In 1985, Kummer & Goodall pleaded for an ecology of intelligence and proposed that innovations might be a good way to measure cognition in the wild. Counts of innovation per taxonomic group are now available in hundreds of avian and primate species, as are counts of tactical deception, tool use and social learning. Robust evidence suggests that innovation rate and its neural correlates allow birds and mammals to cope better with environmental change. The positive correlations between taxonomic counts, and the increasing number of cognitive and neural measures found to be associated with ecological variables, suggest that domain general processes might be more pervasive than previously thought in the evolution of intelligence.
innovation rate; tool use; social learning; tactical deception; brain size; general intelligence
We present an exploratory study of forest-living orangutan pantomiming, i.e. gesturing in which they act out their meaning, focusing on its occurrence, communicative functions, and complexities. Studies show that captive great apes may elaborate messages if communication fails, and isolated reports suggest that great apes occasionally pantomime. We predicted forest-living orangutans would pantomime spontaneously to communicate, especially to elaborate after communication failures. Mining existing databases on free-ranging rehabilitant orangutans' behaviour identified 18 salient pantomimes. These pantomimes most often functioned as elaborations of failed requests, but also as deceptions and declaratives. Complexities identified include multimodality, re-enactments of past events and several features of language (productivity, compositionality, systematicity). These findings confirm that free-ranging rehabilitant orangutans pantomime and use pantomime to elaborate on their messages. Further, they use pantomime for multiple functions and create complex pantomimes that can express propositionally structured content. Thus, orangutan pantomime serves as a medium for communication, not a particular function. Mining cases of complex great ape communication originally reported in functional terms may then yield more evidence of pantomime.
gesture; pantomime; communication; orangutan; great ape; elaboration
Male bottlenose dolphins in Shark Bay, Western Australia form two levels of alliances; two to three males cooperate to herd individual females and teams of greater than three males compete with other groups for females. Previous observation suggested two alliance tactics: small four to six member teams of relatives that formed stable pairs or trios and unrelated males in a large 14-member second-order alliance that had labile trio formation. Here, we present evidence for a third level of alliance formation, a continuum of second-order alliance sizes and no relationship between first-order alliance stability and second-order alliance size. These findings challenge the ‘two alliance tactics’ hypothesis and add to the evidence that Shark Bay male bottlenose dolphins engage in alliance formation that likely places considerable demands on their social cognition.
alliances; coalitions; social cognition
Animals titrate their behaviour against the level of risk and an individual's conspicuousness should influence decisions such as when to flee and for how long to hide. Conspicuousness will vary with variation in substrate colour. Since hermit crabs frequently change the shells they occupy, shell colour will also influence conspicuousness and to be aware of their conspicuousness would require information on both of these factors to be integrated. Reduced boldness in high-contrast shell and substrate combinations compared with situations of low contrast indicates that hermit crabs are aware of current conspicuousness. Differences between individuals remained consistent across conspicuousness levels indicating the presence of animal personalities.
hermit crab; animal personality; conspicuousness; startle response
Four phylogenetically independent lineages of frogs are currently known to sequester lipid-soluble skin alkaloids for which a dietary source has been demonstrated. We report here a remarkable fifth such instance, in Eleutherodactylus iberia and Eleutherodactylus orientalis, two species of miniaturized frogs of the family Eleutherodactylidae from Cuba. Six pumiliotoxins and two indolizidines were found in E. iberia, one of the smallest frogs in the world and characterized by a contrasting colour pattern for which we hypothesize an aposematic function. Analyses of stomach content indicated a numerical prevalence of mites with an important proportion of oribatids—a group of arthropods known to contain one of the pumiliotoxins detected in E. iberia. This suggests that miniaturization and specialization to small prey may have favoured the acquisition of dietary skin alkaloids in these amphibians.
amphibia; Eleutherodactylidae; Eleutherodactylus iberia; Eleutherodactylus orientalis; Cuba; pumiliotoxins
Since its discovery in the winter of 2005–2006, white-nose syndrome (WNS) has killed over one million little brown bats (Myotis lucifugus) in the American northeast. Although many studies have reported die-offs of bats at winter hibernacula, it is important to understand how bat mortality linked to WNS at winter hibernacula affects bat activity levels in their summer ranges. In the summer (May–August) of 2007, 2008 and 2009, we recorded echolocation calls to determine bat activity at sites along the Hudson River, NY (within approx. 100 km of where WNS was first reported). We documented a 78 per cent decline in the summer activity of M. lucifugus, coinciding with the arrival and spread of WNS. We suggest that mortality of M. lucifugus in winter hibernacula is reflected by reduced levels of activity in the summer and that WNS affects the entire bat population of an area, and not only individual hibernacula.
white-nose syndrome; Myotis lucifugus; bats; summer activity; bat mortality
Parasitic infection can modify host mobility and consequently their dispersal capacity. We experimentally investigated this idea using the ciliate Paramecium caudatum and its bacterial parasite Holospora undulata. We compared the short-distance dispersal of infected and uninfected populations in interconnected microcosms. Infection reduced the proportion of hosts dispersing, with levels differing among host clones. Host populations with higher densities showed lower dispersal, possibly owing to social aggregation behaviour. Parasite isolates that depleted host populations most had the lowest impact on host dispersal. Parasite-induced modification of dispersal may have consequences for the spatial distribution of disease, host and parasite genetic population structure, and coevolution.
dispersal; infection; microcosm
The evolutionary stability of honest signalling by offspring is thought to require that begging displays be costly, so the costs and benefits of begging—and whether they are experienced individually or by the whole brood—are crucial to understanding the evolution of begging behaviour. Begging is known to have immediate individual benefits (parents distribute more food to intensely begging individuals) and delayed brood benefits (parents increase provisioning rate to the brood), but the possibility of delayed individual benefits (previous begging affects the current distribution of food) has rarely, if ever, been researched. We did this using playback of great tit Parus major chick begging and a control sound from either side of the nest. Male parents fed chicks close to the speaker more when great tit chick begging, but not other stimuli, was played back. In contrast, there was no effect of playback at the previous visit on the chicks that male parents fed. We have thus demonstrated an immediate individual benefit to begging, but found no evidence of a delayed individual benefit in this species.
begging; honest signalling; costs; benefits; playback; great tits
During the course of a telemetry study on three species of Australian frogs (Litoria caerulea, Litoria dahlii and Cyclorana australis), we found that many of the surgically implanted transmitters had migrated into the bladder. We subsequently implanted small beads into L. caerulea and they were expelled from the body in 10–23 days. Beads implanted into cane toads (Rhinella marina) to document the process were either expelled or were enveloped into the bladder. This appears to be a unique pathway for expulsion of foreign objects from the body, and suggests that caution should be employed in telemetry studies when interpreting the separation of some animals from their transmitters as a mortality event.
anuran amphibians; trans-bladder expulsion; radio telemetry; Hylidae; Bufonidae
Ant–fungus associations are well known from attine ants, whose nutrition is based on a symbiosis with basidiomycete fungi. Otherwise, only a few non-nutritional ant–fungus associations have been recorded to date. Here we focus on one of these associations involving Allomerus plant-ants that build galleried structures on their myrmecophytic hosts in order to ambush prey. We show that this association is not opportunistic because the ants select from a monophyletic group of closely related fungal haplotypes of an ascomycete species from the order Chaetothyriales that consistently grows on and has been isolated from the galleries. Both the ants' behaviour and an analysis of the genetic population structure of the ants and the fungus argue for host specificity in this interaction. The ants' behaviour reveals a major investment in manipulating, growing and cleaning the fungus. A molecular analysis of the fungus demonstrates the widespread occurrence of one haplotype and many other haplotypes with a lower occurrence, as well as significant variation in the presence of these fungal haplotypes between areas and ant species. Altogether, these results suggest that such an interaction might represent an as-yet undescribed type of specific association between ants and fungus in which the ants cultivate fungal mycelia to strengthen their hunting galleries.
ant–fungus association; Cordia nodosa; Chaetothyriales; Hirtella physophora; myrmecophyte; population structure
An intriguing new annelid, Teuthidodrilus samae (Annelida, Cirratuliformia) gen. and sp. nov., was observed and collected during deep-water column exploration of the western Celebes Sea. The Celebes Sea is a deep pocket basin, effectively isolated from surrounding deep water, and is part of the Coral Triangle, a focal area for conservation because of its high diversity and unique geological history. Collected specimens reached 94 mm in length and possessed 10 anterior appendages that were as long or longer than the body. Two characters distinguish T. samae from other polychaetes: notochaetae forming broad, concavo-convex paddles and six pairs of free-standing, oppositely branched nuchal organs. Phylogenetic analysis of five genes and a 29-character morphological matrix showed that T. samae is an acrocirrid (primarily benthic polychaetes) belonging to the morphologically diverse swimming clade. Pelagic animals within primarily benthic clades are of particular interest in evolutionary biology, because their adaptations to life in the water column inform us of the evolutionary possibilities and constraints within the clade and indirectly of the selective pressures at work in this unfamiliar habitat. This new genus illustrates how much we have to learn about even the large, abundant inhabitants of deep-pelagic communities.
Acrocirridae; Celebes Sea; pelagic; Polychaeta; Teuthidodrilus samae
Precopulatory mate guarding (PCMG) is generally assumed to be costly for both sexes. However, males may gain by displaying long-lasting mate guarding under strong male–male competition. Surprisingly, the potential for females to benefit from being held by males has been largely overlooked in previous studies. In Gammarus pulex, an amphipod crustacean, PCMG lasts several weeks, yet females are described as bearing only cost from such male mating strategy. We investigated potential female benefits by assessing the effect of mate guarding on her intermoult duration. Unpaired females had longer intermoult duration than paired females. Intermoult duration clearly decreased when paired females engaged in early and long-lasting mate guarding. In addition, short intermoults and long-lasting mate guarding had no effect on egg number. These results highlight a potential benefit associated with PCMG for G. pulex females, suggesting that the strength of an intersexual conflict over its duration may be overestimated.
amplexus; precopula; intermoult duration; sexual conflict
Acoustic signals during intrasexual interactions may help receivers to establish the cost and benefits of engaging in a confrontation versus avoiding the cost of escalation. Although birdsong repertoires have been previously suggested as providing information during agonistic encounters, the cost (time/neural resources) of assessing large repertoires may decrease the efficiency of the signal for mutual assessment. Acoustic-structural features may, therefore, be used to enable a fast and accurate assessment during this kind of encounters. Recently, it has been suggested that the consistency of songs may play a key role during intrasexual interactions in bird species. Using a playback experiment in a colour-ringed great tit population, we tested the hypothesis that songs differing in consistency may elicit a differential response, indicating that the signal is salient for the receivers. Great tit males clearly responded more aggressively towards highly consistent songs. Our findings, together with previous evidence of increased song consistency with age in the great tit, suggest that song consistency provides information on experience or dominance in this species, and this phenomenon may be more widespread than currently acknowledged.
birdsong; song consistency; sexual selection; intrasexual; interactions
This study aimed to clarify whether infants and preschool children show susceptibility to contagious yawning, a well-known effect that has been demonstrated experimentally in older children and adults by exposing them to video sequences showing yawns. In a first study, parents kept a log of their child's yawns for a one week period. None of the log entries reported any contagious yawns by the children. Although less frequent than in older children and adults, spontaneous yawning by infants and preschoolers showed the typical morning, post-wakening peak, and an increase before bedtime in the evening. In an experimental study, infants and preschoolers watched a presentation that included many images of yawning and a repeated video clip of their own mother yawning, but there was no evidence of contagious yawning. The results suggest that, even when witnessing yawns by someone with whom they have a strong and positive emotional relationship, very young children do not show contagious yawning.
yawning; contagious yawning; infants; children; video; model
Traditional models of sexual selection posit that male courtship signals evolve as indicators of underlying male genetic quality. An alternative hypothesis is that sexual conflict over mating generates antagonistic coevolution between male courtship persistence and female resistance. In the scarabaeine dung beetle Onthophagus taurus, females are more likely to mate with males that have high courtship rates. Here, we examine the effects of exposing females to males with either high or low courtship rates on female lifetime productivity and offspring viability. Females exposed to males with high courtship rates mated more often and produced offspring with greater egg–adult viability. Female productivity and lifespan were unaffected by exposure to males with high courtship rates. The data are consistent with models of sexual selection based on indirect genetic benefits, and provide little evidence for sexual conflict in this system.
female choice; genetic benefits; sexual conflict; courtship rate
The 5th Symposium on Frugivores and Seed Dispersal, held in Montpellier (France), 13–18 June 2010, brought together more than 220 researchers exemplifying a wide diversity of approaches to the study of frugivory and dispersal of seeds. Following Ted Fleming and Alejandro Estrada's initiative in 1985, this event was a celebration of the 25th anniversary of the first meeting in Veracruz, Mexico. Frugivory and seed dispersal are active research areas that have diversified in multiple directions since 1985 to include evolution (e.g. phylogenetic diversity and dispersal adaptations), physiology (e.g. sensory cues and digestion), landscape ecology (movement patterns), molecular ecology (e.g. gene flow, genetic diversity and structure), community ecology (e.g. mutualistic interaction networks) and conservation biology (effects of hunting, fragmentation, invasion and extinction), among others. This meeting provided an opportunity to assess conceptual and methodological progress, to present ever more sophisticated insights into frugivory in animals and dispersal patterns in plants, and to report the advances made in examining the mechanisms and consequences of seed dispersal for plants and frugivores.
conservation biology; dispersal; fragmentation; frugivory; hunting; movement ecology
Penguins (Sphenisciformes) inhabit some of the most extreme environments on Earth. The 60+ Myr fossil record of penguins spans an interval that witnessed dramatic shifts in Cenozoic ocean temperatures and currents, indicating a long interplay between penguin evolution and environmental change. Perhaps the most celebrated example is the successful Late Cenozoic invasion of glacial environments by crown clade penguins. A major adaptation that allows penguins to forage in cold water is the humeral arterial plexus, a vascular counter-current heat exchanger (CCHE) that limits heat loss through the flipper. Fossil evidence reveals that the humeral plexus arose at least 49 Ma during a ‘Greenhouse Earth’ interval. The evolution of the CCHE is therefore unrelated to global cooling or development of polar ice sheets, but probably represents an adaptation to foraging in subsurface waters at temperate latitudes. As global climate cooled, the CCHE was key to invasion of thermally more demanding environments associated with Antarctic ice sheets.
counter-current; fossil; penguin; thermoregulation
The tendency of females to mate with multiple males is often explained by direct and indirect benefits that could outweigh the many potential costs of multiple mating. However, behaviour can only evolve in response to costs and benefits if there is sufficient genetic variation on which selection can act. We followed 108 mating chases of 85 North American red squirrels (Tamiasciurus hudsonicus) during 4 years, to measure each female's degree of multiple male mating (MMM), and used an animal model analysis of our multi-generational pedigree to provide what we believe is the first estimate of the heritability of MMM in the wild. Female red squirrels were highly polyandrous, mating with an average of 7.0 ± 0.2 males on their day of oestrus. Although we found evidence for moderate levels of additive genetic variation (CVA = 5.1), environmental variation was very high (CVE = 32.3), which resulted in a very low heritability estimate (h2 < 0.01). So, while there is genetic variation in this trait, the large environmental variation suggests that any costs or benefits associated with differences among females in MMM are primarily owing to environmental and not genetic differences, which could constrain the evolutionary response to natural selection on this trait.
animal model; heritability; genetic variation; multiple mating; polyandry; promiscuity
While studies of sexual selection focus primarily on female choice and male–male competition, males should also exert mate choice in order to maximize their reproductive success. We examined male mate choice in mosquitofish, Gambusia holbrooki, with respect to female size and female dominance. We found that the number of mating attempts made by a male was predicted by the dominance rank of females in a group, with dominant females attracting more mating attempts than subordinates. The number of mating attempts made by males was independent of the female size. The observed bias in the number of mating attempts towards dominant females may be driven either by straightforward male mate choice, since dominance and female fecundity are often closely related, or via the dominant females mediating male mating behaviour by restricting their access to subordinate females.
mate choice; fitness; dominance; hierarchy; fecundity
Lever systems within a skeleton transmit force with a capacity determined by the mechanical advantage, A. A is the distance from input force to a joint, divided by the distance from the joint to the output force. A lever with a relatively high A in static equilibrium has a great capacity to generate force but moves a load over a small distance. Therefore, the geometry of a skeletal lever presents a trade-off between force and speed under quasi-static conditions. The present study considers skeletal dynamics that do not assume static equilibrium by modelling kicking by a locust leg, which is powered by stored elastic energy. This model predicts that the output force of this lever is proportional to A, but its maximum speed is independent of A. Therefore, no trade-off between force and velocity exists in a lever system with spring-mass dynamics. This demonstrates that the motion of a skeleton depends on the major forces that govern its dynamics and cannot be inferred from skeletal geometry alone.
functional morphology; biomechanics; skeleton; locomotion
Because arbuscular mycorrhizal fungal (AMF) species differ in stimulating the growth of particular host plant species, AMF species may vary in their effects on plant intra-specific competition and the self-thinning process. We tested this hypothesis using a microcosm experiment with Medicago sativa L. as a model plant population and four AMF species. Our results showed that the AMF species Glomus diaphanum stimulated host plant growth more than the other three AMF species did when the plants were grown individually. Glomus diaphanum also induced the highest rate of mortality in the self-thinning plant populations. We also found a positive correlation between mortality and growth response to colonization. Our results demonstrate that AMF species can affect plant mortality and the self-thinning process by affecting plant growth differently.
arbuscular mycorrhizal fungi; Medicago sativa L.; plant mortality; plant self-thinning
Human land-use effects on species populations are minimized in protected areas and population changes can thus be more directly linked with changes in climate. In this study, bird population changes in 96 protected areas in Finland were compared using quantitative bird census data, between two time slices, 1981–1999 and 2000–2009, with the mean time span being 14 years. Bird species were categorized by distribution pattern and migratory strategy. Our results showed that northern bird species had declined by 21 per cent and southern species increased by 29 per cent in boreal protected areas during the study period, alongside a clear rise (0.7–0.8°C) in mean temperatures. Distribution pattern was the main factor, with migratory strategy interacting in explaining population changes in boreal birds. Migration strategy interacted with distribution pattern so that, among northern birds, densities of both migratory and resident species declined, whereas among southern birds they both increased. The observed decline of northern species and increase in southern species are in line with the predictions of range shifts of these species groups under a warming climate, and suggest that the population dynamics of birds are already changing in natural boreal habitats in association with changing climate.
birds; boreal; climate change; decrease; protected areas
Siblings within the same family often differ dramatically in phenotype. Some differences are attributable to initial maternal handicaps (birth or hatching asynchrony, differences in egg or neonate size and hormonal or antioxidant titre); but differences among siblings may also arise from differences in the brood-rearing environment that offspring experience. Here, I use a model system—a long-term study of nestlings in an altricial bird—to study how an initial maternal handicap, hatching asynchrony, regulates the effective social environment of siblings in the same family as measured by offspring survival. The interaction of family size and structure generated wide differences in the effective environments of siblings living in the same physical space (a nest), and reared by the same parents, in the same family structure. Social rank was the key component of the unshared environment of contemporary siblings, and was alone sufficient to generate near-maximal differences in offspring performance. Nestlings sitting side-by-side effectively lived in different worlds.
social rank; birth rank; hatching asynchrony; maternal effect; shared and non-shared environment; effective and objective environment
The Karner blue butterfly (KBB), Lycaeides melissa samuelis, is a federally protected taxon whose relationship to the Melissa blue, Lycaeides melissa, has been a point of contention during the 66 years since the KBB was first described. Using a large population-genomic dataset and a model of population divergence with migration, we investigated the relationship between the KBB and L. melissa, as well as the relationship between L. melissa and a third taxon, Lycaeides idas. We report that gene flow between the KBB and L. melissa is low, and comparable to gene flow between L. melissa and L. idas. Considering this population-genetic evidence, we conclude that the KBB is a unique evolutionary lineage that should be recognized as Lycaeides samuelis.
conservation genetics; gene flow; Lycaeides melissa samuelis; population genomics; next-generation sequencing
The endosymbiotic bacterium Wolbachia pipientis manipulates host reproduction by rendering infected males reproductively incompatible with uninfected females (cytoplasmic incompatibility; CI). CI is believed to occur as a result of Wolbachia-induced modifications to sperm during maturation, which prevent infected sperm from initiating successful zygote development when fertilizing uninfected females' eggs. However, the mechanism by which CI occurs has been little studied outside the genus Drosophila. Here, we show that in the sperm heteromorphic Mediterranean flour moth, Ephestia kuehniella, infected males transfer fewer fertile sperm at mating than uninfected males. In contrast, non-fertile apyrene sperm are not affected. This indicates that Wolbachia may only affect fertile sperm production and highlights the potential of the Lepidoptera as a model for examining the mechanism by which Wolbachia induces CI in insects.
endosymbiont; sperm heteromorphism; Lepidoptera; spermatogenesis