A flyway perspective on food resource abundance in a long-distance migrant, the Eurasian teal (Anas crecca)

Two frequent assumptions about the evolution of long-distance migration in birds are that they travel long distances annually to reach food-rich areas for breeding, and that they time their migratory journey to be at staging sites when the latter provide the best feeding conditions. These assumptions have rarely been properly tested, and there is no study in which a species’ major food types have been measured by standardized methods throughout a flyway and over a large part of the year. We here present such data for Eurasian teal (Anas crecca), converted to a common energetic currency, and collected at wintering, spring staging and breeding sites. Teal did not time migration to maximize local food abundance; most birds left wintering and spring staging sites before a sharp increase in invertebrate food abundance occurred. On the other hand, hatching of ducklings coincided with a peak in invertebrate food abundance on boreal breeding lakes. Mean overall food abundance (invertebrates and seeds combined) did not differ between wintering sites in southern France and breeding sites in northern Sweden at the time of breeding. Our results are inconsistent with the hypothesis that long-distance migration in dabbling ducks has evolved because adult birds gain an immediate pay-off in increased food abundance by flying north in spring. However, our data confirm a selective advantage for breeding at higher latitudes, because hatching of ducklings may coincide with a peak in invertebrate emergence and because longer days may increase the duration of efficient foraging.     

Capital and income breeding and the evolution of colony founding strategies in ants

Ability to store resources that will be used for reproduction represents a potential life history adaptation because storage permits feeding and reproduction to be decoupled spatially and/or temporally. The two ends of a continuum involve acquiring all resources prior to reproduction (capital breeding) or acquiring all resources during the reproductive period (income breeding). Traditional life history theory examines tradeoffs between costs and benefits of such strategies, but this theory has not been integrated into life history studies of ants, even though founding queens have the analogous strategies of fully claustral (capital breeding) and semi-claustral (income breeding). This study demonstrates that facultatively semi-claustral queens of the seed-harvester ant Pogonomyrmex desertorum exhibit phenotypic plasticity during colony founding because unfed queens produced few, small minims, whereas ad libitum fed queens produced larger, heavier minims and additional brood. Fed queens also lost less mass than unfed queens despite their producing more brood. Overall, foraging provides queens with a suite of benefits that likely offset potential negative effects of foraging risk. Life history studies across a diverse array of taxa show that capital breeding is consistently associated with low availability and/or unpredictability of food, i.e., environmental conditions that favor prepackaging of reproductive resources. Such a broad and consistent pattern suggests that similar factors favored the evolution of fully claustral (capital breeding) colony founding in ants. Overall, these data suggest that ant researchers should revise their conventional view that fully claustral colony founding evolved because it eliminated the need for queens to leave the nest to forage. Instead, colony founding strategies should be examined from the perspective of environmental variation, i.e., availability and predictability of food. I also provide a functional scenario that could explain the evolution of colony founding strategies in ants. Received 16 November 2005; revised 1 March 2006; accepted 29 March 2006.     

Spatial-concentration effects and the importance of local enhancement in the evolution of colonial breeding in seabirds

The information center hypothesis (ICH) suggests that birds breed in colonies because this behavior favors information exchange at the colony about the location of food patches. However, the complex suite of behaviors the ICH requires implies that information center following is more likely to evolve after colonial breeding has become established than to promote its initial development. A simpler hypothesis to explain the evolution of colonial breeding is that coloniality concentrates foragers in space, which leads to more rapid discovery of food patches and, by means of local enhancement, more efficient transfer of information about the location of patches than if foragers bred in a dispersed fashion. To assess the effects of breeding dispersion on foraging success, I simulated the foraging behavior of cliff-breeding seabirds (nesting either solitarily or colonially) searching for patchily distributed prey. Results show that colonial breeding is favored when food patches are sufficiently large or short-lived that competition for food is ameliorated. Conversely, dispersed nesting is favored when patches are small or long-lived. Individuals playing a colonial breeding strategy can invade a population of solitarily breeding birds, and once a colonial breeding strategy becomes established, it generally is resistant to invasion. These findings suggest that the spatial-concentration model is a plausible mechanism for the initial development of coloniality.     

The influence of snow conditions on the date of breeding of wading birds in north-east Greenland

The young of wading birds that breed in some parts of the arctic hatch when their insect food supply is most abundant. This appears to be the ultimate factor determining the date of egg-laying. In various other arctic birds the date of breeding may be influenced, directly or indirectly, by the time at which snow or ice melts. The Joint Biological Expedition to N.E. Greenland 1974 studied Ringed plovers Charadrius hiaticula , Dunlins Calidris aplina , Sanderlings Calidris alba , Knots Calidris canutus , and Turnstones Arenaria interpres breeding in various areas within 80 km of each other.
In 1974 the thaw was two or three weeks later than normal in the region. The time at which snow cleared and the dates of breeding of the waders differed considerably between valleys: breeding was delayed in valleys that cleared late. In all areas egg-laying ceased early in July, probably because chicks hatching from eggs laid later would not be ready to migrate by the end of the summer. Hence the mean date and length of the egg-laying period were determined by the date of snow clearance from the breeding areas.
Studies of the potential food resources indicate that the waders in N.E. Greenland do not appear to time their breeding so that the young hatch at some peak of food abundance. Sufficient food resources are probably available over a long period of time.     

Breeding success and lutein availability in great tit (Parus major)

The relationship among temporal variation in the availability of carotenoid-rich food, tissue carotenoid levels and breeding success are poorly known. We studied how diet quality and quantity affect the carotenoid profile and fledging success of great tit (Parus major) nestlings along a pollution gradient. We found declining seasonal trend in lutein concentration of caterpillars, which may be the explanation for the declining trend in nestlings' lutein concentration of plasma with season, despite the increase in caterpillar biomass. This may be because the biomass of most lutein-rich caterpillars (autumnal moths) decreased and less lutein-rich caterpillars (sawflies) increased during the breeding season. The temporal difference in occurrence of different caterpillar species means that peak lutein availability does not coincide with peak caterpillar abundance. However the positive association between total larval biomass and the number of great tit fledglings may suggest that fledging success depends more on total caterpillar availability than on lutein concentration of caterpillars.     

Sterols and steroids in a freshwater crustacean (Proasellus meridianus): hormonal response to nutritional input

For crustaceans that eat shredded plant material in freshwater habitats, the amount and the composition of food greatly vary over time because of the seasonal succession of plant fragments and algal biomass. The acquisition of elements necessary for growth, immune defense, and reproduction depends largely on this variation in food type and availability. In particular, sterols that are required as cellular membrane components and as precursors of ecdysteroids (molting hormones) must be acquired through food because crustaceans do not synthesize the steroid nucleus de novo. The present study examined the possible link between nutrition, sterols, and ecdysteroids in an isopod, Proasellus meridianus. In a first step, quantitative and qualitative analyses of sterols of P. meridianus were performed by gas‐chromatography/mass spectrometry. The results suggested that members of P. meridianus are able to convert dietary plant sterols into cholesterol required for growth and reproduction. In a second step, by manipulating food availability and using an enzyme immuno‐assay, we showed that ecdysteroid content in males and females (ovigerous or not) of P. meridianus decreased significantly after a starvation period. A nutritional input following this starvation period triggered an increase in the ecdysteroid contents of these animals. The comparable ecdysteroid responses to food pulses in males and females suggested that a nutritional control on steroid hormones regulated growth or gametogenesis rather than egg maturation. Thus, it appears that P. meridianus possesses an efficient stop‐and‐go endocrine system that may have been selectively favored in response to seasonal pulses of food.     

Nightjars may adjust breeding phenology to compensate for mismatches between moths and moonlight

Phenology match–mismatch usually refers to the extent of an organism's ability to match reproduction with peaks in food availability, but when mismatch occurs, it may indicate a response to another selective pressure. We assess the value of matching reproductive timing to multiple selective pressures for a migratory lunarphilic aerial insectivore bird, the whip‐poor‐will (Antrostomus vociferus). We hypothesize that a whip‐poor‐will's response to shifts in local phenology may be constrained by long annual migrations and a foraging mode that is dependent on both benign weather and the availability of moonlight. To test this, we monitored daily nest survival and overall reproductive success relative to food availability and moon phase in the northern part of whip‐poor‐will's breeding range. We found that moth abundance, and potentially temperature and moonlight, may all have a positive influence on daily chick survival rates and that the lowest chick survival rates for the period between hatching and fledging occurred when hatch was mismatched with both moths and moonlight. However, rather than breeding too late for peak moth abundance, the average first brood hatch date actually preceded the peak moth abundance and occurred during a period with slightly higher available moonlight than the period of peak food abundance. As a result, a low individual survival rate was partially compensated for by initiating more nesting attempts. This suggests that nightjars were able to adjust their breeding phenology in such a way that the costs of mismatch with food supply were at least partially balanced by a longer breeding season.     

Warming climate advances breeding and improves synchrony of food demand and food availability in a boreal passerine

Global climate change affects ecosystems via several trophic levels. We investigated changes in the timing of breeding in the willow tit (Poecile montanus) and timing of its caterpillar food resource in relation to warming springs in a boreal forest. We used generalized linear mixed effect models to study the importance of synchrony between the timing of breeding in willow tits and the caterpillar food availability on the breeding success, measured as nestling survival rate and mean nestling weight. Both the timing of breeding and the timing of the caterpillar peak advanced during the last decades, and were well explained by spring temperatures. Unlike in most passerine populations studied, synchrony has improved with rising spring temperatures. However, it had only a modest although statistically significant positive influence on breeding success. Spring temperatures do not seem to be used as cues for the timing of caterpillar food availability by willow tits. We conclude that responses to climatic warming seem to be population, species and habitat specific, necessitating research in a wide range of taxa in different climatic zones.     

The relevance of food peak architecture in trophic interactions

Phenological shifts and associated changes in the temporal match between trophic levels have been a major focus of the study of ecological consequences of climate change. Previously, the food peak has been thought to respond as an entity to warming temperatures. However, food peak architecture, that is, timings and abundances of prey species and the level of synchrony between them, determines the timing and shape of the food peak. We demonstrate this with a case example of three passerine prey species and their predator. We explored temporal trends in the timing, height, width, and peakedness of prey availabilities and explained their variation with food peak architecture and ambient temperatures of prebreeding and breeding seasons. We found a temporal match between the predator's breeding schedule and food availability. Temporal trends in the timing of the food peak or in the synchrony between the prey species were not found. However, the food peak has become wider and more peaked over time. With more peaked food availabilities, predator's breeding success will depend more on the temporal match between its breeding schedule and the food peak, ultimately affecting the timing of breeding in the predator population. The height and width of the food peak depended on the abundances and breeding season lengths of individual prey species and their reciprocal synchronies. Peakednesses of separate prey species' availability distributions alone explained the peakedness of the food peak. Timing and quantity of food production were associated with temperatures of various time periods with variable relevance in different prey species. Alternating abundances of early and late breeding prey species caused high annual fluctuation in the timing of the food peak. Interestingly, the food peak may become later even when prey species' schedules are advanced. Climate warming can thus produce unexpected changes in the food availabilities, intervening in trophic interactions.     

It’s All in the Timing: Birth Seasonality and Infant Survival in <Emphasis Type="BoldItalic">Eulemur rubriventer</Emphasis>

Highly seasonal breeding has been considered one of the keys to understanding Malagasy primate socioecology. Strict seasonal breeding may be particularly critical for Malagasy primates because they live in such energetically challenging seasonal environments. Lemurs also live in highly unpredictable environments, and there is growing evidence that reproductive timing may be mediated by additional factors, suggesting that more relaxed breeding seasonality is adaptive in some cases. I tested the adaptive breadth of the birth peak in Eulemur rubriventer, which breed in several different months. I describe reproduction in the species by determining the timing and extent of the birth season (period in which all births occur) and birth peak (period in which the majority of births occur); test whether relaxed reproductive seasonality might increase reproductive success by comparing infant mortality within and outside the birth peak; and model the extent to which fruit availability has an influence on the timing of reproduction. I collected birth data on 5 groups in 2003–2005, which I combined with demographic data that D. Overdorff collected from 5 focal groups and additional censused groups between 1988 and 1996. Thirty births occurred in 8 different months. Births were significantly seasonal, with a unimodal birth peak in late August/September/October, and a mean birth date of October 11. Twenty-three births (76.7%) occurred within 54 d (14.79%) of the year. No births occurred May–July, indicating that conceptions did not occur from late December through late February, and cycling (estimated using gestation length) did not occur until ca. 101 d after the austral summer solstice (December 21). Of 22 infants followed regularly, 18 were born in the birth peak, of which 2 died (11%). All 4 infants born out of season died. Based on fruit availability, I calculated a Theoretical Overlap index (T), which indicated a 3-mo window with optimal food conditions for reproduction. This window corresponded to the timing and breadth of the birth peak in Eulemur rubriventer. These results indicate that a breeding season >3 mo within a given year is not adaptive in the species, likely due in large part to the availability of fruit during key reproductive stages, particularly before breeding.     

Breeding seasonality among long-tailed macaques,Macaca fascicularis,in Peninsular Malaysia

The females among more than 7000 long-tailed macaques, Macaca fascicularis,that had recently been trapped for biomedical research were examined for pregnancy. The animals were taken from the wild in the north of Peninsular Malaysia during the period 1978 to 1979 and in the south from 1978 to 1982. Both samples showed some pregnancies in all months of the year and a peak during the months of May to July. Additional peaks in other months in some years showed that, while the midyear pregnancy peak was significant, it was not as marked a phenomenon as is the breeding peak in many other species. This might be due to the complexity and unpredictability of both the climate and the macaques’ food supply in Peninsular Malaysia. The additional peaks also illustrated the pitfalls of generalizing about breeding seasonality on the basis of small samples.     

Assessment of territory quality and its effects on breeding success in a migrant passerine, the Wheatear Oenanthe oenanthe

This paper examines how a returning migrant assesses the quality of an area as a breeding territory before the period of peak food demand and how effective the assessment is in terms of breeding success. Male Wheatears Oenanthe oenanthe return from Africa to choose territories in the Breckland of eastern England about March, females arriving shortly after males. The food supply was predictable: prey densities during the breeding period (egg-laying to chick independence) were strongly correlated with prey densities at the same sites during the period of arrival and territory establishment. Prey densities were also related to vegetation structure, averaging highest on short turf. Male arrival date and territory size were not significantly related to prey density but were strongly related to vegetation structure, implying that birds used vegetation as an indirect clue to prey availability. Neither territory size nor nest spacing appeared to affect nest losses caused by predators. The major variations in number of young fledged (other than predation) were caused by the number of nestlings hatched and presence of a second brood. Both early arrival and an early first brood improved first-brood success and were necessary for a second brood. Not all birds which arrived early bred early enough for a second brood. First-brood hatching date was strongly negatively correlated with pre-breeding prey availability but not significantly related to vegetation structure. Thus by using vegetation as a clue to habitat quality, some pairs suffered reduced breeding success. This result implies that birds may not be able adequately to assess prey density directly at the time of territory establishment. The critical period for food availability may not be the period of peak demand (nestling period), when food is relatively abundant, but is probably the pre-breeding period when females must accumulate reserves for eggs and when the food supply is poor. Food supply during this period may determine the timing of breeding and the ability to rear a second brood, and may thus have a greater effect on breeding success.     

Why do territorial male Tengmalm's owls fail to obtain a mate?

Non-breeding may occur because non-breeders are immature or somehow physiologically incapable of breeding, or because of a lack of resources (e.g. food resources, mating partners) needed to breed. There is, however, a lack of experimental evidence on whether bachelor males possessing territories and nest-sites are able to breed when supplemented with extra food or provided with mating partners. In vole-eating Tengmalm's owl, Aegolius funereus, we provided supplementary food and transferred females in nest-boxes of non-breeding males. Bachelor males that we supplemented with food did not attract mates at a higher frequency than unfed control males, which suggests that a lack of food did not influence the ability to attract a mating partner. In contrast, bachelor males presented with a female seemed to breed more frequently than bachelor males in the control group without mate addition. This suggests that scarcity of females may be an important reason for the high proportion of non-breeding males in the population (c. 25%) and excludes the possibility that non-breeding males are physiologically unable to breed. The operational sex ratio of the owl population at the time of mating may be male-biased, and some males may thus remain unpaired. Habitat and nest-box quality also seemed to be lower among bachelors than among breeding males. Received: 22 September 1997 / Accepted: 29 December 1997     

Food availability in beach and marsh habitats and the size of the fiddler crab claw,a sexually selected weapon and signal

Males of the sand fiddler crab Uca pugilator possess a greatly enlarged claw that is used as a weapon in ritualized contests for control of breeding burrows and is waved to attract females to breeding burrows. Approximately 5400 crabs were collected along the Atlantic coast of North America at 14 localities, all of which had both beach and salt marsh habitats. Five measurements were made on each claw. Principal components analysis was used to generate a single measure of claw size from the seven correlated measures and scores of the claw. Carapace width was measured to index body size. Claw size was greater in beach than marsh habitats, controlling for body size. However, body size did not differ by habitat type. Claw size was also greater in laboratory‐reared males receiving more food, suggesting that differential access to food could influence habitat‐associated differences in claw size. Chlorophyll a concentration and total organic content, reflecting, respectively, the abundance of benthic algae and other food, were greater in beach than marsh habitats. Moreover, feeding opportunities were greater in the wetter beach habitat because crabs there, but not in marsh habitat, can feed at breeding burrows. Adult fiddler crabs continue to molt and grow in both body and claw size as they age. Energetic investment in the claw relative to the body is plastic. It appears that the availability of food can affect the amount of energy invested in the claw.     

Range sizes and habitat use of non‐breeding Crested Caracaras in Florida

In Florida, habitats that include the breeding territories of Crested Caracaras (Caracara cheriway) are protected, but non‐breeding individuals may be vulnerable because they may occupy different areas and habitats. We captured and radio‐tagged 58 non‐breeding caracaras in Florida from July 2006–March 2009, determined their locations during weekly flights, and used GIS and compositional analysis to evaluate range sizes and habitat use. Non‐breeding caracaras (N = 58) ranged five times more widely during breeding seasons (N = 573 locations) than during non‐breeding seasons (N = 592 locations), and ranged >250 times more widely than breeding caracaras that defend territories year‐round. The large ranges of non‐breeders suggest they may be searching for and evaluating prospective territories or breeding opportunities (territory prospecting). Pasture occupied by cattle was the most used habitat relative to availability and was used more than pasture without cattle, likely because insects associated with cattle are an important food source for caracaras. Cattle numbers in Florida are declining and, because both breeding and non‐breeding caracaras primarily occupy pasture, this may present difficulties for long‐term management. Citrus groves were also used more than expected given availability by non‐breeding caracaras, but are rarely included in nesting territories. Because pasture and citrus were often adjacent, we suggest that citrus groves may function as refugia from socially dominant breeding caracaras. Conservation and recovery efforts for Florida's caracara population are needed throughout the range of non‐breeders, and should include management that ensures availability of habitat matrices of cattle pasture and citrus groves.     

BREEDING ECOLOGY OF TURNSTONES ARENARIA INTERPRES AT HAZEN CAMP, ELLESMERE ISLAND, N.W.T.

Turnstones arrive on Ellesmere Island in late May or early June. Pair-formation takes place during migration, or after arrival in courtship groups along the beaches, or on the nest territory, depending on weather conditions. Pair-formation was not observed at Hazen Camp in 1966 as most birds were already paired when observations began on 3 June. The preferred nesting habitat was Dryas-hummocked tundra closely associated with a marsh, stream, or pond. A census area of 240 ha supported 13 breeding pairs, possibly 14; the total number of pairs breeding in the Hazen Camp study area was estimated to be 70 (3.04 pairs/km²). Egg-laying began on 10 June, with 46% of the first eggs laid 13–15 June. 62% of the sets were completed between 19 and 21 June. Both sexes incubated, the female regularly and the male sporadically. Hatching was also well synchronised; most clutches hatched between 7 and 14 July. Nest success was high (79%). After hatching territories dissolved and family groups moved freely over the tundra, concentrating at ponds where food was readily available for the young. Both adults attended the young during the pre-fledging period, but the females apparently departed long before the young fledged. Males left once the young could fly and the adult fall migration was complete by early August with the exception of late breeders. Most of the young departed in the second half of August. Fall migration is complete by late August or early September. The breeding season appears to be timed so that the young are raised when food is most abundant. Food supply (dipterous insects, especially adult chironomids) was highest from 8 to 12 July at the peak of the hatching period. While food supply for the young declined during the growing period, the early departure of half the adult population, and family movements over the tundra, appeared to reduce the food demand correspondingly. Food appeared to influence the distribution of breeding pairs markedly, restricting them to the vicinity of marshes, streams, and ponds. Territoriality displayed by Turnstones is believed to be closely associated with the protection of the nest against predators and adult food requirements during incubation; it also seems likely that territory has at least a local effect in regulating the number of breeding pairs.     

SHORT NOTES

Earlé, R. A. 1981. Factors governing avian breeding in Acacia savanna, Pietermaritzburg, Part 3: Breeding success, recruitment and clutch size. Ostrich 52:235-243.

The clutch size and breeding success of eight species of birds were monitored over a two year period in Acacia savanna. The mean clutch size fluctuated within the breeding season and four patterns of clutch size variation were noted. A smaller proportion of large clutches were laid in the 1979 season when less food was available. Breeding success was higher in the dry 1979 season but fewer breeding attempts were made and the overall recruitment was thus lower. At the peak of the breeding season, in mid-November, breeding success was at its lowest. Predation on nests was the greatest factor reducing breeding success although rain and strong winds affected some species.     

OBSERVATIONS ON THE BREEDING OF THE PALILA PSITT1ROSTRA BA1LLEUI OF HAWAII

The behavioural ecology and breeding biology of the endangered Palila Psittirostra bailleui was studied from 1971 to 1975. The most intensive breeding occurred from June to August, and coincided with peak production of mamane Sophora chrysophylla seeds, the bir?s major food source. The Palila was able to make adjustments in its breeding to compensate for yearly differentiation in the timing and abundance of this food supply. Sexual chasing and courtship feeding were the most frequently encountered pre-nesting behaviours. Territory was a mate-defended area, which later in the nesting sequence was confined to the nest site. A total of 26 nests was found; most were placed on larger branches of mamane trees. Nest construction occurred primarily in the morning hours and lasted up to 20 days. Both sexes took part in nest construction, albeit the male role was minimal. Unless the nest was placed in the terminal fork of a tree, it usually contained a large stick base. The modal clutch size was two; eggs were laid early in the morning and in all cases one per day. Incubation sometimes began with the first egg and lasted 15–16 days. Only the female incubated, and she covered the eggs for about 75% of the daylight hours and throughout the night. Egg hatching was asynchronous, with the first young emerging early in the morning and the second not until later that same day. Only the female brooded, and the rate declined until day 15 when essentially it stopped. Both parents fed the young by regurgitation, and the number of feedings per hour decreased slightly over the nestling period. It is thought that insects and finely masticated plant material formed the bulk of the nestling diet until about day 5 when mamane seeds became important. Helpers were found at one nest. Young developed slowly and did not leave the nest until 21–27 days old. It is believed that these prolonged nestling periods were able to evolve because of the (former) absence of ground predators. After fledging, young remained with their parents for at least 30 days. Productivity was regulated by small clutch size, low population numbers and by the length of an individual nesting sequence (in that a pair could potentially raise only one brood each year). The primary reason for the endangered status of this bird appears to be the effect of habitat alteration upon a specialist, coupled with the fact that the small effective breeding population and low dis-persability of the species may have resulted in decreased genetic fitness.     

Reproductive biology of the rock shrimp Sicyonia dorsalis (Decapoda: Penaeoidea) from the southeastern coast of Brazil

Summary

Sicyonia dorsalis is not commercially exploited in Brazil, but it forms a critical link in marine food chains by converting detritus food sources into microorganism biomass that is available for higher trophic levels. We examined the reproductive biology of this species, sampled for five and a half years in a tropical locality, the southeastern coast of São Paulo (23° S), Brazil. Monthly samples were taken from January 1998 to June 2003 at depths of 5 to 45 m. Degree of ovarian development was used to examine breeding in adult females. Recruitment was estimated by changes in proportion of individuals in juvenile size classes of carapace length (CL #8.2 mm) in relationship to the total number of individuals. The largest immature female measured 9.2 mm CL, but 75% of females were smaller than 7.3 mm CL. The largest immature male measured 6.8 mm CL. The relative frequency of reproductive females was negatively correlated with the relative frequency of recruitment (p <0.001). Reproductive females were found in every season, with peaks in December-February following the recruitment peak in March-June, and a second peak of spawning in May-July and of recruitment in September-November. The continuous reproductive pattern observed for S. dorsalis, combined with the hypothesis that during spring and summer the phytoplankton production is higher, when the South Atlantic central water mass intrudes into the region during upwelling events, suggests that food availability for protozoeal and mysis larvae may be an important selective factor shaping the seasonal breeding pattern for S. dorsalis. The classical paradigm of continuous reproduction at lower latitudes, with increased seasonality of the breeding period at higher latitudes, seems to be valid for this species.     

Weak breeding seasonality of a songbird in a seasonally arid tropical environment arises from individual flexibility and strongly seasonal moult

In some tropical birds, breeding seasonality is weak at the population level, even where there are predictable seasonal peaks in environmental conditions. It therefore remains unclear whether individuals are adapted to breeding at specific times of the year or flexible to variable environmental conditions. We tested whether the relative year‐round breeding activity of the Common Bulbul Pycnonotus barbatus arises due to within‐individual variability in breeding dates. We collected data from 827 birds via mist‐netting over 2 years with corresponding local weather data. We used a combination of climate envelope and generalized linear mixed models to explore how the timing of breeding is influenced by time of year, individual variation, rainfall and temperature in a West African savannah where seasonal precipitation determines annual variation in environmental conditions. We also pooled 65 breeding records from 19 individuals recorded between 2006 and 2017 based on brood patch occurrence and behavioural observation to compare within‐individual and population variability in breeding dates. We show that the breeding dates of individuals may be as variable as for the population as a whole. However, we observed a seasonal peak in juvenile occurrence that varies significantly between years. Models suggest no relationship between nesting and moult, and within‐year variation in rainfall and temperature, and birds were unlikely to breed during moult but may do so afterwards. Moult was very seasonal, correlating strongly with day length. We suggest that because environmental conditions permit year‐round breeding, and because reproductive output is subject to high predation risk, there is probably a weak selection for individuals to match breeding with variable peak conditions in the environment. Instead, moult, which always occurs annually and successfully, is probably under strong selection to match variable peak conditions in the environment so that long‐term survival ensures future reproduction.