Prior residency as a key determinant of social dominance in the willow tit (Parus montanus)

Summary We studied how age, body size and prior residency affected social dominance in the willow tit (Parus montanus) groups. The contribution of each variable was experimentally tested in unisexual two-bird trials, in which the birds were matched for all variables except the one studied. Large birds were dominant over smaller ones (Fig. 1). The effect of body size was more prominent in males than in females. Age had no influence on dominance. Residents became dominant more often than newcomers (Fig. 2). Adulthood or larger body size did not override the advantage of prior residency (Fig. 2). Therefore, the proximate reason for the age-dependent dominance seen in natural willow tit flocks is most likely the prior residency advantage of the adults. Factors connected with fighting ability (body size and age) seem to be less important than the time of establishment of rank, which may reflect the importance of resource value differences between residents and newcomers in this context. The advantage of residency might make it advantageous to be a member of a flock even as a subordinate, rather than being solitary.     

Extra-pair paternity, offspring mortality and offspring sex ratio in the socially monogamous coal tit (Parus ater)

Females of many socially monogamous bird species commonly engage in extra-pair copulations. Assuming that extra-pair males are more attractive than the females’ social partners and that attractiveness has a heritable component, sex allocation theory predicts facultative overproduction of sons among extra-pair offspring (EPO) as sons benefit more than daughters from inheriting their father’s attractiveness traits. Here, we present a large-scale, three-year study on sex ratio variation in a passerine bird, the coal tit (Parus ater). Molecular sexing in combination with paternity analysis revealed no evidence for a male-bias in EPO sex ratios compared to their within-pair maternal half-siblings. Our main conclusion, therefore, is that facultative sex allocation to EPO is absent in the coal tit, in accordance with findings in several other species. Either there is no net selection for a deviation from random sex ratio variation (e.g. because extra-pair mating may serve goals different from striving for ‘attractiveness genes’) or evolutionary constraints preclude the evolution of precise maternal sex ratio adjustment. It is interesting to note that, however, we found broods without EPO as well as broods without mortality to be relatively female-biased compared to broods with EPO and mortality, respectively. We were unable to identify any environmental or parental variable to co-vary with brood sex ratios. There was no significant repeatability of sex ratios in consecutive broods of individual females that would hint at some idiosyncratic maternal sex ratio adjustment. Further research is needed to resolve the biological significance of the correlation between brood sex ratios and extra-pair paternity and mortality incidence, respectively.     

Experimental and natural changes in the peacock's (Pavo cristatus) train can affect mating success

Petrie et al. (1991) demonstrated a correlation between the degree of elaboration of peacocks' trains and their mating success, and also showed that this correlation occurred because females preferred to mate with the male that had the most elaborate train of those sampled on the lek. Although these data suggest that female choice is responsible for non-random mating in this species, they do not conclusively show that train morphology is the cue that females respond to, because they do not rule out the possible influence of another unidentified variable which is correlated with train elaboration. This paper presents an experimental test of the importance of the peacock's train in determining male mating success. If the number or arrangement of eye-spots in the peacock's train influences mating success, then changing the number of eye-spots should change mating success. This prediction was tested in an experiment where the trains of male peafowl (Pavo cristatus) were manipulated by removing a number of eye-spots between mating seasons. Peacocks with eye-spots removed showed a significant decline in mating success between seasons compared with a control group. This result, together with the observational data, supports the hypothesis that the peacock's train has evolved, at least in part, as a result of female choice.     

Changes in singing behavior of male black-capped chickadees (Parus atricapillus) following mate removal

We removed the mates of ten male black-capped chickadees (Pares atricapillus) during the nest-building period to determine the effect of female presence on dawn singing. During the first dawn chorus following mate removal, males sang significantly longer, increased movement within their territory, and increased the percentage of their territory covered while singing. After the female was returned, these parameters returned to the pre-removal values. Males did not alter the frequency range or modal frequency of their songs when the mate was removed, nor did they change the degree of frequency shifting in the fee-bee song. We conclude that dawn singing in the black-capped chickadee acts, in part, as an intersexual signal, and that the behavior of frequency shifting in the song may be directed more toward rival males than females. Correspondence to: K. Otter     

Brood size manipulation affects frequency of second clutches in the blue tit

The reproductive trade-off hypothesis predicts that the investment made in current reproduction determines the breeders’ future fitness as a consequence of intra-or inter-generational reproductive costs. Long-lived species are expected to favour their own reproductive value at the expense of their offspring, hence incurring in inter-generational costs, whereas short-lived species are expected to invest in the current breeding attempt even at the expense of their own survival, thus incurring in intra-generational costs. We tested whether intensity of current reproductive effort has intra-or inter-generational costs in a short-lived bird, the blue tit Parus caeruleus, with a brood size manipulation experiment. We expected more intra-generational (parental reproduction and/or survival) than inter-generational (offspring quality and survival) reproductive costs. We found that parental effort, measured as the hourly rate of parental visits to nests, increased gradually with experimental manipulation. Brood size manipulation resulted in a gradual increase in the number of fledglings per nest from reduced to increased treatments. We found an effect of the manipulation on the probability of making a second clutch, with adults rearing enlarged broods being less likely to undertake such a second reproduction during the season compared to those rearing control or decreased broods. We found no evidence of other reproductive costs; neither as adult weight after manipulation, apparent parental local survival, apparent offspring local survival or local recruitment. Although the results seem to support the a priori expectations, alternative explanations are discussed.Communicated by M. Soler     

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     

Indirect genetic benefits of polyandry in a spider with direct costs of mating

The search for the evolutionary explanation of polyandry is increasingly focused on direct and indirect selection on female resistance. In a polyandrous spider Stegodyphus lineatus, males do not provide material benefits and females are resistant to remating. Nevertheless, polyandrous females may obtain indirect genetic benefits that offset the costs associated with multiple mating. We manipulated the opportunity for females to select between different partners and examined the effect of female mating history (mated once, mated twice, or rejected the second male) on offspring body mass, size, condition, and survival under high- and low-food rearing regimens. We found that multiple mating, not female choice, results in increased female offspring body mass and condition. However, these effects were present only in low-food regimen. We did not find any effects of female mating history on male offspring variables. Thus, the benefits of polyandry depend not only on sex, but also on offspring environment. Furthermore, the observed patterns suggest that indirect genetic benefits cannot explain the evolution of female resistance in this system.     

Soft song in song sparrows: response of males and females to an enigmatic signal

Low-amplitude “soft song” is used by a variety of songbirds; in some species during aggressive encounters, in others during courtship, and yet others in both these contexts. In song sparrows (Melospiza melodia), soft song has thus far been observed only in aggressive encounters, where its production is a more reliable predictor of attack than any other signaling behavior. We used song playback to test the response of both male and female song sparrows to soft song. The design of the playback experiments took into account the existence of two classes of soft song: crystallized soft song, which consists of song types also found in the broadcast repertoire, and warbled soft song, which consists of less-structured song types not found in the broadcast repertoire. Female song sparrows responded with significantly less courtship display to the playback of crystallized soft song than to that of normal broadcast song, and response to warbled soft song was if anything lower than to that of crystallized soft song. Male song sparrows responded equally aggressively to normal broadcast song as to crystallized soft song, and equally aggressively to warbled soft song as to crystallized soft song. The female results support the conclusion that neither form of soft song functions in courtship. The male results suggest that the reliability of soft song as a signal of aggressive intent is not maintained by a receiver retaliation rule.     

Escape take-off strategies in birds: the significance of protective cover

Avian escape strategy is highly dependent on the ability to fly, and the success rate of raptor attacks is reduced if the prey gets fully airborne. Therefore, when escaping from predator attacks, the initial take-off is crucial and a rapid take-off, high velocity, and high angle of ascent maximize the chance of survival. However, due to the laws of gravity, birds face a trade-off between maximizing its linear acceleration and maximizing its rate of climb when taking off. The optimal policy between velocity and angle of ascent when a bird escapes from an attacking predator might depend on the detailed nature of the predator’s attack, the proximity to cover, and the presence of conspecifics. Many small birds rely on a quick dash to protective cover. In this study, we examine how the availability of protective cover affects take-off strategy in birds. Male great tits (Parus major) were subjected to a surprise attack by a model predator either in the presence or absence of protective cover. When attacked by the predator, great tits took off and perched in the cover when it was available. Birds subjected to the predator attack in the absence of cover took off in 10° higher angle of ascent and flew faster in the start of the take-off, compared to birds that took off in the presence of cover. Thus, this study for the first time shows that a bird do trade off speed with angle of ascent in an adaptive way, depending on the presence of protective cover.     

Effects of testosterone on male-male competition and male-female interactions in blue tits

In temperate-zone birds, testosterone (T) influences male behavior during the breeding season. The elevation of plasma levels of T to a breeding baseline is necessary for basic reproductive behaviors, but it is still unclear whether variation in T levels above this critical threshold influences the intensity of these behaviors. Such a relationship between T and sexually selected traits is a critical assumption of the immunocompetence handicap hypothesis. We here experimentally elevated T levels in blue-tit males above the natural mean (T-males) during the period of nest building and egg laying, without manipulating hormone levels during chick feeding. T-males neither interacted more often with other males, nor did they respond more aggressively to a dummy intruder, compared to males with control implants. T-males did not guard their social mates more closely, but they were more likely to interact with potential extra-pair mates. Females mated to T-males did not change their behavior during egg laying and the treatment did not significantly affect male and female feeding rates. Despite this, nests of T-males produced larger and heavier fledglings in one study year. Our observations suggest that T levels above the natural mean during the mating period do not increase aggressive or territorial behavior in male blue tits. However, if females perceived T-males as high-quality mating partners, superior offspring development in nests of T-males might be caused by higher maternal investment. Hence, male behaviors involved in mate attraction may have been influenced by T levels above the natural mean.Communicated by R. Gibson     

Determinants of parental effort: a behavioural study in the Eurasian kestrel,Falco tinnunculus

We recorded behaviour of kestrels (Falco tinnunculus) in western Finland during the courtship (1988–1992), incubation (1989–1991), early nestling (age of young 1–2 weeks, 1989–1992) and late nestling stages (3–4 weeks, 1989–1991) to examine determinants of their parental effort (PE). In males, PE was estimated as the hunting effort (the proportion of budget time spent in flight-hunting) and in females as the food provisioning rate (number of prey items delivered to the nest per hour). The following predictions derived from the parental investment theory were examined. (1) Parents rearing large clutches and broods should invest more in breeding than do parents rearing small clutches and broods. The hunting effort of parents did not increase with clutch or brood size, but males tending large broods had a higher prey delivery rate than males tending small broods (Figs 1–2). (2) PE of parents should increase in the course of the breeding season. In males, this was true only between the incubation and early nestling phases (Fig. 3). (3) The early pairs should invest more in breeding than late ones. This tended to be true during the early (for males) and late nestling phases (for females) (Fig. 4). (4) There should be a negative correlation between PE of mates within pairs, but no evidence for such adjustment was found (Fig. 5). (5) Females mated with bright-coloured attractive males should show higher PE than females mated with dull-coloured males but our results were inconsistent with this prediction. We conclude that PE decisions of kestrels are mainly based on cost-benefit estimates of residual reproductive value, rather than on current investment indicators, like clutch or brood size. This might be beneficial in environments with highly variable survival prospects of offspring caused by pronounced among-year variation in abundance of the main food (microtine rodents). The results also show that hypotheses explaining variation in PE in the short term are not necessarily valid for long-term PE, e.g. tending clutches or broods, which also reflects the demands of female and young.     

Does colour matter? The importance of colour in avoidance learning, memorability and generalisation

Aposematic species exploit the ability of predators to associate, for example, conspicuous colouration with the unprofitability of prey. We tested the importance of colour for avoidance learning, memory and generalisation in wild-caught great tits (Parus major). First, we determined the birds’ initial colour preferences for red, yellow, orange and grey artificial prey items. The birds showed some preferences, as they were more willing to eat grey prey as their first choice, but these were not strong preferences. We then trained birds to discriminate red, yellow or variable (red and yellow) signals from grey where colours signalled palatable and unpalatable food. In general, the birds learned the discrimination task equally well, irrespective of which colours signalled unpalatability, and subsequently remembered the distinction between previously palatable and previously unpalatable colours in the memorability test. We did not find strong evidence that variability in the signal affected learning or memory. Our results suggest that, in a task where birds must discriminate between palatable and unpalatable prey, it does not matter which specific colour signals unpalatability, although this might be context-dependent. To study whether training also affects responses to unconditioned stimuli, we included orange prey items in the memorability test. Although orange had been palatable in the initial preference test, the birds ate fewer orange prey items after they had been trained to avoid red, yellow or both colours (variable signal) as unpalatable prey, but did not change their preference when trained that these colourful signals were palatable. This indicates that generalisation occurred more readily after a negative experience than a positive experience, a situation that would potentially allow imperfect mimicry to occur.     

Modeling and analysis of nest-site selection by honeybee swarms: the speed and accuracy trade-off

Nest-site selection in honeybees is a process of social decision making in which the scout bees in a swarm locate several potential nest sites, evaluate them, and select the best one by means of competitive signaling. We develop a model of this process and validate that the model possesses the key features of the bees' decision-making process, as revealed by prior empirical studies. Next, we use the model to study the “design” of the nest-site selection process, with a focus on how certain behavioral parameters have been tuned by natural selection to achieve a balance between speed and accuracy. First, we study the effects of the quorum threshold and the dance decay rate. We show that evolution seems to have settled on values for these two parameters that seek a balance between speed and accuracy of decision making by minimizing the time needed to achieve a consensus and maximizing the probability that the best site is chosen. Second, we study the adaptive tuning of the tendency of bees to explore for vs be recruited to a site. We show that this tendency appears to be tuned to regulate the positive feedback process of recruitment to ensure both a reasonably rapid choice and a low probability of a poor choice. Finally we show that the probability of choosing the best site is proportional to its quality, but that this proportionality depends on its quality relative to other discovered sites.

Rank-dependent hoarding effort in willow tits (<Emphasis Type="Italic">Parus montanus</Emphasis>): a test of theoretical predictions

Many birds and mammals store energy as hoarded food supplies. A supply of stored food is beneficial during periods when food is scarce, but building up and managing such a supply also entails costs. The optimal number of caches will be reached when the net benefit is at its maximum. If dominants can steal more stored food from subordinates than the other way around, the optimum will differ between these categories. A previous theoretical model of hoarding in groups with dominant and subordinate members produced three testable predictions: (1) hoarders should store more food as anticipated future conditions get worse; (2) subordinate flock members should store more food than dominants; and (3) dominants should increase hoarding relatively more than subordinates as conditions get worse. Here we present a field experiment on willow tits (Parus montanus) designed to test these predictions. We found support for all three. Hoarding increased as conditions got worse, subordinates stored at a higher rate than dominants, and dominants increased their hoarding effort relatively more than subordinates as conditions worsened. These results support the incorporation of information on dominance and food availability into models predicting food storage behaviour.Communciated by J. Dickinson     

Diet-induced chemical phytomimesis by twig-like caterpillars of Biston robustum Butler (Lepidoptera: Geometridae)

Summary. Polyphagous caterpillars of the giant geometer Biston robustum resemble the twigs of their respective food sources in color and shape. Common predatory ants, including Lasius and Formica, were often observed to freely prowl directly on caterpillars bodies, even after antennal contact. This suggests that the cuticular chemicals of the caterpillars resemble those of the twigs of the foodplants, so we analyzed both by GC and GC-MS. The chemical compositions differed among caterpillars fed on a cherry, Prunus yedoensis, a chinquapin Castanopsis cuspidata, and a camellia Camellia japonica. The cuticular chemicals of the caterpillars resembled those of their corresponding food sources. When the caterpillar diets were switched from the cherry to camellia or chinquapin at the 4^th instars, the caterpillars cuticular chemicals changed after molting to resemble those of their respective foods. Caterpillars also changed their cuticular chemicals when they perched on cherry twigs and fed on camellia or chinquapin leaves, but not when they perched on camellia or chinquapin twigs and fed on cherry leaves. The chemical similarities between the caterpillars and the twigs were due to the digestion of host leaves, which indicates that this is a diet-induced adaptation.     

Eggs of cabbage root fly stimulate conspecific oviposition: Evaluation of the activity and determination of an egg-associated compound

Summary. Previous studies indicated the presence of antennally-active compounds in extracts of eggs laid by female cabbage root flies, Delia radicum, that stimulated oviposition by conspecific females. We confirmed that previously laid D. radicum eggs stimulated oviposition by other D. radicum females, in a dose-dependent manner. Methanol extracts of conspecific eggs stimulated oviposition by females D. radicum, whereas egg extracts of D. antiqua and Psila rosae had no effect. Electrophysiological recordings from the tarsal sensilla of D. radicum females indicated that neurones of the C₅ sensillum responded to the egg extracts from both D. radicum and D. antiqua, but not P. rosae. Chemical analysis revealed that the extract of eggs from D. radicum contained the thia-triaza-fluorene compound, 1,2-dihydro-3-thia-4,10,10b-triaza-cyclopenta[.a.]fluorene-1-carboxylic acid (CIF-1), an oviposition stimulant found previously only in cruciferous plants. Another potentially active component has yet to be identified.     

Tardy females,impatient males: protandry and divergent selection on arrival date in the two sexes of the barn swallow

Protandry reflects the earlier arrival of males than females to the site of reproduction. Such protandry is hypothesised to arise from sex differences in costs and benefits of early arrival. I investigated temporal patterns of arrival date of male and female barn swallows Hirundo rustica and temporal patterns of selection to test the hypothesis that sex differences in selection account for sex differences in arrival date. Mean arrival date of male barn swallows but not of females advanced during the last 33 years, giving rise to an increasing sex difference in arrival date. Early arrival was favoured by increasingly better survival in males, while females showed an opposite pattern that did not reach significance, although the effect differed between sexes. Early arrival increased fecundity in both sexes equally.The sex difference in viability selection in relation to arrival date changed from positive to negative as the degree of protandry increased in recent years, although there was no similar significant relationship for fecundity selection. Furthermore, sex differences in viability selection in a given year affected the degree of protandry in the following year through differential survival of certain phenotypes over others. Finally, temporal changes in sex difference in viability selection and protandry were related to an increase in the interval between first and second clutches, as the duration of the breeding season increased because of climatic amelioration. These findings suggest that arrival date is under divergent selection in the two sexes, providing a mechanism for the evolution of protandry.     

The benefits of multiple mating to female seaweed flies, Coelopa frigida (Diptera: Coelpidae)

Female seaweed flies, Coelopa frigida, have the potential to benefit from mating more than once. Single matings result in low fertility so females may benefit directly from multiple copulations by sperm replenishment. A chromosomal inversion associated with larval fitness, with heterokaryotypic larvae having higher viability than homokaryotypes, means that polyandrous homokaryotypic females have a higher probability of producing genetically fit offspring than monandrous homokaryotypic females. We allowed females to mate only once, repeatedly four times to the same male, or polyandrously four times to four different males. Multiply mated and polyandrous females laid more eggs and produced more offspring than singly mated and monandrous females, respectively. Polyandrous females laid more eggs, had higher egg-to-adult survival rates and produced more offspring than repeatedly mated females. Fertility rates did not differ between treatments. The observed fitness patterns therefore resulted from increased oviposition through multiple mating per se, and a further increase in oviposition coupled with higher egg-to-adult offspring survival benefits to polyandry. Daily monitoring of individual females over their entire life spans showed that multiple copulations induced early oviposition, with polyandrous females ovipositing earlier than repeatedly mated females. Singly mated and polyandrous females incurred a longevity cost independent of egg production, whereas repeatedly mated females did not. This suggests that repeatedly mating with the same male may counteract a general cost of mating. Longevity, however, was not correlated with overall female fitness. Our data are discussed in the overall context of the seaweed fly mating system.Communicated by G. Wilkinson     

Pyrrolizidine alkaloids between plants and insects: A new chapter of an old story

Summary Among alkaloids the pyrrolizidine alkaloids (PAs) play a unique role in the interactions between plants and adapted insects. InSenecio spp. (Asteraceae) PAs are synthesized in the roots as alkaloidN-oxides which are specifically translocated into shootsvia the phloem-path and channeled to the preferred sites of storage (e.g. inflorescences) where they are stored in the cell vacuoles. In differentSenecio spp. senecionineN-oxide is produced as the common product of biosynthesis, which subsequentlyvia a number of simple but specific reactions is transformed into typical speciesspecific PA-patterns. Insects from diverse taxa sequester PAs for their own defense. Lepidopterans (e.g. arctiids such asTyria jacobaeae andCreatonotos transiens) may hydrolyze plant acquired ester-PAs and convert the resulting necine base into insect-specific PAs by esterification with an acid of their own metabolism. Adapted arctiids and the grasshopperZonocerus take up PAs in the state of the tertiary amine.N-Oxides are reduced in the guts prior to uptake. In the bodies the tertiary PAs are rapidlyN-oxidized by a specific mixed-function oxigenase and are maintained in theN-oxide state. The importance of the reversible interconversion of the nontoxicN-oxide (pro-toxine) into the toxic tertiary alkaloid is discussed as the specific feature of PAs in plant-insect interactions.     

Reduced costs of mixed-species pairings in flycatchers: by-product or female strategy?

Heterospecific matings are generally assumed to be unconditionally disadvantageous due to reduced viability or fertility of hybrid offspring. For female collared flycatchers (Ficedula albicollis) mated to male pied flycatchers (Ficedula hypoleuca), the cost of heterospecific pair formation is reduced due to high levels of conspecific extra-pair paternity and a male-biased offspring sex ratio. In order to investigate whether these cost-reducing mechanisms are the result of female mating strategies, rather than being a by-product of species incompatibilities, we manipulated the plumage of male collared flycatchers before pair formation to make them resemble male pied flycatchers. Since species incompatibilities are absent in this design, any systematic effect of manipulation on sex ratio or paternity would indicate a role of female mating strategy. Paternity was determined by means of a likelihood approach that controls the errors made in assigning a chick to be ‘within-pair’ or ‘extra-pair’. Neither the sex ratio nor the male share of paternity was affected by the manipulation in a systematic manner. We therefore conclude that our experimental data provide no support for the suggestion that female behavioural strategies are markedly adjusted in response to formation of mixed-species pairs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.