Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

A dusk chorus effect in a nocturnal bird: support for mate and rival assessment functions

The dawn chorus is a striking feature of spring mornings and a characteristic behaviour of many bird species, particularly the passerines. Dawn singing has been considered a reliable signal of male quality for mate and rival assessment. Singing is presumed to be relatively costly at dawn both because air temperatures are relatively low and because birds have not fed overnight. Models of optimal daily routine predict the existence of a “dusk chorus” in nocturnal birds, although this prediction has received little empirical attention. Nocturnal birds at dusk may be energy-limited because of a lack of daytime feeding, and singing at dusk may thus ensure signal reliability. Here, we used an observational and experimental approach to study vocal behaviour at dusk and dawn in a nocturnal raptor, the little owl Athene noctua. We assess whether male little owls adjust their vocal behaviour according to feeding stage (i.e. period of the night), ambient air temperature and territorial context (i.e. spontaneous calling behaviour vs elicited calling by intrusion). Across different temperatures, we find that both spontaneous vocal activity and inter-individual variability in call duration increased at dusk, clearly indicating a dusk chorus phenomenon. Results from playback presentations suggest that food, rather than air temperature, is likely to be more constraining at dusk. We discuss how comparing dusk and dawn choruses in nocturnal and diurnal species can provide insights into both mechanistic and functional aspects of signalling behaviour.     

Determinants of social dominance and inheritance of agonistic behavior in an island population of silvereyes,Zosterops lateralis

Summary We examined components of agonistic behavior and dominance in parents and offspring of silvereyes on Heron Island, Great Barrier Reef, to discover their determinants. The data were collected over four years on 1,235 juveniles and their parents. Of four behavioral characters used to compare juveniles and their parents, bill clattering was seen significantly more among young from female parents that exhibited bill clattering. The total absence of submissive behavior and the total absence of aggressive behavior were shown more often than not by young from dominant male parents and subordinate female parents, respectively. Young fledging early in the breeding season tended to be more dominant than those fledging late in the season. The number of brood-mates or the local nest density did not affect the dominance of young. Dominance status appears to be little affected by the performance of parents.     

The “dawn chorus” and incubation in the coot (Fulica atra L.)

Summary It has been proposed that both acoustic and non-acoustic factors could explain the dawn peak of territorial advertisement in birds. The coot is a territorial waterbird which does not use vocal advertisement either to attract mates or defend territory boundaries. In spite of the absence of acoustic factors affecting the time budgeting of advertisement, this species shows a dawn peak of display and aggressive behaviours. This trend occurs both in the presence and absence of non-territorial prospecting birds, but the trends are quantitatively different in these two cases (Fig. 1). Males participate in display and fighting more often than females, probably because of their greater size and effectiveness in defence. This asymmetry in defence abilities governs the time budgeting of incubation, the male tending to take time off the nest when territorial defence is most necessary (Fig. 2).     

Experimental manipulation of the dawn and dusk chorus in the blackbird Turdus merula

Summary We present experimental and correlational data that demonstrate that both the nutritional state of the male and his mate's fertility affect the timing and intensity of dawn and dusk song in the blackbird (Turdus merula). Food supplementation leads to paired males singing earlier, for longer, and at higher peak rate at both dusk and dawn succeeding the treatment. We argue that the timing, not just the intensity, of song conveys information about the male's phenotypic condition. Both dawn and dusk song start earlier and have longer duration as a male's mate reaches peak fertility indicating that, in agreement with literature on the great tit (Parus major), song at this time has particular significance in mate guarding for paired males of monogamous species. We discuss the relative importance of these factors as functional explanations for the dawn chorus. Offprint requests to: I.C. Cuthill     

Conservation decision-making in large state spaces

When looking for the best course of management decisions to efficiently conserve metapopulation systems, a classic approach in the ecology literature is to model the optimisation problem as a Markov decision process and find an optimal control policy using exact stochastic dynamic programming techniques. Stochastic dynamic programming is an iterative procedure that seeks to optimise a value function at each timestep by evaluating the benefits of each of the actions in each state of the system defined in the Markov decision process.Although stochastic dynamic programming methods provide an optimal solution to conservation management questions in a stochastic world, their applicability in metapopulation problems has always been limited by the so-called curse of dimensionality. The curse of dimensionality is the problem that adding new state variables inevitably results in much larger (often exponential) increases in the size of the state space, which can make solving superficially small problems impossible. The high computational requirements of stochastic dynamic programming methods mean that only simple metapopulation management problems can be analysed. In this paper we overcome the complexity burden of exact stochastic dynamic programming methods and present the benefits of an on-line sparse sampling algorithm proposed by Kearns, Mansour and Ng (2002). The algorithm is particularly attractive for problems with large state spaces as the running time is independent of the size of the state space of the problem. This appealing improvement is achieved at a cost: the solutions found are no longer guaranteed to be optimal.We apply the algorithm of Kearns et al. (2002) to a hypothetical fish metapopulation problem where the management objective is to maximise the number of occupied patches over the management time horizon. Our model has multiple management options to combat the threats of water abstraction and waterhole sedimentation. We compare the performance of the optimal solution to the results of the on-line sparse sampling algorithm for a simple 3-waterhole case. We find that three look-ahead steps minimises the error between the optimal solution and the approximation algorithm. This paper introduces a new algorithm to conservation management that provides a way to avoid the effects of the curse of dimensionality. The work has the potential to allow us to approximate solutions to much more complex metapopulation management problems in the future.     

Singing performance,frequency matching and courtship success of long-tailed manakins (Chiroxiphia linearis)

We examined the relationship between song performance and courtship success in the lek-mating long-tailed manakin (Chiroxiphia linearis) of Costa Rica. Teams of male manakins form cooperative partnerships consisting of a dominant alpha male and a beta male partner with a variable number of subordinate affiliates. Each alpha and beta partnership performs a joint song virtually in unison, which attracts females to their lek arena. We quantified four aspects of song performance, including synchrony, degree of consistency in the alpha male's singing, degree of consistency in the beta male's singing, and degree of sound frequency matching between partners. When all four variables were included in a discriminant function analysis, average frequency matching alone was useful in discriminating between teams with high rates of visitation by females and those with low visitation rates. Only frequency matching was correlated with the rate at which females visited display lek arenas. We propose that the degree of frequency matching provides an unambiguous ideal against which dual-male courtship performance can be assessed. This aspect of song performance may provide information useful to females in assesing a potential mate's ability to form a cooperative long-term partnership.     

Orientation in captive migratory and sedentary Australian silvereyes <Emphasis Type="Italic">Zosterops lateralis</Emphasis> (Zosteropidae)

The orientation of two closely related subspecies of Australian silvereye Zosterops lateralis was studied in captivity over 14 months. Migratory silvereyes Z. lateralis lateralis showed significant directional preferences during the spring and autumn migration periods and also displayed orientated behaviour during the breeding period. In contrast, the non-migratory subspecies Z. lateralis familiaris did not display any significant directional preferences at any time. This is the first time that the orientation behaviour of a migratory and non-migratory subspecies has been compared over the duration of an annual cycle, both during and outside the migratory periods. The results suggest that migratory silvereyes possess an endogenous program determining the timing and direction of autumn and spring migration and that this program is unique to the migratory subspecies. This is also the first comparison of this nature on southern hemisphere birds and demonstrates that the migratory behaviour of southern hemisphere migrants may be more similar to that of northern hemisphere migrants than previously thought.Submitted to Behavioral Ecology and Sociobiology: 23 Jan 2006.     

Mate preferences by male guppies (Poecilia reticulata) in relation to the risk of sperm competition

We investigated male mate preferences in relation to the perceived risk of sperm competition in the guppy (Poecilia reticulata), a freshwater fish with a promiscuous mating system. Our laboratory experiments revealed that male mate choice behaviour is not influenced by the presence of rival males that are merely in close proximity to a potential mate, as there was no significant difference in the amount of time that males spent with females that were recently either alone or in close proximity to four rival males. Male mate choice behaviour was, however, strongly influenced by the presence of rival males in a second experiment, where those rivals were permitted to copulate with one of the females. In that situation, males spent significantly more time with, and directed significantly more sigmoid courtship displays toward, females that they had recently seen alone compared to females they had seen receiving forced copulations from up to four rival males. Our results therefore indicate that male guppies are sensitive to the risk of sperm competition and alter their mate choice behaviour in an adaptive fashion.Communicated by K. Lindström     

Assessments and decisions in Metellina segmentata (Araneae: Metidae): evidence of a pheromone involved in mate guarding

Assessments and decision-making underlying the initiation of mate guarding in a common web-building spider, Metellina segmentata, are examined in a series of field and laboratory studies. Adult males do not build webs but wander in search of females and mating opportunities. Adult males then wait at the edge of the webs of females and guard them prior to courtship and mating. Guarded females were heavier, larger and carried more mature eggs than solitary females. An active process of information gathering is apparent from introductions of males to the webs of females. Males make accurate assessments about female quality, even in the absence of the resident female. Cues involving web architecture are not used. Males may assess pheromonal cues on the web of the female in deciding whether to guard or abandon a female.     

Selective phonotaxis by male wood frogs (<Emphasis Type="Italic">Rana sylvatica</Emphasis>) to the sound of a chorus

Frogs and toads commonly form large choruses around suitable breeding habitat during the mating season. Although often regarded as a constraint on the acoustic behavior of signalers and receivers, the sounds of a chorus could also serve as an acoustic beacon that allows some frogs to locate the breeding aggregation. Attraction to chorus sounds might be particularly important for explosively breeding frogs. In these species, which often mate just one or a few days during the year, the timing and location of breeding aggregations can be unpredictable because their formation often depends on local climatic factors, such as rainfall or a change in temperature. I used laboratory playback experiments to test the hypothesis that male wood frogs (Rana sylvatica), an explosively breeding frog, exhibit positive phonotaxis toward the sound of a conspecific chorus. Males were released at the center of a rectangular arena with a speaker positioned in each corner facing toward the release point. In a single-stimulus experiment, more males approached a speaker broadcasting a conspecific chorus than the three silent speakers in the arena. In a two-stimulus experiment, more males approached a speaker broadcasting a conspecific chorus compared to the two silent speakers or a fourth speaker simultaneously broadcasting the spectrally overlapping sound of a heterospecific (R. septentrionalis) chorus. These results are consistent with the hypothesis that male wood frogs could use the sound of a chorus as a beacon to locate a short-lived breeding aggregation.     

Determinants of chorus tenure in barking treefrogs (Hyla gratiosa)

In all species of anuran amphibians studied to date, male mating success is positively correlated with male chorus tenure (the number of nights that a male is present in breeding aggregations), yet males spend only a small portion of the total breeding season in choruses. Three hypotheses might explain abbreviated chorus tenure in anuran amphibians: (1) mortality, (2) energy limitations, and (3) movement among choruses. I tested these hypotheses in a study of the barking treefrog (Hyla gratiosa), a species with very abbreviated chorus tenure. Mortality accounted for the short tenures of some males: an estimated 20% of males died while calling in the chorus. Energy limitations were an important determinant of chorus tenure. Males lost weight over successive nights in the chorus, and males with longer chorus tenures were in better condition on their first night in the chorus, lost condition more slowly, and were in poorer condition on their last night in the chorus than were males with shorter chorus tenures. Males that I fed crickets as they left the chorus returned sooner and for more nights than did control (unfed) males. Movement between ponds did not explain abbreviated chorus tenure: less than 16% of the males called at more than one pond, and these males increased their chorus tenures by a median of only 2.5 nights by calling in another chorus. Thus, abbreviated chorus tenure in H. gratiosa appears to be primarily the result of energy limitations, with mortality playing a secondary role.     

From eavesdropping on performance to copying the behavior of others: a review of public information use

Public information (PI), a form of indirect social information, is used by individuals to estimate the quality of environmental parameters. It can be acquired in two ways. One way is by noting the performance of others. The other way it can be acquired is by noting the behavioral decisions of other individuals. Performance-based PI has been observed most often in the context of food and breeding patch estimation, as well as by individuals eavesdropping on contests between others in the context of assessing the fighting ability of opponents and the quality of mates. Evidence for the acquisition of PI from behavioral decisions of others comes mostly from studies of copying behavior, although recent work suggests that it also occurs when individuals estimate the tendency of others to be altruistic. PI use appears to be widespread across many taxa, although most work has been conducted on birds and fish. Absent from the literature are clear examples of PI use in mammals. The use of PI appears to often depend on its cost of acquisition and whether it contradicts an individual’s personal prior experience. PI can be an important benefit of associating with others.     

Exploring the effects of reductions in krill biomass in the Southern Ocean on blue whales using a state-dependent foraging model

Many species of baleen whales were hunted to near extinction in the Southern Hemisphere. The recovery of these populations will be affected by the availability of krill, a major dietary component, in the Southern Ocean. We combine a novel energetics model for baleen whales with a state dependent foraging model to explore the impacts of an expanding krill fishery on baleen whales. We parameterize the model for blue whales, but with simple modifications it could be applied to most baleen whales. We predict that an expanding fishery will have a small but significant impact on the blue whale population through decreased birth rates. However, spreading the catch limit throughout the range of krill can reduce these effects. In addition, whales may be able to reduce these impacts through adaptive changes in foraging behavior. The relationship between krill abundance and blue whale foraging and reproductive success is nonlinear, such that larger reductions in krill biomass, potentially following a loss of sea ice due to climate change, could have a much larger negative impact on the recovery of blue whales.     

The influence of potential reproductive rate and variation in mate quality on male and female choosiness in the sand goby, Pomatoschistus minutus

Recent models of choosiness in mate choice have identified two particularly important factors: the potential reproductive rate (PRR) of the choosing sex relative to that of the chosen sex, and the variation in quality of potential mates. This experimental study tested how these factors affected choosiness in male and female sand gobies, Pomatoschistus minutus. We manipulated relative PRR by means of water temperature, and mate quality by means of body length. The choosing male or female was offered a choice between two mates with either a small or a large difference in body length representing a small or a large variation in mate quality. Choosiness was measured as (1) preference for the larger mate, and (2) as whether or not spawning occurred with the smaller mate, while the larger mate was visible but screened off. We found that females preferred large males, and that their level of choosiness was affected by variation in male quality, but not by their own relative PRR. Males, on the other hand, seemed unselective in all treatments and were in general more likely than females to spawn with their provided partner. This suggests that in the sand goby, variation in male mate quality has a greater influence than relative PRR on facultative changes in female choosiness. However, a general difference in PRR between males and females may be one important factor explaining the observed sex difference in choosiness. Received: 17 April 2000 / Revised: 24 June 2000 / Accepted: 17 July 2000     

Long-term mate and territory fidelity in neotropical buff-breasted wrens (Thryothorus leucotis)

Among sedentary species, mate and site fidelity may be imposed by year-round territoriality and high annual survivorship, both of which theoretically lead to few opportunities to move between mates and breeding territories. We examined mate and territory fidelity in sedentary neotropical buff-breasted wrens (Thryothorus leucotis) in relation to the opportunities to divorce (vacancies on neighbouring territories), breeding experience and site of origin (within vs outside local breeding population). Overall, buff-breasted wrens showed high rates of mate and territory fidelity between years, with individuals significantly more faithful to their territories than to their partners. Mate and territory fidelity was not forced upon buff-breasted wrens, as most individuals had opportunities to switch mates and territories. However, this was not uniform among all individuals. Individuals that had bred together displayed 100% mate and territory fidelity despite the fact that males and females in almost all experienced pairs had opportunities to switch mates and territories. Local recruits (philopatric offspring that acquired territories in the local breeding population) also displayed mate and territory fidelity, but fidelity occurred because recruits rarely had the opportunity to divorce prior to initiating their first breeding attempt. By contrast, almost half of all territorial individuals that originated outside the local breeding population (immigrants) had opportunity to divorce, with approximately two thirds of them switching mates and territories. Divorces occurred within the first 5 months of pairing, suggesting that immigrants may have sampled mates and territories before forming permanent partnerships.     

Breeding synchrony and paternity in the barn swallow (Hirundo rustica)

The barn swallow (Hirundo rustica) is a socially monogamous passerine which usually breeds in colonies where extra-pair copulations are frequent. Males intensively guard their mates during the female fertile period. Since males are more likely to be available for extra-pair copulations when their mate is not fertile, synchrony in timing of breeding may affect paternity of individual males. In this study, we analysed the change in mate-guarding rate by males in relation to the fertility condition of the female, and the relationships between breeding synchrony and density with paternity in first broods of 52 male barn swallows. Paternity (proportion of nestlings fathered in own brood) was assessed by typing of three highly polymorphic microsatellite loci. Mate guarding by males peaked during the fertile period of their mates. Paternity increased as breeding synchrony in the colony increased. Paternity of barn swallows is positively associated with the degree of exaggeration of male tail ornaments. The relationship between male ornamentation and paternity was partly mediated by an effect of ornament size on breeding synchrony. We suggest that females might delay breeding with low-quality males to enhance their opportunities for being fertilised by high-quality extra-pair males. Received: 16 January 1998 / Accepted after revision: 25 October 1998     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

Tactics, effectiveness and avoidance of mate guarding in the blue-footed booby (Sula nebouxii)

We examined the dynamics and avoidance of mate guarding, by males and females, in the blue-footed booby, in which the two social mates are usually simultaneously present on the territory but each of them is unmonitored by the other for one-quarter of its time. Both sexes were promiscuous and liable to switch mates. Cuckolded individuals did not increase their overall presence on the territory, but in response to the extra-pair (EP) courtships of their mates, both sexes doubled their rate of intra-pair (IP) courtship and sometimes showed aggression. The male or female's presence depressed the social mate's EP activity, but intra-pair courtship had no such effect, tending even to propitiate that EP activity. Similarly, when females responded to their social mates' EP courtship with approach or aggression, disruption of EP activity was short-lived. Promiscuous females modified their diurnal pattern of attendance, as if attempting to sidestep monitoring by their mates, but cuckolded males matched the modification. Both sexes tended to perform their EP activities at a distance when their mates were present, possibly to evade monitoring or disruption by their mates. Male and female boobies cannot monitor their mates continuously, they do little to facultatively adjust their presence on territory to the risk of infidelity, and their immediate responses to overt infidelity have only the briefest impact; but the information they acquire while monitoring their mates may be critical to constraining their mates' infidelity and also to calibrating their own reproductive investment.     

The significance of male-male aggression for yearling dispersal in roe deer (Capreolus capreolus)

The significance of adult buck aggression as a proximate factor driving yearling male dispersal in roe deer, Capreolus Capreolus, was studied in Sweden during the dispersal season (April–June) in 1988–1991. In roe there is a well established positive correlation between yearling antler size and dispersal tendency. I tested the prediction that yearlings with large antlers will experience more agonism from adult bucks than those with smaller antlers. Additionally, I studied the relationship between antler size and level of sexual maturity, by recording date of velvet shedding, and sampling testes weights from culled yearlings during the breeding season. The amount of agonism experienced by yearlings was positively correlated with the size of their antlers during April and May, whereas in June no such relationship was found. Aggression towards all yearlings tended to increase as the season progressed. A positive correlation between antler size and testes weight was evident, and the larger the antlers of yearlings the earlier the velvet shedding occured. These results indicate: firstly, that adult buck agonism is an important proximate factor underlying yearling male dispersal; and secondly, that agonism is mainly directed towards the sexually most mature yearlings. This, in turn, suggests mate competition to be the ultimate cause of male-male agonism, and thus male dispersal in this species.