Controls over reproductive phenology among ungulates: allometry and tropical‐temperate contrasts

Ungulates inhabiting high latitudes schedule the timing of conceptions so that offspring are born during the most favourable nutritional conditions for reproductive success. The optimal period for births is less reliably predictable in tropical and subtropical savanna environments where plant growth is governed by rainfall, suggesting that reproductive phenology could be influenced more proximately by resources affecting the body condition of females around the time of conceptions. To assess how these controls operate, we compared the timing of births and conceptions among tropical and subtropical savanna ungulates with the patterns shown by ungulates in northern temperate or subarctic latitudes. The association between the timing of births and the onset of plant growth early in the growing season is less consistent among tropical savanna ungulates than among ungulates inhabiting northern temperate environments, and apparently subject to other influences affecting vegetation phenology. Nevertheless, birth peaks seem to coincide with the time of the year when forage quality is expected to be best for offspring survival and growth for most tropical or subtropical ungulates with gestation periods shorter than a year. When gestation time exceeds one year, proximal effects of nutritional conditions around the time of conceptions apparently become overriding and birth synchrony with early season plant growth is no longer effective. Proximate nutritional influences on conceptions may also govern the somewhat diffuse spread of births shown by ungulate populations in equatorial latitudes where photoperiod cues controlling oestrus and mating cannot be used to schedule the later timing of births.     

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.     

Rainfall influences on ungulate population abundance in the Mara-Serengeti ecosystem

1. Rainfall is the prime climatic factor underpinning the dynamics of African savanna ungulates, but no study has analysed its influence on the abundance of these ungulates at monthly to multiannual time scales. 2. We report relationships between rainfall and changes in age- and sex-structured abundances of seven ungulate species monitored monthly for 15 years using vehicle ground counts in the Maasai Mara National Reserve, Kenya. 3. Abundance showed strong and curvilinear relationships with current and cumulative rainfall, with older topi, Damaliscus korrigum (Ogilby); warthog, Phacochoerus aethiopicus (Pallas); waterbuck, Kobus ellipsyprimnus (Ogilby); and impala, Aepyceros melampus (Lichtenstein) responding to longer lags than younger animals, portraying carryover effects of prior habitat conditions. 4. The abundances of newborn calves were best correlated with monthly rainfall averaged over the preceding 5-6 months for topi, waterbuck, warthog, and 2 months for the migratory zebra Equus burchelli (Gray), but with seasonal rainfall averaged over 2-5 years for giraffe, Giraffa camelopardalis (L.); impala; and kongoni, Alcelaphus busephalus (Pallas). The cumulative late wet-season rainfall was the best predictor of abundance for quarter- to full-grown animals for most species. Monthly rainfall exerted both negative and positive effects on the abundances of zebra, impala and waterbuck. Ignoring age, both sexes responded similarly to rainfall. 5. Births were strongly seasonal only for warthog and topi, but peaked between August and December for most species. Hence abundance was strongly seasonal for young topi and warthog and the migratory zebra. Pronounced seasonality in births for warthog and topi obliterated otherwise strong relationships between abundance and rainfall when both month and rainfall were included in the same model. Aggregated density produced relationships with rainfall similar to those for fully grown animals, emphasizing the necessity of demographic monitoring to reliably reveal rainfall influences on ungulate abundance in the Mara. 6. Strong relationships between abundance and rainfall suggest that rainfall underpins the dynamics of African savanna ungulates, and that changes in rainfall due to global warming may markedly alter the abundance and diversity of these mammals. Ungulates respond to rainfall fluctuations through movements, reproduction or survival, and the responses appear independent of breeding phenology and synchrony, dietary guild, or degree of water dependence. Newborns and adults have contrasting responses to rainfall. Males and females respond similarly to rainfall when age is ignored.     

Birth seasonality and calf mortality in a large population of Asian elephants

In seasonal environments, many species concentrate their reproduction in the time of year most likely to maximize offspring survival. Asian elephants (Elephas maximus) inhabit regions with seasonal climate, but females can still experience 16‐week reproductive cycles throughout the year. Whether female elephants nevertheless concentrate births on periods with maximum offspring survival prospects remains unknown. We investigated the seasonal timing of births, and effects of birth month on short‐ and long‐term mortality of Asian elephants, using a unique demographic data set of 2350 semicaptive, longitudinally monitored logging elephants from Myanmar experiencing seasonal variation in both workload and environmental conditions. Our results show variation in birth rate across the year, with 41% of births occurring between December and March. This corresponds to the cool, dry period and the beginning of the hot season, and to conceptions occurring during the resting, nonlogging period between February and June. Giving birth during the peak December to March period improves offspring survival, as the odds for survival between age 1 and 5 years are 44% higher for individuals born during the high birth rate period than those conceived during working months. Our results suggest that seasonal conditions, most likely maternal workload and/or climate, limit conception rate and calf survival in this population through effects on maternal stress, estrus cycles, or access to mates. This has implications for improving the birth rate and infant survival in captive populations by limiting workload of females of reproductive age. As working populations are currently unsustainable and supplemented through the capture of wild elephants, it is imperative to the conservation of Asian elephants to understand and alleviate the effects of seasonal conditions on vital rates in the working population in order to reduce the pressure for further capture from the wild.     

Rainfall extremes explain interannual shifts in timing and synchrony of calving in topi and warthog

We tested the hypothesis that ungulates time and synchronize births to match gestation and lactation with peak food availability and quality in seasonal environments, using ground counts of topi and warthog conducted over 174 months (July 1989–December 2003) in the Mara–Serengeti ecosystem. During this 15-year period, 2,725 newborn and 45,574 adult female topi and 933 newborn and 7,831 adult warthogs were recorded. Births were distinctly synchronized in both species but far less so than in ungulates in temperate regions. Extreme droughts delayed onset and reduced synchrony of calving and natality rates but high rainfall advanced onset and increased synchrony of calving and natality rates in both species, supporting the seasonality hypothesis. Annual shifts in birth peaks were significantly negatively correlated with the preceding wet season rainfall. Varying the timing and synchrony of births and natality rates are widespread but little understood adaptations of ungulates to climatic extremes. Climate change heightens the need for advancing this understanding because increasing frequency and severity of droughts is likely to decouple phenology of breeding in seasonally breeding ungulates from that in their food plants. Similar studies of African ungulates are either extremely rare or non-existent. New approaches to estimating the time of peak births and its confidence limits and the degree of synchrony of breeding are also presented.     

Reassessing the determinants of breeding synchrony in ungulates

Predicting the consequences of climate change is a major challenge in ecology and wildlife management. While the impact of changes in climatic conditions on distribution ranges has been documented for many organisms, the consequences of changes in resource dynamics for species' overall performance have seldom been investigated. This study addresses this gap by identifying the factors shaping the reproductive synchrony of ungulates. In temporally-variable environments, reproductive phenology of individuals is a key determinant of fitness, with the timing of reproduction affecting their reproductive output and future performance. We used a satellite-based index of resource availability to explore how the level of seasonality and inter-annual variability in resource dynamics affect birth season length of ungulate populations. Contrary to what was previously thought, we found that both the degree of seasonal fluctuation in resource dynamics and inter-annual changes in resource availability influence the degree of birth synchrony within wild ungulate populations. Our results highlight how conclusions from previous interspecific analyses, which did not consider the existence of shared life-history among species, should be treated with caution. They also support the existence of a multi-faceted link between temporal variation in resource availability and breeding synchrony in terrestrial mammals, and increase our understanding of the mechanisms shaping reproductive synchrony in large herbivores, thus enhancing our ability to predict the potential impacts of climate change on biodiversity.     

Birth seasonality in the early Spanish-Mexican colonists of California (1769-1898)

The pattern of birth seasonality in California's early Spanish-Mexican colonists between 1769 and 1898 was reconstructed using genealogical data for progeny of 657 marriages. The monthly distribution of the 3,824 births in this sample shows a strong seasonal pattern, with spring and fall peaks (corresponding to peaks in conceptions during July and February) and a low point in October. This seasonal reproductive pattern is the result of a complicated set of interactions among environmental, physiological, and cultural variables. California's strongly developed winter rainfall pattern and the 19th-century agricultural cycle clearly influenced the seasonal pattern of marriages and births in this agrarian society. Several historical processes interacted with these environmental and economic factors to transform the seasonal birth pattern of the early colonists. Through time the birth pattern becomes less variable and the birth maximum shifts from spring to early winter. This appears to be, at least in part, a result of changes in labor patterns and an increase in average parity. These data suggest a multifactorial explanation for birth seasonality, in which the timing of conceptions and births is influenced by both environmental and socioeconomic factors.     

Ungulate Reproductive Parameters Track Satellite Observations of Plant Phenology across Latitude and Climatological Regimes

The effect of climatically-driven plant phenology on mammalian reproduction is one key to predicting species-specific demographic responses to climate change. Large ungulates face their greatest energetic demands from the later stages of pregnancy through weaning, and so in seasonal environments parturition dates should match periods of high primary productivity. Interannual variation in weather influences the quality and timing of forage availability, which can influence neonatal survival. Here, we evaluated macro-scale patterns in reproductive performance of a widely distributed ungulate (mule deer, Odocoileus hemionus) across contrasting climatological regimes using satellite-derived indices of primary productivity and plant phenology over eight degrees of latitude (890 km) in the American Southwest. The dataset comprised > 180,000 animal observations taken from 54 populations over eight years (2004–2011). Regionally, both the start and peak of growing season (“Start” and “Peak”, respectively) are negatively and significantly correlated with latitude, an unusual pattern stemming from a change in the dominance of spring snowmelt in the north to the influence of the North American Monsoon in the south. Corresponding to the timing and variation in both the Start and Peak, mule deer reproduction was latest, lowest, and most variable at lower latitudes where plant phenology is timed to the onset of monsoonal moisture. Parturition dates closely tracked the growing season across space, lagging behind the Start and preceding the Peak by 27 and 23 days, respectively. Mean juvenile production increased, and variation decreased, with increasing latitude. Temporally, juvenile production was best predicted by primary productivity during summer, which encompassed late pregnancy, parturition, and early lactation. Our findings offer a parsimonious explanation of two key reproductive parameters in ungulate demography, timing of parturition and mean annual production, across latitude and changing climatological regimes. Practically, this demonstrates the potential for broad-scale modeling of couplings between climate, plant phenology, and animal populations using space-borne observations.     

Changing vulnerability to predation related to season and sex in an African ungulate assemblage

The consequences of predation for prey population dynamics depend on the extent to which this mortality is predisposed by malnutrition or senescence, or additive in the sense that animals that would otherwise not have died at that time were killed. In places lacking effective predators, few adult ungulates die during the summer or wet season months when food is plentifully available. Hence the seasonal distribution of predator kills as well as the age and sex classes of the prey indicates the extent to which malnutrition contributes to mortality as well as other influences on vulnerability. Using records of animal deaths assembled over 35 years in South Africa's Kruger National Park, these patterns were investigated for 12 ungulate species forming the prey of lions, and for three other large predators with respect to one prey species. Buffalo, kudu and giraffe were more strongly represented in kills made during the late dry season, while wildebeest and zebra made relatively greater contributions during the wet season. Impala, waterbuck, warthog and rarer antelope species became more prominent in kills during transitional periods between seasons. Five prey species showed an elevation in representation of males in lion kills during the mating season, as well as impala for all predator species. Females were more prominently represented in kills during the time of late gestation and parturition for three prey species. Hence reproductive activities as well as changing vegetation cover and food resources affected vulnerability to predation. Shifts in susceptibility to predation over the seasonal cycle corresponded with rainfall‐related variation in the annual representation of these ungulate species in lion kills. The availability of vulnerable prey species, age and sex classes at different stages of the seasonal cycle helps maintain a high abundance of lions. These factors contribute to the strong additive impact that predation has had on the abundance of some of these ungulate populations.     

Dynamics of births and juvenile recruitment in Mara–Serengeti ungulates in relation to climatic and land use changes

Natality and recruitment govern animal population dynamics, but their responses to fluctuating resources, competition, predation, shifting habitat conditions, density feedback and diseases are poorly understood. To understand the influences of climatic and land use changes on population dynamics, we monitored monthly changes in births and juvenile recruitment in seven ungulate species for 15 years (1989–2003) in the Masai Mara Reserve of Kenya. Recruitment rates declined for all species but giraffe, likely due to habitat alteration and increasing vulnerability of animals associated with recurrent severe droughts, rising temperatures, unprecedentedly strong and prolonged El Niño-Southern Oscillation (ENSO) episodes, expansion of settlements, cultivation and human population growth in pastoral ranches adjoining the reserve. Birth rate showed strong and humped relationships with moving averages of monthly rainfall, whereas recruitment responded strongly to cumulative past rainfall. Increasing livestock incursions into the reserve depressed recruitment rate for quarter-grown topi. Expansion of pastoral settlements depressed birth rate in impala, zebra and giraffe. Frequent ENSO-related droughts caused progressive habitat desiccation and hence nutritional shortfalls for ungulates. The responses to climatic, land use and resource influences did not reflect body size, migratory or resident lifestyle, dietary guild, digestive physiology or degree of synchrony of breeding of the ungulate species.     

Annual Variation in Ecology and Reproduction of Wild Simakobu (Simias concolor)

Seasonal breeding in primates is related to the degree of environmental seasonality, particularly the availability and predictability of food. Southeast Asian species in general show moderate birth seasonality due to either low environmental seasonality or unpredictable fluctuations of mast-fruiting food resources. One Southeast Asian primate, the simakobu (Simias concolor), however, has been reported to be a strict seasonal breeder with births occurring in June and July only. It is unclear whether these observations are characteristic of the species or result from a sampling bias. To address this question, we documented the annual distribution of 11 births in eight groups of simakobu over two consecutive years at Pungut, an undisturbed site on Siberut Island, Indonesia. We assessed annual variation in ecology and reproduction via rainfall, temperature, food availability, feeding time, physical condition, conceptions, and births. Mean monthly temperature was nearly constant (26.3–27.1?°C), and monthly precipitation always high (219–432?mm). Although simakobu foods were abundant year-round, there were two fruit-feeding peaks in June and September. In contrast to previous reports, we documented births in 7?mo. Most births occurred in October (45?%), the wettest month of the year, and most conceptions in March and April, following a peak in unripe fruit availability. Although sample sizes are very small, females seemed to conceive when their physical condition was best, suggesting that simakobu time conceptions flexibly to the recovery of energy reserves. Across study sites, births occurred in 10 calendar months, indicating that simakobu reproduction is not strictly seasonal.     

Reproductive parameters of wild Trachypithecus leucocephalus: seasonality,infant mortality and interbirth interval

Understanding the reproductive parameters of endangered primate species is vital for evaluating the status of populations and developing adequate conservation measures. This study provides the first detailed analysis of the reproductive parameters of wild white‐headed langurs (Trachypithecus leucocephalus), based on demographic data collected over an 8‐year period in the Nongguan Karst Hills in Chongzuo County, Guangxi, China. From 1998 to 2002, a total of 133 live births were recorded in the population based on systematic censuses. Births occurred throughout the year, but the temporal pattern was highly correlated with seasonal variation in temperature and rainfall, with the birth peak coinciding with the dry and cold months of November–March. The average birthrate was 0.47±0.13 births per female per year and mortality for infants younger than 20 months was 15.8%. From 1998 to 2006, 14 females gave birth to 41 infants in four focal groups. The average age at first birth for female langurs was 5–6 years (n=5) and the interbirth interval (IBI) was 23.2±5.2 months (median=24.5 months, n=27). Infants are weaned at 19–21 months of age. The IBI for females with infant loss before weaning was significantly shorter than those for females whose infants survived. It appears that birth seasonality in the white‐headed langurs is influenced by seasonal changes in food availability. The timing of conceptions was found to coincide with peak food availability. The reproductive parameters for white‐headed langurs reported here are quite similar to those reported for other colobine species. One major difference is our observation of lower infant mortality in Trachypithecus. Am. J. Primatol. 71:558–566, 2009. © 2009 Wiley‐Liss, Inc.     

Trade-off between resource seasonality and predation risk explains reproductive chronology in impala

We investigated the variation in birth synchrony displayed by impala Aepyceros melampus populations across their distribution from southern to eastern Africa. Our analysis was based on field data from Chobe National Park in Botswana and Mala Mala Private Game Reserve in South Africa (4 and 13 years of monitoring, respectively). We compared our results with those from other studies conducted across the impala species range. Impala lambing was highly synchronized in Chobe with 90% of lambs born within 2 weeks in mid-November. Variation in rainfall in the preceding wet season explained 74% of variation in the dates of the first lamb observation in Mala Mala. In Chobe, the earliest birth peak occurred after the highest rainfall and the body condition of lambs in that cohort was also best for both males and females. No association was found between the lunar cycle and the estimated onset of the conception period, despite previous studies having found an association between the lunar cycle and the rutting behaviour in males. On a regional scale, impalas in areas with a marked dry season (several months with no rain) tend to synchronize births with the onset of the rains, when grass quality is highest. Number of months with rain explained 78% of the regional variation in birth synchrony. Neither latitude nor total rainfall contributed significantly to a stepwise multiple regression model. These data support the theory that impalas synchronize births in areas with a highly seasonal food supply, and temporally space births in less seasonal (equatorial) areas to reduce predation risks.     

Sources of variation in interbirth intervals among captive bonnet macaques (Macaca radiata)

Female fitness is a function of variation in the length of females' reproductive careers, the viability of their offspring, and the frequency with which they give birth. Infant loss shortens interbirth intervals in most primate species, but we know considerably less about other factors that contribute to variation in the length of interbirth intervals within groups. In one large captive group of bonnet macaques, maternal parity, age, experience, family size, and recent reproductive history are all associated with variation in the length of intervals that follow the birth of surviving infants. Primiparous females have the longest interbirth intervals, while multiparous females who have produced surviving infants in the past and have raised their last infant successfully have the shortest interbirth intervals. Infant sex and maternal rank have no direct effect upon the length of interbirth intervals. One of the underlying causes of variation in the length of interbirth intervals after surviving births seems to be variation in the timing of conceptions among females. Females who conceive early in the mating season tend to have shorter interbirth intervals than other females. However, females who are multiparous, experienced, and have recently raised infants have late conceptions and short interbirth intervals.     

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.     

Resource availability and roosting ecology shape reproductive phenology of rain forest insectivorous bats

Bats in temperate and subtropical regions typically synchronize birth of a single young with peaks in resource availability driven by local climate patterns. In tropical rain forest, insects are available throughout the year, potentially allowing departures from seasonal monoestry. However, reproductive energy budgets may be constrained by the cost of commuting to foraging grounds from distant roosts. To test these hypotheses, we simultaneously tracked female reproductive activity of 11 insectivorous bat species, insect biomass, and local weather variables for 20 months in a Malaysian rain forest. Five species roost in forest structures and hence have low commuting costs, whereas six species depend on caves, which are limited in the landscape, and are presumed to incur higher commuting costs to foraging sites. Monthly insect biomass was positively correlated with monthly rainfall, and there was a significant relationship between insect biomass and lactation in cave‐roosting but not forest‐roosting species. Cave‐roosting species were seasonally monoestrus, with parturition confined to a two‐month period, whereas in forest‐roosting species, pregnancy and lactation were recorded throughout the year. Our results suggest that the energetic costs of commuting from roosts to foraging grounds shape annual reproductive patterns in tropical rain forest insectivorous bats. Ongoing changes in forest landscapes are likely to increase these costs for cave‐roosting bats, further restricting reproductive opportunities. Climate change is projected to influence the timing of rainfall events in many tropical habitats, which may disrupt relationships between rainfall, insect biomass, and bat reproductive timing, further compromising reproductive success.     

Interannual variability in fruit abundance and the reproductive seasonality in Sumatran Long-tailed macaques (Macaca fascicularis)

In an environment with a seasonal food supply, most primate species show birth peaks which precede the peak food period by some two to five months. Sumatran Long-tailed macaques (Macaca fascicularis), however, showed birth peaks during or after the fruit peak. Years of high birth rates and early birth peaks alternated with years of low birth rates and late peaks. The timing of births was strongly influenced by a female's condition, which depends on food supply and her previous reproductive history. Pregnant females were more active than other females, whereas females with young infants were less active.
The unusual timing of births is ascribed to the unpredictability of the height of the annual fruit peak. This hypothesis is supported by the reproductive patterns of other South-east Asian primates and by a model comparing the two types of reproductive timing. Further differences between the two strategies of reproductive timing are predicted.     

Climate and Species Richness Predict the Phylogenetic Structure of African Mammal Communities

We have little knowledge of how climatic variation (and by proxy, habitat variation) influences the phylogenetic structure of tropical communities. Here, we quantified the phylogenetic structure of mammal communities in Africa to investigate how community structure varies with respect to climate and species richness variation across the continent. In addition, we investigated how phylogenetic patterns vary across carnivores, primates, and ungulates. We predicted that climate would differentially affect the structure of communities from different clades due to between-clade biological variation. We examined 203 communities using two metrics, the net relatedness (NRI) and nearest taxon (NTI) indices. We used simultaneous autoregressive models to predict community phylogenetic structure from climate variables and species richness. We found that most individual communities exhibited a phylogenetic structure consistent with a null model, but both climate and species richness significantly predicted variation in community phylogenetic metrics. Using NTI, species rich communities were composed of more distantly related taxa for all mammal communities, as well as for communities of carnivorans or ungulates. Temperature seasonality predicted the phylogenetic structure of mammal, carnivoran, and ungulate communities, and annual rainfall predicted primate community structure. Additional climate variables related to temperature and rainfall also predicted the phylogenetic structure of ungulate communities. We suggest that both past interspecific competition and habitat filtering have shaped variation in tropical mammal communities. The significant effect of climatic factors on community structure has important implications for the diversity of mammal communities given current models of future climate change.     

Seasonality at the equator: isotope signatures and hormonal correlates of molt phenology in a non-migratory Amazonian songbird

Background

Birds, across their annual cycle, progress through sequences of life-history stages such as reproduction and molt. The mechanisms that control annual avian itineraries involve endocrine responses triggered by seasonal environmental factors, including changes in resource availability and/or photoperiod. However, at equatorial latitudes birds are exposed to different degrees of seasonality, and the mechanisms underlying phenology of birds near the equator remain less explored. We studied the silver-beaked tanager, an endemic Amazonian songbird, from an equatorial lowland population. Remarkably, in this species, song behavior has been shown to be seasonally aligned to minimal changes in day length near the equator. Here, we aimed to further explore the phenology of silver-beaked tanagers by assessing shifts of food sources utilization as potential ultimate factors. We measured triple isotopic tracers of carbon (δ¹³C), nitrogen (δ¹⁵N) and sulphur (δ³⁴S) in blood and feathers of birds throughout a whole year. In addition, we assessed the degree of seasonality in the molting activity, in relation to circulating levels of corticosterone, as well as to testosterone as a proxy of the reproductive condition of males.

Results

There was important seasonal variation of δ³⁴S values in relation to rainfall patterns and changes in estuarine water composition. Despite the seasonal rainfall, we found no substantial variation in the foraging ecology of birds over seasons. This was accompanied by uniform levels of corticosterone throughout the year, probably associated with the absence of drastic seasonal resource shortages. Even so, silver-beaked tanagers showed a marked seasonal molting schedule, which was related to variation in the circulating levels of both corticosterone and testosterone.

Conclusions

These findings suggest that foraging niche is not life history stage-dependent in silver-beaked tanagers, and highlight rainfall as an important environmental cue for bird phenology. Our stable isotope results encourage further studies addressing the influence of estuarine water dynamics on bird timing. In addition, the results suggest a primary role of steroid hormones in regulating seasonal life history stages under the absence of a marked photoperiod. Contrary to what might be expected for a tropical songbird, our physiological data in silver-beaked tanagers do not support reproduction-molt overlapping.

     

Effects of annual rainfall and habitat types on the body mass of impala (Aepyceros melampus) in the Zambezi Valley, Zimbabwe

Body mass is often considered as a good indicator of body condition of individuals in ungulates, hence of their fitness, and thus, may be used as an index to monitor the status of populations subject to harvesting schemes. Here, we report the influence of annual rainfall (a proxy for primary production) and habitat on the body mass of impala in a population cropped for meat in a communal area of Zimbabwe. We analyzed the data from 2 contrasted years for rainfall, in two different habitats. In the good year (i.e. high annual rainfall) impala were heavier than in the poor year, and adult females seemed to be less affected than males by variation in primary production. We show that adult males were suffering from a seasonal decrease in body mass, supposedly linked to the rut, particularly in good habitat. Overall, the habitat effect appeared to be dominated by the rainfall effect, and this may be due to the very high animal densities in the good habitat, i.e. fast resource depletion. Our results also suggest that males (juvenile and adult) are more susceptible to changes in food resource abundance and quality than females, which supports previous studies on sexually dimorphic and polygynous species.