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Experimentally transplanting guppies to evolve in a novel, predator-free environment reveals that the direction of plasticity in gene expression is usually opposite to the direction of adaptive evolution; that is, those genes whose expression changes are disadvantageous are more strongly selected upon than those whose changes are advantageous. Effect of phenotypic plasticity on selection Phenotypic plasticity may have a role in the evolutionary adaptation of populations to changed environmental conditions, but it has not been clear whether it facilitates or hinders adaptation. These authors allow guppies to evolve in a novel predator-free environment and look at the resulting changes in gene expression. They see that the genes that evolve expression differences are not those that were adaptively phenotypically plastic, but those whose plasticity of gene expression was non-adaptive in the ancestral population. In other words, genes whose expression reacts to the environment in an advantageous way tend not to evolve in response to selection, whereas those whose expression changes are disadvantageous are strongly selected on. Phenotypic plasticity is the capacity for an individual genotype to produce different phenotypes in response to environmental variation 1 . Most traits are plastic, but the degree to which plasticity is adaptive or non-adaptive depends on whether environmentally induced phenotypes are closer or further away from the local optimum 2 , 3 , 4 . Existing theories make conflicting predictions about whether plasticity constrains or facilitates adaptive evolution 4 , 5 , 6 , 7 , 8 , 9 , 10 , 11 , 12 . Debate persists because few empirical studies have tested the relationship between initial plasticity and subsequent adaptive evolution in natural populations. Here we show that the direction of plasticity in gene expression is generally opposite to the direction of adaptive evolution. We experimentally transplanted Trinidadian guppies ( Poecilia reticulata ) adapted to living with cichlid predators to cichlid-free streams, and tested for evolutionary divergence in brain gene expression patterns after three to four generations. We find 135 transcripts that evolved parallel changes in expression within the replicated introduction populations. These changes are in the same direction exhibited in a native cichlid-free population, suggesting rapid adaptive evolution. We find 89% of these transcripts exhibited non-adaptive plastic changes in expression when the source population was reared in the absence of predators, as they are in the opposite direction to the evolved changes. By contrast, the remaining transcripts exhibiting adaptive plasticity show reduced population divergence. Furthermore, the most plastic transcripts in the source population evolved reduced plasticity in the introduction populations, suggesting strong selection against non-adaptive plasticity. These results support models predicting that adaptive plasticity constrains evolution 6 , 7 , 8 , whereas non-adaptive plasticity potentiates evolution by increasing the strength of directional selection 11 , 12 . The role of non-adaptive plasticity in evolution has received relatively little attention; however, our results suggest that it may be an important mechanism that predicts evolutionary responses to new environments.

Species richness is greatest in the tropics, and much of this diversity is concentrated in mountains. Janzen proposed that reduced seasonal temperature variation selects for narrower thermal tolerances and limited dispersal along tropical elevation gradients [Janzen DH (1967) Am Nat 101:233–249]. These locally adapted traits should, in turn, promote reproductive isolation and higher speciation rates in tropical mountains compared with temperate ones. Here, we show that tropical and temperate montane stream insects have diverged in thermal tolerance and dispersal capacity, two key traits that are drivers of isolation in montane populations. Tropical species in each of three insect clades have markedly narrower thermal tolerances and lower dispersal than temperate species, resulting in significantly greater population divergence, higher cryptic diversity, higher tropical speciation rates, and greater accumulation of species over time. Our study also indicates that tropical montane species, with narrower thermal tolerance and reduced dispersal ability, will be especially vulnerable to rapid climate change.

The impact of anthropogenic climate change on terrestrial organisms is often predicted to increase with latitude, in parallel with the rate of warming. Yet the biological impact of rising temperatures also depends on the physiological sensitivity of organisms to temperature change. We integrate empirical fitness curves describing the thermal tolerance of terrestrial insects from around the world with the projected geographic distribution of climate change for the next century to estimate the direct impact of warming on insect fitness across latitude. The results show that warming in the tropics, although relatively small in magnitude, is likely to have the most deleterious consequences because tropical insects are relatively sensitive to temperature change and are currently living very close to their optimal temperature. In contrast, species at higher latitudes have broader thermal tolerance and are living in climates that are currently cooler than their physiological optima, so that warming may even enhance their fitness. Available thermal tolerance data for several vertebrate taxa exhibit similar patterns, suggesting that these results are general for terrestrial ectotherms. Our analyses imply that, in the absence of ameliorating factors such as migration and adaptation, the greatest extinction risks from global warming may be in the tropics, where biological diversity is also greatest.

Body size has long been recognized to play a key role in shaping species interactions. For example, while small species thrive in a diversity of environments, they typically lose aggressive contests for resources with larger species. However, numerous examples exist of smaller species dominating larger species during aggressive interactions, suggesting that the evolution of traits can allow species to overcome the competitive disadvantage of small size. If these traits accumulate as lineages diverge, then the advantage of large size in interspecific aggressive interactions should decline with increased evolutionary distance. We tested this hypothesis using data on the outcomes of 23,362 aggressive interactions among 246 bird species pairs involving vultures at carcasses, hummingbirds at nectar sources, and antbirds and woodcreepers at army ant swarms. We found the advantage of large size declined as species became more evolutionarily divergent, and smaller species were more likely to dominate aggressive contests when interacting with more distantly-related species. These results appear to be caused by both the evolution of traits in smaller species that enhanced their abilities in aggressive contests, and the evolution of traits in larger species that were adaptive for other functions, but compromised their abilities to compete aggressively. Specific traits that may provide advantages to small species in aggressive interactions included well-developed leg musculature and talons, enhanced flight acceleration and maneuverability, novel fighting behaviors, and traits associated with aggression, such as testosterone and muscle development. Traits that may have hindered larger species in aggressive interactions included the evolution of morphologies for tree trunk foraging that compromised performance in aggressive contests away from trunks, and the evolution of migration. Overall, our results suggest that fundamental trade-offs, such as those associated with body size, are more likely to break down over evolutionary time, changing the rules that govern species interactions and structure ecological communities.

The empirical study of natural selection reveals that adaptations often involve trade‐offs between competing functions. Because natural selection acts on whole organisms rather than isolated traits, adaptive evolution may be constrained by the interaction between traits that are functionally integrated. Yet, few attempts have been made to characterize how and when such constraints are manifested or whether they limit the adaptive divergence of populations. Here we examine the consequences of adaptive life‐history evolution on locomotor performance in the live‐bearing guppy. In response to increased predation from piscivorous fish, Trinidadian guppies evolve an increased allocation of resources toward reproduction. These populations are also under strong selection for rapid fast‐start swimming performance to evade predators. Because embryo development increases a female’s wet mass as she approaches parturition, an increased investment in reproductive allocation should impede fast‐start performance. We find evidence for adaptive but constrained evolution of fast‐start swimming performance in laboratory trials conducted on second‐generation lab‐reared fish. Female guppies from high‐predation localities attain a faster acceleration and velocity and travel a greater distance during fast‐start swimming trials. However, velocity and distance traveled decline more rapidly over the course of pregnancy in these same females, thus reducing the magnitude of divergence in swimming performance between high‐ and low‐predation populations. This functional trade‐off between reproduction and swimming performance reveals how different aspects of the phenotype are integrated and highlights the complexity of adaptation at the whole‐organism level.

Ecologically similar species often compete aggressively for shared resources. These interactions are frequently asymmetric, with one species behaviorally dominant to another and excluding it from preferred resources. Despite the potential importance of this type of interference competition as a source of selection, we know little about patterns of trait divergence between dominant and subordinate species. We compiled published data on phylogenetically independent, closely related species of North American birds where one species was consistently dominant in aggressive interactions with a congeneric species. We then compared the body size, breeding phenology, life history, ecological breadth, and biogeography of these species. After accounting for body size and phylogeny, we found repeated patterns of trait divergence between subordinate and dominant species within genera. Subordinate species that migrated seasonally arrived 4-7 days later than dominants on their sympatric breeding grounds, and both resident and migratory subordinates initiated breeding 7-8 days later than their dominant, sympatric congeners. Subordinate species had a 5.2% higher annual adult mortality rate and laid eggs that were 0.02 g heavier for their body mass. Dominant and subordinate species used a similar number of different foods, foraging behaviors, nest sites, and habitats, but subordinates were more specialized in their foraging behaviors compared with closely related dominant species. The breeding and wintering ranges of subordinate species were 571 km farther apart than the ranges of dominant species, suggesting that subordinate species migrate greater distances. Range sizes and latitudinal distributions did not consistently differ, although subordinate species tended to breed farther north or winter farther south. These results are consistent with dominant species directly influencing the ecological strategies of subordinate species (via plastic or genetically based changes), either by restricting their access to resources or simply through aggression. Alternatively, these ecological traits may covary with patterns of behavioral dominance, with no direct consequences of interactions. Regardless of the mechanism, recognizing that the relative position of a species within a dominance hierarchy is correlated with a suite of other ecological and fitness related traits has far-reaching implications for the mechanisms underlying species distributions and the structure of biological assemblages.

Aggressive interactions among closely related species are common, and can play an important role as a selective pressure shaping species traits and assemblages. The nature of this selective pressure depends on whether the outcomes of aggressive contests are asymmetric between species (i.e., one species is consistently dominant), yet few studies have estimated the prevalence of asymmetric versus symmetric outcomes to aggressive contests. Here we use previously published data involving 26,212 interactions between 270 species pairs of birds from 26 taxonomic families to address the question: How often are aggressive interactions among closely related bird species asymmetric? We define asymmetry using (i) the proportion of contests won by one species, and (ii) statistical tests for asymmetric outcomes of aggressive contests. We calculate these asymmetries using data summed across different sites for each species pair, and compare results to asymmetries calculated using data separated by location. We find that 80% of species pairs had aggressive outcomes where one species won 80% or more of aggressive contests. We also find that the majority of aggressive interactions among closely related species show statistically significant asymmetries, and above a sample size of 52 interactions, all outcomes are asymmetric following binomial tests. Species pairs with dominance data from multiple sites showed the same dominance relationship across locations in 93% of the species pairs. Overall, our results suggest that the outcome of aggressive interactions among closely related species are usually consistent and asymmetric, and should thus favor ecological and evolutionary strategies specific to the position of a species within a dominance hierarchy.

Widespread recognition of the importance of biological studies at large spatial and temporal scales, particularly in the face of many of the most pressing issues facing humanity, has fueled the argument that there is a need to reinvigorate such studies in physiological ecology through the establishment of a macrophysiology. Following a period when the fields of ecology and physiological ecology had been regarded as largely synonymous, studies of this kind were relatively commonplace in the first half of the twentieth century. However, such large‐scale work subsequently became rather scarce as physiological studies concentrated on the biochemical and molecular mechanisms underlying the capacities and tolerances of species. In some sense, macrophysiology is thus an attempt at a conceptual reunification. In this article, we provide a conceptual framework for the continued development of macrophysiology. We subdivide this framework into three major components: the establishment of macrophysiological patterns, determining the form of those patterns (the very general ways in which they are shaped), and understanding the mechanisms that give rise to them. We suggest ways in which each of these components could be developed usefully.

Physical barriers to gene flow were once viewed as prerequisites for adaptive evolutionary divergence. However, a growing body of theoretical and empirical work suggests that divergence can proceed within a single population. Here we document genetic structure and spatially replicated patterns of phenotypic divergence within a bird species endemic to 250 km2 Santa Cruz Island, California, USA. Island scrub-jays (Aphelocoma insularis) in three separate stands of pine habitat had longer, shallower bills than jays in oak habitat, a pattern that mirrors adaptive differences between allopatric populations of the species' mainland congener. Variation in both bill measurements was heritable, and island scrub-jays mated nonrandomly with respect to bill morphology. The population was not panmictic; instead, we found a continuous pattern of isolation by distance across the east–west axis of the island, as well as a subtle genetic discontinuity across the boundary between the largest pine stand and adjacent oak habitat. The ecological factors that appear to have facilitated adaptive differentiation at such a fine scale—environmental heterogeneity and localized dispersal—are ubiquitous in nature. These findings support recent arguments that microgeographic patterns of adaptive divergence may be more common than currently appreciated, even in mobile taxonomic groups like birds.

Life history theory predicts that parents should value their own survival over that of their offspring in species with a higher probability of adult survival and fewer offspring. We report that Southern Hemisphere birds have higher adult survival and smaller clutch sizes than Northern Hemisphere birds. We subsequently manipulated predation risk to adults versus offspring in 10 species that were paired between North and South America on the basis of phylogeny and ecology. As predicted, southern parents responded more strongly to reduce mortality risk to themselves even at a cost to their offspring, whereas northern parents responded more strongly to reduce risk to their offspring even at greater risk to themselves.