Antiphonal duetting and sex hormones in the tropical bush shrike Laniarius funebris (HARTLAUB).

We investigated changes in antiphonal duetting with phases of reproduction and circulating levels of luteinizing hormone, testosterone, and estradiol in slate-colored boubous (Laniarius funebris) breeding in aviaries. Frequency of overall male singing did not vary with reproductive phase while frequencies of female singing and female vocal responses to male song were reduced during incubation and feeding of nestlings. This resulted in significant changes in frequency of duetting. Males sang the sexual song type M1 more often during courtship and nest building than during the nestlings phase. Their territorial song types M2 and M4 did not vary with breeding phase. Females were less responsive to M1 during incubation and to M2 during the nest building and nestlings than during the courtship phase. Plasma levels of luteinizing hormone and testosterone increased in males from the prebreeding to the courtship phase. While testosterone decreased already during nest building and remained low during subsequent phases of reproduction, luteinizing hormone decreased during incubation and feeding of nestlings. Female luteinizing hormone levels were highest during nest building. Female estradiol levels decreased from nest building to incubation and increased again during subsequent nest building. Female testosterone levels were low but not basal and did not vary with phase. Neither the overall male and female singing frequencies nor the frequencies of male song types were correlated with hormonal state. However, female participation in territorial duets M4 correlated positively with their testosterone levels. It is suggested that in this monogamous, duetting species with prolonged pairbonds behavioral cues between the mates are more important than the hormonal state in control of male and female singing.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sound production and associated behavior in a cichlid fish,Cichlasoma centrarchus. II. Breeding pairs

Synopsis Pairs of Cichlasoma centrarchus were observed daily in the laboratory. Both males and females made sounds during a breeding cycle but all sounds were aggressive in context; no sounds were heard to accompany courtship. Males made more sounds before spawning than afterwards and these were associated with territorial defense and with establishment of dominance over the female. Females produced more sounds after spawning than before, most in the context of brood defense but some toward the male during pre-spawning nest preparation. Prior to spawning, the number of sounds made by the males toward their mates increased but the aggressive actions accompanying them became less intense. No such inverse correlation of agonistic intensity with number of sounds made was found for the females. From this study and earlier ones by the author it was concluded that sound in this species is a threat display which 1) provides an expression for agonism alternative to the performance of actions which could injure the female or drive her away, and 2) lessens the risk of injury to male or female during territory or brood defense.     

Female Sexual Interference in the Smooth Newt,Triturus vulgaris vulgaris

Competition between unmated females for males was observed in the smooth newt, T. vulgaris, during the early part of the breeding season. Females interfered at the spermatophore-transfer stage of courtship, preventing a rival female from picking up the sperm mass. Interference often resulted in the rival female abandoning the courtship encounter. In many of the encounters (70%), the interfering female was inseminated. However, interference between females was not observed after the commencement of ovulation. By contrast, the incidence of sexual interference between males was low during the preovulatory period and increased significantly post-ovulation. Sexual interference in newts and salamanders has previously only been observed between males. During the greater part of the breeding season, when females are ovipositing, eggs are the limiting resource and males compete for access to receptive females. However, this study suggests that early in the breeding season, prior to commencement of ovulation, the limiting resource may temporarily be sperm, due to the physiological constraint of spermatophore production. This may lead to female competition for mates.     

Seasonal sex role changes in the blenniid Petroscirtes breviceps, a nest brooder with paternal care

The plasticity of the sex roles in the blenniid fish Petroscirtes breviceps , a nest brooder with exclusive paternal care, was studied throughout an 8 month breeding season. Males performed most courtships early and late in the breeding season, whereas females performed most in the middle of the season. These results indicated that the sex of individuals initiating courtship changed seasonally, with courtship role reversal in the middle of the season. Intrasexual aggression in both sexes occurred much more frequently in mid-season than in the early and late seasons. Males frequently fought when available nest sites were limited, regardless of the presence of females, suggesting that males competed for nests in order to qualify to mate (resource competition). In contrast, courting females fought only in mid-season, when females' relative success in entering nests decreased, indicating that females competed for limited mating opportunities (mating competition). The reversed courtship roles and female mating competition in mid-season suggested that the sex roles in P. breviceps changed seasonally from the conventional roles to reversed roles and back again during one breeding season. This study provides the first empirical evidence of multiple changes in the sex roles of animals within a breeding season.     

Mate Choice in Captive American Kestrels,Falco sparverius,Parasitized by a Nematode,Trichinella pseudospiralis

The American kestrel (Falco sparverius) — Trichinella pseudospiralis host-parasite association was used as a laboratory model to study the effect of a nematode infection on (1) mate choice of female kestrels and (2) competitive abilities of male kestrels during the breeding season. Females were given the opportunity to choose between infected and uninfected males. There was no significant difference in the proportion of females that chose uninfected (n = 18) over infected (n = 16) males. Eight of 11 competitions between infected and uninfected males exposed to a caged female and a nest box were dominated by uninfected males. Female kestrels seemed to base their choices on the intensity of display effort by the males; the experimental level of infection did not affect male courtship behaviour. Although T. pseudospiralis is known to debilitate American kestrels, parasitized males in mate-choice tests may have diverted energy from less important functions to court the females.     

Dangerous love? Predation risk does not affect female mate choice in blue‐black grassquits

Predation risk may be an important factor affecting female mate choice. Hypothetically, females could choose extravagantly ornamented males that survive in high predation risk environments. However, this decision could be different if choosing a conspicuous male under high predation risk is costly for females or results in reduced offspring survival. In such contexts, females could become indifferent to male quality or prefer inconspicuous males. We tested this idea using captive blue‐black grassquits (Volatinia jacarina, Linnaeus, 1766), a species in which males perform conspicuous leap displays coupled with songs during the breeding season, which presumably subjects females and offspring to higher predation risk. Females were placed in an arena with speakers on opposite sides emitting male courtship songs. One speaker emitted songs at a high rate (proxy for a conspicuous male) while the other speaker broadcast songs at a low rate (proxy for a less conspicuous male). While the female evaluated the two male songs, a third speaker emitted vocalizations characterizing three levels of risk: adult predator, nest predator, and no‐risk control. Females showed no preference for either male stimuli across the predation risk treatments. This lack of preference relative to frequency of male vocal displays suggests that leap‐song frequency is not used by females during mate choice. We suggest that in addition to its role in courtship, male grassquit displays also signal status to other males when competing for territories. Thus, we propose that predation risk does not directly influence blue‐black grassquit intersexual selection and that females in this species may exercise indirect mate choice, choosing social mates based on male ability to establish and defend a territory, and relying secondarily upon other aspects of male display attributes, such as its visual components.     

Seasonal Changes in Courtship Song and the Medial Preoptic Area in Male European Starlings (Sturnus vulgaris)

In male starlings (Sturnus vulgaris) courtship song plays a critical role in mate attraction. During the breeding season courtship song occurs prior to copulation and appears to reflect male sexual arousal. Outside the breeding season starlings sing, but song appears unrelated to reproduction. The aromatization of testosterone (T), likely within the medial preoptic nucleus (POM), is critical for the expression of male sexual arousal. The present study was performed to determine whether seasonal changes in the POM might relate to seasonal changes in courtship singing behavior in male starlings. T concentrations, the volume of the POM, and aromatase within the POM were examined both during and outside of the breeding season in male starlings. Song was also recorded at these times both with and without a female present. The POM was largest and contained dense aromatase immunostaining only during the spring breeding season, when T concentrations were highest and males responded to a female with an increase in courtship song. Outside the breeding season the volume of the POM was small, T concentrations were low, and males displayed no changes in song expression in response to female conspecifics. Song bout length was positively related to POM volume, and males sang longer songs in spring. Only males with nestboxes in spring responded to a female, and the POM tended to be larger in these males, suggesting that nestbox possession might influence neuroplasticity within the POM. Overall, the findings suggest that T-dependent plasticity and aromatase activity within the POM might regulate courtship singing in a wild songbird.     

Breeding ecology of Kori Bustard Ardeotis kori strunthiunculus in the Serengeti National Park

The breeding ecology of the Kori Bustard Ardeotis kori strunthiunculus was studied in the plains of the Serengeti National Park, Tanzania in 2014 and 2015. Random transects were used to search for male courtship displays, nests, chicks and subadults. GPS satellite collars were used to locate nesting females. Linear regression analyses and post hoc tests were used to determine the predictors that contributed most to the variation of the dependent variables (courtship display, nest, chicks and subadults). The results indicate that courtship behaviour peaks during the short dry and short rainy season before the peaks in nests and chicks. The highest nest frequency was found in short grass habitats. Female Kori Bustard may undergo repeated nestings within a single breeding season. The adult sex ratio was female skewed during the breeding season. The Kori Bustard breeding season in the Serengeti plains is relatively long, lasting for almost nine months, and taking place during all seasons except for the long dry season. We recommend that management authorities conduct assessments of Kori Bustard recruitment as well as habitat suitability in the Serengeti ecosystem to develop future conservation strategies.     

Seasonal changes in courtship song and the medial preoptic area in male European starlings (Sturnus vulgaris)

In male starlings (Sturnus vulgaris) courtship song plays a critical role in mate attraction. During the breeding season courtship song occurs prior to copulation and appears to reflect male sexual arousal. Outside the breeding season starlings sing, but song appears unrelated to reproduction. The aromatization of testosterone (T), likely within the medial preoptic nucleus (POM), is critical for the expression of male sexual arousal. The present study was performed to determine whether seasonal changes in the POM might relate to seasonal changes in courtship singing behavior in male starlings. T concentrations, the volume of the POM, and aromatase within the POM were examined both during and outside of the breeding season in male starlings. Song was also recorded at these times both with and without a female present. The POM was largest and contained dense aromatase immunostaining only during the spring breeding season, when T concentrations were highest and males responded to a female with an increase in courtship song. Outside the breeding season the volume of the POM was small, T concentrations were low, and males displayed no changes in song expression in response to female conspecifics. Song bout length was positively related to POM volume, and males sang longer songs in spring. Only males with nestboxes in spring responded to a female, and the POM tended to be larger in these males, suggesting that nestbox possession might influence neuroplasticity within the POM. Overall, the findings suggest that T-dependent plasticity and aromatase activity within the POM might regulate courtship singing in a wild songbird.     

Behavioral responses of males to estradiol-treated females in the budgerigar (<Emphasis Type="Italic">Melopsittacus undulatus</Emphasis>)

We investigated the effects of estradiol-treated females on the behavior of male budgerigars. In comparison to control females, females given implants of estradiol showed elevated nest behavior and darker cere color, which are characteristics of breeding females. After we confirmed the efficacy of estradiol treatment on behavior and morphology, each female was paired with a male mate. Males paired with estradiol-treated females showed more courtship behavior (auditory and visual display, and courtship feeding) to their mates than males paired with control females. These data indicate that budgerigar females with a high estrogen level enhance males' courtship behavior. Since males did not show response to estradiol-treated females soon after females were introduced, effects of estradiol-treated female budgerigars may be mediated by the endocrine system, rather than wholly by the nervous system, of males. Electronic Publication     

Alternative reproductive tactics and courtship in the common goby

In the marine common goby Pomatoschistus microps , males either build nests and attract females to spawn or act as sneakers, forcing themselves into other males' nests during mating, to fertilize some of the eggs in the clutch. The reproductive tactics of small males were studied in the presence of surplus nest material and an excess of gravid females, while a larger, nestholding male spawned with a female. Of 24 small males, 13 only attempted to sneak, seven only courted females and/or spawned in their own nest, and four performed both behaviours. The small males who attracted females successfully were significantly larger than the ones who failed to do so. Females had been courting the small male more often when he subsequently spawned in his own nest, than when he did not get a female. The small males did not court the females very frequently, and their courtship rate did not depend on whether or not they attracted females. Instead, female courtship behaviour may be a cue for a male either to spawn as a nest holder or become a sneaker. The study pointed out the importance of female behaviour for male reproductive tactics, which could be more important in fish than is known currently.     

Female preference of the gobiid fish Eviota abax in relation to elongation of the first dorsal spine in males

The significance of elongation of the first and second spines of the first dorsal fin in male Eviota abax (Gobiidae) was examined in relation to female preference. Reproductive behavior occurred in three phases: male approach to female home, male leading female to nest, and male entering nest. During each phase, the female either accompanied the male or departed. The ratio of positive female mating responses to males before the elongated male dorsal spines were clipped and thereafter was compared in an aquarium. Females preferred male elongated spines in the first phase of courtship, but not in the second phase . Because females may be more exposed to predators during the second phase, owing to conspicuous male courtship, their greater receptivity to elongated dorsal spines in the first phase may decrease predation pressures during subsequent phases of courtship. Electronic Publication     

Male-male Competition and Female Mate Choice through Courtship Display in the Territorial Damselfish Stegastes nigricans

Both sexes of the herbivorous damselfish Stegastes nigricans maintain individual feeding territories. These territories are distributed contiguously, forming distinct colonies. Females visit male territories to spawn, and eggs are guarded by males until hatching. Male-male competition and female mate choice were studied in two colonies of different size compositions. Only larger individuals bred in both colonies. Some males in the large colony, that were larger than the breeding males in the small colony, did not succeed in reproducing probably because of severe attacks by the larger males while courting. However, females did not choose large size among breeding males. The most important male characteristic in female choice was the frequency of courtship displays in both colonies. Females in the large colony chose males mainly on the basis of the frequency of displays conducted in the females' territories, whereas females in the small colony chose males on the basis of the frequency of displays conducted in the males' territories. This difference may be a result of the difference in colony size. The distances between females' and males' territories were much greater in the large colony, and, because females cannot see courtship displays conducted in distant male territories, males in the large colony may have had to visit female territories frequently in order to conduct courtship near the females.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.     

Mate choice in Darwin's Finches

Female Geospiza conirostris on Isla Genovesa, Galapagos, pair preferentially with males who have had previous breeding experience. They choose mates on the basis of courtship behaviour and black adult plumage. By mating with experienced black males, they gain a fitness advantage in terms of fledgling production and recruitment of young into the breeding population. Behavioural signs of past breeding experience and black plumage are reliable age- and condition-dependent traits. We suggest that females use conspicuous black plumage to identify old males at a distance, then interactions through courtship to modify initial assessments. Females paired with inferior males may increase the genetic quality of their offspring by extra-pair copulations; results of heritability analysis of morphology are consistent with this suggestion. Females change mates at a frequency of 12–27% per breeding season. They re-pair with males who are generally old, experienced, and hold territories adjacent to the deserted male. Females that re-pair gain a benefit, whereas males who are deserted within a breeding season incur a cost of more than 50% of their future potential production for that season. We conclude that females in choosing males seek reliable indicators of potential parental care, and in addition they may seek indicators of genetic quality.     

Undirected Song Encourages the Breeding Female Zebra Finch to Remain in the Nest

Wild zebra finches sing frequently during the breeding season, but the vast majority of song is of the undirected song type that is not directed at any individual, and the function of which is obscure — it appears to be ignored by all potential recipients. It is sung close to the nest-site, has a peak in production during the egg-laying period, and diminishes thereafter. The incidence of undirected song is positively correlated with extra-pair courtship, a finding consistent with the hypothesis that it is a means of advertising availability for extra-pair matings. Typically, undirected song occurred outside the nest when the female was inside, and a positive relationship was found between the amount of singing given by the male during the 5-min interval immediately after the female entered the nest and the time she subsequently spent inside the nest. Keeping the partner inside the nest during her fertile period is an advantage to the male: it serves as a form of paternity protection against other males and it allows him opportunities to pursue his own extra-pair matings. Occupancy of the nest during laying is also a means of guarding against intraspecific brood parasitism, which was high at this colony.     

Mate Choice and Mating Pattern in a Stream Goby of the Genus Rhinogobius

The mate choice and mating pattern of a benthic goby Rhinogobius sp. CB (cross band type) were investigated in the Kamo River, Shikoku, Japan. During the breeding season, gravid females assumed a nuptial color and either males or females initiated a courtship display. Males preferentially courted a female of similar size to lead her to his nest, whereas females courted more frequently when they encountered a large male. Eggs in any one nest were always at the same developmental stage. Sampling data of nesting males and females indicated that, in more than half the nests, males gathered more than one female before spawning. In some nests with eggs, two or three females had spent ovaries, indicating that the eggs were laid by multiple females within a short span of time. However, a comparison between the total number of eggs which females would spawn in one nest and the number of eggs actually deposited suggested that eggs were contributed by one female in most nests. This low level of polygyny in spite of multiple female availability is attributed to a limited available spawning area of the nest.     

Parental defence in Long-eared Owls Asio otus: effects of breeding stage, parent sex and human persecution

The responses of male and female Long-eared Owls to a human approaching the nest were examined. Each nest was only tested once. Nest defence increased significantly throughout the breeding season because older chicks were defended more strongly than younger chicks and eggs. No correlations were however found between defence intensity and laying date, clutch- or brood-size. These results generally do not support either the renesting-potential hypothesis or the positive reinforcement hypothesis but are in accordance with both the age-investment hypothesis and the vulnerability hypothesis. Females defended nestlings more often and more vigorously than did males. Given division of labour between sexes and the related reversed sexual dimorphism in size, female Long-eared Owls may be more willing to engage in nest defence simply because they are always nearer eggs and chicks during all breeding stages and are larger/heavier than males; consequently they may allocate more time and energy to this activity. Finally, owls experiencing higher levels of human persecution took smaller risks when defending nests than owls breeding in an undisturbed area.