Comparing food limitation among three stages of nesting: supplementation experiments with the burrowing owl

Food availability is an important limiting factor for avian reproduction. In altricial birds, food limitation is assumed to be more severe during the nestling stage than during laying or incubation, but this has yet to be adequately tested. Using food‐supplementation experiments over a 5‐year period, we determined the degree and timing of food limitation for burrowing owls (Athene cunicularia) breeding in Canada. Burrowing owls are an endangered species and food limitation during the nestling stage could influence reproductive performance of this species at the northern extent of their range. Supplemented pairs fledged on average 47% more owlets than unfed pairs, except during a year when natural food was not limiting (i.e., a prey irruption year). The difference in fledgling production resulted from high nestling mortality in unfed broods, with 96% of all nestling deaths being attributed to food shortage. Supplemental feeding during the nestling period also increased fledgling structural size. Pairs fed from the start of laying produced the same number of hatchlings as pairs that received no supplemental food before hatch. Furthermore, pairs supplemented from egg laying to fledging and pairs supplemented during the nestling period alone had the same patterns of nestling survival, equal numbers of fledglings, and similar fledgling mass and structural size. Our results provide empirical support for the hypothesis that the nestling period is the most food‐limited phase of the breeding cycle. The experimental design we introduce here could be used with other altricial species to examine how the timing of food limitation differs among birds with a variety of life‐history strategies. For burrowing owls, and other species with similar life histories, long‐term, large‐scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation.     

Effects of food abundance, density and climate change on reproduction in the sparrowhawk Accipiter nisus

The reproductive success of predators depends on abiotic environmental conditions, food abundance and population density, and food abundance, density and their interactions may respond to changes in climatic conditions. Timing of reproduction by five of the eight numerically most common prey of the sparrowhawk Accipiter nisus advanced significantly since 1971, during a period of temperature increase. There was no evidence that mean laying date or any other reproductive parameter of sparrowhawks changed consistently during the study period 1977–1997. Laying date advanced and percentage of unsuccessful female sparrowhawks decreased with beech mast in the current year, an index of food abundance for avian prey. Mean laying date of sparrowhawks was advanced in warmer springs, and although mean clutch size was not larger in warm than in cold springs, mean brood size of successful pairs and breeding success increased in such springs, showing that sparrowhawks enjoyed a fitness gain when reproducing early. The timing of sparrowhawk reproduction with respect to the peak in abundance of fledgling prey increased, from a good match between mean timing of fledging by prey and maximum demand for food by the predator in 1977, to reproduction occurring later than the peak in fledging prey availability in 1997. The size of the breeding population of sparrowhawks was not predicted by mean spring temperature, the size of the breeding population the previous year or beech mast crop. The size of the post-breeding population was predicted by size of the breeding and post-breeding population the previous year and by the proportion of unsuccessful females the current year. These findings imply that sparrowhawks did not respond to change in climate, although climate changed the timing of reproduction by the main prey species.     

Effects of a food supplementation experiment on reproductive investment and a post-mating sexually selected trait in magpies Pica pica

Food availability is an important factor affecting breeding success in birds. Food supplementation experiments in birds have in general focused on the effects on reproductive success in terms of female investment (laying date, clutch size, egg size), however, it is also known that the estimation of mate quality based on sexually selected signals influences female reproductive investment. In the particular case of magpies, females use nest size, a post-mating sexually selected signal, to assess male's likelihood to invest in reproduction, and accordingly adjust reproductive investment (clutch size). Then, the possible effects of food supplementation on female reproductive investment could be mediated by other variables related to parental quality, such as nest size in magpies. In the present study, we explore if higher food availability in a magpie territory affected both male sexually selected traits (i.e. nest size) and female reproductive investment (laying date, egg size, clutch size). We performed a food supplementation experiment in which we experimentally increased food availability in several magpie territories, keeping others as controls. In food-supplemented territories, males built significantly larger nests and females significantly increased egg size by 4.1% compared to control females. Results suggest that the continuous provisioning of protein rich food allowed magpie females to increase egg size. However, laying date and clutch size did not differ between control and food-supplemented magpie pairs. Food availability also affected the relationship between female reproductive investment and nest size. In control territories, females decreased their egg size in response to a larger nest, whereas a tendency for the opposite relationship was revealed in food-supplemented territories. We discuss the possibility that magpie females adopt different strategies for reproductive investment according to food availability.     

Physiological ecology of Mediterranean blue tits (Parus caeruleus L.): effects of ectoparasites (Protocalliphora spp.) and food abundance on metabolic capacity of nestlings

The consequences of nest ectoparasites, such as Protocalliphora larvae, on nestling birds have been the subject of numerous studies. Despite observed reductions in mass and hematocrit of chicks from parasitized nests, no studies have found any effect of Protocalliphora on nestling survival, suggesting that fitness consequences of Protocalliphora are either weak or occur after fledging. From experiments on the metabolic performance of chicks, we found that parasitized chicks suffer from reduced thermogenic and metabolic capacities as a result of decreased mass and hematocrit. Hence, Protocalliphora may potentially affect nestling survival after fledging, when energetically costly activities such as flight and moult are undertaken. Previous studies have demonstrated an increase in parental feeding rate to compensate for the detrimental consequences of parasite infestation. We tested whether parasite effects on nestling aerobic capacity were dependent on food availability during the feeding period. Measures of caterpillar densities and experimental manipulations of parasite loads allowed us to investigate relationships among host, parasite, and environment. A positive relationship between chick aerobic and thermogenic performances and caterpillar density suggests that negative effects of parasitism may be offset by increased food availability. This study provides the first measurement of the effects of an ectoparasite on metabolic competence in wild birds and documentation of the effect of food availability on ectoparasite virulence using a quantitative measure of food abundance.     

Post‐fledging survival of Little Owls Athene noctua in relation to nestling food supply

Among the range of determinants of post‐fledging survival in altricial birds, the energy supply to the growing juveniles is likely to play a central role. However, the exact mechanisms shaping post‐fledging survival are poorly understood. Using a food supplementation experiment, we determined the effect of variation in food supply on the survival of juvenile Little Owls Athene noctua from hatching to 2 months post‐fledging. Experimental broods were food‐supplemented for 36 days during the nestling and the early post‐fledging period. The fate of 307 juveniles (95 of them provided with extra food) was determined by nest monitoring and radiotelemetry. In unsupplemented birds, the rates of survival measured at 5‐day intervals were lowest during the nestling stage, remained low during the early post‐fledging stage and steadily increased after about 2 weeks post‐fledging. Food supplementation substantially increased nestling survival, but we detected no direct treatment effect on post‐fledging survival. Instead, we found a strong indirect effect of food supplementation, in that fledglings of good physical condition had markedly higher chances of surviving the post‐fledging period compared with those in poor condition. Experimental food supplementation increased survival over the first 3 months from 45% to 64.6%. This suggests that energy reserves built up during the nestling stage influence post‐fledging survival and ultimately parental reproductive output. The low nestling and post‐fledging survival shows that the early life‐history stages constitute a crucial bottleneck of reproductive ecology in Little Owls. The strong treatment effects on the number of independent offspring indicate that natural variation in food supply is an important determinant of spatio‐temporal patterns in Little Owl demography.     

Reproductive measures and chick provisioning of Cory's Shearwater Calonectris diomedea borealis in the Azores

Capsule Timing of breeding influenced wing-length at fledging, and egg size may be an indicator of fledging weight and the amount of food received by chicks.

Aims To investigate chick growth, temporal patterns of chick food provisioning and the importance of indices of parental condition or quality, egg size and hatching date, to predict nestling body mass and wing-length at fledging, and compare breeding and chick feeding characteristics between colonies in the northeast Atlantic.

Methods A survey of Cory's Shearwater nests was carried out at Vila islet. A sample of 52 chicks, ringed and weighed at hatching, was selected to study chick growth and food provisioning.

Results Hatching success (51%) was much lower than fledging success (87%). Both hatching date and egg size contributed to explain wing-length at fledging, but hatching date, which was negatively correlated with wing-length at fledging, had the most important contribution (22%). There was some indication that egg size may explain variation in fledging weight and the amount of food received by chicks. Food delivery and feeding frequency of chicks varied throughout the chick development stage and three phases were distinguished: (1) 0–29 days, the highest feeding frequency values and a linear increase in food delivery; (2) 30–69 days, an oscillation in food delivery and medium feeding frequencies; (3) 70–90+ days, a sharp decrease in both food delivery and feeding frequency.

Conclusion Variation in food availability did not seem sufficient to override the overall importance of indices of parental quality in determining reproductive measures and chick provisioning. Breeding and feeding characteristics were similar between colonies in the northeast Atlantic, with variability in chick provisioning higher further south.     

Effect of food availability on nestling growth and fledging success in manipulated pallid swift broods

Growth rate and fledging success were assessed in natural and manipulated broods of the pallid swift Apus pallidus. Daily measurements of chick mass, wing length, and insect abundance allowed us to examine the short-term variation of chick growth in relation to food availability.
The number of fledged nestlings increased with brood size. Wing length and body mass were slightly but significantly smaller in larger broods, and the nestlings of enlarged broods needed longer to fledge. We discuss how these differences could influence survival after fledging.
Hatching asynchrony caused a significant difference in growth among siblings, and the difference between the oldest and youngest chick was greater in larger broods.
Chick growth was independent of daily food availability. We suggest that this was due to an increased effort of the parents at their expense, when food availability was poor.
The ability of this species to raise an additional chick is in line with most findings on birds, but partially in contrast with results for the common swift in which, at least during poor seasons, the additional nestling caused an increased mortality and lowered the reproductive success.     

Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus,Parus major)

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first‐egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak‐beech forest, and found that both advanced their mean first‐egg dates by 11–12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2–3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70–75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within‐individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.     

Fruit abundance may fine-tune timing of reproduction of the Chilean Elaenia (Elaenia chilensis), a long-distance migratory bird in South America

Timing of reproduction has a great impact on the breeding success of birds because a mismatch with the moment when environmental conditions are warm or when food is most plentiful can reduce nestling survival and increase the energetic cost of parental care. Consequently, birds synchronize gonadal maturation with the most favourable environmental conditions, using photoperiod changes throughout the year as an initial proximal cue. Additionally, non-photic cues, such as temperature and food abundance, may be necessary to fine-tune reproductive timing. However, the influence of non-photic cues on finely tuning reproductive timing is not yet fully understood for migratory birds. Here, we evaluate how much non-photic cues influence the reproductive timing of the Chilean Elaenia Elaenia chilensis, a long-distance migrant that reproduces in the Andean–Patagonian Forest. We assessed associations of mean temperature, ripe fruit and arthropod abundances with the number of nests in the laying period, and also with the number of nests with hatchlings. In both analyses we used cross-correlations and partial least squares path modelling. Mean temperature was not consistently associated with the breeding phenology of Chilean Elaenias. The increase in number of nests in the laying period was preceded by the increase in caterpillar abundance and coincided with the increase in ripe fruit abundance. The timing of nests with hatchlings matched with the timing of highest ripe fruit abundance. Both types of food could contribute to the beginning of reproduction of birds and be used as proximal cues by Chilean Elaenias. Ripe fruits would also be beneficial for nestling growth because parents feed them with fruits, and might potentially play an ultimate role in reproduction. Because ripe fruit abundance was related to the egg-laying and hatching stages, it may be the main cue used by Chilean Elaenias to fine-tune reproductive timing. These findings allow advances in our understanding of the importance of non-photic cues in the reproductive phenology of migratory birds and also generalize our knowledge among regions and taxa, as most studies on this topic focus on the Northern Hemisphere.     

Begging in Common Redstart Nestlings: Scramble Competition or Signalling of Need?

Begging behaviour by the young affects parental food distribution among nestlings of altricial birds. We present an analysis of two types of begging behaviour (assuming the front nest positions and gaping) based on videotaped natural nestling feeding in European common redstart (Phoenicurus phoenicurus). We test whether these types of begging support the predictions of two mathematical models: scramble competition with competitive asymmetries between nestlings [Anim. Behav. 27 (1979) 1210] or honest signalling model [Nature 352 (1991) 328]. None of the measured variables of nestling or parental behaviour were affected by body weight differences between siblings. In contrast, both gaping and nest positioning were affected by individual differences in nestling hunger. In agreement with the honest signalling model, hungrier nestlings gaped with higher probability and started to gape sooner after the arrival of the parent than did their less hungry nestmates. Those nestlings with the shortest latency to gape also received food more often. Nest positioning was related to nestling hunger in a way unforeseen by the existing models. The intervals between nestling position changes were several times longer than the intervals between parental feeding visits, and parents preferred to feed nestlings in front positions, so nestlings in front positions were always less hungry than nestlings in back. Hence the pattern of movements influenced the feeding decision in favour of the more satiated nestlings and acted against the effect of gaping. Nestling movement seemed to be caused by the less hungry nestlings moving actively from front to rear positions. Low mortality of individual nestlings within broods that survived to fledging and small within‐brood variation in fledging weights indicated low competition among nestmates. We suggest that there are two behavioural mechanisms that contribute to the equalization of fledging weights in common redstart nestlings: the signalling of need through gaping and the regular turnover of nestlings at front positions.     

Fitness consequences of timing of breeding in birds: date effects in the course of a reproductive episode

In seasonal environments, avian reproductive performance almost generally declines in the course of the season. Quantifying the associated fitness consequences of timing of breeding, i.e. of date‐related factors, is important for understanding the evolution of temporal patterns in avian life‐histories and for predicting consequences of climate change. The seasonal decline can also be caused by an effect of parental quality: individuals with high phenotypic quality may breed early. The results of existing experimental studies investigating whether date or quality effects cause the seasonal decline are inconsistent, indicating that both mechanisms might be involved. However, it remains unclear to what extent the confounding effect of quality occurs and what the fitness consequences of timing per se over a whole breeding episode are. In a cross‐fostering experiment using the barn swallows’ second broods we evaluated the causes for the seasonal decline in reproductive performance for three distinct periods of a reproductive attempt, the early nestling period, the late nestling period and the post‐fledging period, and we assessed the overall fitness consequences of timing per se. A seasonal decline in juvenile feather growth rate was mainly due to date effects in the late nestling period, although we determined quality effects during early nestling development. Date effects on survival were present in the post‐fledging period, but not in the nestling period. The decline in feather length due to date effects in the nestling period accounted for 9% of the seasonal decline in post‐fledging survival, whereas date effects arising only in the post‐fledging period caused 91% of the decline. These results suggest that date effects increase in the course of a reproductive episode. Thus, the benefits of an early timing of breeding can be quantified only when considering also the post‐fledging period. We suggest that the timing of breeding evolved through a trade‐off between date‐related benefits and quality‐related costs of early breeding.     

The importance of a main dish: nestling diet and foraging behaviour in Mediterranean blue tits in relation to prey phenology

Insectivorous birds rely on a short period of food abundance to feed their young; they must time their reproduction to match the timing of Lepidoptera larvae, their main prey. Apart from the net result (i.e. birds are timed or mistimed with respect to the food's peak), an important aspect is the possible influence of other factors, such as the seasonality of the environment or the abundance and diversity of species contributing to the caterpillar peak, on birds' phenology and their ability to cope well with unpredictable food supplies. In a 2-year study, we explored the seasonal variation of nestling diet in Mediterranean blue tits Cyanistes caeruleus and how reproductive parameters (nestling condition, provisioning rates) are affected by the phenology and composition of food. We also examined the influence of the synchrony between offspring needs and caterpillar peak in shaping the composition of the nestlings' diet. We found that the effect of synchrony on nestling condition varied between years which may be partially due to differences in food peak attributes. The adequacy of birds' timing in relation to prey phenology affected foraging decisions; those birds that were not able to correctly adjust their timing were forced to rely on less preferred prey (tortricids). In this sense, we found that relative contribution of tortricids (smaller caterpillars but easier to get) and noctuids (preferred prey but more difficult to find) to the diet influenced nestling condition and parental provisioning effort; parents performed fewer feeding events and reared heavier nestlings as the contribution of noctuids to the diet increased. The relationship between the proportion of caterpillars and nestling mass was curvilinear, whereas that parameter was negatively affected by the percentage of pupae. Our results show how changes in diet composition may contribute to explain the effect of mismatching on birds' breeding performance.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.     

Survival of Herring Gull Larus argentatus chicks: an experimental analysis of the need for early breeding

In 1985 and 1990, we manipulated the hatching date of Herring Gulls Larus argentatus by exchanging complete clutches between pairs which had started egg laying on different dates in the season. The aim was to investigate whether the observed seasonal decline in Hedging success can be explained by laying date. In both years, fledging success of advanced pairs followed the natural seasonal trend, supporting the date hypothesis. However, in both years, Hedging success of delayed pairs remained as high as that of control pairs with the same original hatching date as the foster parents, supporting the parental quality hypothesis. These data suggest that the delayed pairs may have compensated for less favourable environmental conditions. Although conspecific predation was the main direct cause of chick mortality, food shortage probably indirectly determined the mortality rate. A model of fledging success in relation to the timing of egg laying predicts that when food is abundant the optimal hatching date coincides with the mean hatching date of the colony (synchronization advantage). When food is less abundant, early breeders have the highest fledging success.     

Testing sources of variation in nestling‐stage nest success of Florida Scrub‐Jays in suburban and wildland habitats

Human modification of habitats can reduce reproductive success by providing novel cues to which birds may respond with behaviors that are actually maladaptive in those environments. Ad libitum human‐provided foods may provide the perception that urban habitats are food‐rich even as natural food availability decreases. Similarly, human activity may increase the perception that predation risk is high even as natural predators may decrease in abundance. In response, birds may reduce parental care with a subsequent cost to successful reproduction. Florida Scrub‐Jays (Aphelocoma coerulescens) in suburban areas have lower nest success during the nestling period than do wildland jays, possibly the result of such maladaptive responses, but maybe because of ecological differences with wildlands. We manipulated adult perception of predation risk and the availability of nestling foods in suburban and wildland areas to determine if these factors influenced parental care and nestling begging, and if the behavioral responses of adults influence nest survival during the nestling stage. Experimentally increasing perception of predation risk reduced parental care by both suburban and wildland females, but did not influence care by males. Increasing food availability, but not predation risk, had little influence on parental care, but resulted in decreased nestling begging rates and an increase in the frequency (pitch) of begging calls in both habitats. However, neither parental care nor food availability influenced nest survival during the nestling stage. Instead, the presence of helpers was the most important variable in nest survival analyses, suggesting that habitat‐specific differences in nest survival during the nestling stage were not simply the result of maladaptive parental behavior or shortage of nestling food resources in the suburban habitat. The lack of helpers combined with ecological differences, such as the abundance of nest predators, may be why fewer nests of Florida Scrub‐Jays survive during this stage in suburban areas.     

Ural owl sex allocation and parental investment under poor food conditions

Parents are expected to overproduce the less costly sex under poor food conditions. The previously regular 3-year cycle in the abundance of voles, the main prey of the Ural owl, Strix uralensis, temporarily disappeared in 1999–2001. We studied Ural owls' parental feeding investment and sex allocation during these poor-quality years. We sexed hatchlings and embryos in unhatched eggs of all 131 broods produced during these years. Population wide, the owls produced significantly more males (56%). The parental food investment in the brood was estimated by sorting out the prey remains in the bottom of nest boxes. Food delivered to 83 broods without chick mortality showed no clear sex-specific investment. Nestling mortality was equal in both sexes. Thus, evidence for an investment-driven sex allocation is weak. Neither laying date, brood size nor the female's condition correlated with offspring sex ratios. In these poor years, parents provided less food per chick and the fledging weight of daughters was reduced more than the weight of sons compared with years of high food abundance (1983 and 1986). We discuss, in relation to published studies, the possibility of a sex-allocation scenario where, under poor food conditions, a daughter's long-term fitness is reduced more than a son's.     

Breeding time,food supply and fitness components of Blue Tits Parus caeruleus in Mediterranean habitats

We analysed the relationship between the timing of food availability and within-season variation of both reproductive success and nestling body size of Blue Tits Parus caeruleus in Mediterranean habitats. Synchronization between food supply and reproduction was expected to be positively related to fitness components. We measured deviation from maximum food supply using a parameter that we called “time-lag”, which quantifies the degree of synchronization between the date of maximum food requirements by the nestlings and the date of maximum caterpillar supply in the habitat. This parameter was expected to be related to reproductive success as measured by the number and body-condition of fledglings. The predictions were that time-lag should be negatively correlated with the proportion of nestlings raised to fledging and the size of the fledglings. These predictions have been tested in different types of habitat. The results demonstrate that caterpillar supply during a critical nestling period can have a strong influence on fitness components. As predicted, synchronization with caterpillar supply is positively related to the number and body size of fledglings. Since there is large between-habitat variation in the timing of food supply, the key issue seems to be the adjustment to local patterns of food availability.     

Do parents play a role in the timing and process of fledging by nestling Mountain Bluebirds?

What causes young birds to leave nests remains unclear for almost all altricial species. For many years, the assumption was that parents often controlled the time of fledging by coaxing young from nests, e.g., by holding food within view, but out of reach, of nestlings. This assumption, though, was based solely on scattered anecdotal reports of such behavior. We used continuous video‐recording of nests to assess the role of parents, if any, in the timing and process of fledging of cavity‐nesting Mountain Bluebirds (Sialis currucoides). We placed perches ~50 cm in front of nest‐box entrances to give parents ample opportunity to display food to nestlings. We found no evidence that parents routinely initiated the fledging process. On the day of fledging, parents did not perch on supplemental perches with food more often, or for longer periods of time, than on the day before fledging. Also, after going to nest‐box entrances, parents never held food away from a nestling reaching for the food. Parents were usually absent (16 of 19 cases) when the first nestling fledged. In the remaining three cases, a parent perched with food in view of a nestling for 8, 15 and 65 s, respectively, just before that nestling fledged. Although these might have appeared to be attempts at coaxing, in each case, the parent was encountering, for the first time, a nestling partially emerging from the nest entrance. Parents may simply have hesitated to approach nests because the nestling's position prevented parents from delivering food in the normal manner. Finally, the rate at which parents fed nestlings on the day of fledging did not differ from the rate the day before, suggesting that parents do not try to use hunger to induce fledging. Our results are consistent with previous research suggesting that, in Mountain Bluebirds, it is a nestling that initiates fledging, typically when it reaches some threshold state of development.     

Reproductive parameters in relation to food supply in the whiskered tern ( Chlidonias hybrida )

It is generally accepted that breeding terns are sensitive to food supply and that their reproductive effort could be substantially affected by the availability and access to resources. In this study we examined reproductive parameters in the whiskered tern Chlidonias hybrida in relation to food supply during the courtship feeding period (food brought by males to females) over a 2-year period (2004–2005). We also studied whether the condition of the nestlings [body condition index (BCI)] was related to a proxy of the reproductive investment of the adults (the clutch size) during a season of food shortage. Behavioural observations showed a decrease in the intensity of male courtship feeding between years (2004 > 2005), and a strong shift in the relative abundance of the two prey groups (invertebrates/fish; invertebrate prey dropped from 88.0 to 49.3%) brought by males. This change in food delivery rates did not result in a delay in laying, but there was a significant difference in mean clutch size between years (2.71 ± 0.49 eggs in 2004 and 2.05 ± 0.78 eggs in 2005) without any within-year variation in relation to the laying dates. The egg size (volume and length) was related to the year (2004 < 2005), suggesting a trade-off in the quantity and the quality of eggs between the two seasons. We also found no evidence that the investment in a large clutch affected nestling BCI in the course of the food shortage season. Since many pairs (about 60%) interrupted breeding during the incubation stage, we assumed that parents that succeeded in rearing nestlings in these conditions were probably ‘high-quality’ individuals. Our results therefore showed that whiskered terns are sensitive to the varying food conditions they experienced throughout the courtship period. The diversity of prey types could be a key factor in the reproductive investment of this tern species.     

Annual variation in the reproductive hormone and behavior rhythm in a population of the Asian short-toed lark: Can spring temperature influence activation of the HPG axis of wild birds?

Although a number of studies have demonstrated a correlation between ambient temperature and the timing of reproductive behavior in many bird species, the relationships between temperature, activation of the hypothalamus–pituitary–gonad (HPG) axis, laying, and hatching in free-living birds, remain unclear. We investigated the relationships between spring temperature, reproductive hormones, and behavior, in a population of the Asian short-toed lark (Calandrella cheleensis) on the Inner Mongolian Grasslands in 2014, 2015 and 2016. LH and T levels peaked earliest in the year with the highest April temperature (2014) and latest in the year with the lowest April temperature (2016), and rose faster in 2014 than in 2015 or 2016. Laying and hatching occurred earliest in 2014 and latest in 2016. E2 also peaked earlier in 2014 than in the other two years but there was no significant difference in peak E2 levels among the three years. The peak of hatching only coincided with the peak in grasshopper nymph (the main food of nestlings) abundance in 2015 and the nestling fledging rate in 2015 was significantly higher than that in 2014 and 2016. We also conducted a controlled experiment on the effect of temperature on plasma LH, T and E2 levels in wild-caught larks, which shows that mean plasma LH, T and E2 levels in the 16 °C group all peaked 4 days earlier than in the 13 °C group. All these results suggest that activation of HPG endocrine axis in the Asia short-toed lark population is closely related to ambient temperature, and that this subsequently influences the timing of laying and hatching. The HPG axis' endocrine function is a physiological factor that mediates effects of ambient temperature on the timing of breeding. Temperature induced annual variation in the timing of hatching was, however, insufficient to synchronize the nestling period with the annual peak in grasshopper nymph abundance. Unusually warm, or cold, spring temperatures advanced, or delayed, grasshopper nymph development to an extent that exceeded the Asian short-toed lark's capacity to advance or delay the onset of breeding and reproductive success was lower in the two years of the study in which the nestling period was asynchronous with the peak of grasshopper nymph abundance. Long-term research on the relationship between the timing of reproduction and population demographics of the Asian short-toed lark is required to understand the effects of climate change on this species and on temperate birds in general.