Fledgling survival increases with development time and adult survival across north and south temperate zones

Slow life histories are characterized by high adult survival and few offspring, which are thought to allow increased investment per offspring to increase juvenile survival. Consistent with this pattern, south temperate zone birds are commonly longer‐lived and have fewer young than north temperate zone species. However, comparative analyses of juvenile survival, including during the first few weeks of the post‐fledging period when most juvenile mortality occurs, are largely lacking. We combined our measurements of fledgling survival for eight passerines in South Africa with estimates from published studies of 57 north and south temperate zone songbird species to test three predictions: (1) fledgling survival increases with length of development time in the nest; (2) fledgling survival increases with adult survival and reduced brood size controlled for development time; and (3) south temperate zone species, with their higher adult survival and smaller brood sizes, exhibit higher fledgling survival than north temperate zone species controlled for development time. We found that fledgling survival was higher among south temperate zone species and generally increased with development time and adult survival within and between latitudinal regions. Clutch size did not explain additional variation, but was confounded with adult survival. Given the importance of age‐specific mortality to life history evolution, understanding the causes of these geographical patterns of mortality is important.     

Corticosterone,testosterone and life-history strategies of birds

Steroid hormones have similar functions across vertebrates, but circulating concentrations can vary dramatically among species. We examined the hypothesis that variation in titres of corticosterone (Cort) and testosterone (T) is related to life-history traits of avian species. We predicted that Cort would reach higher levels under stress in species with higher annual adult survival rates since Cort is thought to promote physiological and behavioural responses that reduce risk to the individual. Conversely, we predicted that peak T during the breeding season would be higher in short-lived species with high mating effort as this hormone is known to promote male fecundity traits. We quantified circulating hormone concentrations and key life-history traits (annual adult survival rate, breeding season length, body mass) in males of free-living bird species during the breeding season at a temperate site (northern USA) and a tropical site (central Panama). We analysed our original data by themselves, and also combined with published data on passerine birds to enhance sample size. In both approaches, variation in baseline Cort (Cort0) among species was inversely related to breeding season length and body mass. Stress-induced corticosterone (MaxCort) also varied inversely with body mass and, as predicted, also varied positively with annual adult survival rates. Furthermore, species from drier and colder environments exhibited lower MaxCort than mesic and tropical species; T was lowest in species from tropical environments. These findings suggest that Cort0, MaxCort and T modulate key vertebrate life-history responses to the environment, with Cort0 supporting energetically demanding processes, MaxCort promoting survival and T being related to mating success.     

Prediction of mean adult survival rates of southern African birds from demographic and ecological covariates

Estimates of annual survival rates of birds are valuable in a wide range of studies of population ecology and conservation. These include modelling studies to assess the impacts of climatic change or anthropogenic mortality for many species for which no reliable direct estimates of survival are available. We evaluate the performance of regression models in predicting adult survival rates of birds from values of demographic and ecological covariates available from textbooks and databases. We estimated adult survival for 67 species using dead recoveries of birds ringed in southern Africa and fitted regression models using five covariates: mean clutch size, mean body mass, mean age at first breeding, diet and migratory tendency. Models including these explanatory variables performed well in predicting adult survival in this set of species, both when phylogenetic relatedness of the species was taken into account using phylogenetic generalized least squares (51% of variation in logit survival explained) and when it was not (48%). Two independent validation tests also indicated good predictive power, as indicated by high correlations of observed with expected values in a leave‐one‐out cross validation test performed using data from the 67 species (35% of variation in logit survival explained), and when annual survival rates from independent mark–recapture studies of 38 southern African species were predicted from covariates and the regression using dead recoveries (48%). Clutch size and body mass were the most influential covariates, both with and without the inclusion of phylogenetic effects, and a regression model including only these two variables performed well in both of the validation tests (39 and 48% of variation in logit survival explained). Our regression models, including the version with only clutch size and body mass, are likely to perform well in predicting adult survival rate for southern African species for which direct survival estimates are not available.     

Sperm competition in tropical versus temperate zone birds

Sperm competition represents an important component of post-copulatory sexual selection. It has been argued that the level of sperm competition declines in birds towards the equator. However, to date, sperm competition estimates have been available mainly for avian species inhabiting the northern temperate zone. Here we apply a novel approach, using the coefficient of between-male variation (CV_bm) in sperm size as an index for sperm competition risk, in a comparative analysis of 31 Afrotropical and 99 northern temperate zone passerine species. We found no difference in sperm competition risk between the two groups, nor any relationship with migration distance. However, a multivariate model indicated that sperm competition risk was highest in species with a combination of low body mass and few eggs per clutch. The effect of clutch size was most pronounced in tropical species, which indicates that sperm competition risk in tropical and temperate species is differently associated with particular life-history traits. Although tropical species had lower sperm competition risk than temperate zone species for overlapping clutch sizes, the idea of a generally reduced risk of sperm competition in tropical birds was not supported by our analysis.     

Evolution of clutch size in cavity-excavating birds: the nest site limitation hypothesis revisited

There are two major competing hypotheses for variation in clutch size among cavity-nesting species. The nest site limitation hypothesis postulates that nesting opportunities are more limited for weak excavators, which consequently invest more in each breeding attempt by laying larger clutches. Alternatively, clutch size may be determined by diet; the clutch sizes of strong excavators may be smaller because they are able to specialize on a more seasonally stable prey. We built a conceptual model that integrated hypotheses for interspecific variation in clutch size and tested it with comparative data on life-history traits of woodpeckers (Picidae) and nuthatches (Sittidae). In most analyses, diet explained more variation in clutch size among species than did propensity to excavate. Migratory status was positively associated with clutch size but was difficult to distinguish from diet since resident species consumed more bark beetles (a prey available in winter) and had smaller clutches than migratory species. The literature suggests that cavities are not limited in natural, old-growth forests. Although our data do not rule out nest site limitation, we conclude that annual stability of food resources has a larger impact on the evolution of clutch sizes in excavators than does limitation of nest sites.     

Life-history of two African Sylvia warblers: low annual fecundity and long post-fledging care

The investigation of factors that cause differences in life-history traits between temperate and tropical birds is often hampered by a lack of knowledge about tropical species. Even within the well-known warblers of the genus Sylvia , which include resident species from temperate and tropical regions as well as migrants, there are few data from tropical species. We investigated the breeding biology of the tropical species Sylvia lugens and S. boehmi in a 2-year study in Kenya. Both species had a clutch size of 2.0 and laid c.  3.7 clutches per year. Breeding was characterized by long incubation periods ( S. lugens 14.5 days, S. boehmi 15.0 days), long nestling periods (16.0 and 12.9 days, respectively) and high predation rates (Mayfield nest success S. lugens 33.2%, S. boehmi 19.4%). Annual fecundity was 2.3 fledglings in S. lugens and 1.4 fledglings in S. boehmi . After fledging, the young birds were fed for 37.5 days ( S. lugens ) and 58.5 days ( S. boehmi ) (time to independence) and they stayed in their parents' territory for days or weeks, even after feeding had stopped. Fledgling survival until independence was 55.4% in S. lugens and 69.2% in S. boehmi . In general, S. lugens and S. boehmi have smaller but more numerous clutches, longer developmental periods, higher nest predation rates, lower annual fecundity and longer post-fledging care than their temperate congenerics.     

Offspring fitness and individual optimization of clutch size

Within-year variation in clutch size has been claimed to be an adaptation to variation in the individual capacity to raise offspring. We tested this hypothesis by manipulating brood size to one common size, and predicted that if clutch size is individually optimized, then birds with originally large clutches have a higher fitness than birds with originally small clutches. No evidence was found that fitness was related to the original clutch size, and in this population clutch size is thus not related to the parental capacity to raise offspring. However, offspring from larger original clutches recruited better than their nest mates that came from smaller original clutches. This suggests that early maternal or genetic variation in viability is related to clutch size.     

Apparent survival of tropical birds in a wet,premontane forest in Costa Rica

Despite the importance of tropical birds in the development of life history theory, we lack information about demographic rates and drivers of population dynamics for most species. We used a 7‐year (2007–2013) capture‐mark‐recapture dataset from an exceptionally wet premontane forest at mid‐elevation in Costa Rica to estimate apparent survival for seven species of tropical passerines. For four of these species, we provide the first published demographic parameters. Recapture probabilities ranged from 0.21 to 0.53, and annual estimates of apparent survival varied from 0.23 to 1.00. We also assessed the consequences of inter‐annual variation in rainfall on demographic rates. Our results are consistent with inter‐annual rainfall increasing estimates of apparent survival for two species and decreasing estimates for three species. For the three species where we could compare our estimates of apparent survival to estimates from drier regions, our estimates were not consistently higher or lower than those published previously. The temporal and spatial variability in demographic rates we document within and among species highlights the difficulties of generalizing life history characteristics across broad biogeographic gradients. Most importantly, this work emphasizes the context‐specific role of precipitation in shaping tropical avian demographic rates and underscores the need for mechanistic studies of environmental drivers of tropical life histories.     

Lower annual fecundity in long-distance migrants than in less migratory birds of temperate Europe

We use widely supported handbook data on annual fecundity (clutch size × annual number of normal broods) to obtain indications related to the relative costs of long-distance migration compared to lower levels of migratoriness. Our examples show that the yearly production of eggs in congeneric passerines of similar size from temperate Europe is lower in long-distance migrants than in their less migratory relatives. The same tendency shows up in a more heterogeneous sample of non-passerines. In most passerines and in one among five pairs of non-passerines, this is due to longer breeding periods allowing a higher number of clutches in the less migratory species in spite of a tendency towards larger clutches in the passerine long-distance migrants. If both migratory types have only one clutch (as in one species pair of the passerines and in four non-passerine pairs) the trend towards larger clutches was reversed between the two types. The higher fecundity of the less migratory species suggests that wintering under harsh conditions may be more expensive than trans-Sahara migration among similar species. In keeping with this result, Ortolan and Rock Buntings (Emberiza hortulana and E. cia) breeding syntopically (and, in the 1980s, still in relatively stable populations) in an inner Alpine valley provide a well-studied example illustrating the high longevity of the migrants compared to the relatively short life span of the residents.     

REPRODUCTIVE COST,AGE-SPECIFIC SURVIVAL AND A COMPARISON OF THE REPRODUCTIVE STRATEGY IN TWO EUROPEAN TITS (GENUS PARUS)

Theoretical analyses of optimal reproductive rates usually assume a trade-off between offspring production and parental survival. This study verified a survival cost for willow tit males; nonbreeding males survived better than males attending a brood. Theory also predicts a smaller clutch size in birds that are less successful in transforming reproductive investments into mature offspring. As predicted, we found that crested tits, suffering a higher nest predation rate, laid smaller clutches than willow tits. The generally lower survival rate of willow tit adults may largely be attributed to their higher reproductive commitment (larger willow tit clutch size), because no significant interspecific survival difference remained between nonbreeding males. Finally, in willow tits we found a positive correlation between average clutch size and juvenile survival rate (density-dependent) the ensuing year, suggesting that willow tits may adjust clutch size in response to changing survival prospects for their young by using the breeding density as a cue.     

Nesting success of understory forest birds in central Panama

Greater nest predation in tropical than temperate birds has been hypothesized to be a primary selective force generating latitudinal differences in avian life history traits. Few extensive data sets, however, have been available from tropical forests to compare with data from temperate forests. To increase the amount of empirical information available for addressing issues related to the evolution of life history traits of tropical birds, we measured the nesting success of understory birds in lowland forest of central Panama. We found and monitored the fates of 696 nests of 71 species over two breeding seasons. Daily nest predation rates for the ten species for which we obtained the largest samples ranged from 1.6 to 8.3%, equivalent to a loss of 43 to 92% of nests. These values overlapped extensively the range of daily predation rates experienced by ecologically similar species in North America. Proportion of nests fledging young, estimated with the Mayfield method, was significantly lower in tropical (range: 8 to 57%) than temperate (27 to 60%) species. Nesting success in Panama varied among years, however, being greater in 1996 than 1997. In 1996, nesting success was similar to that of species breeding in forest fragments of midwestern North America. When compared with success of nests in large, contiguous forest tracts of North America, however, tropical avian nesting success was consistently lower by approximately 23%. We conclude that nesting success in central Panama may be poor in most breeding seasons, but also may be punctuated by occasional years of relatively exceptional success, a possibility heretofore unappreciated because of a general paucity of data from the tropics. Furthermore, our results indicate substantial variation in levels of nesting success among species, and almost no variation in clutch size. Such large interspecific variation, as well as potentially large annual variation, in nesting success does not support the hypothesis that uniformly low levels of nesting success select for small tropical clutch sizes.     

Do avian cooperative breeders live longer?

Cooperative breeding is not common in birds but intriguingly over-represented in several families, suggesting that predisposing factors, similar ecological constraints or a combination of the two facilitate the evolution of this breeding strategy. The life-history hypothesis proposes that cooperative breeding is facilitated by high annual survival, which increases the local population and leads to a shortage of breeding opportunities. Clutch size in cooperative breeders is also expected to be smaller. An earlier comparative analysis in a small sample of birds supported the hypothesis but this conclusion has been controversial. Here, I extend the analysis to a larger, worldwide sample and take into account potential confounding factors that may affect estimates of a slow pace of life and clutch size. In a sample of 81 species pairs consisting of closely related cooperative and non-cooperative breeders, I did not find an association between maximum longevity and cooperative breeding, controlling for diet, body mass and sampling effort. However, in a smaller sample of 37 pairs, adult annual survival was indeed higher in the cooperative breeders, controlling for body mass. There was no association between clutch size and cooperative breeding in a sample of 93 pairs. The results support the facilitating effect of high annual survival on the evolution of cooperative breeding in birds but the effect on clutch size remains elusive.     

Mass gained during breeding positively correlates with adult survival because both reflect life history adaptation to seasonal food availability

Both mass (as a measure of body reserves) during breeding and adult survival should reflect variation in food availability. Those species that are adapted to less seasonally variable foraging niches and so where competition dominates during breeding, will tend to have a higher mass increase via an interrupted foraging response, because their foraging demands increase and so become more unpredictable. They will then produce few offspring per breeding attempt, but trade this off with higher adult survival. In contrast, those species that occupy a more seasonal niche will not gain mass because foraging remains predictable, as resources become superabundant during breeding. They can also produce more offspring per breeding attempt, but with a trade-off with reduced adult survival. We tested whether the then predicted positive correlation between levels of mass gained during seasonal breeding and adult survival was present across 40 species of tropical bird measured over a 10-year period in a West African savannah. We showed that species with a greater seasonal mass increase had higher adult survival, controlling for annual mass variation (i.e. annual variation in absolute food availability) and variation in the timing of peak mass (i.e. annual predictability of food availability), clutch size, body size, migratory status and phylogeny. Our results support the hypothesis that the degree of seasonal mass variation in birds is probably an indication of life history adaptation: across tropical bird species it may therefore be possible to use mass gain during breeding as an index of adult survival.     

A meta‐analysis of global avian survival across species and latitude

Tropical birds are purported to be longer lived than their temperate counterparts, but it has not been shown whether avian survival rates covary with latitude worldwide. Here, we perform a global‐scale meta‐analysis of 949 estimates from 204 studies of avian survival and demonstrate that a latitudinal survival gradient exists in the northern hemisphere, is dampened or absent for southern hemisphere species, and that differences between passerines and nonpasserines largely drive these trends. We also show that while extrinsic factors related to climate were poor predictors of apparent survival compared to latitude alone, the relationship between apparent survival and latitude is strongly mediated by intrinsic traits – large‐bodied species and species with smaller clutch size had the highest apparent survival. Our findings reveal that differences among intrinsic traits and whether species were passerines or nonpasserines surpass latitude and its underlying climatic factors in explaining global patterns of apparent avian survival.     

Functional linkages for the pace of life, life-history, and environment in birds

For vertebrates, body mass underlies much of the variation in metabolism, but among animals of the same body mass, metabolism varies six-fold. Understanding how natural selection can influence variation in metabolism remains a central focus of Physiological Ecologists. Life-history theory postulates that many physiological traits, such as metabolism, may be understood in terms of key maturational and reproductive characteristics over an organism's life-span. Although it is widely acknowledged that physiological processes serve as a foundation for life-history trade-offs, the physiological mechanisms that underlie the diversification of life-histories remain elusive. Data show that tropical birds have a reduced basal metabolism (BMR), field metabolic rate, and peak metabolic rate compared with temperate counterparts, results consistent with the idea that a low mortality, and therefore increased longevity, and low productivity is associated with low mass-specific metabolic rate. Mass-adjusted BMR of tropical and temperate birds was associated with survival rate, in accordance with the view that animals with a slow pace of life tend to have increased life spans. To understand the mechanisms responsible for a reduced rate of metabolism in tropical birds compared with temperate species, we summarized an unpublished study, based on data from the literature, on organ masses for both groups. Tropical birds had smaller hearts, kidneys, livers, and pectoral muscles than did temperate species of the same body size, but they had a relatively larger skeletal mass. Direct measurements of organ masses for tropical and temperate birds showed that the heart, kidneys, and lungs were significantly smaller in tropical birds, although sample sizes were small. Also from an ongoing study, we summarized results to date on connections between whole-organism metabolism in tropical and temperate birds and attributes of their dermal fibroblasts grown in cell culture. Cells derived from tropical birds had a slower rate of growth, consistent with the hypothesis that these cells have a slower metabolism. We found that dermal fibroblasts from tropical birds resisted chemical agents that induce oxidative and non-oxidative stress better than do cells from temperate species, consistent with the hypothesis that birds that live longer invest more in self-maintenance such as antioxidant properties of cells.     

Different responses to cold weather in two pied flycatcher populations

We compared how breeding parameters differ according to prevailing weather conditions between a marginal, subarctic (69°N) and temperate (61°N) population of the pied flycatcher Ficedula hypoleuca, a small migratory insectivorous passerine. We predicted that the effects of weather on breeding performance (clutch size, hatching success, nestling growth, fledging success) would be greater at northern latitudes, where the weather conditions are more extreme and unpredictable. We found that the breeding parameters, except clutch size, were not, however, inferior in the north. Northern birds, unlike the southern ones, responded to colder conditions by laying smaller clutches and maintaining a larger energy reserve (indicated by higher female body mass and higher levels of subcutaneous fat). If a cold spell occurred during the nestling period, southern flycatchers had 5–10% lower fledging success than the northern ones. Our results indicate that in the north, the breeding individuals coped with cold and variable weather better than the individuals in the southern population. This could be adaptive, because at high latitudes there is a higher probability of cold weather at the time of breeding.     

Monthly survival of African Sylvia warblers in a seasonally arid tropical environment

Latitudinal gradients of life-history traits in animals are thought to be shaped by environmental variables. For example, it has been suggested that the increase in avian clutch size from the tropics towards the northern temperate regions is caused by a reduced survival of adult birds in the north due to increasing environmental seasonality. However, the tropical savannahs of East Africa show pronounced seasonality in resources caused by distinct rainy and dry seasons. This raises the question of whether survival and other life-history traits of birds living in these tropical savannahs are influenced by this seasonality, making them more similar to northern temperate species. We used 2-year monthly resighting data, a multistate modelling approach and the program MARK to test whether survival, transition probabilities between breeding states and other life-history traits of two resident Kenyan Sylvia species (Aves: Passeriformes: Sylviidae) are shaped by seasonality of rainfall in their environment. Contradicting our hypotheses, the two species showed only very slight influence of seasonality of rainfall on their survival. Survival in the dry months was hardly lower than in the rainy months. The species in the more seasonal environment ( S. boehmi , annual survival 71%) survived as well as the one in the more constant environment ( S. lugens , 56%). The observed survival rates correspond well to other life-history traits of the two species and are of similar magnitude to survival rates of other tropical passerines. This implies that either seasonality is not the driving force behind the life-history traits of the two species or the birds do not experience their environment as seasonal, as might be suggested by fluctuations in rainfall.     

A comparison of life‐history and parental care in temperate and tropical wrens

Comparing closely related species that live in different environments is a powerful way to understand selective pressures that influence life‐history evolution. We examined a suite of life‐history traits and parental care in neotropical buff‐breasted wrens Cantorchilus leucotis and north‐temperate Carolina wrens Thryothorus ludovicianus (Family Troglodytidae), to test hypotheses about life‐history evolution. As expected, buff‐breasted wrens exhibited smaller clutch sizes and higher annual adult survival than Carolina wrens. We found minimal support for the nest predation hypothesis, as nest survival and age‐corrected provisioning rates to whole broods were similar between species, and number of breeding attempts and breeding season length were greater in temperate wrens. Critical predictions of the food limitation hypothesis were not supported; in particular age‐corrected provisioning rates per nestling were higher in the tropical than temperate species. The adult survival and offspring quality hypothesis garnered the most support, as buff‐breasted wrens exhibited greater age‐corrected provisioning rates per nestling, a longer nestling period, longer re‐nesting intervals following nest success, and lower annual fecundity than Carolina wrens. Despite similarly prolonged breeding seasons, reproductive strategies differ between species with buff‐breasted wrens investing considerably in single broods to optimize first‐year survival and Carolina wrens investing in multiple small broods to optimize annual fecundity.     

Clutch size variation in passerine birds: The nest predation hypothesis

Summary The hypothesis that a negative relationship exists between clutch size and the probability that the nest will be robbed is tested, using data for passerine birds given in the literature. The data for four separate groups of species, viz. hole-nesters, semi hole-nesters and open-nesters nesting above and on the ground, respectively, were examined in relation to geographical gradients and seasonal and annual variation. In general, the data analysis results support the hypothesis, but cannot yet be considered as proven. More data on the riks of nest predation are needed. The most serious discrepancy is that for the Fieldfare, the protection from predation provided by nesting in colonies does not seem to be accompanied by a corresponding increase in clutch size. However, the clutch size of the Brambling, a species with seeks out such colonies for its breeding sites, does tend to increase in these hatitats.The pattern of clutch size variation was similar for the two groups of hole-nesting species, but differed significantly from that found for the two groups of open-nesters. The difference in the clutch size variation of the two groups of open-nesting species predicted from the hypothesis, viz. that, in northern regions, both latitudinal and altitudinal increases in clutch size should more commonly be found for those species which nest on the ground, compared to those nesting above ground level, was confirmed.A seasonal decrease in clutch size, in temperate regions, was found to be typical for species whose nests are subject to relatively little predation, particularly for those species which have a short breeding season. Advantages of laying small clutches, resulting in fewer nestlings than the number which would be possible for the parents to rear successfully, are discussed, and a simple model is presented which does not assume that nest predation is dependent on clutch size.     

Interspecific associations between circulating antioxidant levels and life-history variation in birds

Antioxidants play an important role in protecting tissues against aging-associated oxidative damage and are thus prime candidates for relating physiological mechanisms to variation in life histories. We measured total antioxidant capacity, antioxidant response to stress, and levels of uric acid, vitamin E, and four carotenoids in 95 avian species, mostly passerines from Michigan or Panama. We compared antioxidant measures to seven variables related to life histories (clutch size, survival rate, incubation period, nestling period, basal metabolic rate, body mass, and whether the species lived in a tropical or temperate climate). Life-history-related traits varied over at least three statistically independent axes. Higher antioxidant levels were generally characteristic of more rapid development, lower survival rate, smaller body size, larger clutch size, and higher mass-adjusted metabolic rate, but the relationships of particular antioxidants with individual life-history traits showed considerable complexity. Antioxidant-life history associations differed between tropical and temperate species and varied with respect to taxonomic sampling. Vitamin E showed few relationships with life-history traits. Overall, our results partly support the hypothesis that antioxidant levels evolve to mirror free radical production. Clearly, however, the complex patterns of physiological diversification observed here result from the interplay of many factors, likely including not just investment in somatic maintenance but also phylogenetic constraint, diet, and other aspects of ecology.