Two different jumping mechanisms of water striders are determined by body size.

Current theory for surface tension-dominant jumps on water, created for small- and medium-sized water strider species and used in bioinspired engineering, predicts that jumping individuals are able to match their downward leg movement speed to their size and morphology such that they maximize the takeoff speed and minimize the takeoff delay without breaking the water surface. Here, we use empirical observations and theoretical modeling to show that large species (heavier than ~80 mg) could theoretically perform the surface-dominated jumps according to the existing model, but they do not conform to its predictions, and switch to using surface-breaking jumps in order to achieve jumping performance sufficient for evading attacks from underwater predators. This illustrates how natural selection for avoiding predators may break the theoretical scaling relationship between prey size and its jumping performance within one physical mechanism, leading to an evolutionary shift to another mechanism that provides protection from attacking predators. Hence, the results are consistent with a general idea: Natural selection for the maintenance of adaptive function of a specific behavior performed within environmental physical constraints leads to size-specific shift to behaviors that use a new physical mechanism that secure the adaptive function.

Antarctic skuas recognize individual humans.

Recent findings report that wild animals can recognize individual humans. To explain how the animals distinguish humans, two hypotheses are proposed. The high cognitive abilities hypothesis implies that pre-existing high intelligence enabled animals to acquire such abilities. The pre-exposure to stimuli hypothesis suggests that frequent encounters with humans promote the acquisition of discriminatory abilities in these species. Here, we examine individual human recognition abilities in a wild Antarctic species, the brown skua (Stercorarius antarcticus), which lives away from typical human settlements and was only recently exposed to humans due to activities at Antarctic stations. We found that, as nest visits were repeated, the skua parents responded at further distances and were more likely to attack the nest intruder. Also, we demonstrated that seven out of seven breeding pairs of skuas selectively responded to a human nest intruder with aggression and ignored a neutral human who had not previously approached the nest. The results indicate that Antarctic skuas, a species that typically inhabited in human-free areas, are able to recognize individual humans who disturbed their nests. Our findings generally support the high cognitive abilities hypothesis, but this ability can be acquired during a relatively short period in the life of an individual as a result of interactions between individual birds and humans.

Assessing dietary specialization to inform the conservation of the fairy pitta (Pitta nympha), an endangered vermivore.

Quantifying the diet of endangered species is crucial for conservation, especially for diet specialists, which can be more susceptible to environmental changes. The vulnerable fairy pitta (Pitta nympha) is considered a specialist that primarily feeds its nestlings with earthworms. However, there have been few studies of the nestling diet provisioned by parents, and no assessments of earthworm proportion in the diet of adults. Our study aimed to fill these gaps, shedding light on crucial factors for conservation. Combining new observations with existing literature, we confirmed a consistent dominance of earthworms in the nestling diet, regardless of rainfall, nestling age, and time of day. We extrapolated the total earthworm consumption during a breeding event, accounting for potential variation in the availability of earthworms and their prevalence in the adult diet. We used literature-based earthworm densities in pitta habitats and our estimates of family earthworm consumption to calculate the habitat area that could provide a pitta family with the number of earthworms consumed during a breeding event. The predictions matched observed pitta home range sizes when assumed that the adult diet is comprised of approximately 70% earthworms. The results highlight the importance of earthworm-rich habitats for conservation planning of the fairy pitta. To mitigate the effects of habitat destruction, we discuss conservation practices that may involve enhancing earthworm abundance in natural habitats and providing vegetation cover for foraging pittas in adjacent anthropogenic habitats rich in earthworms. To guide conservation efforts effectively, future studies should investigate whether previously reported breeding in developed plantation habitats is due to high earthworm abundance there. Future studies should also quantify correlations between local earthworm densities, home range size, and the breeding success of the fairy pitta.

Escape behaviors in prey and the evolution of pennaceous plumage in dinosaurs.

Numerous non-avian dinosaurs possessed pennaceous feathers on their forelimbs (proto-wings) and tail. Their functions remain unclear. We propose that these pennaceous feathers were used in displays to flush hiding prey through stimulation of sensory-neural escape pathways in prey, allowing the dinosaurs to pursue the flushed prey. We evaluated the escape behavior of grasshoppers to hypothetical visual flush-displays by a robotic dinosaur, and we recorded neurophysiological responses of grasshoppers' escape pathway to computer animations of the hypothetical flush-displays by dinosaurs. We show that the prey of dinosaurs would have fled more often when proto-wings were present, especially distally and with contrasting patterns, and when caudal plumage, especially of a large area, was used during the hypothetical flush-displays. The reinforcing loop between flush and pursue functions could have contributed to the evolution of larger and stiffer feathers for faster running, maneuverability, and stronger flush-displays, promoting foraging based on the flush-pursue strategy. The flush-pursue hypothesis can explain the presence and distribution of the pennaceous feathers, plumage color contrasts, as well as a number of other features observed in early pennaraptorans. This scenario highlights that sensory-neural processes underlying prey's antipredatory reactions may contribute to the origin of major evolutionary innovations in predators.

Long-term memory of experienced jays facilitates problem-solving by naïve group members in the wild.

Long-term memory affects animal fitness, especially in social species. In these species, the memory of group members facilitates the acquisition of novel foraging skills through social learning when naïve individuals observe and imitate the successful foraging behavior. Long-term memory and social learning also provide the framework for cultural behavior, a trait found in humans but very few other animal species. In birds, little is known about the duration of long-term memories for complex foraging skills, or the impact of long-term memory on group members. We tested whether wild jays remembered a complex foraging task more than 3 years after their initial experience and quantified the effect of this memory on naïve jay behavior. Experienced jays remembered how to solve the task and their behavior had significant positive effects on interactions by naïve group members at the task. This suggests that natural selection may favor long-term memory of solutions to foraging problems to facilitate the persistence of foraging skills that are specifically useful in the local environment in social birds with long lifespans and overlapping generations.

Genetic composition of communal roosts of the Eurasian Magpie (Pica pica) inferred from non-invasive samples.

Many animal species form communal roosts in which they aggregate and sleep together. Several benefits of communal roost have been suggested, but due to lack of data on relatedness among group members, it is unknown whether these benefits can be amplified by the formation of kin-based communal roosts. We investigate the genetic composition of two winter roosts of Eurasian Magpies (Pica pica), using microsatellite markers on non-invasive samples. Using permutation tests by reshuffling the alleles presented in the roosts, we determined that individuals in the communal roosts of magpies were not more related than expected by chance, suggesting that kinship may not be a driving force for the formation of communal roosts in magpies. However, the pairwise relatedness and estimated relationship based on a maximum likelihood approach revealed that the roosts involve both kin and non-kin. Relatedness coefficients varied widely within a roost, indicating that family subgroups form a small proportion of the total number of birds in a roost. Our results suggest that ecological benefits of communal roost in animals are sufficient for the evolution of communal roosts without any involvement of kinship.

Reformulation of Trivers-Willard hypothesis for parental investment.

The Trivers-Willard hypothesis (TWH) plays a central role in understanding the optimal investment strategies to male and female offspring. Empirical studies of TWH, however, yielded conflicting results. Here, we present models to predict optimal comprehensive multi-element parental strategies composed of primary sex ratio, brood size, resource allocation among offspring, and the resultant secondary sex ratio. Our results reveal that the optimal strategy depends on sex differences in the shape of offspring fitness function rather than in fitness variance. Also, the slope of the tangent line (through the origin) to the offspring fitness function can be used to predict the preferred offspring sex. We also briefly discuss links between the model and the empirical research. This comprehensive reformulation of TWH will offer a thorough understanding of multi-element parental investment strategies beyond the classical TWH.

Social behavior mediates the use of social and personal information in wild jays.

The factors favoring the evolution of certain cognitive abilities in animals remain unclear. Social learning is a cognitive ability that reduces the cost of acquiring personal information and forms the foundation for cultural behavior. Theory predicts the evolutionary pressures to evolve social learning should be greater in more social species. However, research testing this theory has primarily occurred in captivity, where artificial environments can affect performance and yield conflicting results. We compared the use of social and personal information, and the social learning mechanisms used by wild, asocial California scrub-jays and social Mexican jays. We trained demonstrators to solve one door on a multi-door task, then measured the behavior of naïve conspecifics towards the task. If social learning occurs, observations of demonstrators will change the rate that naïve individuals interact with each door. We found both species socially learned, though personal information had a much greater effect on behavior in the asocial species while social information was more important for the social species. Additionally, both species used social information to avoid, rather than copy, conspecifics. Our findings demonstrate that while complex social group structures may be unnecessary for the evolution of social learning, it does affect the use of social versus personal information.

Wild birds recognize individual humans: experiments on magpies, Pica pica.

The ability to distinguish among heterospecific individuals has been reported in only a few animal species. Humans can be viewed as a special type of heterospecifics because individuals differ widely in behavior, ranging from non-threatening to very threatening toward animals. In this study, we asked whether wild magpies can recognize individual humans who had accessed their nests. We compared the behavior of breeding pairs toward individual humans before and after the humans climbed up to the birds' nests, and also toward climbers and non-climbers. We have evidence for (i) aggressive responses of the magpie pairs toward humans who had repeatedly accessed their nests (climbers) and a lack of response to humans who had not accessed the nest (non-climbers); (ii) a total lack of scolding responses toward climbers by magpie pairs whose nests had not been accessed; (iii) a selective aggressive response to the climber when a climber and a non-climber were presented simultaneously. Taken together, these results suggest that wild magpies can distinguish individual humans that pose a threat to their nests from humans that have not behaved in a threatening way. The magpie is only the third avian species, along with crows and mockingbirds, in which recognition of individual humans has been documented in the wild. Here, we propose a new hypothesis (adopted from psychology) that frequent previous exposure to humans in urban habitats contributes to the ability of birds to discriminate among human individuals. This mechanism, along with high cognitive abilities, may predispose some species to learn to discriminate among human individuals. Experimental tests of these two mechanisms are proposed.

Size-assortative mating and sexual size dimorphism are predictable from simple mechanics of mate-grasping behavior.

BACKGROUND: A major challenge in evolutionary biology is to understand the typically complex interactions between diverse counter-balancing factors of Darwinian selection for size assortative mating and sexual size dimorphism. It appears that rarely a simple mechanism could provide a major explanation of these phenomena. Mechanics of behaviors can predict animal morphology, such like adaptations to locomotion in animals from various of taxa, but its potential to predict size-assortative mating and its evolutionary consequences has been less explored. Mate-grasping by males, using specialized adaptive morphologies of their forelegs, midlegs or even antennae wrapped around female body at specific locations, is a general mating strategy of many animals, but the contribution of the mechanics of this wide-spread behavior to the evolution of mating behavior and sexual size dimorphism has been largely ignored. RESULTS: Here, we explore the consequences of a simple, and previously ignored, fact that in a grasping posture the position of the male's grasping appendages relative to the female's body is often a function of body size difference between the sexes. Using an approach taken from robot mechanics we model coercive grasping of females by water strider Gerris gracilicornis males during mating initiation struggles. We determine that the male optimal size (relative to the female size), which gives the males the highest grasping force, properly predicts the experimentally measured highest mating success. Through field sampling and simulation modeling of a natural population we determine that the simple mechanical model, which ignores most of the other hypothetical counter-balancing selection pressures on body size, is sufficient to account for size-assortative mating pattern as well as species-specific sexual dimorphism in body size of G. gracilicornis. CONCLUSION: The results indicate how a simple and previously overlooked physical mechanism common in many taxa is sufficient to account for, or importantly contribute to, size-assortative mating and its consequences for the evolution of sexual size dimorphism.

Evolution of switchable aposematism: insights from individual-based simulations.

Some defended prey animals can switch on their normally hidden aposematic signals. This switching may occur in reaction to predators' approach (pre-attack signals) or attack (post-attack signals). Switchable aposematism has been relatively poorly studied, but we can expect that it might bring a variety of benefits to an aposmetic organism. First, the switching could startle the predators (deimatism). Second, it could facilitate aversive learning. Third, it could minimize exposure or energetic expense, as the signal can be switched off. These potential benefits might offset costs of developing, maintaining and utilizing the switchable traits. Here we focused on the third benefit of switchability, the cost-saving aspect, and developed an individual-based computer simulation of predators and prey. In 88,128 model runs, we observed evolution of permanent, pre-attack, or post-attack aposematic signals of varying strength. We found that, in general, the pre-attack switchable aposematism may require moderate predator learning speed, high basal detectability, and moderate to high signal cost. On the other hand, the post-attack signals may arise under slow predator learning, low basal detectability and high signal cost. When predator population turnover is fast, it may lead to evolution of post-attack aposematic signals that are not conforming to the above tendency. We also suggest that a high switching cost may exert different selection pressure on the pre-attack than the post-attack switchable strategies. To our knowledge, these are the first theoretical attempts to systematically explore the evolution of switchable aposematism relative to permanent aposematism in defended prey. Our simulation model is capable of addressing additional questions beyond the scope of this article, and we open the simulation software, program manual and source code for free public use.

Ecological niche partitioning in a fragmented landscape between two highly specialized avian flush-pursuit foragers in the Andean zone of sympatry.

In the Andes, pairs of ecologically similar species are often separated by narrow elevational sympatry zones but the mechanisms mediating sympatry are not fully understood. Here, we describe niche partitioning within a sympatry zone in a fragmented Andean landscape between two closely related flush-pursue species: a high-elevation montane forest dweller, (Myioborus melanocephalus), and a mid-elevation montane forest dweller, (M. miniatus). As all flush-pursuers use very similar hunting techniques involving visual displays to flush and pursue insects in air, and benefit from being the "rare predators", ecological sorting between species in sympatry zones should allow their co-existence. We found that both species occupied vegetation resembling their typical allopatric habitats: a mosaic of pastures, clearings, and shrubs with small proportion of high trees for M. melanocephalus, and dense high forests with high proportion of trees, lower irradiance and higher humidity for M. miniatus. M. melanocephalus often foraged in bushes and at lower heights, whereas M. miniatus often foraged in tree crowns. The two species differed relatively little in their foraging technique. These results demonstrate how ecological sorting permits species of divergent elevational distributions and habitats to successfully coexist in sympatric zones where habitat diversity allows both species to find their preferred habitat.

Water strider females use individual experience to adjust jumping behaviour to their weight within physical constraints of water surface tension.

Different species of water striders match leg speeds to their body sizes to maximize their jump take off velocity without breaking the water surface, which might have aided evolution of leg structures optimized for exploitation of the water surface tension. It is not understood how water striders achieve this match. Can individuals modify their leg movements based on their body mass and locomotor experience? Here we tested if water striders, Gerris latiabdominis, adjust jumping behaviour based on their personal experience and how an experimentally added body weight affects this process. Females, but not males, modified their jumping behaviour in weight-dependent manner, but only when they experienced frequent jumping. They did so within the environmental constraint set by the physics of water surface tension. Females' ability to adjust jumping may represent their adaptation to frequent increases or decreases of the weight that they support as mating bouts, during which males ride on top of females, start or end, respectively. This suggests that natural selection for optimized biomechanics combined with sexual selection for mating adaptations shapes this ability to optimally exploit water surface tension, which might have aided adaptive radiation of Gerromorpha into a diversity of semiaquatic niches.

Trails of ants converge or diverge through lens-shaped impediments, resembling principles of optics.

Analogies across disciplines often indicate the existence of universal principles such as optimization, while the underlying proximate mechanisms may differ. It was reported recently that trails of ants refract at the border of substrates, on which walking speeds differ. This phenomenon is analogous to the travel-time-minimizing routes of light refracting at the borders between different media. Here, we further demonstrate that ant tracks converge or diverge across lens-shaped impediments similar to light rays through concave or convex optical lenses. The results suggest that the optical principle of travel time reduction may apply to ants. We propose a simple mathematical model that assumes nonlinear positive feedback in pheromone accumulation. It provides a possible explanation of the observed similarity between ant behavior and optics, and it is the first quantitative theoretical demonstration that pheromone-based proximate mechanisms of trail formation may produce this similarity. However, the future detailed empirical observations of ant behavior on impediment edges during the process of pheromone trail formation are needed in order to evaluate alternative explanations for this similarity.

Increased female resistance to mating promotes the effect of mechanical constraints on latency to pair.

Size-assortative mating, defined as a positive linear association of body size between members of mating pairs, can arise from mechanical constraints on pairing efficiency, particularly when mating success is affected by males' mate-grasping force. In this context, female resistance is predicted to have an important role in changing the threshold force necessary for males to hold females, thereby contributing to the effect of mechanical constraints. Thus, increased female resistance is expected to increase the paring success of an optimally sized male relative to the female body size (sexual size ratio = male body size/female body size = 0.86), which leads to positive size-assortative mating. However, very little is known about the extent to which female resistance affects mechanical constraints on mate grasping. Here, using the water strider Gerris gracilicornis (Hemiptera: Gerridae), we tested whether the level of female resistance affected the relationship between the sexual size ratio and latency to pair. We found that optimally sized males mated sooner than other males when females resisted a male's mating attempts. When females did not resist, an effect of sexual size ratio on latency to pair was not found. Our results thus imply that increased female resistance to male mating attempts may strengthen the pattern of size-assortative mating. We provide clear empirical evidence that female resistance to mating influences the effect of mechanical constraints on size-assortative mating under sexual conflict. This result further suggests that patterns of size-assortative mating can be altered by a variety of ecological circumstances that change female resistance to mating in many other animal species under sexual conflict.

Vigilance responding to number of conspecifics among mixed groups of cranes in demilitarized zone.

Numerous studies have addressed antipredatory benefits of mixed-species flocks of foragers, but studies on individual's vigilance as a function of group size are limited. In the Cheolwon area of the Korean Demilitarized Zone, vigilance of the subordinate White-naped cranes (Grus vipio) in 11 groups composed of conspecifics and the dominant Red-crowned cranes (Grus japonensis) was examined. Vigilance correlated negatively with group size due to negative correlation with the number of conspecifics, but not the dominant heterospecifics. This is consistent with the hypothesis that a decrease in vigilance in larger groups is due to antipredatory benefits from increased predator detection in larger groups (associated with the presence of a larger number of conspecifics). This suggested that the mechanism leads to canceling out of the otherwise expected antipredatory benefits to the subordinate species from the increased predator detection by larger group size (associated with larger number of dominants). This is one of only a few behavioral studies of these endangered crane species in the relatively inaccessible wintering area of international importance in the areas of high conservation value.

Directional raids by army ants as an adaption to patchily distributed food: a simulation model.

A typical colony of Neotropical army ants (subfamily Ecitoninae) regularly raids a large area around their bivouac by forming a narrow directional column that can reach up to one hundred meters in length. The raid is finished and then relaunched 12-17 times, each time toward different orientation. After completing all bouts the colony relocates to a new area. A hypothetical alternative to this foraging mode is raiding radially and symmetrically by expanding the search front in every direction like a circular bubble. Using an existing agent-based modeling software that simulates army ants' behavior, we compared the two possible modes of foraging in different food distributions. Regardless of the food patch abundance, the radial raiding was superior to the directional raiding when food patches had low quality, and the directional raiding was favorable when the patches were rich. In terms of energy efficiency, the radial raiding was the better strategy in a wide range of conditions. In contrast, the directional raiding tended to yield more food per coverage area. Based on our model, we suggest that the directional raiding by army ants is an adaptation to the habitats with abundance of high-quality food patches. This conclusion fits well with the ecology of army ants.

Defense sequestration associated with narrowing of diet and ontogenetic change to aposematic colours in the spotted lanternfly.

Bright colours in distasteful prey warn off predators, but processes associated with ontogenetic acquisition of warning colours and distasteful compounds have been studied in only a few organisms. Here, we study spotted lanternflies (Lycorma delicatula; Fulgoridae) that change to red colouration when they narrow their host plant preferences to primarily the tree of heaven (Ailanthus altissima; Simaroubaceae), which is chemically defended by quassinoids. In experiments, we showed that birds taste-avoided lanternflies collected on Ailanthus but not those collected on the secondary hosts. Birds also taste-avoided seeds infused with ailanthone, the main quassinoid sequestered from Ailanthus by lanternflies as shown through mass spectrometry analyses. Hence, the narrowing of host preferences by lanternflies synchronizes the timing of change to red colour with the acquisition of quassinoid defenses. A schematic graphical population-level model of these processes is provided. This is the first report of quassinoid sequestration by insects and the first evidence that Simaroubaceae plants provide defensive chemicals to insects. This is the first report of a fulgoroid insect sequestering identified chemical defenses. The results highlight the importance of the pan-tropical taxon Fulgoridae for evolutionary biology of complex aposematic strategies and for understanding the links between timing of defense sequestration, timing of host plant preference shifts, and timing of colour change.

Impact of land reclamation and agricultural water regime on the distribution and conservation status of the endangered Dryophytes suweonensis.

Knowledge about the distribution and habitat preferences of a species is critical for its conservation. The Suweon Treefrog (Dryophytes suweonensis) is an endangered species endemic to the Republic of Korea. We conducted surveys from 2014 to 2016 at 890 potentially suitable sites across the entire range of the species in South Korea. We then assessed whether D. suweonensis was found in the current and ancestral predicted ranges, reclaimed and protected areas, and how the presence of agricultural floodwater affected its occurrence. Our results describe a 120 km increase in the southernmost known distribution of the species, and the absence of the species at lower latitudes. We then demonstrate a putative constriction on the species ancestral range due to urban encroachment, and provide evidence for a significant increase in its coastal range due to the colonisation of reclaimed land by the species. In addition, we demonstrate that D. suweonensis is present in rice fields that are flooded with water originating from rivers as opposed to being present in rice fields that are irrigated from underground water. Finally, the non-overlap of protected areas and the occurrence of the species shows that only the edge of a single site where D. suweonensis occurs is legally protected. Based on our results and the literature, we suggest the design of a site fitting all the ecological requirements of the species, and suggest the use of such sites to prevent further erosion in the range of D. suweonensis.

Microbial abundance on the eggs of a passerine bird and related fitness consequences between urban and rural habitats.

Urban environments present novel and challenging habitats to wildlife. In addition to well-known difference in abiotic factors between rural and urban environments, the biotic environment, including microbial fauna, may also differ significantly. In this study, we aimed to compare the change in microbial abundance on eggshells during incubation between urban and rural populations of a passerine bird, the Eurasian Magpie (Pica pica), and examine the consequences of any differences in microbial abundances in terms of hatching success and nestling survival. Using real-time PCR, we quantified the abundances of total bacteria, Escherichia coli/Shigella spp., surfactin-producing Bacillus spp. and Candida albicans on the eggshells of magpies. We found that urban magpie eggs harboured greater abundances of E. coli/Shigella spp. and C. albicans before incubation than rural magpie eggs. During incubation, there was an increase in the total bacterial load, but a decrease in C. albicans on urban eggs relative to rural eggs. Rural eggs showed a greater increase in E. coli/Shigella spp. relative to their urban counterpart. Hatching success of the brood was generally lower in urban than rural population. Nestling survival was differentially related with the eggshell microbial abundance between urban and rural populations, which was speculated to be the result of the difference in the strength of the interaction among the microbes. This is the first demonstration that avian clutches in urban and rural populations differ in eggshell microbial abundance, which can be further related to the difference in hatching success and nestling survival in these two types of environments. We suggest that future studies on the eggshell microbes should investigate the interaction among the microbes, because the incubation and/or environmental factors such as urbanization or climate condition can influence the dynamic interactions among the microbes on the eggshells which can further determine the breeding success of the parents.