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.     

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.     

Differences between temperate and tropical birds in seasonal acclimatization of thermoregulatory traits

Phenotypic flexibility can be an important determinant of fitness in variable environments. The climatic variability hypothesis (CVH) predicts that phenotypic flexibility in thermoregulatory traits will be greater in temperate species than tropical species as a means of coping with increased temperature seasonality at higher latitudes. However, support for the CVH has been mixed, and recent studies suggest that tropical birds are capable of substantial phenotypic flexibility. To test the generality of the CVH, we used flow‐through respirometry to quantify seasonal acclimatization in thermoregulatory traits in suites of temperate (n = 6) and tropical (n = 41) birds. We used W/S ratios (winter/summer trait values) to quantify the direction and magnitude of seasonal change (W/S ratio of 1 means no seasonal change). Temperate species exhibited coordinated changes in thermoregulatory traits in winter, including large increases in thermoneutral zone (TNZ) breadth and reductions in heat loss below the lower limit of the TNZ. Conversely, tropical species exhibited idiosyncratic seasonal thermoregulatory responses, and mean W/S ratios were close to 1 for all traits, indicative of little seasonal change and consistent with predictions of the CVH. Nevertheless, mean W/S ratios did not differ significantly between temperate and tropical species for either M_b or BMR, demonstrating that tropical birds can also exhibit substantial thermoregulatory flexibility. Our results highlight the need for complementary acclimation experiments to determine if latitudinal differences in seasonal acclimatization are due to inherent differences in capacity for flexibility.     

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.     

Stress responses in tropical sparrows: comparing tropical and temperate Zonotrichia

Seasonal modulation of the adrenocortical response appears to be ubiquitous in mid- to high-latitude vertebrates but has not been investigated in tropical vertebrates. Previous studies demonstrate that temperate passerines show seasonality in corticosterone secretion and corticosteroid binding globulin capacities. We examined seasonal and sex differences in the stress response in an equatorial population of Zonotrichia capensis, the only Zonotrichia that breeds in the tropics, and compared the results with those of northern Zonotrichia. Seasonal differences in tropical Zonotrichia would presumably be independent of photoperiod and thus directly related to such activities as reproduction and feather molt. In addition, we investigated the possible role of binding globulin as a sex steroid binding globulin, as suggested for temperate passerines. Similar to northern congeners, Z. capensis show seasonal modulation in total corticosterone and binding globulin capacity with higher levels during breeding than molt. However, unlike many temperate passerines, there are no sex differences in corticosterone secretion or binding globulin capacity. Furthermore, the seasonal differences in total corticosterone diminish when the free levels are calculated. The contrast between equatorial and northern congeners indicates factors such as breeding environment and life-history strategy may play important roles in shaping stress response in these species.     

Cellular Metabolic Rate Is Influenced by Life-History Traits in Tropical and Temperate Birds

In general, tropical birds have a “slow pace of life,” lower rates of whole-animal metabolism and higher survival rates, than temperate species. A fundamental challenge facing physiological ecologists is the understanding of how variation in life-history at the whole-organism level might be linked to cellular function. Because tropical birds have lower rates of whole-animal metabolism, we hypothesized that cells from tropical species would also have lower rates of cellular metabolism than cells from temperate species of similar body size and common phylogenetic history. We cultured primary dermal fibroblasts from 17 tropical and 17 temperate phylogenetically-paired species of birds in a common nutritive and thermal environment and then examined basal, uncoupled, and non-mitochondrial cellular O₂ consumption (OCR), proton leak, and anaerobic glycolysis (extracellular acidification rates [ECAR]), using an XF24 Seahorse Analyzer. We found that multiple measures of metabolism in cells from tropical birds were significantly lower than their temperate counterparts. Basal and uncoupled cellular metabolism were 29% and 35% lower in cells from tropical birds, respectively, a decrease closely aligned with differences in whole-animal metabolism between tropical and temperate birds. Proton leak was significantly lower in cells from tropical birds compared with cells from temperate birds. Our results offer compelling evidence that whole-animal metabolism is linked to cellular respiration as a function of an animal’s life-history evolution. These findings are consistent with the idea that natural selection has uniquely fashioned cells of long-lived tropical bird species to have lower rates of metabolism than cells from shorter-lived temperate species.     

Seasonality of peak metabolic rate in non‐migrant tropical birds

Birds that are year‐round residents of temperate and tropical regions have divergent life histories. Tropical birds have a slower ‘pace of life’, one characteristic of which includes lower peak metabolic rate and daily activity levels. Temperate resident birds are faced with seasonal variation in thermogenic demand. This challenge is met with seasonally increased peak metabolic rate during winter. These thermogenic demands are much lower in birds that are year‐round tropical residents. By measuring peak (summit) metabolic rate in tropical and temperate resident bird species during summer and winter, we asked whether tropical birds exhibit seasonality in peak metabolic rate, and if the direction of seasonality differs between tropical and temperate species. We measured summit metabolism in seven tropical and one temperate species during the winter and during the summer breeding season to test the hypothesis that summit metabolism of tropical residents would change seasonally. We consider whether metabolic seasonality is associated with breeding season for tropical species. We found that summit metabolism was significantly greater during the summer for most tropical residents, while the temperate resident matched several previous reports with higher summit metabolism in winter. We conclude that metabolic seasonality occurs in tropical residents and differs from temperate residents, suggesting that breeding during the summer may be driving relatively higher metabolism as compared to winter thermogenesis in temperate birds.     

Patterns of seasonal and yearly mass variation in West African tropical savannah birds

Birds in the northern hemisphere usually increase mass reserves in response to seasonal low temperatures and shorter day length that increase foraging unpredictability and so starvation risk. In the lowland tropics, relatively low temperatures and short day lengths are absent and so the risk of starvation may be reduced, leading to much smaller seasonal effects on mass. Nevertheless, other factors such as high temperatures and water and food availability may vary greatly between tropical wet and dry seasons, leading to variable starvation risk and seasonal mass effects. Using data collected from 47 species of birds caught over a 10‐year period in a tropical savannah region in West Africa we tested for seasonal variation in mass in response to a predictable, strongly seasonal tropical climate. Many species (91%) showed seasonal variation in mass, and this was often in a clear annual pattern that was constant across the years. Many species (89%) varied their mass in response to seasonally predictable rainfall. Annual variation in mass was also important (45% of species). Relatively few species (13%) had a seasonal pattern of mass variation that varied between years. Feeding guild or migratory status was not found to affect seasonal or annual mass variation. Seasonal mass change was on average 8.1% across the 21 species with a very large sample size and was comparable with both northern and southern temperate species. Our study showed that biologically significant consistent seasonal mass variation is common in tropical savannah bird species, and this is most likely in response to changing resource availability brought about by seasonal rainfall and the interrupted foraging response due to the constraints of breeding.     

Differences in leaf phenology between juvenile and adult individuals of two tree species in a seasonally dry tropical woodland

In temperate forests, juvenile trees anticipate leaf phenology compared to adults, thus avoiding shading and herbivory. This is also expected to occur in seasonal tropical forests due to intense herbivory and shading during the rainy season; however, the anticipation of leaf phenology by juveniles in seasonal tropical forests has yet to be demonstrated. Stem‐succulent species are expected to be prone to juvenile phenological anticipation because these species are able to use water stored in their stems for leaf flushing in the dry season. We investigated this hypothesis by comparing leaf phenology (bud break, leaf expansion) of juveniles and adults of two species with contrasting wood densities in the transition between dry and rainy seasons in a tropical dry woodland. We also investigated the level of light limitation that juveniles experience in the rainy season. Both species exhibited bud break during the dry season, but only expanded their leaves with the occurrence of the first rains. In general, the stem‐succulent species had a more precocious bud break; however, anticipation by juveniles occurred only in the species with more dense wood. Canopy openness was lower than in temperate deciduous forests, but the fact that the full expansion of leaves occurred only with rainfall indicates that bud break in anticipation of canopy closure contributes only to keeping leaf photosynthetic balance from going negative, and not to higher carbon gain. The importance of anticipated budding for escaping herbivory remains an alternative explanation in need of investigation.     

SOME ASPECTS OF THE BEHAVIOUR OF THE WATTLED PLOVER AFRIBYX SENEGALLUS (LINNAEUS)

The question of how aridity might influence avian clutch size, through the influences of rainfall seasonality and environmental stochasticity (unpredictability), has received little attention. A marked east-west gradient in aridity across South Africa provides a unique opportunity to test for such influences. Using an extensive collection of nest records for 106 terrestrial bird species from the South African Nest Record Card Scheme, we tested three predictions related to rainfall seasonality and stochasticity. Analyses were conducted at two levels, the first examining each species independently, and the second grouping species into five dietary guilds. The first prediction, that clutch size should generally increase with higher rainfall seasonality (i.e. higher seasonal fluctuation of food availability), was supported, particularly in the most arid environments where food abundance is more closely linked to rainfall. Controlling for rainfall seasonality, the second prediction, that clutch size should generally decrease as a bet-hedging strategy in arid, stochastic environments, was also supported. Although the timing of the rainy season differs among regions in South Africa (winter, early summer, later summer, year-round), birds primarily nest during spring. The relative timing of rainfall and breeding is expected to have different consequences for seasonal variation in clutch size among rainfall regions. The third prediction, of different patterns of seasonal variation in clutch size between rainfall regions, was also supported. In the winter and early-summer rainfall regions, early-nesting birds (breeding with or soon after the rains) generally had a larger clutch size than late-nesting birds. In the late-summer rainfall region, early-nesting birds (breeding well before the rains) had a smaller clutch size than late-nesting birds.     

Timing of breeding in variable environments: tropical birds as model systems

Animals need to adjust reproductive decisions to environmental seasonality. In contrast to species from the well-studied temperate zones, little is known for tropical birds about the environmental cues that stimulate reproductive activity and the physiological mechanisms that regulate reproduction. I am investigating the environmental and endocrine mechanisms that underlie the timing of reproduction in spotted antbirds from the near-equatorial Panamanian rainforest and in small ground finches from the equatorial arid Galápagos islands. Spotted antbirds live in a fairly predictable seasonal environment and show regular changes in gonad sizes and some reproductive hormones. Despite the small annual variation in photoperiod close to the equator, these birds can measure slight photoperiodic increases and use it to initiate reproductive activity. Spotted antbirds also respond to seasonal changes in food availability, which allows them to flexibly adjust gonad growth to environmental conditions. Testosterone is involved in regulating song and aggressive behavior in these year-round territorial birds, although it can remain at low plasma levels throughout the year. In contrast, small ground finches exposed to a rather unpredictable climate on Galápagos appear to grow their gonads whenever heavy rains fall and have regressed gonads during other times of the year. The lack of a physiological preparation for the breeding season and their response to short-term cues related to rainfall indicate a striking flexibility in the regulation of breeding in small ground finches. I suggest that tropical birds can serve as model systems to study the physiological adaptations to different environments. Unraveling the neuroendocrine mechanisms behind the flexibility in reproductive timing will clarify whether differences found between temperate and tropical birds represent variations of the same basic mechanism or instead reflect a fundamental divergence in physiological control systems.     

Rainfall and labile carbon availability control litter nitrogen dynamics in a tropical dry forest

N cycling in tropical dry forests is driven by rainfall seasonality but the mechanisms involved are not well understood. We studied the seasonal variation in N dynamics and microbial biomass in the surface litter of a tropical dry forest ecosystem in Mexico over a 2-year period. Litter was collected at 4 different times of the year to determine changes in total, soluble, and microbial C and N concentrations. Additionally, litter from each sampling date was incubated under laboratory conditions to determine potential C mineralization rate, net N mineralization, net C and N microbial immobilization, and net nitrification. Litter C concentrations were highest in the early-dry season and lowest in the rainy season, while the seasonal changes in N concentrations varied between years. Litter P was higher in the rainy than in the early-dry season. Water-soluble organic C (WSOC) and water-soluble N concentrations were highest during the early- and late-dry seasons and represented up to 4.1 and 5.9% of the total C and N, respectively. NH₄⁺ and NO₃⁻ showed different seasonal and annual variations. They represented an average 23% of soluble N. Microbial C was generally higher in the dry than in the wet seasons, while microbial N was lowest in the late-dry and highest in the early-rainy seasons. Incubations showed that lowest potential C mineralization rates and C and N microbial immobilization occurred in rainy season litter, and were positively correlated to WSOC. Net nitrification was highest in rainy season litter. Our results showed that the seasonal pattern in N dynamics was influenced by rainfall seasonality and labile C availability, and not by microbial biomass. We propose a conceptual model to hypothesize how N dynamics in the litter layer of the Chamela tropical dry forest respond to the seasonal variation in rainfall.     

Similar survival of birds between wet and seasonally dry Neotropical environments

Studies in the tropics suggest a regional similarity in survival rates of adult birds; however, this literature often overlooks species in semi-arid tropical environments. Bird survival in seasonally dry environments (e.g. seasonally dry tropical forests (SDTFa)) may be lower than that in more constantly wet areas (e.g. tropical rainforests (TRFs)), especially if the birds are negatively affected by seasonal rainfall or food-limitation. However, survival could be similar across these tropical environments, as the asymmetry between young and adult mortality tends to be high in all tropical areas, and the higher risk of mortality in young animals may favour adult survival (residual reproductive investment) regardless of the local climatic conditions. To fill this knowledge gap, we tested the hypothesis that bird survival is similar between seasonally dry (SDTF) and constantly wet (TRF) Neotropical environments. We estimated the apparent survival of 27 South American bird populations from three SDTF areas and 39 populations from a TRF. Apparent survival was estimated from Cormack–Jolly–Seber (CJS) models fitted using a Bayesian structure and the resulting variation in survival rates between study areas and with body mass was explored using a Bayesian phylogenetic mixed model. Apparent annual survival of passerines did not differ between areas (geometrical mean of survival: SDTF = 0.50, 0.56, 0.64; TRF = 0.58), but body mass was positively associated with survival. The variation in bird survival was partially explained by phylogenetic relationships among species. Our results suggest that bird survival is regionally similar in Neotropical forests, despite the climatic variation. We discuss possible physiological and behavioural mechanisms adopted by birds in SDTFs to attenuate effects of environmental seasonality on survival.     

Season of birth affects juvenile survival of giraffe

Variation in timing of reproduction and subsequent juvenile survival often plays an important role in population dynamics of temperate and boreal ungulates. Tropical ungulates often give birth year round, but survival effects of birth season for tropical ungulate species are unknown. We used a population of giraffe in the Tarangire Ecosystem of northern Tanzania, East Africa to determine whether calf survival varied by season of birth. Variation in juvenile survival according to season of birth was significant, with calves born during the dry season experiencing the highest survival probability. Phenological match may confer a juvenile survival advantage to offspring born during the dry season from greater accumulated maternal energy reserves in mothers who conceive in the long rainy season, high-protein browse in the late dry-early short rainy seasons supplementing maternal and calf resources, reduced predation due to decreased stalking cover, or some combination of these. Asynchrony is believed to be the ancestral state of all ungulates, and this investigation has illustrated how seasonal variation in vegetation can affect juvenile survival and may play a role in the evolution of synchronous births.     

Phenology, Lignotubers, and Water Relations of Cochlospermum vitifolium, a Pioneer Tropical Dry Forest Tree in Costa Rica

We examined structural and physiological traits relevant to the phenology of the tropical dry forest (TDF) pioneer tree Cochlospermum vitifolium . Despite marked seasonality in rainfall, meristem activity occurred throughout the year. Leaves were produced almost continuously during the rainy season, while leaf shedding started early during drought, before changes in soil water content were observed. Phenological activity under drought included flowering and fruiting of leafless trees; bud break and shoot extension took place before the end of the dry season. Low wood density of C. vitifolium stems (0.17 g/cm³) and lignotubers (0.14 g/cm³) provided water and starch storage needed to support phenological events such as branch extension, leaf flushing, and reproduction during the dry season, and probably also contributed to survival following mechanical damage and fire, typical of early TDF successional stages. Lignotuber water and starch contents showed substantial seasonal variation, declining from the beginning of the dry season to their lowest levels at the time of reproduction and dry-season flushing. Stems progressively replaced lignotubers as main storage organs as tree size increased. Evidence for a role of water stores in buffering daily water deficits was weak. Leaf water potentials remained above −1.2 MPa and stomatal conductance below 350 mmol/m²/s, suggesting that gas exchange during the rainy season was limited to prevent xylem cavitation. Leaf shedding occurred when early-morning and mid-day Ψ_L converged at the rainy–dry season transition, without changes in lignotuber or soil water content, suggesting that leaves of C. vitifolium are closely tuned to atmospheric drought.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp .     

Seasonal Patterns of Flowering and Fruiting in a Dry Tropical Forest in Jamaica1

Tropical dry forests occupy more area and are more endangered than rainforests, yet their regeneration ecology has received less study and is consequently poorly understood. We recorded the flowering and fruiting phenology of a tropical dry forest in Jamaica over a period of 26 mo within ten 15 × 15‐m plots. Community‐wide recruitment reached a maximum in the wet season, whereas no recruitment occurred during the dry season. We observed a unimodal peak in rainfall and fruit production, and the periodicity and intensity of seed production were significantly correlated with rainfall seasonality (the optimal time for germination). Flowering at the community and system levels lagged behind a significant increase and subsequent decrease in rainfall by 7 and 3 mo, respectively, indicating that the dominant factor controlling flowering periodicity is the passage of the major (4‐mo long) rainy season and changes in soil moisture conditions. Fruiting lagged behind flowering by 2 mo and a significant increase in fruiting occurred 2 mo prior to a significant increase in rainfall. At the population level, a correspondence analysis identified a major dichotomy in the patterns of flowering and fruiting between species and indicated two broad species groups based on their time of peak fruiting and the number of times they were in fruit. These were either individuals which were usually in peak fruit 1–2 mo prior to the start of the major rainy season or those that were in fruit more or less continuously throughout the year with no peak fruiting time. This study supports the view that seasonal variation in rainfall and hence soil water availability constitutes both the proximate and the ultimate cause of flowering periodicity in tropical dry forests.     

A seasonal difference of daily energy expenditure in a free-living subterranean rodent,the silvery mole-rat (Heliophobius argenteocinereus; Bathyergidae)

In seasonal climatic regimes, animals have to deal with changing environmental conditions. It is reasonable to expect that seasonal changes are reflected in animal overall energetics. The relation between daily energy expenditure (DEE) and seasonally variable ecological determinants has been studied in many free-living small mammals; however with inconsistent results. Subterranean mammals, i.e. fossorial (burrowing) mammals which live and forage underground, live in a seasonally and diurnally thermally stable environment and represent a suitable model to test seasonality in DEE in respect to seasonal changes, particularly those in soil characteristics and access to food supply. Both factors are affected by seasonal rainfall and are supposed to fundamentally determine activity of belowground dwellers. These ecological constraints are pronounced in some tropical regions, where two distinct periods, dry and rainy seasons, regularly alternate. To explore how a tropical mammal responds to an abrupt environmental change, we determined DEE, resting metabolic rate (RMR) and sustained metabolic scope (SusMS) in a solitary subterranean rodent, the silvery mole-rat, at the end of dry season and the onset of rainy season. Whereas RMR did not differ between both periods, mole-rats had 1.4 times higher DEE and SusMS after the first heavy rains. These findings suggest that rainfall is an important environmental factor responsible for higher energy expenditure in mole-rats, probably due to increased burrowing activity. SusMS in the silvery mole-rat is comparable to values in other bathyergids and all bathyergid values rank among the lowest SusMS found in endothermic vertebrates.     

Fruiting phenology in a tropical dry evergreen forest on the Coromandel coast of India in relation to plant life-forms,physiognomic groups,dispersal modes,and climatic constraints

The fruiting phenology of 22 woody plant species belonging to 19 families was studied with respect to life-forms, physiognomic groups and dispersal modes, for 1 year at monthly intervals, in a tropical dry evergreen forest at Oorani (12°11′N, 79°57′E) on the Coromandel coast of India. At the community level, bimodal fruiting pattern prevailed, with a major peak in the dry season and a minor one in the early rainy season. An annual fruiting pattern was observed in many species and among the studied species fruiting lasted for 2–9 months. There was no significant difference in the frequency of species at three fruiting stages across the life-form categories and many species of upper and lower canopy trees and lianas were in the ripe fruiting phase during the late dry season. Plant physiognomic groups displayed distinct seasonality in fruiting pattern. The fruit maturation period was much longer for the wet season fruiting brevi-deciduous species than evergreen and deciduous species that fruited during the dry season. The variation in timing of fruiting behaviour among zoochorous species demonstrated less seasonality and zoochorous fruits were available throughout the year. Fruiting in anemochorous species peaked during the driest months and dryness favoured the dissemination of seeds. The fruiting patterns observed in the studied tropical dry evergreen forest across various plant traits were comparable with patterns recorded in other tropical seasonal forests.     

Relating tree growth to rainfall in Bolivian rain forests: a test for six species using tree ring analysis

Many tropical regions show one distinct dry season. Often, this seasonality induces cambial dormancy of trees, particularly if these belong to deciduous species. This will often lead to the formation of annual rings. The aim of this study was to determine whether tree species in the Bolivian Amazon region form annual rings and to study the influence of the total amount and seasonal distribution of rainfall on diameter growth. Ring widths were measured on stem discs of a total of 154 trees belonging to six rain forest species. By correlating ring width and monthly rainfall data we proved the annual character of the tree rings for four of our study species. For two other species the annual character was proved by counting rings on trees of known age and by radiocarbon dating. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall in certain periods of the year, indicating that rainfall plays a major role in tree growth. Three species showed a strong relationship with rainfall at the beginning of the rainy season, while one species is most sensitive to the rainfall at the end of the previous growing season. These results clearly demonstrate that tree ring analysis can be successfully applied in the tropics and that it is a promising method for various research disciplines.     

Climate anomalies affect annual survival rates of swifts wintering in sub‐Saharan Africa

Several species of migratory swifts breed in the Western Palearctic, but they differ in reproductive traits and nonbreeding areas explored in Africa. We examined survival and recapture probabilities of two species of swifts by capture–mark–recapture data collected in northern Italy (Pallid Swift Apus pallidus in Carmagnola, Turin, and Common Swift Apus apus in Guiglia, Modena) in the breeding season (May–July). Apparent survival rates were relatively high (>71%), comparable to other studies of European swifts, but showed marked annual variations. We used geolocators to establish the exact wintering areas of birds breeding in our study colonies. Common Swifts explored the Sahel zone during migration and spent the winter in SE Africa, while the Pallid Swifts remained in the Sahel zone for a longer time, shifting locations southeast down to Cameroun and Nigeria later in winter. These movements followed the seasonal rains from north to south (October to December). In both species, we found large yearly differences in survival probabilities related to different climatic indices. In the Pallid Swift, wintering in Western Africa, the Sahel rainfall index best explained survival, with driest seasons associated with reduced survival. In the Common Swift, wintering in SE Africa, the El Niño–Southern Oscillation (ENSO) cycle performed significantly better than Sahel rainfall or North Atlantic Oscillation (NAO). Extreme events and precipitation anomalies in Eastern Africa during La Niña events resulted in reduced survival probabilities in Common Swifts. Our study shows that the two species of swifts have similar average annual survival, but their survival varies between years and is strongly affected by different climatic drivers associated with their respective wintering areas. This finding could suggest important ecological diversification that should be taken into account when comparing survival and area use of similar species that migrate between temperate breeding areas and tropical wintering areas.