies. Species with narrow geographical distributions were found to produce significantly fewer seeds (per unit measurement) than common species (in four of six studies), but did not differ with respect to breeding system (five of five studies). The majority of traits (including seed size, competitive ability, growth form and dispersal mode) were related to rarity in different ways from one study to the next. The highly context‐dependent nature of most trait relationships with rarity implies that application of knowledge concerning rare–common differences and similarities to management plans will vary substantially for different vegetation types and on different continents. A comparative analysis of distribution patterns in relation to several life‐history and ecological traits among 700 Australian eucalypt species was then performed. A significantly dispro­portionate number of tall species and species with long flowering durations had wide geographical ranges. Trait relationships with distribution were explored further through the development of a methodology incorporating multiple spatial scales. Eight theoretical categories were described illustrating variation in distribution patterns (and hence rarity and commonness) across small, intermediate and large spatial scales, based on the spatial structure of species occurrence across the Australian landscape. Each eucalypt species was placed into a category, and trait variation was explored across all species in relation to distribution patterns across multiple spatial scales. This approach yielded important information about trait relationships with distribution among the eucalypts, linking the spatial structure of points‐of‐occurrence with patterns of rarity and commonness. With the pressing need to protect increasing numbers of threatened species and slow rates of extinction, the development and refinement of a broadly usable methodology for rarity studies that encompasses multiple spatial scales, which can be used for any geographical location, will be useful in both conservation and management.     

Effects of host plant on life‐history traits in the polyphagous spider mite Tetranychus urticae

Studying antagonistic coevolution between host plants and herbivores is particularly relevant for polyphagous species that can experience a great diversity of host plants with a large range of defenses. Here, we performed experimental evolution with the polyphagous spider mite Tetranychus urticae to detect how mites can exploit host plants. We thus compared on a same host the performance of replicated populations from an ancestral one reared for hundreds of generations on cucumber plants that were shifted to either tomato or cucumber plants. We controlled for maternal effects by rearing females from all replicated populations on either tomato or cucumber leaves, crossing this factor with the host plant in a factorial design. About 24 generations after the host shift and for all individual mites, we measured the following fitness components on tomato leaf fragments: survival at all stages, acceptance of the host plant by juvenile and adult mites, longevity, and female fecundity. The host plant on which mite populations had evolved did not affect the performance of the mites, but only affected their sex ratio. Females that lived on tomato plants for circa 24 generations produced a higher proportion of daughters than did females that lived on cucumber plants. In contrast, maternal effects influenced juvenile survival, acceptance of the host plant by adult mites and female fecundity. Independently of the host plant species on which their population had evolved, females reared on the tomato maternal environment produced offspring that survived better on tomato as juveniles, but accepted less this host plant as adults and had a lower fecundity than did females reared on the cucumber maternal environment. We also found that temporal blocks affected mite dispersal and both female longevity and fecundity. Taken together, our results show that the host plant species can affect critical parameters of population dynamics, and most importantly that maternal and environmental conditions can facilitate colonization and exploitation of a novel host in the polyphagous T. urticae, by affecting dispersal behavior (host acceptance) and female fecundity.     

Forest bees are replaced in agricultural and urban landscapes by native species with different phenologies and life‐history traits

Anthropogenic landscapes are associated with biodiversity loss and large shifts in species composition and traits. These changes predict the identities of winners and losers of future global change, and also reveal which environmental variables drive a taxon's response to land use change. We explored how the biodiversity of native bee species changes across forested, agricultural, and urban landscapes. We collected bee community data from 36 sites across a 75,000 km² region, and analyzed bee abundance, species richness, composition, and life‐history traits. Season‐long bee abundance and richness were not detectably different between natural and anthropogenic landscapes, but community phenologies differed strongly, with an early spring peak followed by decline in forests, and a more extended summer season in agricultural and urban habitats. Bee community composition differed significantly between all three land use types, as did phylogenetic composition. Anthropogenic land use had negative effects on the persistence of several life‐history strategies, including early spring flight season and brood parasitism, which may indicate adaptation to conditions in forest habitat. Overall, anthropogenic communities are not diminished subsets of contemporary natural communities. Rather, forest species do not persist in anthropogenic habitats, but are replaced by different native species and phylogenetic lineages preadapted to open habitats. Characterizing compositional and functional differences is crucial for understanding land use as a global change driver across large regional scales.     

Effects of the demographic transition on the genetic variances and covariances of human life‐history traits

The recent demographic transitions to lower mortality and fertility rates in most human societies have led to changes and even quick reversals in phenotypic selection pressures. This can only result in evolutionary change if the affected traits are heritable, but changes in environmental conditions may also lead to subsequent changes in the genetic variance and covariance (the G matrix) of traits. It currently remains unclear if there have been concomitant changes in the G matrix of life‐history traits following the demographic transition. Using 300 years of genealogical data from Finland, we found that four key life‐history traits were heritable both before and after the demographic transition. The estimated heritabilities allow a quantifiable genetic response to selection during both time periods, thus facilitating continued evolutionary change. Further, the G matrices remained largely stable but revealed a trend for an increased additive genetic variance and thus evolutionary potential of the population after the transition. Our results demonstrate the validity of predictions of evolutionary change in human populations even after the recent dramatic environmental change, and facilitate predictions of how our biology interacts with changing environments, with implications for global public health and demography.     

Variability in life‐history and ecological traits is a buffer against extinction in mammals

Anthropogenic degradation of the world's ecosystems is leading to a widespread and accelerating loss of biodiversity. However, not all species respond equally to existing threats, raising the question: what makes a species more vulnerable to extinction? We propose that higher intraspecific variability may reduce the risk of extinction, as different individuals and populations within a species may respond differently to occurring threats. Supporting this prediction, our results show that mammalian species with more variable adult body masses, litter sizes, sexual maturity ages and population densities are less vulnerable to extinction. Our findings reveal the role of local variation among populations, particularly of large mammals, as a buffering mechanism against extinction, and emphasise the importance of considering trait variation in comparative analyses and conservation management.     

A large‐scale latitudinal pattern of life‐history traits in a strictly univoltine damselfly

1. Variation in thermal conditions and season length along latitudinal gradients affect body size‐related traits over different life stages. Selection is expected to optimise these size traits in response to the costs and benefits. 2. Egg, hatchling, larval and adult size in males and females were estimated along a latitudinal gradient of 2730 km across Europe in the univoltine damselfly Lestes sponsa, using a combination of field‐collection and laboratory‐rearing experiments. In the laboratory, individuals were grown in temperatures and photoperiod simulating those at the latitude of origin, and in common‐garden conditions. 3. The size of adults sampled in nature was negatively correlated with latitude. In all populations the females were larger than the males. Results from simulated and common‐garden rearing experiments supported this pattern of size difference across latitudes and between sexes, suggesting a genetic component for the latitudinal size trend and female‐biased size dimorphism. In contrast, hatchling size showed a positive relationship with latitude, but egg size, although differing between latitudes, showed no such relationship. 4. The results support a converse Bergmann cline, i.e. a negative body size cline towards the north. This negative cline in body size is probably driven by progressively stronger seasonal time and temperature constraints towards the higher latitudes and by the obligate univoltine life cycle of L. sponsa. As egg size showed no relationship with latitude, other environmental factors besides temperature, such as desiccation risk, probably affect this trait.     

Environmental,spatial and phylogenetic determinants of fish life‐history traits and functional composition of Australian rivers

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Multimodal signalling in estrildid finches: song,dance and colour are associated with different ecological and life‐history traits

Sexual traits (e.g. visual ornaments, acoustic signals, courtship behaviour) are often displayed together as multimodal signals. Some hypotheses predict joint evolution of different sexual signals (e.g. to increase the efficiency of communication) or that different signals trade off with each other (e.g. due to limited resources). Alternatively, multiple signals may evolve independently for different functions, or to communicate different information (multiple message hypothesis). We evaluated these hypotheses with a comparative study in the family Estrildidae, one of the largest songbird radiations, and one that includes many model species for research in sexual selection and communication. We found little evidence for either joint evolution or trade‐offs between song and colour ornamentation. Some negative correlations between dance repertoire and song traits may suggest a functional compromise, but generally courtship dance also evolved independently from other signals. Instead of correlated evolution, we found that song, dance and colour are each related to different socio‐ecological traits. Song complexity evolved together with ecological generalism, song performance with investment in reproduction, dance with commonness and habitat type, whereas colour ornamentation was shown previously to correlate mostly with gregariousness. We conclude that multimodal signals evolve in response to various socio‐ecological traits, suggesting the accumulation of distinct signalling functions.     

Maintenance of body‐size variation and host range in the orange‐tip butterfly: evidence for a trade‐off between adult life‐history traits

1. The evolution of host range and preference in phytophagous insects is driven by a female's oviposition choice impacting her offspring's fitness. Analysis of the fitness of progeny on different host plants has commonly been restricted to the performance of immature stages. However, since host use can affect adult size, it is important to measure the ongoing effects of host choice on the resulting imagines. 2. The orange‐tip butterfly, Anthocharis cardamines, shows a strong preference for two host plants in Britain, Alliaria petiolata and Cardamine pratensis, which affect body size. Whilst females exhibit a strong positive size–fecundity relation, the impact of body‐size alteration is unknown in males. In this study, fitness effects of host plant choice for male A. cardamines were examined. 3. Males reared on C. pratensis were smaller and emerged earlier than those reared on A. petiolata, and early‐season males were smaller than late‐season ones in the field. Interestingly, regression analysis indicated that the earlier emergence of small males was a host‐mediated rather than a size‐mediated effect. Small size was associated with reduced male dispersal in a semi‐isolated wild population over a 3‐year period. 4. It is proposed that the earlier emergence associated with C. pratensis has evolved in response to depressed dispersal in isolated/semi‐isolated populations associated with this patchily distributed host. We suggest that adult life‐history traits are important for the maintenance of host range in this species, and offer a critique of Courtney's earlier hypothesis that host range is maintained by time‐limited oviposition behaviour.     

Immune deployment increases larval vulnerability to predators and inhibits adult life‐history traits in a dragonfly

While deploying immune defences early in ontogeny can trade‐off with the production and maintenance of other important traits across the entire life cycle, it remains largely unexplored how features of the environment shape the magnitude or presence of these lifetime costs. Greater predation risk during the juvenile stage may particularly influence such costs by (1) magnifying the survival costs that arise from any handicap of juvenile avoidance traits and/or (2) intensifying allocation trade‐offs with important adult traits. Here, we tested for predator‐dependent costs of immune deployment within and across life stages using the dragonfly, Pachydiplax longipennis. We first examined how larval immune deployment affected two traits associated with larval vulnerability to predators: escape distance and foraging under predation risk. Larvae that were induced to mount an immune response had shorter escape distances but lower foraging activity in the presence of predator cues. We also induced immune responses in larvae and reared them through emergence in mesocosms that differed in the presence of large predatory dragonfly larvae (Aeshnidae spp.). Immune‐challenged larvae had later emergence overall and lower survival in pools with predators. Immune‐challenged males were also smaller at emergence and developed less sexually selected melanin wing coloration, but these effects were independent of predator treatment. Overall, these results highlight how mounting an immune defence early in ontogeny can have substantial ecological and physiological costs that manifest both within and across life stages.