Environmental regulation of the reproductive system in a flexibly breeding Sonoran Desert bird, the Rufous-winged Sparrow, Aimophila carpalis

We investigated reproductive regulation in male Rufous-winged Sparrows, Aimophila carpalis, a Sonoran Desert passerine that breeds after irregular summer rains. Field and captive data demonstrate that increased photoperiod stimulates testicular development in March and maintains it until early September. Free-living birds caught in July and placed on captive long days (16L: 8D) maintained developed testes for up to 7 months, and free-living birds caught in September, during testicular regression, redeveloped testes when placed on captive long days, indicating that these birds were still photosensitive. Captive birds on long days maintained testicular development when exposed to temperatures mimicking those occurring during regression in free-living birds. In free-living birds, testicular development was observed during spring and summer, but unless this was associated with rainfall, breeding (indicated by juveniles) did not occur. Large increases in plasma luteinizing hormone (LH) in free-living males were correlated with heavy rainfall in July/August, when the birds bred, and in November, when they did not breed. In captive birds, plasma LH concentrations were unresponsive to photoperiodic changes, but may have responded to social cues. Plasma prolactin concentrations were directly correlated with photoperiod in free-living birds, but an effect of photoperiod on prolactin secretion was not seen in captive birds. It is concluded that male Rufous-winged Sparrows use long photoperiods to stimulate and maintain testicular development, but exposure to long photoperiods does not terminate breeding by inducing absolute photorefractoriness. The specific timing of reproductive behaviors is apparently determined by elevated plasma LH coinciding with long day stimulated gonad development.     

Auditory stimulation of reproductive function in male Rufous-winged Sparrows, Aimophila carpalis

Prolonged exposure to conspecific song stimulates gonadal function and reproductive hormone secretion in female birds but few studies have investigated the physiological effects of conspecific song exposure on males outside of short-term, aggressive interactions. We exposed male Rufous-winged Sparrows, Aimophila carpalis, either to conspecific song (CS Song), to heterospecific song (Black-throated Sparrow, Amphispiza bilineata; HS Song), or to no recorded song (No Song) for 59 consecutive days (two h per day). Birds were exposed to short days (8L:16D) for the first 21 days of treatment and were then transferred to long days (13L:11D) for the remaining 38 days. During long day exposure, CS Song birds experienced faster growth of testes than HS Song and No Song birds. HS Song birds also grew their testes faster than No Song birds. Plasma luteinizing hormone (LH) and testosterone did not differ between CS Song and No Song birds. However, plasma LH was higher in HS Song birds compared to other groups. There were no differences in hypothalamic immunocytochemical labeling for gonadotropin-releasing hormone, its precursor proGnRH, or gonadotropin-inhibitory hormone, nor were there differences in two song control nuclei volumes (HVC and RA) between CS Song and No Song treatment groups. Furthermore, we found no effect of heterospecific song on free-living Rufous-winged Sparrow aggressive behaviors. These data indicate that long-term exposure to auditory stimuli, such as song, can influence the reproductive system of male songbirds and different types of auditory stimuli can have differential effects on reproductive function.     

Plasticity of the Rufous-winged Sparrow, Aimophila carpalis, song control regions during the monsoon-associated summer breeding period

In most temperate zone songbirds, exposure to increasing photoperiod in the spring stimulates the reproductive system and induces reproductive behaviors. Additionally, the brain regions that control singing (song control regions; SCRs) are larger during the breeding season, thus paralleling changes in the activity of the reproductive system. However, in some birds, environmental factors other than photoperiod initiate breeding. For example, free-living male Rufous-winged Sparrows develop their testes in March due to increasing photoperiod, but have relatively low plasma T until after they begin to breed, usually in July, during the monsoon period when day length is declining. We tested the hypothesis that SCRs grow and singing behavior increases after the monsoon rains begin. We captured adult male Rufous-winged Sparrows in July 2002, 7 days before and 20 days after the monsoon rains began, euthanized birds in the field, collected their brains, and measured SCR volumes from sections immunostained for the neuronal marker NeuN. In June and July 2006, we measured song rates in the field before and after the monsoon rains. SCR volumes were larger and singing behavior increased after the onset of the monsoon rains, coinciding with the initiation of breeding. Unlike in other species studied so far, SCR volumes grew as day length was decreasing. Comparative studies utilizing species that do not breed when day length is increasing may provide information on the relative contributions of various environmental factors to SCR neuroplasticity.     

Alula size signals male condition and predicts reproductive performance in an Arctic‐breeding passerine

While studies of achromatic plumage signaling are scarce relative to chromatic ornaments, achromatic ornaments have the potential to act as an efficient form of visual communication due to the highly conspicuous contrast between black and white body regions. Recently, achromatic plumage reflectance has been shown to indicate condition, yet the condition‐dependence of achromatic patch size remains unstudied. Here we show the first evidence that alula size, an achromatic plumage patch, has the potential to signal a male’s condition and predict reproductive performance. In Arctic‐breeding snow buntings Plectrophenax nivalis, the size of the alula simultaneously predicted pre‐breeding physiological health and the number of offspring produced, through an intermediate variable (lay date). Snow buntings appear to pair assortatively; males and females arriving earlier pair together, and changes in body condition over the breeding season are positively related within pairs. We suggest that simple achromatic plumage patches, like alula size, have the potential to act as condition‐dependent signals. Consequently, females may benefit from assessing these signals to reliably evaluate a male’s condition and reproductive potential as a means of maximizing their reproductive success.     

Female survival,lifetime reproductive success and mating status in a passerine bird

In facultatively polygynous birds, secondary females of polygynously mated males typically have reduced annual reproductive success, because polygynous males provide less paternal care than monogamous males. Life history theory predicts that, as a result of increased reproductive investment, secondary females should suffer from reduced survival and lifetime reproductive success, but previous studies provided only weak support for this hypothesis. We used 7 years of data to study the fitness of female collared flycatchers Ficedula albicollis in relation to mating status by estimating survival and lifetime reproductive success. Taking differences in recapture probability into account, a mark-recapture analysis revealed that females observed at least once to breed as secondary female had higher survival than other females. This relationship was not confounded by laying date, because when we assessed the impact of laying date on survival, we found similar survival patterns. Females of polygynous males had reduced breeding success in terms of number of young fledged during the current reproductive event. However, during their lifetime females found at least once in primary or secondary mating status produced significantly more eggs, and at least the same number of fledglings and recruits as monogamous females. Thus, in the collared flycatcher, females of polygynously mated males seem to suffer from mating status during the most recent reproductive event, but considering survival and lifetime reproductive success, the apparently disadvantageous mating event is not necessarily associated with reduced residual reproductive value.     

Arrival fat and reproductive performance in a long-distance passerine migrant

Long-distance passerine migrants deposit substantial fat stores to fuel their migratory journey. Many of those migratory birds arrive at their northerly breeding grounds with larger fat stores than were necessary to reach their breeding area. Both male and female American Redstarts ( Setophaga ruticilla) arrived to breed in Michigan's Upper Peninsula with fat, and females arrived with more fat than males in 2 out of 3 years. We test the hypothesis that migrants arriving at the breeding grounds with more body fat have higher reproductive success than birds arriving with little or no fat. Females, and to a lesser extent males, that arrive with fat experience gains in reproductive performance as evidenced by increased clutch size, egg volume, and nestling mass. The results have implications for understanding how events occurring during one phase of the annual cycle influence survival and/or reproductive performance in subsequent phases.     

Seasonal variability in survivorship of a cooperatively breeding tropical passerine

Survival of tropical passerines is thought to be higher than those in northern temperate regions, but relatively few tropical studies have addressed this issue, particularly in tropical Asia. We examined factors that may have influenced the survival rate of a cooperatively breeding bird, the puff-throated bulbul (Alophoixus pallidus), in an evergreen forest in northeastern Thailand. These factors included year, season (breeding and non-breeding), sex, and presence of helper(s) in a family group. We present evidence of breeding season-dependent survival in a tropical passerine using an information theoretic approach based on both mark-recapture and resighting data collected during 6 years of study. Based on colour-banded adults the annual survival rate did not vary significantly among years (average = 0.85 ± 0.02 SE). The mean lifespan (MLS) for the population was 6.22 ± 4.38 SE years. Survivorship was lower during the breeding season (0.89 ± 0.02 SE) than during the non-breeding season (0.96 ± 0.02 SE). The MLS of males and females was 6.70 ± 7.73 SE and 5.87 ± 4.88 SE years, respectively. The annual survival rate we observed was high compared to the estimates of other tropical and temperate passerines, possibly due to the relatively stable climatic conditions in tropical latitudes and puff-throated bulbuls being generalists that exploit a wide range of food resources both in space and time.     

Long-term brood division and exclusive parental care in a cooperatively breeding passerine

In some species of birds, individual adults feed different subsets of the brood after the young have left the nest. However, few studies have shown that such ‘brood division’ represents long-term separation into subfamilies, rather than short-term biases in care, and the function of brood division is unknown. Most functional hypotheses assume that brood division is an adult adaptation to increase reproductive success; others suggest that fledglings are responsible for brood division, or that brood division is the outcome of conflicts of interest within families. We tested for long-term brood division in the white-browed scrubwren, Sericornis frontalis, a cooperatively breeding passerine, and assessed functional hypotheses, using behavioural, morphological and molecular evidence. We found that 71% of scrubwren broods became divided in the second week after fledging, and remained divided for the following 4-5 weeks of care. Furthermore, most individual young were fed almost exclusively by a single adult, even in undivided broods and broods of one. Brood division did not arise through adults caring for young of a specific sex, nor did genetic paternity explain division. Tests of other hypotheses were indirect, relying for example on testing mechanisms rather than predictions. We rejected the hypotheses that brood division was primarily an adaptation to reduce the mean or variance in predation, or that it was likely to increase the rate at which young were provisioned. Fledglings probably had an active role in maintaining brood division, with fledglings that had been biggest in the nest tending to be fed by adults that had fed nestlings at the greatest rate. This suggests that the most dominant fledgling may sequester the care of the best feeder. We also found evidence that brood division might be an adult adaptation to reduce the effects of sibling competition, or that it might arise through conflicts of interest among adults. We further suggest that social specialization, resulting from learning about a particular individual, might bring additional advantages to long-term brood division. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Demographic response to environmental variation in breeding,stopover and non-breeding areas in a migratory passerine

Demographic rates of migratory species passing through several areas during their annual cycle may be affected by environmental conditions at each of these areas. Recent studies provide evidence that their impact is not necessarily immediate, but can be delayed. We studied survival, reproductive success and arrival date at the breeding grounds of red-backed shrikes Lanius collurio, a trans-Saharan migrant, in relation to weather and vegetation on the breeding grounds, the stopover sites during migration and in the wintering areas. These environmental factors are used as proxy of the shrike’s food supply. We analysed detailed demographic data of some 4,600 individuals from 25 years with multistate capture–recapture and mixed models. Survival probabilities of juveniles and breeders of both sexes varied in parallel across time, suggesting that all cohorts were sensitive to similar causes of mortality. Reproductive performance increased with temperature and decreased with rainfall on the breeding area. Moreover, it increased with vegetation cover in the Sahelian stopover area used on autumn migration suggesting a carry-over effect. Arrival date was negatively affected by spring temperatures in the breeding area. Hence, demographic rates were affected by environmental factors on the breeding grounds, but also outside and elsewhere. This suggests that the shrike’s population dynamics are driven by environmental factors operating at various scales of space and time. However, only a small amount of the temporal variation in demographic rates is explained by the environmental factors considered, suggesting that additional factors, such as those operating during migration, might be important.     

Reproductive photorefractoriness in rams and accompanying changes in the patterns of melatonin and prolactin secretion

Exposure of rams to alternating 16-week cycles of long and short days (16L:8D and 8L:16D) results in periods of testicular regression followed by testicular redevelopment, and there is an inverse relationship between the blood levels of prolactin and testis activity. In this study, two groups of rams were held under long or short day lengths for a period of 94 weeks. When held under either long or short days for more than 16 weeks, the animals showed spontaneous changes in gonadal activity and in the secretion of prolactin, both of which were no longer correlated with the prevailing photoperiod, i.e., they became photorefractory. The photorefractoriness was characterized by cyclical changes in testis function which were independent of day length. The period of these spontaneous cycles was similar during both treatment regimens (long days: 40.9 +/- 1.5 weeks; short days: 38.1 +/- 0.33 weeks), suggesting the presence of an endogenous pacemaker for the reproductive system. The changes in blood prolactin levels during photorefractoriness were no longer correlated with testis activity, and though cyclical, the period lengths differed under the two regimens (long days: 31.8 +/- 1.4 weeks; short days: 48.6 +/- 0.3 weeks). The rates of change in testis function and prolactin secretion were slower during the refractory state than during the photosensitive state. Upon switching the rams to a different photoperiod after the 94 weeks of exposure to fixed day lengths, the rams showed relatively rapid testicular and prolactin responses. Photoperiodic information appears to be relayed to the endocrine system through the daily pattern of melatonin secretion by the pineal. We measured the daily blood levels of melatonin on several occasions during phases of photosensitivity and photorefractoriness in the same group of rams. During the first 21 weeks under both lighting treatments, the rams showed synchronized daily patterns in their blood levels of melatonin, with elevated levels occurring mainly during the daily period of darkness. Similar synchronized daily rhythms were also seen when the rams were switched to a different photoperiod following 94 weeks of exposure to either long or short days. Between Weeks 21 and 94, the daily rhythms of melatonin did not occur consistently in all rams; often, the patterns differed markedly between individual rams held under the same day length and peak levels of melatonin were not always confined to periods of darkness.(ABSTRACT TRUNCATED AT 400 WORDS)     

Low intensity, mixed livestock grazing improves the breeding abundance of a common insectivorous passerine

Livestock grazing is a major driver of ecosystem change and has been associated with significant declines in various bird species in Britain and worldwide. However, there is little experimental evidence to show how grazing affects bird populations. We manipulated livestock densities in a replicated field experiment and found that mixed sheep and cattle grazing, at low intensity, improved the breeding abundance of a common upland passerine, the meadow pipit Anthus pratensis, after two years. Plots stocked with sheep alone (at high or low density) or not stocked at all held fewer pipit territories. Despite a year-on-year decline in pairs of meadow pipits in intensively grazed plots, we found no effect of sheep number on breeding abundance. Our results support the hypothesis that mixed species of herbivores generate greater heterogeneity in vegetation structure, which modifies prey availability, resulting in a greater abundance of birds. The results of our study should inform the management of grassland areas and enhance the abundance of some bird species, particularly in areas that have seen significant shifts from mixed livestock grazing to grazing dominated by single species of animals.     

Fine-scale genetic structure and its consequence in breeding aggregations of a passerine bird

The pattern of fine-scale genetic structure in a population may reflect current biological processes of the species, such as natal dispersal, the breeding system and demography. We investigated the spatial distribution of nests and fine-scale genetic structure during two breeding seasons in a population of a weakly territorial, flock-living passerine bird, the vinous-throated parrotbill, Paradoxornis webbianus . Nest distribution was clustered. There were two peaks of egg laying within each breeding season, and spatial clustering of nests was more extreme during the second peak after controlling for breeding density. The patterns of genetic structure during the breeding season varied with parental sex and season. Genetic structure occurred during the second laying peak: males breeding within 200 m of one another at this time were significantly more closely related than males breeding farther apart. However, no apparent genetic structure was detected in males during the first laying peak or among females in either laying peak. These results reveal male-oriented kin affiliation during part of the breeding season in this species. Furthermore, juvenile recruitment into the winter flocks was positively related to this increased relatedness among males via kin affiliation. This study implies that fine-scale genetic structure during the breeding season could be a factor determining individual fitness and may play an important role in our understanding of the evolution of social systems.     

Growth response, endocrine profiles and reproductive performance of fine-wool ewe lambs treated with ovine prolactin before breeding

Twenty-four 6-mo-old ewe lambs received one of two ovine prolactin (oPRL) treatments 28 d before fall breeding. Beginning on the first day of treatment (Day 0), 12 lambs received a subcutaneous injection (12 ml) of a carrier vehicle (0 mg oPRL) on alternate days for 28 d while 12 lambs received injections containing 5 mg oPRL. On Days 0 and 28, jugular blood was collected from six lambs in each group before treatment and at 30-min intervals for 6 h thereafter. Neither feed intake, efficiency of gain nor animal weights differed (P > 0.20) between groups. One hour after treatment on Day 0, ewe lambs receiving 5 mg oPRL had greater (P < 0.10) serum PRL levels than did controls (121.9 and 61.5 +/- 24.7 ng/ml, respectively). Differences in serum PRL persisted throughout remaining sampling intervals on both Days 0 and 28. Serum samples obtained on alternate days during the 28-d treatment period revealed no differences (P > 0.20) in PRL concentrations between control (48.3 +/- 5.3 ng/ml) and oPRL-treated (55.7 +/- 5.3 ng/ml) ewes. Neither serum insulin nor growth hormone responded (P > 0.05) to exogenous oPRL on either Day 0 or 28. No difference (P > 0.30) in percentage of ewe lambs cycling during treatment or breeding was detected between groups. Subsequent lambing percentages were similar (P > 0.30), with 36.4% of control and 25.0% of oPRL-treated ewes producing offspring. Administering 5 mg oPRL on alternate days for 28 d before breeding did not enhance growth and(or) reproductive performance in virgin ewe lambs.     

Antioxidants in the egg yolk of a wild passerine: differences between breeding seasons

Carotenoids and fat-soluble vitamins play crucial roles in several physiological processes. Yolk carotenoid composition may be influenced by the bird's dietary intake and by the possible discrimination during carotenoid metabolism. Information regarding the pigment composition of passerine eggs is very limited. In the present 2-year study, we determined the carotenoid and vitamin concentrations and the percentage profile of carotenoid components in collared flycatcher (Ficedula albicollis) eggs. The major carotenoid was lutein in both years, followed by cis-lutein, zeaxanthin and beta-carotene as other significant components. Carotenoid concentration was higher in 2000 than in 2001. As caterpillar abundance and ambient temperature was lower in 2001 than in 2000, we suggest that the decreased availability of carotenoid-rich resources and the elevated antioxidant demand of the female restricted the carotenoid allocation to the eggs. Our study is the first to indicate that the concentration and profile of yolk carotenoids may considerably vary in the same wild bird population between breeding seasons with different environmental conditions. Compared to other birds, the yolk of this species was relatively rich in vitamins A and E. Vitamin E concentration was positively correlated with total carotenoid concentration, which may be beneficial for the nestlings as these antioxidants synergistically provide better protection.     

Establishing the breeding provenance of a temperate-wintering North American passerine, the Golden-crowned Sparrow, using light-level geolocation

The migratory biology and connectivity of passerines remains poorly known, even for those that move primarily within the temperate zone. We used light-level geolocators to describe the migratory geography of a North American temperate migrant passerine. From February to March of 2010, we attached geolocator tags to 33 Golden-crowned Sparrows (Zonotrichia atricapilla) wintering on the central coast of California, USA, and recovered four tags the following winter (October to December 2010). We used a bayesian state-space model to estimate the most likely breeding locations. All four birds spent the breeding season on the coast of the Gulf of Alaska. These locations spanned approximately 1200 kilometers, and none of the individuals bred in the same location. Speed of migration was nearly twice as fast during spring than fall. The return rate of birds tagged the previous season (33%) was similar to that of control birds (39%), but comparing return rates was complicated because 7 of 11 returning birds had lost their tags. For birds that we recaptured before spring migration, we found no significant difference in mass change between tagged and control birds. Our results provide insight into the previously-unknown breeding provenance of a wintering population of Golden-crowned Sparrows and provide more evidence of the contributions that light-level geolocation can make to our understanding of the migratory geography of small passerines.     

Short-day experience is not a prerequisite for the termination of photorefractoriness in the reproductive cycle of baya weaver,Ploceus philippinus

In most photoperiodic avian forms (irrespective of temperate or tropical distribution) including the baya weaver,Ploceus philippinus, seasonal reproduction comes to an end due to the development of a photoperiodically controlled photorefractory phase when birds cease to respond to the stimulatory effect of long days. In the present paper photoperiodic control of the termination of photorefractory phase has been examined by studying the effect of short-day exposure lasting 4–6 months on long-day response of birds. Results indicate that unlike in other photoperiodic birds short-day exposure of winter is not a prerequisite for the termination of photorefractory phase in the reproductive cycle of baya weaver. Artificial long days on the other hand hasten the termination of this phase. Refractory phase in baya weaver, therefore, unlike that in temperate forms, is a temporary state resulting most likely from a sequel of physiological events triggered by long days of spring/summer which temporarily mask the photostimulatory response. Spontaneous termination of photorefractoriness in birds of tropical habitats may have a selective value imparting to the reproductive cycle the necessary elasticity for adaptation to diverse ecological conditions.