Size‐dependent costs of migration: Migrant bird species are subordinate to residents,but only at small body sizes

Migrant species are commonly thought to be poor competitors in aggressive interactions with resident species. However, no studies have tested whether this relationship is widespread. Here, we compare the behavioural dominance of closely related species of migratory and nonmigratory birds, testing whether migrants are consistently subordinate to resident species in aggressive contests. We compiled published behavioural dominance data involving migrant and resident congeners, gathering additional data on the body mass and migratory distance of each species. Focal species included a diverse array of birds (28 taxonomic families, 12 orders) from around the world. We found that migrant species are usually subordinate to resident species, but that this relationship disappears at larger body sizes. For smaller birds (<500 g), resident species were behaviourally dominant in 83%–88% of comparisons; for larger birds (>500 g), resident species were dominant in only 25%–30% of comparisons. The relative difference in body mass best predicted dominance relationships among species, with larger species dominant in 80%–84% of comparisons. When migrant and resident masses were equal, however, resident species were still more likely to be dominant in smaller birds, suggesting that other factors may also contribute to the subordinate status of migrants. Overall, our results suggest that in smaller species, the evolution of migration is associated with lighter weights and other traits that compromise the competitive abilities of migrants relative to residents. In contrast, larger species appear able to evolve migration without compromising their size or competitive abilities in aggressive contests, suggesting size‐dependent constraints on the evolution of migration.     

Relative size predicts competitive outcome through 2 million years

Competition is an important biotic interaction that influences survival and reproduction. While competition on ecological timescales has received great attention, little is known about competition on evolutionary timescales. Do competitive abilities change over hundreds of thousands to millions of years? Can we predict competitive outcomes using phenotypic traits? How much do traits that confer competitive advantage and competitive outcomes change? Here we show, using communities of encrusting marine bryozoans spanning more than 2 million years, that size is a significant determinant of overgrowth outcomes: colonies with larger zooids tend to overgrow colonies with smaller zooids. We also detected temporally coordinated changes in average zooid sizes, suggesting that different species responded to a common external driver. Although species‐specific average zooid sizes change over evolutionary timescales, species‐specific competitive abilities seem relatively stable, suggesting that traits other than zooid size also control overgrowth outcomes and/or that evolutionary constraints are involved.     

Insularity and the evolution of melanism,sexual dichromatism and body size in the worldwide‐distributed barn owl

Island biogeography has provided fundamental hypotheses in population genetics, ecology and evolutionary biology. Insular populations usually face different feeding conditions, predation pressure, intraspecific and interspecific competition than continental populations. This so‐called island syndrome can promote the evolution of specific phenotypes like a small (or large) body size and a light (or dark) colouration as well as influence the evolution of sexual dimorphism. To examine whether insularity leads to phenotypic differentiation in a consistent way in a worldwide‐distributed nonmigratory species, we compared body size, body shape and colouration between insular and continental barn owl (Tyto alba) populations by controlling indirectly for phylogeny. This species is suitable because it varies in pheomelanin‐based colouration from reddish‐brown to white, and it displays eumelanic black spots for which the number and size vary between individuals, populations and species. Females are on average darker pheomelanic and display more and larger eumelanic spots than males. Our results show that on islands barn owls exhibited smaller and fewer eumelanic spots and lighter pheomelanic colouration, and shorter wings than on continents. Sexual dimorphism in pheomelanin‐based colouration was less pronounced on islands than continents (i.e. on islands males tended to be as pheomelanic as females), and on small islands owls were redder pheomelanic and smaller in size than owls living on larger islands. Sexual dimorphism in the size of eumelanic spots was more pronounced (i.e. females displayed much larger spots than males) in barn owls living on islands located further away from a continent. Our study indicates that insular conditions drive the evolution towards a lower degree of eumelanism, smaller body size and affects the evolution of sexual dichromatism in melanin‐based colour traits. The effect of insularity was more pronounced on body size and shape than on melanic traits.     

Resource Holding Potential and the Outcome of Aggressive Interactions between Paired Male <Emphasis Type="Italic">Aegus chelifer chelifer</Emphasis> (Coleoptera: Lucanidae) Stag Beetles

Interactions between male stag beetles usually involve aggressive behavior using their long mandibles as weapons to compete with rival males over females. Considerable variation exists within populations in male body size, and may affect their behavior and the outcome of male-male contests. We investigated the aggressive interactions between male Aegus chelifer chelifer, a small tropical stag beetle species. Morphological traits in relation to aggressiveness and the outcome of fights were examined in laboratory-reared beetles. The fight-engagement ratios of major and minor morph males were not significantly different and analyses revealed that the size of body parts had more effect on the fighting success than the weapon part (mandibles). The probability of winning a contest was higher in males with a larger head width (HW), and so HW was considered as the resource holding potential (RHP). No effects of the trait size on the initiation of fights or aggressive intensity was found. Relationships between the fight duration and RHP were not significantly consistent with any assessment strategies, but were close to the mutual assessment model.     

Role of body size in dominance interactions between male water striders, Aquarius paludum

Water striders are a model system for the study of sexual size dimorphism, but the effect of body size on the dominance relationship between individuals has not been experimentally tested. In 34 staged contests between males of the water strider Aquarius paludum, we determined the effect of body size difference between contestants on the outcome of the aggressive interactions. In contests between a large and a small male, the larger individuals won the interactions significantly more often than expected by chance. This is the first experimental evidence for the importance of body size in pair-wise contests among water striders.     

Heterothermy,body size,and locomotion as ecological predictors of migration in mammals

1. Migration is ubiquitous among animals and has evolved repeatedly and independently. Comparative studies of the evolutionary origins of migration in birds are widespread, but are lacking in mammals. Mammalian species have greater variation in functional traits that may be relevant for migration. Interspecific variation in migration behaviour is often attributed to mode of locomotion (i.e. running, swimming, and flying) and body size, but traits associated with the evolutionary precursor hypothesis, including geographic distribution, habitat, and diet, could also be important predictors of migration in mammals. Furthermore, mammals vary in thermoregulatory strategies and include many heterothermic species, providing an alternative strategy to avoid seasonal resource depletion.
2. We tested the evolutionary precursor hypothesis for the evolution of migration in mammals and tested predictions linking migration to locomotion, body size, geographic distribution, habitat, diet, and thermoregulation. We compiled a dataset of 722 species from 27 mammalian orders and conducted a series of analyses using phylogenetically informed models.
3. Swimming and flying mammals were more likely to migrate than running mammals, and larger species were more likely to migrate than smaller ones. However, heterothermy was common among small running mammals that were unlikely to migrate. High-latitude swimming and flying mammals were more likely to migrate than high-latitude running mammals (where heterothermy was common), and most migratory running mammals were herbivorous. Running mammals and frugivorous bats with high thermoregulatory scope (greater capacity for heterothermy) were less likely to migrate, while insectivorous bats with high thermoregulatory scope were more likely to migrate.
4. Our results indicate a broad range of factors that influence migration, depending on locomotion, body size, and thermoregulation. Our analysis of migration in mammals provided insight into some of the general rules of migration, and we highlight opportunities for future investigations of exceptions to these rules, ultimately leading to a comprehensive understanding of the evolution of migration.

     

Rapid evolution by sexual selection in a wild,invasive mammal

Sexual selection theory provides a framework for investigating the evolution of traits involved in attracting and competing for mates. Given the sexual function of such traits, studies generally focus on individual interactions (i.e., displays and contests) in explaining trait origin and persistence. We show that ecological factors can strongly influence the adaptive value of these traits, and changes to these factors can lead to rapid evolutionary change. We compared sexually selected traits in the small Indian mongoose (Urva auropunctata) between their sparsely populated native range and four tropical islands to which they were introduced within the last 150 years and where, due to a lack of interspecific competition and predation, they have become invasive and densely populated. Because of a likely increase in encounter rate, we predicted that selection on long-distance chemical advertisement by males would relax in the introduced range. Accordingly, male, but not female, anal pads (used in scent marking) decreased in size in relation to both time since introduction and population density, and their relationship to body size and condition weakened. Concurrently, as predicted by intensified sperm competition, testis size increased following introduction. The small Indian mongoose thus experienced an inversion in the relative contributions to fitness of two sexual traits, followed by their rapid evolution in line with ecological changes.     

Interactions among moths, crossbills, squirrels, and lodgepole pine in a geographic selection mosaic

Repeated patterns among biological communities suggest similar evolutionary and ecological forces are acting on the communities. Conversely, the lack of such patterns suggests that similar forces are absent or additional ones are present. Coevolution between a seed predator, the red crossbill (Loxia curvirostra complex), and lodgepole pine (Pinus contorta var. latifolia) exemplifies the ecological and evolutionary predictions for coevolving systems. In the absence of another seed predator and preemptive competitor (pine squirrels Tamiasciurus hudsonicus), natural selection by crossbills results in the evolution of larger cones with thicker distal scales, while relaxation of selection by squirrels results in the evolution of cones with more seeds and a greater ratio of seed mass to cone mass. However, in one range, the Little Rocky Mountains, distal scale thickness has diverged as expected but cone size has not. In these mountains seed predation by lodgepole pine cone borer moths (Eucosma recissoriana) was about 10 times greater than in other ranges lacking squirrels. We quantified moth predation and cone traits and found that moths select for smaller cones with fewer seeds. Thus, selection by moths in the Little Rocky Mountains counters both selection by crossbills for large cone size and relaxation of selection by squirrels favoring more seeds per cone and accounts for the relatively small and few-seeded cones in these mountains. It is also apparent that selection by crossbills changes seed defenses in a manner that favors seed predation by moths, whereas selection by squirrels likely reduces such predation. These results demonstrate the importance of considering the evolutionary consequences of community context in locally evolved (coevolved) traits and interactions.     

Deferred agonistic behavior in a long-lived scincid lizard Eumeces laticeps

Summary A laboratory experiment with the broad-headed skink (Eumeces laticeps) involving staged agonistic encounters demonstrates that larger males have an advantage over smaller ones in agonistic bouts. Field data on head wounds produced by intraspecific fighting during the breeding season show a much higher frequency of new wounds among males over 100 mm in snout-vent-length than in smaller males. The significant difference in new-wound frequency strongly suggests avoidance of fights by the small males, which is corroborated by laboratory and field observations. Access by males to reproductively active females depends on the ability to defeat other males in aggressive contests virtually always involving head biting if the males are of nearly equal size. Because the probability of winning agonistic encounters increases with size, young males avoid fights with older males. Aggressive contests with larger males and reproductive attempts other than courtship in the absence of larger males are deferred.Aggressive behavior in E. laticeps may be employed in direct defense of females, but might also be expressed in defense of specific sites and/or territories. In the laboratory, males in their home cages were significantly more likely to win encounters with males of similar size than were males fighting in the home cages of opponents. This suggests that encounter site could be important in determining encounter outcome and that field study of possible site defense or territoriality is needed.     

Fine‐ and Broad‐Scale Approaches to Understanding the Evolution of Aggression in Crickets

Male field crickets frequently engage in agonistic contests to establish dominance in social interactions and gain access to mate attraction territories. Crickets (Orthoptera: Gryllidae) are often used as a model taxon to study aggression, but limited documentation of aggression in some cricket species hinders our understanding of its evolutionary costs and benefits. Our study investigated cricket aggression at two scales: the within‐species scale for two cricket species, Gryllus assimilis and G. veletis, whose aggression had not been adequately documented and the among‐species scale to detect evolutionary patterns in species’ levels of aggression. In both G. veletis and G. assimilis, winners spent more time being aggressive than losers, but they were not larger or heavier. Collectively, our results reveal that G. veletis males are more aggressive than G. assimilis. Male G. veletis had higher aggression scores that male G. assimilis. The majority of G. veletis contests escalated to grappling (a highly aggressive behavior), while less than one quarter of G. assimilis contests escalated to grappling. Further, G. veletis males transitioned between two of the most aggressive behaviors most often while G. assimilis transitioned between two of the least aggressive behaviors most often. We integrate this new information on aggression for G. assimilis and G. veletis with previously documented aggression data for many cricket species to investigate aggression in a broader evolutionary context than previously possible. Within a phylogenetic context, we test the hypothesis that species whose males use burrows from which to call and attract females are more aggressive than species with non‐burrowing males. We found evidence consistent with this hypothesis; species with burrowing males tended to be more aggressive than species with non‐burrowing males. Together, our study provides fine‐scale understanding of aggression in two cricket species and broad‐scale evolutionary context for aggression across cricket species.     

Offenders tend to be heavier: experimental encounters in mangrove-dwelling monitor lizards (<Emphasis Type="Italic">Varanus indicus</Emphasis>)

The evolution of male-biased sexual size dimorphism is often explained by sexual selection providing competitive advantage to the larger males. The aggressive interactions are often dangerous and energy consuming; thus, it is advantageous to reduce the risks by adjusting behavior to correspond with body size as a predictor of fighting success. Organization of contests into distinct phases with the initial displays preceding the real combat allows individuals to assess the body size and strength of the rival. We staged interactions between mangrove-dwelling monitor lizards (Varanus indicus) to uncover the initialization of aggression and factors determining the course of an encounter. The analyses revealed the importance of both absolute and relative body size of encountering males. The attack rate increases with the body weight of the lizard and offenders initializing a contact phase of the fight tend to be the heavier male of the dyad. Regardless of the final outcome of the combat, the results show that only short visual contact provides sufficient information about the body size of the opponent. This enables combatants to determine whether to initiate the fight or not. This finding together with the ethological details of contests provides the first evidence for the ability of mutual assessment in varanids.     

Predation drives recurrent convergence of an interspecies mutualism

Mutualisms are important ecological interactions that underpin much of the world's biodiversity. Predation risk has been shown to regulate mutualism dynamics in species‐specific case studies; however, we lack studies which investigate whether predation can also explain broader patterns of mutualism evolution. We report that fish‐anemone mutualisms have evolved on at least 55 occasions across 16 fish families over the past 60 million years and that adult body size is associated with the ontogenetic stage of anemone mutualisms: larger‐bodied species partner with anemones as juveniles, while smaller‐bodied species partner with anemones throughout their lives. Field and laboratory studies show that predators target smaller prey, that smaller fishes associate more with anemones, and that these relationships confer protection to small fishes. Our results indicate that predation is likely driving the recurrent convergent evolution of fish‐anemone mutualisms and suggest that similar ecological processes may have selected convergence in interspecies interactions in other animal clades.     

Rules of the seed size game: contests between large‐seeded and small‐seeded species

The coexistence of multiple seed size strategies within plant communities have been considered puzzling, based on a theoretical expectation of the existence of an optimal seed size under each set of specific environmental conditions. A model aimed at explaining the coexistence of different seed sizes has been suggested, where a seed size – seed number tradeoff is connected to a tradeoff between competition and colonization, leading to a competitive advantage in larger‐seeded species and a colonization advantage in smaller‐seeded species. Recently an alternative model has been suggested, based on a tradeoff between stress tolerance and fecundity, associated with the variation from large to small seeds. Here, we examine the role of seed size for recruitment in two‐species contests subjected to various treatments. In a garden experiment seeds of 14 plant species were combined pair‐wise into seven pairs, each with one larger‐seeded species and one smaller‐seeded species. Each species‐pair was sown with sparse and dense seed densities and subjected to different treatments of shading and litter. Recruitment was recorded during two years. Our results showed a general advantage of larger‐seeded species over smaller‐seeded species. This seed size advantage increased in treatments with litter, whereas there were minor effects of shade, and no effect of seed density was found. We thus found little support for a density dependent seed size game as assumed in models of a competition‐colonization tradeoff, whereas our results fit well with a model based on a tradeoff between stress tolerance and fecundity. Our experiment provides novel empirical data to theoretical models on co‐existence between multiple seed size strategies.     

Aggression and RHP in the Northern Swordtail Fish, Xiphophorus cortezi: The Relationship Between Size and Contest Dynamics in Male–Male Competition

Current theory predicts that contest outcome, as well as decisions on whether to initiate a contest, escalate during a contest or retreat are decided by asymmetries in resource holding potential (RHP) and/or expected payoffs between contestants. In this investigation, dyadic contests were staged between male swordtail fish (Xiphophorus cortezi) where individuals were paired based on cumulative fight records and were ranked at the end of the trials in order to approximate RHP. Size was the only asymmetry that I did not attempt to control for and as a result, I was able to determine the relationships between size, contest initiation, escalation and outcome. Individuals changed their contest initiation strategy based on their size relative to that of their opponents, and contrary to predictions, the smaller of the two males in each contest was more likely to initiate the conflict than was the larger male. However, the larger of the two males was more likely to win and standard length proved to be a moderate predictor of an individual's final rank. Regardless of size, initiators fared poorly, winning only 31% of the contests. In instances where the smaller males won the contests, they were no more likely to initiate the encounter than was the larger male. However, when small males did win, fights lasted longer, suggesting that in some cases smaller males may be able to outlast their opponents.     

Why are small males aggressive?

Aggression is ubiquitous in the animal kingdom, whenever the interests of individuals conflict. In contests between animals, the larger opponent is often victorious. However, counter intuitively, an individual that has little chance of winning (generally smaller individuals) sometimes initiates contests. A number of hypotheses have been put forward to explain this behaviour, including the "desperado effect" according to which, the likely losers initiate aggression due to lack of alternative options. An alternative explanation suggested recently is that likely losers attack due to an error in perception: they mistakenly perceive their chances of winning as being greater than they are. We show that explaining the apparently maladaptive aggression initiated by the likely loser can be explained on purely economic grounds, without requiring either the desperado effect or perception errors. Using a game-theoretical model, we show that if smaller individuals can accurately assess their chance of winning, if this chance is less than, but close to, a half, and if resources are scarce (or the contested resource is of relatively low value), they are predicted to be as aggressive as their larger opponents. In addition, when resources are abundant, and small individuals have some chance of winning, they may be more aggressive than their larger opponents, as it may benefit larger individuals to avoid the costs of fighting and seek alternative uncontested resources.     

Larger brains spur species diversification in birds

Evidence is accumulating that species traits can spur their evolutionary diversification by influencing niche shifts, range expansions, and extinction risk. Previous work has shown that larger brains (relative to body size) facilitate niche shifts and range expansions by enhancing behavioral plasticity but whether larger brains also promote evolutionary diversification is currently backed by insufficient evidence. We addressed this gap by combining a brain size dataset for >1900 avian species worldwide with estimates of diversification rates based on two conceptually different phylogenetic‐based approaches. We found consistent evidence that lineages with larger brains (relative to body size) have diversified faster than lineages with relatively smaller brains. The best supported trait‐dependent model suggests that brain size primarily affects diversification rates by increasing speciation rather than decreasing extinction rates. In addition, we found that the effect of relatively brain size on species‐level diversification rate is additive to the effect of other intrinsic and extrinsic factors. Altogether, our results highlight the importance of brain size as an important factor in evolution and reinforce the view that intrinsic features of species have the potential to influence the pace of evolution.     

Evolutionary correlations among morphology, habitat use and clinging performance in Caribbean Anolis lizards

A central issue in evolutionary biology concerns whether morphology, performance and habitat use have coevolved. We investigated evolutionary relationships among the size of the subdigital toepad, clinging ability and perch height in 12 species of Caribbean Anolis lizard. Specifically, we predicted that: (1) because larger anole species tend to perch high in the canopy, both toepad area and clinging ability should scale with positive allometry to enable small and large lizards to possess approximately similar ratios of both variables relative to mass; (2) anole species with relatively larger toepads (i.e. size-adjusted) should be relatively better clingers compared with species with relatively small toepads; (3) species that perch high in the canopy should possess relatively large clinging abilities (either on an absolute or a size-adjusted basis). Our first hypothesis was refuted, as both toepad area and clinging ability scaled close to isometry (0.67) relative to mass, indicating that large lizard species have low ratios of clinging ability to mass compared with small lizard species. However, our second and third predictions were confirmed. Anole species with relatively larger toepads were relatively better clingers compared with species with relatively smaller toepads. Anole species that perched high in the canopy (either on an absolute scale or relative to size) tended to have relatively larger toepads and greater clinging capacities compared with species that perched lower in the canopy. These data provide indirect comparative evidence that the evolution of increased toepad size in some anole species is adaptive, by facilitating the occupation of perches high in the canopy.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 389–398.     

Beauty and the beast: mechanisms of sexual selection in humans

Literature in evolutionary psychology suggests that mate choice has been the primary mechanism of sexual selection in humans, but this conclusion conforms neither to theoretical predictions nor available evidence. Contests override other mechanisms of sexual selection; that is, when individuals can exclude their competitors by force or threat of force, mate choice, sperm competition, and other mechanisms are impossible. Mates are easier to monopolize in two dimensional mating environments, such as land, than in three-dimensional environments, such as air, water, and trees. Thus, two-dimensional mating environments may tend to favor the evolution of contests. The two-dimensionality of the human mating environment, along with phylogeny, the spatial and temporal clustering of mates and competitors, and anatomical considerations, predict that contest competition should have been the primary mechanism of sexual selection in men. A functional analysis supports this prediction. Men's traits are better designed for contest competition than for other sexual selection mechanisms; size, muscularity, strength, aggression, and the manufacture and use of weapons probably helped ancestral males win contests directly, and deep voices and facial hair signal dominance more effectively than they increase attractiveness. However, male monopolization of females was imperfect, and female mate choice, sperm competition, and sexual coercion also likely shaped men's traits. In contrast, male mate choice was probably central in women's mating competition because ancestral females could not constrain the choices of larger and more aggressive males through force, and attractive women could obtain greater male investment. Neotenous female features and body fat deposition on the breasts and hips appear to have been shaped by male mate choice.     

The Evolution of Olfactory Capabilities in Wild Birds: A Comparative Study

Modern birds possess an olfactory apparatus similar to that of other vertebrates, yet the major evolutionary forces that drove the evolution of diversity in olfactory capabilities in birds remain elusive. Several non-mutually exclusive evolutionary scenarios for the evolution of olfactory capability in birds have been proposed. Olfactory capability may have evolved due to its role in recognition in social contexts. In addition, olfactory capability may have evolved because it provides a selective advantage in foraging or navigation. Finally, olfactory capability could be favored whenever the ecological conditions of species hindered the use of other senses and/or favored olfaction. Here we evaluate predictions from these hypotheses in a comparative study using a species-level phylogeny of 142 bird species and considering the indirect effects of some predictors on others. We find an interactive effect of aquatic dependence and diet on the size of the olfactory bulb: vegetarian and omnivore birds living in aquatic environments have larger olfactory bulbs than terrestrial birds, whereas species with an animal diet, aquatic and terrestrial species have similar-sized olfactory bulbs. In addition, the size of the olfactory bulb was weakly related with social complexity, with colonial species having relatively larger olfactory bulb than solitary breeders or those forming small aggregations. Our results suggest that the role of foraging in driving enhancements in the avian olfactory apparatus is contingent on ecological conditions that may affect the transmission of odor-based signals. This provides evidence for the largely neglected possibility that the evolution of the olfactory apparatus of birds has been driven by the interaction between ecological and behavioral traits, rather than being solely due to main effects of these traits.     

THE EVOLUTION OF FEMALE BODY SIZE IN RED-WINGED BLACKBIRDS: THE EFFECTS OF TIMING OF BREEDING,SOCIAL COMPETITION,AND REPRODUCTIVE ENERGETICS

We examined opposing selective forces on female body size in the sexually dimorphic red-winged blackbird: social competition favoring larger females, and energetic advantages favoring smaller females. Downhower proposed that selection might drive female birds to be smaller than the optimum for survival, if smaller females were able to exceed their energetic requirements for self-maintenance earlier in the season and therefore breed earlier. Since in most birds the earliest breeders fledge the most young, this could favor the evolution of smaller female size, and therefore contribute to the magnitude of sexual size dimorphism in these birds. We tested this hypothesis in 1987 and 1988 by comparing the size and breeding date of female red-winged blackbirds. Consistent with our preditions, early-nesting females had much higher nesting success, but contrary to prediction, larger females bred earlier. We then examined the effects of female size on competition. If large females have an advantage in social competition, and if competition influences breeding date and reproductive success, then larger females might breed earlier. Primary females, the first females to arrive and nest on a territory, were more aggressive than lower ranked females; more aggressive females settled on better territories and laid earlier than less aggressive females; and larger females were more aggressive. Social competition between females may therefore favor large females. Finally, we tested the prediction that selection favoring large females might be limited by energetic constraints on large females. We found that large females had less fat than small females during breeding, and that the levels of fat that females of a given size carried affected breeding date and egg size. Therefore, social competition may favor large females, but reproductive energetics favoring smaller females may constrain selection for large female body size.