Immigration pattern and success in red squirrels

Summary We studied the characteristics of immigrants and the effects of immigration on reproductive activity and spacing behaviour in red squirrels living in high-quality woodlands. Male immigration peaked in spring, female immigration in autumn. There was no sex bias in dispersal distance of local recruits or in the proportion of male/female immigrants, but more subadults than adults immigrated on the study plots. Hence, hypotheses explaining sex-biased dispersal were irrelevant in explaining immigration patterns in our study populations. Immigrant females were not in breeding condition, nor had they produced a litter prior to immigration. Hence breeding dispersal did not occur. Red squirrels are promiscuous, and females defend intrasexual territories while males have overlapping home ranges with a dominance hierarchy (Wauters et al. 1990; Wauters and Dhondt 1992). Site fidelity is very important to reproductive success and most parents still have a high residual reproductive value after having produced a litter. Under such circumstances, the resident fitness hypothesis (RFH; Anderson 1989) predicts that parents can benefit by forcing emigration of offspring if the latter are likely to find nearby vacancies. The settlement pattern of successful immigrants, which had a higher probability of becoming established when they had high body mass and when they were settling in plots with reduced intrasexual competition, agreed with the RFH and with the proximate dispersal mechanism suggested by Gliwicz (1992), that dispersal tendency in both sexes depends on the degree of intrasexual competition under local conditions. The fact that close inbreeding was never observed could indicate that random immigration of both sexes, within the social environment of a partly territorial, relatively long-lived species, has evolved not only to reduce competition for resources between parents and offspring but also as an inbreeding avoidance mechanism. Correspondence to: L. Wauters     

Evolutionary aspects of perfume collection in male euglossine bees (Hymenoptera) and of nest deception in bee-pollinated flowers

Summary A fascinating pollination system has been evolved between perfume producing flowers and perfume collecting male euglossine bees in the neotropics. Detailed investigations have contributed to an understanding of the interactions between euglossine males and flowers as a pollination system. The role which the collected perfume plays in the reproductive behaviour of euglossine bees is not fully understood. A favoured hypothesis suggests that the collected fragrances are used as precursors for male sex pheromones and thus serve to attract conspecific males or females. It is not known how perfume collection behaviour evolved. Here, an evolutionary approach presents a new hypothesis which suggests that the evolution of perfume collection in euglossine males is based upon pre-existing signals which were attractive to females and males. It is further suggested that, at the evolutionary outset, flowers mimicked nest sites to deceive nest-seeking euglossine bees. In addition, a comparative study was undertaken on the phenomena of nest-mimicking flowers in related bee families.     

Labial gland marking secretion in males of two Scandinavian cuckoo bumblebee species (genusPsithyrus)

Summary Chemical components of the cephalic labial gland secretion, which most likely is used as a precopulatory attracting signal, were identified in males of the cuckoo bumblebeesPsithyrus vestalis (Geoffroy) andP. bohemicus (Seidl.) by gas chromatography-mass spectrometry (GC-MS). The former species has geranylcitronellyl acetate as main component, plus in decreasing amounts geranylcitronellol, an eicosadienal, an eicosenol, and an eicosenal. Another 17 fatty acid derivatives are present in minor amounts. The labial secretion ofP. vestalis differs distinctly in its chemistry from that of all of the 8 other ScandinavianPsithyrus species. In the taxonomically closely related (same subgenus)P. bohemicus, the labial gland secretion contains 16 fatty acid derivatives, of which 12 (75%) are present also in the secretion ofP. vestalis. Thus, both the labial gland secretions, and morphological traits lend support to the hypothesis that the two species share a common ancestor. The reproductive isolation between them is, at least in part, supported by the inclusion of a second biosynthetical pathway (the mevalonic acid pathway producing terpenoids) inP. vestalis, in addition to the pathway common to both species (the polyketide pathway producing fatty acid derivatives).     

Behaviour of stream-dwelling brown trout towards odours present in home stream water

Summary Preference behaviour patterns of wild brown trout (Salmo trutta L.) towards odours in their home-stream water were measured in laboratory experiments. Stream-dwelling, sexually mature trout were captured by electrofishing in different sections of a stream, transferred to a hatchery, and exposed to water sampled from various stream sections and to neutral water, scented by fish captured in the same stream sections. The fish preferred stream water originating from their home stream over that from a neighboring stream. Among water samples from their home stream, they preferred water from the home sections over water from distant sections both upstream and downstream. In most cases they also preferred neutral water scented by other mature fish captured in their home section over water scented by fish from the distant sections. Trout from two neighboring stream sections showed indifferent responses towards water and fish from the adjacent section. The attractive properties of stream water sampled from home sections coincided with those obtained with neutral water scented by fish from the same sites. Accordingly, the attractive components in stream water may be intraspecific odours derived from separate spawning demes of fish present in local areas of the stream.     

Social influence on the correlation between behaviours in young-of-the-year perch

The connection between risk-taking behaviour and exploratory behaviour in young-of-the-year perch (Perca fluviatilis) was studied in aquarium experiments to see whether individual behaviour patterns could be identified in this species and also to investigate how individual behaviour is influenced by their social environment. Risk-taking was defined as the time spent foraging in an open area vs hiding in the vegetation in the presence of a piscivore. Explorative behaviour was measured as latency to enter a passage leading to an unknown area. Groups of four fish were used for the observations, and both behaviours measured were positively correlated with the mean scores of these behaviours in the other group members. Risk-taking and explorative behaviours were correlated only when data was adjusted for the behaviour of the other group members. Individuals that spent more time in the open than their companions also tended to be faster than the others to enter the passage to the unknown area and vice versa. The results indicate that there are consistent individual differences in boldness in perch, but also that behaviour could be modified according to the behaviour of group members.     

The tactics of mutual mate choice and competitive search

A model of mutual mate choice is described, formulated as a dynamic game, which yields predictions about mating behaviour under the influence of time constraints, choice costs and competition for mates. These variables were examined because they may result in a change in the distribution of qualities among unmated individuals of both sexes over the course of the breeding season. The model predicts that mutual choice gives rise to assortative mating, although high costs of choice and/or inaccurate assessment both lead to lower overall correlations between the qualities (or the attractiveness) of mates. When all individuals are present from the start of the breeding season, the correlation between the qualities of individuals pairing at a given time declines throughout the season, so that mates are more closely matched among individuals who pair early than among those who pair late (and extra-pair copulation may thus be more common among the latter). Delayed arrival of lower-quality individuals may, however, lead to an increase in this correlation with time during the early part of the season. The mean quality of unmated males and females declines over time, because more attractive individuals tend to mate sooner. As a result of this decline, and because of time constraints, superior individuals become less choosy as the season progresses. If choice is costly, however, then inferior individuals become more selective with time during the early part of the season, and the level of choosiness peaks later for such individuals. Received: 28 March 1996 / Accepted after revision: 21 September 1996     

Deception by helpers in cooperatively breeding white-winged choughs and its experimental manipulation

White-winged choughs (Corcorax melanorhamphos) are obligate cooperative breeders, living in groups which may contain up to 20 birds. Although breeding is dominated by a single pair, all birds contribute to rearing young, including the provisioning of nestlings. However, some birds which have carried food to the nest, even to the point of placing the food in the gaping mouth of a nestling, consume the food themselves rather than provision the nestlings. Birds which fail to feed nestlings are typically young, and are only likely to fail to deliver food when they cannot be observed by other group members. Birds which have just failed to deliver food are more likely to engage in alternative helping behaviours such as allopreening the nestlings than are helpers which have just delivered food in the conventional manner. Failure to deliver food is almost eliminated when foraging constraints are experimentally reduced by supplemental feeding of the group. Collectively these observations suggest that young white-winged choughs act deceptively by simulating helping behaviours without sacrificing food supplies. Received: 24 January 1997 / Accepted after revision: 6 June 1997     

How Kentish plovers,<Emphasis Type="Italic"> Charadrius alexandrinus</Emphasis>, cope with heat stress during incubation

Biparental incubation is frequent among shorebirds and is expected when the survival prospects of offspring increase relative to uniparental incubation. To understand why this occurs, it is important to identify the factors that constrain uniparental incubation. It is assumed that birds choose nesting sites that provide an appropriate microclimate for incubation. Many shorebirds nest in sites with no or little cover, where ambient temperatures at ground level might be >50°C during very hot days. Shorebirds nest in exposed sites because predation risk on incubating adults is higher in covered sites. In hot environments, incubating shorebirds might experience heat stress in exposed sites, and this may compromise nesting success if adults are unable to attend their nests continuously, limiting the possibilities of uniparental incubation and thus the expression of a sexual conflict over incubation. The operative temperatures of Kentish plovers (Charadrius alexandrinus) were recorded in exposed and covered sites, and the thermal behaviour and incubating tactics of pair members were studied in a hot environment. During the hottest part of the day, there was a difference of 10–15°C in the operative temperatures of plovers between covered and exposed sites. Plovers in covered sites did not exhibit any thermoregulatory behaviour indicative of thermal stress, probably because the thermal range encountered by them in such places during most of the daytime was close to the thermo-neutral zone. The frequency with which plovers in exposed sites exhibited thermoregulatory behaviour was related to ambient temperature. Under very hot conditions, incubating birds were probably unable to maintain homeostasis for long periods and pair members resorted to shortening incubation bouts. Female Kentish plovers mainly incubate in the daytime and males during the night. However, the probability of diurnal incubation by males increased with ambient temperature in exposed nests, but not in covered ones. In fact, the frequency of participation in diurnal incubation by males was greater in exposed than in covered sites, suggesting that the participation of males in diurnal incubation may be related to the inability of females to stay at the nest during long periods when the ambient temperature is high. Even after resorting to shortened incubation bouts, the plovers may be unable to attend their nests continuously during heat waves, and the nests may be deserted. The propensity of plovers to desert their nests was affected by proximity to water, with nests located close to water being deserted less frequently. It seems likely that susceptibility to thermal stress changed in relation to proximity to water because in sites close to water it was possible to belly-soak, which would allow a more continuous nest attendance. Therefore, despite the adoption of behavioural solutions to face heavy heat loads, nesting success was vulnerable to these solutions because heat stress during extended periods may constrain parental nest attendance, and this may limit the opportunities for sexual conflicts over incubation.Communicated by J. Graves     

Futurecoast: Predicting the future coastal evolution of England and Wales

Analysis of shoreline change is often based exclusively upon the littoral cell concept and modelling of hydrodynamic processes. The Futurecoast study has considered fresh approaches to assessing shoreline evolution, which have been used to provide an analysis of future long-term evolution for the entire shoreline of England and Wales. This has been based upon an improved understanding of coastal systems and their behavioural characteristics. The study has included a range of supporting studies, focussing upon maximizing use of existing information and experience. A number of additional data sets have also been produced. The integration of leading expertise from different areas of coastal research to collectively consider this information has been the foundation for the study. The key outputs from this research are: (1) Improved understanding of coastal behaviour; (2) Assessment of future shoreline evolution; (3) Supporting information and data; (4) Delivery of results on an interactive CD-ROM.     

Social foraging and dominance relationships: the effects of socially mediated interference

In socially foraging animals, it is widely acknowledged that the position of an individual within the dominance hierarchy of the group has a large effect upon its foraging behaviour and energetic intake, where the intake of subordinates can be reduced through socially mediated interference. In this paper, we explore the effects of interference upon group dynamics and individual behaviour, using a spatially explicit individual-based model. Each individual follows a simple behavioural rule based upon its energetic reserves and the actions of its neighbours (where the rule is derived from game theory models). We show that dominant individuals should have larger energetic reserves than their subordinates, and the size of this difference increases when either food is scarce, the intensity of interference suffered by the subordinates increases, or the distance over which dominant individuals affect subordinates increases. Unlike previous models, the results presented in this paper about differences in reserves are not based upon prior assumptions of the effects of social hierarchy and energetic reserves upon predation risk, and emerge through nothing more than a reduction in energetic intake by the subordinates when dominants are present. Furthermore, we show that increasing interference intensity, food availability or the distance over which dominants have an effect also causes the difference in movement between ranks to increase (where subordinates move more than dominants), and the distance over which dominants have an effect changes the size of the groups that the different ranks are found in. These results are discussed in relation to previous studies of intra- and interspecific dominance hierarchies.