Reproductive stage and phonotactic preferences of female midwife toads (<Emphasis Type="Italic">Alytes cisternasii</Emphasis>)

We studied the phonotactic response of Alytes cisternasii females at different reproductive stages (before and after oviposition) toward two male call characteristics (call rate and frequency) differing in variability, as well as in the strength of female preference. We measured female phonotactic response over several repetitions by means of the frequency of selection of the population-preferred alternative, consistency of choice, frequency of reciprocal calling activity, frequency of null trials, and average approach time. Our results showed that female mating preferences in A. cisternasii are highly influenced by the variability of male call characteristics, while only female motivation is related to reproductive stage.Communicated by T. Czeschlik     

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     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

Mate guarding,copulation strategies and paternity in the sex-role reversed,socially polyandrous red-necked phalarope<Emphasis Type="Italic"> Phalaropus lobatus</Emphasis>

In a recent review, Westneat and Stewart (2003) compiled evidence that extra-pair paternity results from a three-player interaction in which sexual conflict is a potent force. Sequentially polyandrous species of birds appear to fit this idea well. Earlier breeding males may attempt to use sperm storage by females to obtain paternity in their mates subsequent clutches. Later-breeding males may consequently attempt to avoid sperm competition by preferring to pair with previously unmated females. Females may bias events one way or the other. We examined the applicability of these hypotheses by studying mating behavior and paternity in red-necked phalaropes (Phalaropus lobatus), a sex-role reversed, socially polyandrous shorebird. Male red-necked phalaropes guarded mates more strongly than other shorebirds. Males increased within-pair copulation attempts during their mates fertile period, and maintained or further increased attempts towards the end of laying, suggesting an attempt to fertilize the females next clutch; these attempts were usually thwarted by the female. Paired males sought extra-pair copulations with females about to re-enter the breeding pool. Multilocus DNA fingerprinting showed that 6% of clutches (4/63) each contained one chick sired by a male other than the incubator, producing a population rate of these events of 1.7% (n=226 chicks). Male mates had full paternity in all first clutches (n=25) and 15 of 16 monogamous replacement clutches. In contrast, 3 of 6 clutches of second males contained extra-pair young likely fathered by the females previous mate. Previously mated female phalaropes may employ counter-strategies that prevent later mating males from discriminating against them. The stability of this polyandrous system, in which males provide all parental care, ultimately may depend on females providing males with eggs containing primarily genes of the incubating male, and not a previous mate.Communicated by M. Webster     

Effects of kinship on territorial conflicts among groups of lions,<Emphasis Type="Italic"> Panthera leo</Emphasis>

Inclusive fitness theory predicts that cost of tolerant behaviour during competitive interactions is lower for relatives than for nonrelatives. Many studies have examined the effect of relatedness on behaviour within social groups. In contrast, kin selection acting among groups has received less attention. The genetic structure of African lion (Panthera leo) populations creates a strong possibility that kin selection among groups modifies behaviour during group conflicts. We used playback experiments and genetic data to investigate the importance of relatedness during simulated territorial disputes in lions. However, we found no effect of relatedness on territorial behaviour. Degree of relatedness did not affect the decision to approach simulated intrusions, nor did it affect the behaviour during approaches. The decision to approach was instead affected by position within the territory and consecutive playback number (a measure of habituation). For playbacks that did elicit an approach, the speed of response was not detectably affected by relatedness, but was affected by odds (the ratio of residents to intruders), number of intruders, number of bouts, presence of cubs, position within the territory, temperature and playback number. Although responses were unaffected by relatedness, it remains possible that other aspects of behaviour during natural encounters among prides are affected by kin selection.Communicated by L. Sterck     

Manipulation of nest-box density affects extra-pair paternity in a population of blue tits (<Emphasis Type="Italic">Parus caeruleus</Emphasis>)

We tested the effect of manipulation of breeding density on the occurrence of extra-pair paternity in a blue tit (Parus caeruleus) population during two consecutive years. In a homogeneous oak forest, nest-box manipulation provided a high density plot (plot A, 1.10 and 1.32 pairs/ha) and a low density plot (plot B, 0.43 and 0.46 pairs/ha). Microsatellite analysis on 91 broods revealed a higher proportion of extra-pair paternity in broods in plot A (mean of 17.2%) than in plot B (mean of 11.4%). A correlative approach showed that the proportion of extra-pair young in broods was affected by the number of breeding neighbours within 100 m around the nest-box, by the distance to the nearest breeding neighbour and by an additional plot effect. However, the nearest neighbours accounted for only 39.3% of extra-pair paternities and distance to extra-pair fathers was significantly higher than the nearest neighbour distance in both plots. This implies that the effect of density on the occurrence of extra-pair paternities is associated with active female choice to enhance the brood fitness. Although there were more extra-pair young in broods when density was high, the number of extra-pair fathers did not increase and stayed close to one. We suggest that density increases the cost of mate guarding by males, thereby increasing the possibility for females to solicit extra-pair paternities to the cuckolding male they have chosen. Finally, we discuss why correlatives approaches do not always show evidence for an effect of breeding density on extra-pair paternity occurrence.Communicated by M. Soler     

Dufour gland secretion in the harvester ant genus <Emphasis Type="Italic">Pogonomyrmex</Emphasis>

Summary. Dufour gland secretions of the harvester ants Pogonomyrmex barbatus, P. rugosus and P. maricopa were investigated. The glandular blends exhibit species specific patterns, but more importantly, based on indiviual profiles of Dufour gland contents of colony members it is possible to separate the colonies from each other in each species. It is possible that these collective colony specific cues serve as longer lasting trunk route markers. Behavioral tests demonstrate that workers of P. rugosus prefer homing routes marked by members of their own colony to routes marked by a foreign conspecific colony.     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

Effects of body mass,age, dominance and parasite load on foraging time of bighorn rams,<Emphasis Type="Italic"> Ovis canadensis</Emphasis>

In sexually dimorphic ungulates, males generally spend less time foraging than females, possibly because of difference in body mass or because of the energetic requirements of lactation. The relationship between body size and foraging time has received little attention at the intra-specific level, because few studies have documented activity budgets for individuals of known size. Bighorn rams are a good model to explore how body mass affects foraging time, because they range in mass from 55 to 140 kg. We examined how the foraging time of bighorn rams varied according to individual characteristics. We observed rams in a marked population and constructed time budgets during the 3 months preceding the rut. We determined ram social rank based on agonistic encounters and collected fecal samples to count lungworm larvae. Time spent foraging was negatively correlated with body mass. After accounting for age differences, larger rams spent less time foraging and more time lying than smaller rams. Among rams aged 6–12 years, dominants spent less time feeding than subordinates, while fecal output of lungworm larvae was negatively correlated with foraging time for rams of all ages. Body mass accounts for much of the individual variation in foraging time, suggesting that sexual dimorphism is important in explaining differences in feeding time between males and females.Communicated by P. Heeb     

Genetic mating system and timing of extra-pair fertilizations in the Kentish plover

It is still unclear why females in many bird species pursue extra-pair copulations. Current hypotheses focus mainly on indirect benefits such as obtaining particular good genes for their offspring or maximizing genetic compatibility between themselves and the father of their offspring. Supporting the latter, a recent study of shorebirds suggests that extra-pair matings may function to avoid the negative effects of genetic similarity between mates. Here, we further investigate genetic parentage in the Kentish plover, Charadrius alexandrinus, a shorebird with a highly variable social mating system. DNA fingerprinting revealed that most pairs were genetically monogamous: 7.9% of the broods (7/89) contained extra-pair young, comprising 3.9% of all chicks (9/229). These cases represented, however, three alternative reproductive behaviors: extra-pair paternity, quasi-parasitism (extra-pair maternity) and intraspecific brood parasitism. This is the first study showing the occurrence of all three behaviors in one shorebird species. We also found that extra-pair fertilizations (extra-pair paternity and quasi-parasitism) were more frequent later in the breeding season. There was no consistent relationship between genetic similarity of mates and laying date; the pattern, as well as the degree of genetic similarity, differed among breeding sites within the study population.Communicated by M. Webster