Testosterone and year‐round territoriality in tropical and non‐tropical songbirds

Past studies have suggested a fundamental difference in testosterone concentrations between tropical and northern latitude male birds, with the convention being that males in the tropics express much lower levels of testosterone. However, recent comparative studies have shown that tropical males with a short and synchronous breeding season (i.e. a breeding season typical of northern species) express maximum testosterone levels similar to those of northern latitude birds. Here, we ask the converse: do northern latitude songbirds that express a defining life‐history characteristic typical of the tropics, i.e. year‐round territoriality, have an annual testosterone profile similar to that of tropical songbirds? For the few year‐round territorial species for which data are available, we found that seasonal testosterone profiles and seasonal maxima in plasma testosterone were similar between males of tropical and non‐tropical species. For example, males of both groups expressed seasonal maxima during the period when females were fertile, and testosterone levels at this time were similar. In contrast, this and other studies show that species with seasonal territories typically express maximum testosterone levels earlier in the breeding cycle, when territories are first being established. Taken together, we suggest that specific life‐history traits may play a more important role in determining testosterone profiles of tropical and non‐tropical birds than breeding latitude and encourage further studies to allow for more formal comparisons.     

Short‐term changes in body condition in relation to habitat and rainfall abundance in American redstarts Setophaga ruticilla during the non‐breeding season

In migratory birds, environmental conditions during the stationary period of the non‐breeding season are crucial to consider because they ultimately affect the fitness of individuals by influencing their subsequent migration, breeding success and survival. Although a few studies have investigated the influence of non‐breeding habitat on the capacity of individuals to cope with long‐term seasonal rainfall fluctuations, it remains unknown how habitat quality and variations in rainfall abundance–at a monthly scale–interact to affect non‐breeding condition of migrating birds. In this study, we examined the influence of monthly changes in rainfall abundance on body condition of non‐breeding female redstarts Setophaga ruticilla living either in a high quality habitat (mangrove) or in a low quality habitat (scrub). Body condition of both mangrove and scrub redstarts showed important variations over the study period, demonstrating for the first time that body condition of non‐breeding female redstarts can change rapidly in response to short‐term fluctuations in rainfall. Importantly, we found that female redstarts living in mangrove were usually in better condition during periods of low rainfall compared to females living in scrub. However, body condition did not differ between mangrove females and scrub females during an episode of frequent, heavy precipitation. Importantly, our study also demonstrated that the duration of a perturbation is an important determinant of body condition in redstarts since a prolonged drought resulted in similar low body condition for birds from both habitats. Age was not correlated with body condition whatever the habitat and the rainfall conditions. Our results demonstrate that high quality habitat can temporarily reduce the deleterious effect of a short‐term drought on body condition, but also, that a habitat of low quality does not constrain individuals when climatic conditions are optimal.     

High rates of extra‐pair paternity in two equatorial populations of rufous‐collared sparrow,Zonotrichia capensis

The latitudinal increase in extra‐pair paternity (EPP) rates in birds suggests broad selective benefits to low EPP rates in the tropics. However, we have few EPP data from tropical birds, particularly from species with close relatives at high latitudes. Here, we report EPP rates in two resident equatorial populations of rufous‐collared sparrow Zonotrichia capensis, a genus well‐represented at high latitudes. We found 64% and 60% of broods contained extra‐pair offspring, and 42% and 52% of all young were extra‐pair. EPP rates were similar in these populations, despite clear differences in elevation, temperature, rainfall, and breeding season length. These findings provide evidence that EPP rates in tropical birds can be as high as those observed in temperate birds, and suggest that the selective pressures acting on EPP rates vary markedly across tropical birds.     

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.     

Diversity of insect‐induced galls along a temperature– rainfall gradient in the tropical savannah region of the Northern Territory,Australia

Evidence regarding the effect of temperature and rainfall on gall‐inducing insects is contradictory: some studies indicate that species richness of gall‐inducing insects increases as environments become hotter and drier, while others suggest that these factors have no effect. The role of plant species richness in determining species richness of gall‐inducing insects is also controversial. These apparent inconsistencies may prove to be due to the influence of soil fertility and the uneven distribution of gall‐inducing insect species among plant taxa. The current study tested hypotheses about determinants of gall‐inducing insect species richness in a way different to previous studies. The number of gall‐inducing insect species, and the proportion of species with completely enclosed galls (more likely to give protection against heat stress and desiccation), were measured in replicate plots at five locations along a 500‐km N‐S transect in the seasonal tropics of the Northern Territory, Australia. There is a strong temperature–rainfall gradient along this transect during the wet season. Plant species lists had already been compiled for each collection plot. All plots were at low elevation in eucalypt savannah growing on infertile soils. There was no evidence to suggest that hot, dry environments in Australia have more gall‐inducing insect species than cooler, wetter environments, or that degree of enclosure of galls is related to protecting insects from heat stress and desiccation. The variable number of gall‐inducing insect species on galled plant species meant that plant species richness did not influence gall species richness. Confirmation is still required that low soil fertility does not mask temperature–rainfall effects and that galls in the study region are occupied predominantly in the wet season, when the temperature–rainfall gradient is most marked.     

Effects of El Niño Southern Oscillation on avian breeding phenology

Aim

Climate oscillations are known to influence the reproductive phenology of birds. Here, we quantify the effects of cyclic climatic variation, specifically El Niño Southern Oscillation (ENSO), on birds that breed opportunistically. We aim to show how inter‐decadal climate fluctuations influence opportunistic breeding. This knowledge is essential for tracking the phenological responses of birds to climate change.

Location

Temperate and arid Australia.

Methods

We assessed variation in egg‐laying (start, peak, conclusion, length) during the three phases of ENSO (El Niño, La Niña and Neutral) for 64 temperate and 15 arid region species using ~80,000 observations. Linear mixed‐effect models and analysis of variance were used to (1) determine if, on average within each region, egg‐laying dates differed significantly among species between Neutral‐El Niño and Neutral‐La Niña phases, and (2) assess how La Niña and El Niño episodes influence egg‐laying in birds which breed early in the year.

Results

During La Niña phases, which are characterized by mild/wet conditions, most bird species in the temperate and arid regions exhibited longer egg‐laying periods relative to Neutral phases. However, there was substantial variation across species. This effect was strongly seasonal; species breeding in spring experienced the greatest increases in egg‐laying periods during La Niña. Further, we found only small differences in peak egg‐laying dates during Neutral and La Niña in the arid region; suggesting that hot temperatures may constrain breeding regardless of rainfall. The effects of El Niño on breeding phenology were not consistent in the temperate and arid regions and may be confounded by highly mobile species opportunistically moving and breeding with localized rainfall during dry periods.

Main conclusions

In both arid and temperate regions, increased rainfall associated with La Niña phases positively influences avian breeding, and likely recruitment. However, dry El Niño phases may not have the dramatic impacts on breeding phenology that are commonly assumed.

     

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.     

No downregulation of immune function during breeding in two year‐round breeding bird species in an equatorial East African environment

Some equatorial environments exhibit substantial within‐location variation in environmental conditions throughout the year and yet have year‐round breeding birds. This implies that breeding in such systems are potentially unrelated to the variable environmental conditions. By breeding not being influenced by environmental conditions, we become sure that any differences in immune function between breeding and non‐breeding birds do not result from environmental variation, therefore allowing for exclusion of the confounding effect of variation in environmental conditions. This create a unique opportunity to test if immune function is down‐regulated during reproduction compared to non‐breeding periods. We compared the immune function of sympatric male and female chick‐feeding and non‐breeding red‐capped Calandrella cinerea and rufous‐naped larks Mirafra africana in equatorial East Africa. These closely‐related species occupy different niches and have different breeding strategies in the same grassland habitat. Red‐capped larks prefer areas with short grass or almost bare ground, and breed during low rainfall periods. Rufous‐naped larks prefer areas of tall grass and scattered shrubs and breed during high rainfall. We measured the following immune indices: nitric oxide, haptoglobin, agglutination and lysis, and measured total monthly rain, monthly average minimum (T_min) and maximum (T_max) temperatures. Contrary to our predictions, we found no down‐regulation of immune function during breeding; breeding birds had higher nitric oxide than non‐breeding ones in both species, while the other three immune indices did not differ between breeding phases. Red‐capped larks had higher nitric oxide concentrations than Rufous‐naped larks, which in turn had higher haptoglobin levels than red‐capped larks. T_max was higher during breeding than during non‐breeding for red‐capped larks only, suggesting potential confounding effect of T_max on the comparison of immune function between breeding and non‐breeding birds for this species. Overall, we conclude that in the two year‐round breeding equatorial larks, immune function is not down‐regulated during breeding.     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

Short‐distance partial migration of Neotropical birds: a community‐level test of the foraging limitation hypothesis

Partial migration of tropical birds was long believed to be driven by variation in food abundance. Recent evidence from a partially‐migratory species suggests that in contrast, limited foraging opportunities at high elevations during severe wet season storms drives the most metabolically‐challenged individuals down to elevations where rainfall is lighter. Here, I test community‐level predictions of this hypothesis by examining the relationship between high‐elevation rainfall in the second half of the year and counts of migrant birds in lowland forest during late December each year from 1990–2009. I contrast results derived from analysis of all migrant species with both analyses of only the frugivorous migrants, and analyses of resident species. Counts of migrant species were on average positively associated with montane rainfall with differences of up to 72% in the numbers of birds counted in drier or wetter years. Frugivores and smaller birds responded more strongly to variation in rainfall compared to the broader migrant species pool. Interestingly, counts of resident species were also higher following wetter montane wet seasons. Results of analyses exploring the cause of resident responses were not consistent with climatic effects on breeding productivity or short‐term weather effects on detectability. Results were, however, consistent with cryptic down‐slope migration of individuals breeding at higher elevations augmenting lowland resident populations in wet years. These results suggest that changes in rainfall amount, storm intensity, and timing of severe weather events would lead to large increases in or losses of an important behaviour.     

The role of El Niño–Southern Oscillation in the dynamics of a savanna large herbivore population

Our understanding of large‐scale climatic phenomena and dynamics of large herbivore populations comes principally from research in northern regions with temperate, seasonal climate and animal communities with relatively low species diversity. To assess the generality of that perspective, we investigated effects of El Niño–Southern Oscillation (ENSO) on population dynamics of African buffalo Syncerus caffer inhabiting a semi‐arid savanna with variable rainfall. We used linear and nonlinear‐threshold models to investigate relationships between population parameters and explanatory variables affecting forage conditions (seasonal rainfall, Southern Oscillation Index [SOI]). El Niño‐related droughts in 1982–1983 and 1991–1992 were associated with strongly negative population change, a pattern expected to coincide with a decrease in normally high and constant adult survival. Consistent with that nonlinear pattern, we detected threshold relationships between wet‐season rainfall and population change. Juvenile recruitment was described best by linear relationships with dry‐season. Because ENSO operates primarily through wet‐season rainfall, whereas population dynamics were also related to dry‐season rainfall, SOI did not have the predictive ability of individual weather components.     

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.     

Nest survival in year‐round breeding tropical red‐capped larks Calandrella cinerea increases with higher nest abundance but decreases with higher invertebrate availability and rainfall

Nest survival is critical to breeding in birds and plays an important role in life‐history evolution and population dynamics. Studies evaluating the proximate factors involved in explaining nest survival and the resulting temporal patterns are biased in favor of temperate regions. Yet, such studies are especially pertinent to the tropics, where nest predation rates are typically high and environmental conditions often allow for year‐round breeding. To tease apart the effects of calendar month and year, population‐level breeding activity and environmental conditions, we studied nest survival over a 64‐month period in equatorial, year‐round breeding red‐capped larks Calandrella cinerea in Kenya. We show that daily nest survival rates varied with time, but not in a predictable seasonal fashion among months or consistently among years. We found negative influences of flying invertebrate biomass and rain on nest survival and higher survival of nests when nests were more abundant, which suggests that nest predation resulted from incidental predation. Although an increase in nest predation is often attributed to an increase in nest predators, we suggest that in our study, it may be caused by altered predator activity resulting from increased activity of the primary prey, invertebrates, rather than activity of the red‐capped larks. Our results emphasize the need to conduct more studies in Afro‐tropical regions because proximate mechanisms explaining nest predation can be different in the unpredictable and highly variable environments of the tropics compared with the relatively predictable seasonal changes found in temperate regions. Such studies will aid in better understanding of the environmental influences on life‐history variation and population dynamics in birds.     

Seasonal changes in mixed‐species bird flocks and antipredator information

Animals acquire information produced by other species to reduce uncertainty and avoid predators. Mixed‐species flocks (MSFs) of birds are ubiquitous in forest ecosystems and structured, in part, around interspecific information transfer, with “nuclear” species providing information that other species eavesdrop on. We hypothesized that in a seasonal tropical forest, the amount of information produced by birds about predation would be dynamic and particularly would decrease inside MSFs when the nuclear species leave MSFs to breed. We obtained baseline information on MSF encounter rate and species composition along established sampling routes over 9 months near the Sino‐Vietnamese border. We also conducted three experiments to quantify information produced by different species in response to typical predator encounters, including a moving predator stimulus presented inside of MSFs, and a stationary predator model presented both inside and outside of MSFs. MSFs were much less frequent in the breeding season with fewer individuals of the nuclear species, David's Fulvetta (Alcippe davidi), participating, though the diversity of other species remained stable. Fulvettas were the dominant producer of alarm‐related information both to the moving and stationary stimuli in MSFs and were also among the most active mobbers to stimuli presented outside of MSFs. In the breeding season, they tended to call less to the moving stimulus, and substantially fewer individuals responded to the in‐flock stationary stimulus. Other species increased their own information production at stationary predator stimuli (inside and outside of MSFs) during the breeding season, perhaps due to their increased investment in offspring during this time. Yet even during the breeding season, David's Fulvetta remained the highest producer of information about predators in MSFs. Hence, while we show that information production in MSFs can be somewhat dynamic, we describe a continually asymmetric communication system, in which a nuclear species is important to the whole community.     

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.     

Adapting to the unpredictable: reproductive biology of vertebrates in the Australian wet-dry tropics

In the wet-dry tropics of northern Australia, temperatures are high and stable year-round but monsoonal rainfall is highly seasonal and variable both annually and spatially. Many features of reproduction in vertebrates of this region may be adaptations to dealing with this unpredictable variation in precipitation, notably by (i) using direct proximate (rainfall-affected) cues to synchronize the timing and extent of breeding with rainfall events, (ii) placing the eggs or offspring in conditions where they will be buffered from rainfall extremes, and (iii) evolving developmental plasticity, such that the timing and trajectory of embryonic differentiation flexibly respond to local conditions. For example, organisms as diverse as snakes (Liasis fuscus, Acrochordus arafurae), crocodiles (Crocodylus porosus), birds (Anseranas semipalmata) and wallabies (Macropus agilis) show extreme annual variation in reproductive rates, linked to stochastic variation in wet season rainfall. The seasonal timing of initiation and cessation of breeding in snakes (Tropidonophis mairii) and rats (Rattus colletti) also varies among years, depending upon precipitation. An alternative adaptive route is to buffer the effects of rainfall variability on offspring by parental care (including viviparity) or by judicious selection of nest sites in oviparous taxa without parental care. A third type of adaptive response involves flexible embryonic responses (including embryonic diapause, facultative hatching and temperature-dependent sex determination) to incubation conditions, as seen in squamates, crocodilians and turtles. Such flexibility fine-tunes developmental rates and trajectories to conditions--especially, rainfall patterns--that are not predictable at the time of oviposition.     

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.     

The effects of rainfall on different components of seasonal fecundity in a tropical forest passerine

Seasonal fecundity is a composite metric that is determined by component parameters such as clutch size, nest survival and re‐nesting probability. Many of these component parameters are known to vary with environmental conditions, in particular rainfall prior to or during the breeding season. In some species, seasonal fecundity is positively related to rainfall, but little is known about which component parameters of seasonal fecundity respond most strongly to rainfall. We used intensive nest monitoring of a multi‐brooded tropical forest passerine, the Montserrat Oriole Icterus oberi, to examine the effects of rainfall during the pre‐breeding season on component parameters of annual fecundity. We monitored all nests of a total of 42 pairs over 5 years in which rainfall varied substantially. We then related clutch size, nest survival, onset and length of the breeding season, re‐nesting probability and re‐nesting interval to pre‐breeding season rainfall using generalized linear mixed models that accounted for random variation across sites and individual pairs, and incorporated other variables known to affect the response. Higher pre‐breeding season rainfall led to an increase in clutch size and a decrease in re‐nesting interval, but nest survival, re‐nesting probability and length of the breeding season were not affected by variation in rainfall. The onset of the breeding season was delayed in very dry years. We conclude that higher rainfall is likely to increase food availability and thus body condition of female Montserrat Orioles, leading to an increase in fecundity due to larger clutch sizes.     

Rainfall as a breeding stimulus and clutch size determinant in South African arid-zone birds

Rainfall is known to be a stimulus for initiating breeding in arid-zone birds, but its influence on clutch size is unknown for most southern African birds. The breeding activity of nine out of 11 species studied in the arid Nama Karoo biome of South Africa increased markedly after appreciable rainfall. Some resident insectivores did breed in the absence of rainfall, even under drought conditions, but also laid eggs within one week of small rain showers. Nomadic granivores required more substantial rainfall to stimulate what was usually a synchronized, population-wide breeding response. The timing and length of the breeding season was dependent on the integrated effect that rainfall and temperature had on the growing season of the vegetation. Over 50% of the 11 species studied showed evidence of an increase in clutch size following large rainfall events. Grey-backed Finchlark Eremopterix verticalis began laying larger clutches within one week of a rainfall stimulus, illustrating how rapid this response can be.     

Evidence of postbreeding prospecting in a long‐distance migrant

1. Organisms assess biotic and abiotic cues at multiple sites when deciding where to settle. However, due to temporal constraints on this prospecting, the suitability of available habitat may be difficult for an individual to assess when cues are most reliable, or at the time they are making settlement decisions. For migratory birds, the postbreeding season may be the optimal time to prospect and inform settlement decisions for future breeding seasons.
2. We investigated the fall movements of flammulated owls (Psiloscops flammeolus) within breeding habitat after fledglings had gained independence and before adults left for migration. From 2013 to 2016, we trapped owls within a breeding population wherein all nesting owls and their young have been banded since 1981. We used stable isotopes in combination with mark–recapture data to identify local individuals and differentiate potential prospecting behavior from other seasonal movements such as migration or staging.
3. We commonly captured owls in the fall—predominantly hatch‐year owls—that were not known residents of the study area. Several of these nonresident owls were later found breeding within the study area. Stable isotope data suggested a local origin for virtually all owls captured during the fall.
4. Our results suggest that hatch‐year flammulated owls, but also some after‐hatch‐year owls, use the period between the breeding season and fall migration to prospect for future breeding sites. The timing of this behavior is likely driven by seasonally variable costs associated with prospecting.
5. Determining the timing of prospecting and the specific cues that are being assessed will be important in helping predict the extent to which climate change and/or altered disturbance regimes will modify the ecology, behavior, and demographics associated with prospecting.