Successful Colonization of a Novel Urban Environment is Associated with an Urban Behavioural Syndrome in a Reed‐Nesting Waterbird

Urbanization is currently considered one of the most rapid types of global environmental change. More and more wildlife species are subject to increasing anthropogenic disturbance, but only some of them adapt to become successful urban exploiters. Although a number of fixed life history characteristics may enhance population persistence in an urbanized landscape, behavioural traits are thought to be critical for colonization of a novel urban environment. Within the class of birds, behavioural responses to urban environment have been assessed mostly for passerine species. Here, I examined behavioural alterations in a recently founded urban population of a reed‐nesting rallid species, the Eurasian coot Fulica atra. I found that urban coots adapted to nest in anthropogenically transformed littoral zones with scarce or no emergent vegetation, which precluded efficient nest concealment. Despite this, urban individuals spent, on average, less time incubating due to more frequent bouts off the nest than suburban conspecifics. This pattern was likely possible due to the relaxation of predation pressure in the colonized urban environment. Urban birds were also more aggressive, approached a human intruder at the nest more closely and were more likely to actively defend their nests. Finally, they had a reduced level of physiological stress (as measured with heterophil/lymphocyte ratio) when compared with suburban individuals. In conclusion, the behaviour of coots from the recently founded urban population was consistent with behavioural syndromes found in typical urban exploiters. The results support the central role of behavioural adaptation in successful colonization of novel environments by wildlife.     

Extirpation of alternative prey during a small rodent crash

Summary We document two episodes, in different years, of Barn Owls (Tyto alba) preying on a winter population of Burrowing Owls (Athene cunicularia) on a southern California island. The predation in each case followed a marked shift in the diet of the Barn Owls, due to the cyclic decline of their normal small mammal prey. Heavy predation in the first year resulted in the extirpation of the Burrowing Owls on the island. Such heavy predation on alternative prey species is commonly reported in cyclic predator-prey systems, however this is the first documented case of extirpation of the alternative prey. Complete elimination of any prey species by terrestrial predators is, in fact, very rare.     

Habitat utilisation and nesting behaviour of two sympatric weavers in Mbalmayo District,Cameroon

Avian ecology and behaviour data provides crucial information for bird conservation and management, but are still missing for many species, particularly in developing countries. This study examines habitat utilisation, nesting behaviour, anthropogenic disturbances and impacts on agricultural crops of Village Weaver Ploceus cucullatus and Vieillot’s Black Weaver Ploceus nigerrimus in Mbalmayo District, Cameroon. Abundance was estimated by nest counts and point counts of birds at 40 previously located colonies, where also tree metrics and anthropogenic disturbances were measured. In total, more than 1 700 birds of the two species were estimated within a 9 km2 study area, divided into two blocks and five different habitat types. Village Weavers were generally more abundant, especially in the urban habitat, whereas Vieillot’s Black Weavers were most numerous in the rural habitat. Among the tree species used for nesting, mango Mangifera indica and African border tree Newbouldia laevis had the highest nest counts, which also was positively related to canopy cover diameter. Traffic noise disturbance was associated with lower bird and nest counts, whereas the opposite was observed on farming disturbance. The relative impacts on agricultural crops were estimated for African oil palm Elaeis guineensis (38%), maize Zea mays (36%), Musa spp. (18%) and Mangifera indica (8%). The implications for long-term coexistence between farmland birds and humans are discussed, and the challenge is to minimise both kinds of bird–human interactions, i.e. anthropogenic disturbances and agricultural crop damage.     

Wintering Snowy Owls Bubo scandiacus integrate plumage colour,behaviour and their environment to maximize efficacy of visual displays

For a comprehensive understanding of the evolution of animal signals, it is necessary to understand how the performance of visual displays is maximized to get the most possible attention from receivers. We assessed whether the white plumage of Snowy Owls Bubo scandiacus functioned as a social signal and, if so, how coloration and behavioural adaptations enhance signal efficacy. Signalling theory predicts that: (1) the colour properties of plumage should vary across the body, with the brightest parts being those involved in visual display performance; (2) specific displays calling attention to or enhancing detection or conspicuousness to conspecifics should be evident; and (3) location of the signallers should be such that signal efficacy is optimized. All three predictions were supported. The brightest areas of the plumage (particularly the face, throat and upper breast) were always unspotted, and white is particularly effective in open habitats characteristic of this species. The birds displayed a specific posture and orientated toward the sun preferentially on sunny days, and Owls with the whitest (least spotted) plumage displayed more and signalled more frequently from perches on the ground, where albedo from the snow may enhance the visual display. Snowy Owls integrate coloration, behaviour and environment through habitat selection to maximize the efficacy of their visual displays.     

Decline of birds in a human modified coastal dune forest landscape in South Africa

Previous studies demonstrate that old-growth forest remnants and vegetation regenerating after anthropogenic disturbance provide habitat for birds in a human modified coastal dune forest landscape in northern KwaZulu-Natal, South Africa. However, occurrence does not ensure persistence. Based on a 13-year monitoring database we calculated population trends for 37 bird species and general trends in overall bird density in different vegetation types. We evaluated species' characteristics as covariates of population trend and assessed changes in rainfall and proportional area and survey coverage per vegetation type. 76% of species assessed have declined, 57% significantly so at an average rate of 13.9% per year. Overall, bird density has fallen at 12.2% per year across old-growth forest and woody regenerating vegetation types. Changes in proportional area and coverage per vegetation type may partly explain trends for a few species but are unlikely to account for most. Below average rainfall may have contributed to bird declines. However, other possibilities warrant further investigation. Species with larger range extents tended to decline more sharply than did others, and these species may be responding to environmental changes on a broader geographical scale. Our results cast doubt on the future persistence of birds in this human modified landscape. More research is needed to elucidate the mechanisms driving population decline in the study area and to investigate whether the declines identified here are more widespread across the region and perhaps the continent.     

Behavioural differences between two captive populations of red jungle fowl (Gallus gallus) with different genetic background,raised under identical conditions

Ex situ conservation of threatened species may lead to behavioural adaptation, which can affect success of reintroduction attempts. In previous studies, we investigated the effects of captivity on the behaviour of red jungle fowl (Gallus gallus) and found that captive populations differed behaviourally as well as genetically. The aim of the present study was to compare the behaviour of two of the previously studied populations, raised under identical conditions. Eggs were collected from birds at Copenhagen zoo (Cop) and Götala research station (Got) and were incubated and hatched together. Twenty-eight birds (16 Got and 12 Cop) were reared together and tested in eight different behavioural tests, measuring different aspects of fear-related behaviours as well as exploratory and social behaviours. The study revealed several differences in fear-related behaviours between the populations but none in exploratory or social behaviours. In general, one of the populations (Cop) showed more intense fear behaviours than the other (Got), which instead were less fearful in their behaviours. This indicates that breeding animals in captivity may lead to behavioural modifications, which can affect the outcome of reintroductions. The results further suggest that fear-related behaviours are dependent on the genetic background of the animals while social behaviours may be more influenced by the social environment. Since fear-related behaviours, such as predator avoidance and fear of humans, are essential for a life in the wild, these aspects are crucial for the breeding of animals in captivity for conservation purposes.     

Interactions between Males of the Lizard Liolaemus tenuis: Roles of Familiarity and Memory

For territorial organisms, recognition of familiar individuals can reduce the frequency and intensity of aggressive encounters (‘dear enemy’ phenomenon), stabilize social systems, and reduce the cost of territory maintenance. Here, we investigated the behavioural events displayed during contests between familiar and unfamiliar individuals in the lizard Liolaemus tenuis (Liolaemidae), a species in which males are territorial. The behaviours recorded were attack, warning, evasion, and submission, and the latencies to the first aggressive (attack or warning) behaviour. Additionally, we assessed the ability of individuals to remember a familiar conspecific after a period without social interaction. Individual males reduced and delayed aggressive behaviour directed towards socially familiar individuals compared with unfamiliar ones. These results suggest that males distinguished between familiar and unfamiliar conspecific males and are in agreement with the ‘dear enemy’ phenomenon. Other behaviours were similar in the contests between familiar and unfamiliar individuals. Recognition of familiar conspecifics was lost after 20 d without social interactions. This may be relevant for interactions with floater males or with neighbours that lose their territory and subsequently attempt to fight for their ex‐neighbour's territory.     

Consistent avoidance of human disturbance over large geographical distances by a migratory bird

Recent work on animal personalities has demonstrated that individuals may show consistent behaviour across situations and contexts. These studies were often carried out in one location and/or during short time intervals. Many animals, however, migrate and spend their life in several geographically distinct locations, and they may either adopt behaviours specific to the local environment or keep consistent behaviours over ecologically distinct locations. Long-distance migratory species offer excellent opportunities to test whether the animals maintain their personalities over large geographical scale, although the practical difficulties associated with these studies have hampered such tests. Here, we demonstrate for the first time consistency in disturbance tolerance behaviour in a long-distance migratory bird, using the common crane Grus grus as an ecological model species. Cranes that hatched in undisturbed habitats in Finland choose undisturbed migratory stop-over sites in Hungary, 1300-2000 km away from their breeding ground. This is remarkable, because these sites are not only separated by large distances, they also differ ecologically: the breeding sites are wooded bogs and subarctic tundra, whereas the migratory stop-over sites are temperate zone alkaline grasslands. The significance of our study goes beyond evolutionary biology and behavioural ecology: local effects on behaviour may carry over large distances, and this hitherto hidden implication of habitat selection needs to be incorporated into conservation planning.     

The assessment of fear in domestic hens exposed to a looming human stimulus

The behavioural and physiological responses of hens exposed to a slowly approaching human being were assessed using remote observation and radio telemetry of heart rate. Forty isolated hens of two strains and from two rearing environments were used. The sequence of behavioural changes included looking around, ceasing feeding, head shaking or complete withdrawal and finally crouching or escape behaviour. The heart rate rose from a mean level of 303 beats/min when the birds were at rest, to 465 beats/min when the cage was opened and the bird caught and held at the conclusion of the approach. The light hybrid strain showed more pronounced behavioural responses and a greater proportional rise in heart rate in the course of the approach than the medium hybrid birds, consistent with the induction of a higher level of fear. An environmental effect was also observed, pen-reared birds were more disturbed than caged ones. The similarity of the time course of the physiological and behavioural measures supports the concept of fear as an intervening variable which has simultaneous effects on heart rate and behaviour. Both may be effective in assessing fear, and they should be regarded as complementary rather than alternative measures.     

Wood mice aggressiveness and flight response to human handling: Effect of individual and environmental factors

Numerous studies have examined human disturbance repercussions on wildlife, mainly focused on the effects on behaviour, reproductive success and population dynamics. However, few studies have addressed the behaviour of prey species during and after human capture and handling and how this may correlate to individual characteristics or variation in their physical environment they inhabit. We explored wood mouse’s fleeing and aggressive behaviours in response to captures by human in their natural habitat. Eighty‐seven wood mice were caught using Sherman live traps. For each trapped individual, aggressiveness was measured as the total number of bites inflicted upon the investigator during handling time. Afterwards, each mouse was released in a two‐metre radius partially covered vegetation area that allowed visual mouse tracking by the observer and flight behaviour was registered by individual one‐zero focal sampling technique during 2 min. Both aggressiveness and fleeing behaviour were analysed regarding individual (sex, reproductive status, age) and environmental factors (habitat and season). Males, adults and breeding individuals showed heightened aggression levels. Higher aggressiveness levels were found in wood mice occupying scrubland and during summer and autumn. The flight response was exclusively explained by reproductive status, whereby breeding individuals spent more time on fast escape than nonbreeding ones. These results indicate that both individual and environmental factors seem to influence defensive behaviours in the wood mouse during and after being captured by a human. Since human disturbance shares many aspects with the predation risk, behavioural responses found to captures may likely be influenced by previous experience of individuals with predators as well as to seasonal and habitat features conditioning predators’ densities but also protection against them.     

Embryonic temperature shapes behavioural change following social experience in male leopard geckos, Eublepharis macularius

Individual variation in behavioural expression and behavioural plasticity exist in all species, and early experiences are critical determinants of both. The leopard gecko is a lizard with temperature-dependent sex determination, and in this species, embryonic incubation temperature (IncT) affects the display of social behaviours. For example, adult males hatched from eggs incubated at an IncT that produces predominantly males (male-biased IncT, 32.5°C) are more aggressive and less sexually active than males hatched from eggs at an IncT that produces predominantly females (female-biased IncT, 30°C). It is not known, however, whether IncT influences behavioural plasticity in adulthood. We assessed whether adult males hatched from eggs incubated at female- and male-biased IncTs showed different changes in territorial behaviour (scent marking in an empty test arena), anticipatory behaviours (activity, scent marking and tail vibrations in response to cues that predict the introduction of a female) and courtship behaviour following social interactions with females. We found that heterosexual social experiences increased territorial behaviour and the display of anticipatory behaviours in males from the female-biased IncT but not in males from the male-biased IncT. This difference was not due to differences in the amount of social experiences acquired, and the greater change in anticipatory behaviour in males from the female-biased IncT suggest that they acquired sexual conditioning more readily. Differences in sex steroid concentrations or neural metabolism caused by IncT could underlie this difference. These results highlight the profound implications of maternal nest site selection for offspring phenotype and plasticity in this species.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Moderate Human Disturbance of Rain Forest Alters Composition of Fruiting Plant and Bird Communities

Forests worldwide are experiencing rapid environmental change due to human activity. We aimed to increase understanding of anthropogenic impacts on community composition and species interactions. In a natural experiment, we asked whether subsistence human land use has altered the community composition of a Neotropical rain forest on the island of Tobago, in the West Indies. We surveyed fruiting plants and birds in three adjacent habitat types that varied in level of disturbance, and used multivariate analyses to determine whether changes in the plant community were associated with differences in avifauna composition. The three forest habitats had similar plant and bird diversities, yet markedly different species compositions and abundances. Primary forest had the most diverse plant community, while disturbed habitats had a more homogeneous plant composition. Primary and disturbed forest had distinct community compositions, with canopy cover and the relative abundance of plant types explaining 83 percent of the variation in bird species assemblages. Seemingly moderate human disturbance has led to substantial changes in the plant and bird assemblages of Tobago's rain forest, outside of a protected reserve. Our study highlights the direct links between human disturbance and the structure of rain forests, underscoring the impact of even moderate activity on community composition.     

Use of artificial arenas to predict the social organisation of halictine bees: Data for fourteen species from Chile

I studied the behavioural profiles of fourteen species of Chilean halictine bee using the circle tube apparatus. Interactions were classified as aggressive, avoidance or cooperative. One species, Corynura chloris, is believed to be semisocial and its behaviour was largely aggressive, Ruizantheda mutabilis is known to be communal and its behaviour was predominantly cooperative whereas Penapis toroi is a member of a subfamily from which only solitary behaviour is known and it primarily exhibited avoidance behaviours. As a result, I suggest that the relative frequency of these different behaviours may be useful in establishing the social organisation of species. Based upon comparisons of behavioural profiles, I predict the social organizations of the remaining species and suggest that at least some of the bees in the populations of Ruizantheda proxima, Caenohalictus dolator, Ca. species A and Pseudagapostemon pississi investigated exhibit communal behaviour; those of Co. patagonica and Co. herbsti likely have a reproductive division of labour, probably within semisocial societies, and that the populations of Co. corinogaster, Co. melanocladius, Ca. pygosinuatum, Ca. cuprellus and Lasioglossum aricense studied are probably solitary. Phylogenies suggest that solitary behaviour in Ca. pygosinuatum and Ca. cuprellus is reversed from communal behaviour and solitary behaviour in L. aricense may be a reversal from eusociality. The response “back and follow” is not indicative of dominance or cooperation as it correlated only with the frequency of avoidance interactions among the taxa studied. I discuss the utility of the circle tube apparatus in the identification of populations worthy of more detailed sociobiological investigation. Received 10 August 2005; revised 8 March 2006; accepted 21 March 2006.     

Evaluating cropland in the Canadian prairies as an ecological trap for the endangered Burrowing Owl Athene cunicularia

Anthropogenic development may influence the choices animals make and their resulting reproductive success and survival. If such choices are maladaptive, the impact of anthropogenic change can be catastrophic to small or declining populations. Over the past century, Canada's prairie landscapes have been altered dramatically, with over two-thirds of its native grasslands now having been converted to cropland. The decline of the endangered Burrowing Owl Athene cunicularia population is assumed to have resulted from this landscape change, yet no causal link has been demonstrated. One hypothesis to explain this population decline is that owls get caught in an ecological trap, whereby they prefer to establish nests at the start of each breeding season in landscapes that later confer lower reproductive success. Agricultural landscapes represent a plausible potential ecological trap because the short and sparse vegetation in annual crops (seeded each spring) is predicted to be attractive nesting/foraging habitat for Burrowing Owls when they arrive from northward migration, yet crops become substantially taller and denser over the growing season so prey are predicted to become less accessible by the time broods have hatched. We tested this ecological trap hypothesis in a 3-year study, involving 379 Burrowing Owl pairs, across the agricultural landscapes of southern Alberta and Saskatchewan. In support of the hypothesis, Burrowing Owls did prefer to settle in breeding home-ranges that contained a higher proportion of cropland, and their prey-delivery rates during brood-rearing were lower at nests with a higher proportion of cropland growing in the surrounding landscape. However, in contradiction to a key prediction, the number of fledglings produced (range = 0–9) was higher, not lower, for pairs with more actively growing cropland in their landscapes. Therefore, the decline of the Burrowing Owl in Canada does not appear to result from cropland forming an ecological trap during the breeding season. We also found a significant positive relationship between the amount of summer fallow within Burrowing Owl home-ranges and the quantity of vertebrate prey delivered to the nest and the number of juveniles fledged, highlighting the importance of this declining land use in raptor conservation.     

Interspecific variation in the response of fish to anthropogenic noise

1. Elevated levels of anthropogenic noise, especially those observed through boating activity, can negatively impact fish species, but it remains unclear which species are most affected and which behavioural metrics are best used in assessing fish responses to underwater noise. The effects of boat sounds on freshwater species are of particular interest because freshwater environments are less studied than the marine realm despite comparably high levels of biodiversity.
2. In the current study, we examine the behavioural responses to boat noise in two freshwater species that differ in their hypothesised response to sound inputs: the spottail shiner (Notropis hudsonius), a species with known hearing specialisations, and the bluegill sunfish (Lepomis macrochirus), a species with more generalised hearing capabilities. Fish were presented with boat noise in a laboratory setting, and their swimming, escape and foraging behaviours were assessed to examine differential responses in relation to hypothesised hearing abilities.
3. Both species showed a decrease in general swimming behaviours but an increase in erratic movements in response to boat noise, indicative of stress responses for both species. Despite the similarities in response based on swimming behaviours however, only spottail shiners exhibited true escape responses to the onset of the noise stimulus, suggesting a more extreme reaction in the species with a more refined hearing ability.
4. Taken together, these results show that freshwater fish can respond to increased levels of anthropogenic noise, but that the severity of the response may differ based on auditory structures and therefore presumed hearing ability. The differences seen between behavioural metrics used (swimming vs. escape responses) also demonstrate how care must be taken in choosing a metric when developing exposure guidelines for underwater sound exposures, as different metrics could lead to differential impact assessments.

     

A meta‐analysis of human disturbance impacts on Antarctic wildlife

Evidence‐based assessments are increasingly recognized as the best‐practice approach to determine appropriate conservation interventions, but such assessments of the impact of human disturbance on wildlife are rare. Human disturbance comprises anthropogenic activities that are typically non‐lethal, but may cause short‐ and/or longer‐term stress and fitness responses in wildlife. Expanding human activity in the Antarctic region is of particular concern because it increases the scope and potential for increased human disturbance to wildlife in a region that is often thought of as relatively untouched by anthropogenic influences. Here, we use a meta‐analytical approach to synthesise research on human disturbance to wildlife over the last three decades in the Antarctic and sub‐Antarctic region. We combine data from 62 studies across 21 species on the behavioural, physiological and population responses of wildlife to pedestrian, vehicle and research disturbances. The overall effect size indicated a small, albeit statistically significant negative effect of disturbance (?0.39; 95% CI: ?0.60 to ?0.18). Negative effects were found for both physiological and population responses, but no evidence was found for a significant impact on wildlife behavioural responses. Negative effects were found across pedestrian, vehicle and research disturbances. Significant and high among‐study heterogeneity was found in both disturbance and response sub‐groups. Among species, it remains unclear to what extent different forms of disturbance translate into negative population responses. Most current guidelines to limit wildlife disturbance impacts in Antarctica recommend that approaches be tailored to animal behavioural cues, but our work demonstrates that behavioural changes do not necessarily reflect more cryptic, and more deleterious impacts, such as changes in physiology. In consequence, we recommend that pedestrian approach guidelines in the Antarctic region be revisited. Due to the high heterogeneity in effects, management guidelines for different sites and species will need to be developed on a case‐by‐case basis, ideally in conjunction with carefully designed experiments. Guidelines to reduce the impact of research activities per se require development to reduce the potential impacts of conducting research. We identify research questions that, if answered, will further improve the evidence base for guidelines to manage human disturbance in Antarctica.     

Disturbance-dependent Yellow-legged Gull (Larus michahellis) predation on Larid chicks decreases with chick age

Predation is one of the key factors shaping the dynamics of animal populations. In birds, nest loss due to predation can be a significant cause of low reproductive success. Ground-nesting birds are among the bird groups most susceptible to predation, mainly because their nests are easily accessible to a broad suite of potential predators. For these birds, anthropogenic disturbances can generate changes in nest predation risk by altering their antipredator behaviour and also by altering the behaviour of the predator species, i.e. the predator becoming much more aware of predation opportunities due to frequent disturbances and/or motivated to repeat predation attempts when some are successful. To date, most previous studies investigating this have focused on a single effect, either predation or disturbance, on chick survival. It remains unknown how the risk of predation with and without disturbance varies with chick age. In this study, we used behavioural observations to assess how the interaction between predators and disturbance affects predation risk in chicks and how this interacts with chick age. Specifically, we investigated the effect of disturbance caused by humans and stray dogs on the predation of Slender-billed Gull Chroicocephalus genei chicks by Yellow-legged Gulls Larus michahellis, and whether this depended on the age of the chicks. Our results revealed that disturbance had a significant positive effect on predation measures of Slender-billed Gull chicks by Yellow-legged Gulls, but that this effect was mediated both by disturbance type and the age of chicks. Stray dogs entering the colony had a stronger disturbance effect on chicks than passing humans, increasing predation risk by Yellow-legged Gulls. Our results also showed that chick age interacts with disturbance type to determine the predation risk. This is probably mediated by chicks' capacity to escape predation by gathering in a single large crèche that runs into the water when disturbed. To preserve Slender-billed Gull colonies in one of its few remaining breeding sites in Tunisia, and as gulls tend to react even when the disturbance occurs relatively far from the colonies, it is crucial to (1) restrict human access to dikes and islets where large colonies breed and (2) construct artificial islets attractive to gulls and inaccessible to stray dogs.     

Size and sex matter: reproductive biology and determinants of offspring survival in Gazella marica

For prey, many behavioural traits are constrained by the risk of predation. Therefore, shifts between warning and cryptic coloration have been suggested to result in parallel changes in several behaviours. In the present study, we tested whether changes in chromatic contrast among eight populations of the strawberry poison‐dart frog, Dendrobates pumilio, co‐vary with behaviour, as expected if selection is imposed by predators relying on visual detection of prey. These eight populations are geographically isolated on different island in the Bocas del Toro region of Panama and have recently diverged morphologically and genetically. We found that aggression and explorative behaviour were strongly correlated and also that males tended to be more aggressive and explorative if they belonged to populations with conspicuously coloured individuals. We discuss how evolutionary switches between predator avoidance strategies and associated behavioural divergence between populations may affect reproductive isolation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Absence of within-colony kin discrimination in behavioural interactions of swarm-founding wasps

Within-colony kin discrimination has not been demonstrated conclusively for any social insect, perhaps partly because highly polymorphic genetic markers necessary to assess within-colony relatednesses have only recently become available. We use microsatellite loci to investigate within-colony kin discrimination in behavioural interactions in the neotropical multiple-queen wasp, Parachartergus colobopterus. Within-colony kin discrimination would be particularly advantageous in this species since average genetic relatedness among colony members overall is low (0.32 =/- 0.06), compared to the relatedness value between full sisters of 0.75. Using seven colonies of individually marked females, we recorded behavioural interactions that were cooperative (222 grooming, 2438 feeding), aggressive (511 body or wing biting, 240 mandible biting) or neutral (1676 antennating). We expected cooperative behaviours to favour closer kin and aggressive behaviours to be directed towards more distant kin, but found that none of the behaviours we investigated showed discrimination on the basis of relatedness. We could have detected a difference in relatedness values of as little as between 0.03 and 0.12, depending on the behaviour being analysed. Thus, we found no evidence for kin discrimination in within-colony behaviour in this species.     

Effects of Social Structure on the Behaviour and Performance of Alternative Reproductive Phenotypes in Male Rock Shrimp, Rhynchocinetes typus

Males that adopt alternative mating tactics within a conditional strategy often undergo costly morphological changes when switching to the next phenotype during ontogeny. Whether costs of changing to a subsequent reproductive phenotype are outweighed by a higher mating probability may depend on the frequencies of different phenotypes in a group of competitors. Benefits and costs associated with different phenotype frequencies depend on interactions within and between alternative phenotypes, but the underlying behavioural mechanisms have rarely been studied. Herein, we used the rock shrimp Rhynchocinetes typus as a model: ontogenetic male stages of this species differ in morphological and behavioural traits that indicate alternative reproductive phenotypes. The small, subordinate, male stage (typus) develops via several intermediate stages (intermedius) to the dominant male stage (robustus): in competitive interactions the typus males usually employ the sneaking tactic, while the robustus males invariably employ the monopolizing fighter tactic. In laboratory experiments, we manipulated phenotype frequencies to examine whether there are frequency‐dependent effects on searching behaviour, aggressiveness and mating probability. With increasing frequency of robustus males, the rate of aggressive interactions among them increased. Furthermore, robustus males increased walking velocity when more than one robustus male was present. In contrast, typus males did not adjust their searching or aggressive behaviour. The increase of aggressive interactions among robustus males provided more opportunities for typus males to seize a temporarily unguarded female. While typus males exploit fights among robustus males that produce mating opportunities for them, robustus males benefit from typus males, which reveal the presence of receptive females. We suggest that each phenotype benefits from the presence of the other phenotype and suffers costly interference among individuals of the same phenotype. Whether frequency‐dependent effects on the mating probability of subordinates also affect their ontogenetic switchpoint should be examined in future studies.