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     

An experimental study of male adornment in the scarlet-tufted malachite sunbird: II. The role of the elongated tail in mate choice and experimental evidence for a handicap

Summary Scarlet-tufted malachite sunbirds (Neetarinia johnstoni) are endemic to the high altitude zones of East African mountains. On Mount Kenya the males are resident and territorial all year. The females arrive over an extended period at the beginning of the breeding season, after spending the non-breeding season in nomadic flocks at slightly lower elevations. Males are bright iridescent green with elongated central tail feathers, which are displayed during courtship. We examined the role of the tail in mate choice using natural variation and investigated the influence of tail length on male time budgets by experimental manipulation. Territorial males that paired had longer tails than territorial males which remained unpaired. Males with naturally long tails started breeding earlier, resulting in fledglings being produced earlier in the season when the weather was more clement. Birds which bred together in one year were paired together in the next year if they both survived. Thus the male's tail may only be involved as a cue in the formation of new pairs. Males which were already paired were subjected to one of three experimental treatments — having their tail elongated, shortened, or manipulated but kept at the same length. Males with experimentally shortened tails spent more time in flight and hawked for flying insects with a higher efficiency than control males. Both control and elongated tail males reduced the amount of time in flight and had a lower hawking efficiency after manipulation. These results suggest that the long tail of male scarlet-tufted malachite sunbirds is a handicap. The factors influencing the reproductive success of a pair are discussed. Offprint requests to: M.R. Evans     

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.     

Energetic state and the performance of dawn chorus in silvereyes (Zosterops lateralis)

Stochastic dynamic programming (SDP) models predict that males singing to attract a mate should concentrate singing in what has been termed the dawn chorus. This is because male birds should have a variable surplus of fat in the morning that can be used to fuel singing, with the amount of fat available dependent upon such factors as his quality, foraging success and risk of predation. In this manner, the dawn chorus can act as an indicator of male quality in the context of female mate choice. We test a key prediction of SDP models of singing behaviour that males with greater fat levels should sing more. We conducted an experiment where we recorded the dawn chorus of male silvereyes (Zosterops lateralis) on three consecutive days. Each male received supplementary food on the second day, which enabled us to sample his dawn chorus before, during and after food supplementation. We also collected data on the effect of supplementary food on the body mass of silvereyes. As predicted by SDP models, we found that silvereyes sang for a greater proportion of the time after receiving supplementary food. Supplementary food also had a significant effect on the complexity of a male song, indicating that males not only increased the quantity of their song but also the quality of their song when they received extra food. As the provision of supplementary food significantly increased the mass of fed birds, our results support a causal link between male energy reserves and his ability to perform the dawn chorus.     

Female mate choice in the zebra finch — the effect of male beak colour and male song

Studies of female mate preference in zebra finches (Taenopygia guttata) have shown that male beak colour and song rate are important. However, the two characters are correlated. Here the effect of beak colour and song rate on female choice are examined independently. In mate choice tests involving two males, beak colour was manipulated artificially using nail varnish. The results showed that females showed a significant preference for males with a high song rate, but not with a red beak. Females did not prefer males with a red beak if song rate was low and females preferred males with orange beaks who expressed a high song rate. Female preference for males with red beaks was not found when beak and song characters were no longer correlated.     

Pair formation,pre-spawning waiting,and protandry in kokanee,<Emphasis Type="Italic"> Oncorhynchus nerka</Emphasis>

The timing of arrival to breeding areas can have profound effects on reproductive success. Under some conditions (restricted breeding seasons and mating systems characterized by a longer period of mating among males than females), the maximization of mating opportunities by males theoretically selects for the earlier arrival of males than females (a phenomenon called protandry). This study quantifies the relationship between the arrival timing and spawning success of male kokanee (non-anadromous Oncorhynchus nerka). The spawning behavior of kokanee was observed in a large pen and the spawning success of each male was estimated as the number of spawning events he participated in. A male's spawning success depended primarily on his success at pairing with and mate-guarding females, and less on participation in spawning events while unpaired. Males who paired earlier in the season had higher spawning success than males who paired later in the season because they experienced more opportunities to pair with new females. Among males who eventually paired (some males never did), arriving early was correlated with pairing early. However, selection for protandry was weak, largely because early arrival did not guarantee that a male would pair. Pre-spawning waiting by females also weakened the correlation between arrival day and pairing day. The random probability of pairing with respect to arrival day and pre-spawning waiting by females likely explains the weak selection for protandry in kokanee and the low amount of protandry observed in other sockeye salmon (anadromous O. nerka) populations.     

Temporal performance roles during vocal interactions in nightingales (Luscinia megarhynchos B.)

Summary The temporal patterning of vocal interactions between territorial nightingales (Luscinia megarhynchos B). was investigated during nocturnal dyadic singing. We distinguished three forms of temporal performance roles (Table 1; Fig. 3): inserter (preferred song start 0.5–1.0 s after offset of a neighbour's song), overlapper (preferred song start 0.5–1.0 s after onset of a neighbour's song), and Autonomous bird (no evidence for temporal responses adjusted to the timing of a neighbour's singing; Fig. 4). With the beginning of the daily dawn chorus, mutual temporal adjustment of song performances could no longer be ascertained (Fig. 2). To test the flexibility of song onset timing, we presented subjects with three playback programs (I, II, III) simulating a non-flexible conspecific bird. Sequences of songs recorded from a stranger that shared no song types with the subjects were used; intersong pauses were unaltered in I, lengtened in II, and shortened in III. Results confirmed that the timing of song onset can be influenced by the temporal patterning of auditory stimuli (Table 1; Fig. 5): Inserters and overlappers sang at a slower rate during program II than during program I. During program III, which by accelerated stimulus succession caused frequent overlap of songs, inserters and overlappers interrupted their singing behaviour (Fig. 6). This suggests that overlapping, which results in signal interference, is of some functional significance in vocal interactions.     

Calling characteristics of parasitized and unparasitized populations of the field cricket Teleogryllus oceanicus

Summary Examination of three populations of the field cricket Teleogryllus oceanicus revealed the presence of an acoustically-orienting parasitoid fly, Ormia ochracea (Tachinidae), in the population of crickets that has been introduced to the Hawaiian Islands. The cricket is native to Australia and the Pacific, and the fly is native to North America but has also been introduced to Hawaii. Up to 27% of males and 7% of females in Hawaii were infested with fly larvae. Song structure in the parasitized Hawaiian population was distinct from that of the other two groups, with the Hawaiian crickets showing several reduced song parameters. In addition, onset and cessation of calling at dusk and dawn were more abrupt in the Hawaiian population. These results are consistent with selective pressure from the phonotactic flies to decrease risky calling. Silent males were present in all three populations, suggesting that these noncallers may not represent a unique adaptation to the parasitoid. Correspondence to: M. Zuk     

Reproductive strategy and singing activity: blue tit and great tit compared

The costs and benefits of bird song are likely to vary among species, and different singing patterns may reflect differences in reproductive strategies. We compared temporal patterns of singing activity in two songbird species, the blue tit (Cyanistes caeruleus) and the great tit (Parus major). The two species live side by side year round, and they have similar breeding ecology and similar rates of extra-pair paternity. However, they differ in two aspects of reproductive strategy that may have an influence on song output: blue tits are facultatively polygynous and have a fairly short breeding season with almost no second broods, whereas great tits are socially monogamous but more commonly raise second broods. We found that great tit males continued singing at high levels during the egg-laying and incubation periods, while monogamously paired blue tit males strongly reduced singing activity after the first days of egg-laying by their female. Since males of both species sang much more intensely shortly before sunrise than after sunrise, at midday or in the evening, this difference was most conspicuous at dawn. No differences in singing activity were found within species when testing for male age. We suggest that in contrast to blue tits, great tit males continued singing after egg-laying to defend the territory and to encourage the female for a possible second brood.     

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     

Mechanisms of selective attention in grasshopper choruses: who listens to whom?

In many rhythmically signaling species of acoustic insects and anurans, males form choruses at typical population densities. Recent findings that females may prefer leading calls indicate that the timing of a male's signals relative to those of neighboring choruses is an important component of mate attraction. Within a chorus, however, males cannot time their calls such that they lead all neighbors. Hence, they are expected to show selective attention toward only a subset of the group. We used field playback experiments to determine the incidence of and mechanism(s) responsible for selective attention in choruses of the territorial grasshoppers Ligurotettix coquilletti and L. planum. Our data revealed significant selective attention in both species and are inconsistent with either fixed-threshold or fixed-number mechanisms. Rather, regulation of selective attention by a sliding-threshold mechanism is supported. We discuss these results in the context of the evolution of chorus structure. Received: 24 September 1997 / Accepted after revision: 28 February 1998     

Trade-off between mate guarding and mate attraction in the polygynous great reed warbler

Summary In a Swedish population of the polygynous great reed warbler Acrocephalus arundinaceus, about 40% of the males formed pairs with more than one female. Males sang two completely separated types of song: (1) long song when they tried to attract mates and (2) short shong when they guarded a fertile female. As soon as a male had attracted a female he immediately switched from long to short song and apparently guarded her for at least 3 days. Most males left their female and started singing long several days before her fertile period ended. This behavior probably increased the risk of cuckoldry. By assuming that males of polygynous species maximize their fitness over the whole breeding season (i.e., maximize fitness rate), we predicted longer mate guarding periods when the probability of attracting a second female was low and the risk of cuckoldry was high. These predictions were supported by observations. The probability of attracting a second female decreased during the breeding season and, in accordance, the number of days that males sang long song during the primary female's fertile period was negatively correlated with the time of the season. This trend held also for individual males when comparing mating periods of their primary and secondary female. The increasing rate of male intrusions with season may also have contributed to the increasing mate guarding periods. Offprint requests to: D. Hasselquist     

Spike dives of juvenile southern bluefin tuna (<Emphasis Type="Italic">Thunnus maccoyii</Emphasis>): a navigational role?

Tunas make sharp descents and ascents around dawn and dusk called spike dives. We examine spike dives of 21 southern bluefin tuna (Thunnus maccoyii) implanted with archival tags in the Great Australian Bight. Using a new way to categorize this behavior, we show that spike dives are similar among all the fish in the study. The dive profiles are mirror images at dawn and dusk and are precisely timed with respect to sunrise and sunset. We analyze the possible reasons for spike dives, considering the timing of spike dives, the characteristic dive profile, and the tuna's magnetic habitat. In addition, we present anatomical evidence for elaboration of the pineal organ, which is light mediated and has been implicated in navigation in other vertebrates. The new evidence presented here leads us to suspect that spike dives represent a survey related to navigation.     

Close-range acoustic signaling and mate choice in Hawaiian crickets (Gryllidae: Laupala)

Evolution of the mate recognition system (MRS) can play a central role in animal speciation. One dramatic consequence of changes in the MRS is the failure of individuals from divergent lineages to successfully court and mate, thereby reducing gene flow between these groups. Here, we test the role of an acoustic mating signal on mate choice in a Hawaiian cricket genus (Gryllidae: Laupala). Speciation in Laupala is proceeding at an extremely rapid rate, apparently driven by divergence in aspects of the mate recognition system, most conspicuously the pulse rate of male calling song. Previous studies demonstrate that females prefer the pulse rate of a conspecific male’s song when perceived at long range, in laboratory phonotaxis trials. In this study, we examined mate choice in two species that differ dramatically in pulse rate: Laupala paranigra and Laupala kohalensis. We tested the female’s preference in both species for pulse rates at close range, by providing females an opportunity to mate with hybrid males producing a range of intermediate pulse rates. Results of our study demonstrate that while strong behavioral barriers exist between these two species, variation in the pulse rate of male calling song did not predict female mate choice at close range. These results suggest a more complex architecture to mate recognition in Laupala than previously hypothesized.     

Mass-dependent take-off ability in wintering great tits (Parus major): comparison of top-ranked adult males and subordinate juvenile females

When birds are attacked by predators, initial take-off is crucial for survival. Theoretical studies have predicted that predation risk in terms of impaired flight ability increases with body weight. However, studies in which attacks were simulated, and within-individual daily changes in body weight were used to test mass-dependent take-off outside migration period, have so far failed to show an effect of mass on velocity. In this field study I compared the mass/velocity relationships of alarmed adult male and juvenile female great tits, Parus major. Fattening strategies differ among members of the dominance-structured basic flocks of wintering great tits, and dominant individuals often carry significantly less amount of fat reserves than subordinates. Since the range of body weight gain/loss is the least among dominant males, it was expected that impaired flight ability is more likely in lower-ranked female great tits. The results show that the birds differed significantly in their daily increase of relative body weight. Average daily weight increase of adult males was 6.2%, while it was 12.2% in juvenile females. Males were faster than females at take-off both at dawn and at dusk. Flight velocity of males did not differ significantly between dawn and dusk, whereas females took off at a significantly lower speed at dusk than at dawn. The results suggest that the larger fat reserves of subordinate females needed to increase their chances of overwinter survival probably place them at increased risk of predation. Electronic Publication     

The relative roles of predation and sperm competition on the duration of the post-copulatory association between the sexes in the blue crab, Callinectes sapidus

In many species, post-copulatory mate guarding prevents other males from mating with the guarded female. In crabs, males stay with their mates to protect the female from predators because, in some species, mating occurs when she is soft and vulnerable after molting. I tested the relative roles of sperm competition and predation on the duration of the post-copulatory association in the blue crab, Callinectes sapidus. Unpaired females suffered greater predation mortality than paired females and males stayed with the female longer in the presence of predators than in their absence, suggesting that the post-copulatory association protects females during their vulnerable period. However, the association may also occur in blue crabs because of sperm competition since spermathecal contents of females in the field indicate that 12.4% mated twice. Females experimentally mated with two males contained both males ejaculates and each ejaculate had access to the unfertilized eggs, suggesting that the size of a male's ejaculate influences his fertilization rate in a multiply-mated female. Males stayed longest in response to a high risk of sperm competition. Longer post-copulatory associations allowed the first male's ejaculate to harden into a type of sperm plug, which limited the size of a second inseminator's ejaculate in a non-virgin female as compared with a virgin. Males passed larger ejaculates in the presence of rivals and when previous ejaculates were in the female spermathecae, another response to sperm competition. Larger ejaculates may need longer post-copulatory associations before a more effective sperm plug forms. Large males stayed with the female longer, which is consistent with their ability to pass larger ejaculates than small males and suggests that there may be costs to minimizing the duration of the post-copulatory association. In the field, associations last long enough to protect the female during her vulnerable phase and may ensure that the guarding male fertilizes the most eggs in the female, even if she remates. Thus, the post-copulatory association protects female blue crabs from additional inseminators as well as from predators. Received: 23 January 1996 / Accepted after revision: 9 November 1996     

The importance of biparental care in a precocial,monogamous bird,the bar-headed goose (Anser indicus)

Summary This study presents the first direct evidence of benefit derived from the male's parental effort during rearing of young in a monogamous, precocial bird species. We compared mothers accompanied by a mate (paired mothers) with single mothers, some of which were artificial widows, in a semicaptive flock of bar-headed geese (Anser indicus). Paired mothers were less often displaced by other birds and suffered fewer interruptions of brooding. Their goslings spent more time grazing and survived with a higher probability. Although one beneficial result of the male's parental effort may be the continued availability and survival of his mate, a more direct benefit arises from higher offspring survival between hatching and fledging.     

Mate choice by female white-crowned sparrows in a mixed-dialect population

Summary It has been argued that the song dialects of white-crowned sparrows, Zonotrichia leucophrys, may function to restrict gene flow between dialect populations, thus promoting adaptation to local environmental conditions. One proposed mechanism whereby this may occur is if a female mates preferentially with males singing her natal dialect, typically the song type of her father. Previous field and laboratory studies of the role played by song differences in mate choice by female white-crowned sparrows have produced equivocal results. These investigations were conducted in populations in which a single song dialect was predominant. We examined mate choice in a population in which two song dialects are equally common. We found that songs of mates of individual females in successive years were no more likely to be of the same dialect than expected by chance. In addition, individual females did not mate preferentially with males whose songs matched those of their fathers. We conclude that, in this population, females choose mates on the basis of characteristics other than song type.     

Delayed maturation of multiple signals in a migratory songbird

Many animals use multiple signals in sexual communication, but our understanding of the interactions between multiple signals, particularly in inexperienced breeders, is limited. In birds, delayed plumage maturation (DPM) is well documented; young birds appear duller than adults, despite reaching sexual maturity. Age-related changes in song structure are also common in songbirds, though the extent to which songs of yearling males differ from those of adults (delayed song maturation, DSM) and its prevalence in species with DPM is unknown. We tested for DSM in the mate-attraction (repeat) song of a species with dramatic DPM, the American redstart (Setophaga ruticilla). Repeat song structure of territorial yearling and adult males differed significantly, based on discriminant analysis of nine non-collinear song features. Combined with previous evidence of delayed maturation in territorial (serial) song, we provide the first evidence for DSM in different song types used in different behavioral contexts during the breeding season of a migratory songbird. Within adults, variation in repeat song was associated with pairing, earlier onset of breeding, and number of offspring sired, suggesting a potential benefit for expressing more adult-like song. We found no relationship between the expression of adult-like repeat song and plumage in either age class. These results indicate that delayed maturation of repeat song in yearling redstarts is not due to a generalized delay in signal development, and suggest further work is needed to determine whether mate attraction (for both yearling and adult males) provides greater pressure to sound more “adult-like” than to look more “adult-like”.     

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.