Age- and sex-dependent spring arrival dates of Eastern Kingbirds

ABSTRACT.   Among migratory passerines, the first birds to arrive on the breeding grounds are usually older males. Early arrival by older birds may be driven by experience, age-dependent changes in body condition, age-dependent access to resources during the nonbreeding period, or latitudinal segregation by age. Males may arrive earlier than females (protandry) because males maintain better condition due to greater access to resources during the winter, or because selection favors early arriving males that acquire the best territories or experience enhanced mating opportunities. During a 4-yr study (2004–2007) in Oregon, we found that older Eastern Kingbirds ( Tyrannus tyrannus ), regardless of sex, arrived nearly a week before younger birds and that males arrived about 5 d before females. Age- and sex-dependent arrival dates do not appear to be related to differences in body condition, social dominance in winter, or latitudinal segregation, and protandry is unrelated to the ability of early-arriving males to acquire high-quality territories. Instead, we propose that young birds have less to gain from early arrival because of their probable inability to displace experienced birds from prime territories and that protandry evolved due to enhanced mating opportunities for early arriving males that arise from the high rates of extra-pair matings in our population of Eastern Kingbirds.     

Sex-differentiated migration patterns, protandry and phenology in North European songbird populations

This study aims to investigate causes and mechanisms controlling protandrous migration patterns (the earlier breeding area arrival of males relative to females) and inter-sexual differences in timing of migration in relation to the recent climate-driven changes in phenology. Using standardised ringing data from a single site for eight North European migratory passerines collected throughout 22 years, we analysed sex-differentiated migration patterns, protandry and phenology of the entire populations. Our results show protandrous patterns for the first as well as later arriving individuals for all studied species. Males show more synchronous migration patterns compared to females and, hence, first arriving females followed males more closely than later arriving individuals. However, we found no inter-sexual differences in arrival trends as both sexes advance spring arrival over time with the largest change for the first arriving individuals. These findings seem in support of the “mate opportunity” hypothesis, as the arrival of males and females is strongly coupled and both sexes seem to compete for early arrival. Changes in timing of arrival in males and females as a response to climatic changes may influence subsequent mating decisions, with subsequent feedbacks on population dynamics such as reproductive success and individual fitness. However, during decades of consistent earlier spring arrival in all phases of migration we found no evidence of inter-sexual phenological differences.     

Factors influencing plasticity in the arrival‐breeding interval in a migratory species reacting to climate change

Climate change is profoundly affecting the phenology of many species. In migratory birds, there is evidence for advances in their arrival time at the breeding ground and their timing of breeding, yet empirical studies examining the interdependence between arrival and breeding time are lacking. Hence, evidence is scarce regarding how breeding time may be adjusted via the arrival‐breeding interval to help local populations adapt to local conditions or climate change. We used long‐term data from an intensively monitored population of the northern wheatear (Oenanthe oenanthe) to examine the factors related to the length of 734 separate arrival‐to‐breeding events from 549 individual females. From 1993 to 2017, the mean arrival and egg‐laying dates advanced by approximately the same amount (~5–6 days), with considerable between‐individual variation in the arrival‐breeding interval. The arrival‐breeding interval was shorter for: (a) individuals that arrived later in the season compared to early‐arriving birds, (b) for experienced females compared to first‐year breeders, (c) as spring progressed, and (d) in later years compared to earlier ones. The influence of these factors was much larger for birds arriving earlier in the season compared to later arriving birds, with most effects on variation in the arrival‐breeding interval being absent in late‐arriving birds. Thus, in this population it appears that the timing of breeding is not constrained by arrival for early‐ to midarriving birds, but instead is dependent on local conditions after arrival. For late‐arriving birds, however, the timing of breeding appears to be influenced by arrival constraints. Hence, impacts of climate change on arrival dates and local conditions are expected to vary for different parts of the population, with potential negative impacts associated with these factors likely to differ for early‐ versus late‐arriving birds.     

Effects of Spring Temperatures on the Strength of Selection on Timing of Reproduction in a Long-Distance Migratory Bird

Climate change has differentially affected the timing of seasonal events for interacting trophic levels, and this has often led to increased selection on seasonal timing. Yet, the environmental variables driving this selection have rarely been identified, limiting our ability to predict future ecological impacts of climate change. Using a dataset spanning 31 years from a natural population of pied flycatchers (Ficedula hypoleuca), we show that directional selection on timing of reproduction intensified in the first two decades (1980–2000) but weakened during the last decade (2001–2010). Against expectation, this pattern could not be explained by the temporal variation in the phenological mismatch with food abundance. We therefore explored an alternative hypothesis that selection on timing was affected by conditions individuals experience when arriving in spring at the breeding grounds: arriving early in cold conditions may reduce survival. First, we show that in female recruits, spring arrival date in the first breeding year correlates positively with hatch date; hence, early-hatched individuals experience colder conditions at arrival than late-hatched individuals. Second, we show that when temperatures at arrival in the recruitment year were high, early-hatched young had a higher recruitment probability than when temperatures were low. We interpret this as a potential cost of arriving early in colder years, and climate warming may have reduced this cost. We thus show that higher temperatures in the arrival year of recruits were associated with stronger selection for early reproduction in the years these birds were born. As arrival temperatures in the beginning of the study increased, but recently declined again, directional selection on timing of reproduction showed a nonlinear change. We demonstrate that environmental conditions with a lag of up to two years can alter selection on phenological traits in natural populations, something that has important implications for our understanding of how climate can alter patterns of selection in natural populations.     

Carry-Over Effects of Nonbreeding Habitat on Start-to-Finish Spring Migration Performance of a Songbird

For migratory animals, conditions during the nonbreeding period may carry-over to influence spring migration performance. Animals in low-quality habitats are predicted to be in poorer condition, show later migration timing, and travel at slower speeds. This can result in subsequent negative effects on fitness. We tested the hypothesis that nonbreeding season body condition and habitat quality carry-over to affect spring migration performance of a long-distance migratory songbird, the Wood Thrush (Hylocichla mustelina). We tracked individual birds between multiple breeding sites in North America and nonbreeding sites in Central America. First, we compared body condition of nonbreeding birds migrating to the same general region of the breeding range with spring migration performance (timing, speed, and duration) obtained from light-level geolocators. Second, we assessed the Normalized Difference Vegetation Index (NDVI) as a proxy for nonbreeding habitat quality, and predicted that birds from wetter habitat or in wetter years (higher NDVI) would show improved migration performance relative to birds from drier sites. We found no evidence of individual-level carry-over effects of nonbreeding season body condition on spring migration performance. Lower NDVI of nonbreeding habitat resulted in delayed spring migration departure, but this effect disappeared by arrival at breeding sites. Birds occupying drier nonbreeding sites migrated faster and for fewer days, compensating for their relatively late departure. We also documented a broader pattern in NDVI and migration timing and distance, in that birds that occupied the wettest areas in the southern part of the nonbreeding range departed significantly later and migrated farther. Our results suggest that individual carry-over effects of nonbreeding habitat quality may be compensated for by a faster and shorter migration strategy. At a broad scale, consistently later spring timing and longer migration distances were associated with the wettest areas (the highest quality habitats) of the Wood Thrush non-breeding range. This supports the theory that high-quality habitats offset the costs of farther migration, resulting in a leap-frog migration pattern.     

A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds

Although there is substantial evidence that Northern Hemisphere species have responded to climatic change over the last few decades, there is little documented evidence that Southern Hemisphere species have responded in the same way. Here, we report that Australian migratory birds have undergone changes in the first arrival date (FAD) and last date of departure (LDD) of a similar magnitude as species from the Northern Hemisphere. We compiled data on arrival and departure of migratory birds in south‐east Australia since 1960 from the published literature, Bird Observer Reports, and personal observations from bird watchers. Data on the FAD for 24 species and the LDD for 12 species were analyzed. Sixteen species were short‐ to middle‐distance species arriving at their breeding grounds, seven were long‐distance migrants arriving at their nonbreeding grounds, and one was a middle‐distance migrant also arriving at its nonbreeding ground. For 12 species, we gathered data from more than one location, enabling us to assess the consistency of intraspecific trends at different locations. Regressions of climate variables against year show that across south‐east Australia average annual maximum and minimum temperatures have increased by 0.17°C and 0.13°C decade^?1 since 1960, respectively. Over this period there has been an average advance in arrival of 3.5 days decade^?1; 16 of the 45 time‐series (representing 12 of the 24 species studied) showed a significant trend toward earlier arrival, while only one time‐series showed a significant delay. Conversely, there has been an average delay in departure of 5.1 days decade^?1; four of the 21 departure time‐series (four species) showed a significant trend toward later departure, while one species showed a significant trend toward earlier departure. However, differences emerge between the arrival and departure of short‐ to middle‐distance species visiting south‐east Australia to breed compared with long‐distance species that spend their nonbreeding period here. On average, short‐ to middle‐distance migrants have arrived at their breeding grounds 3.1 days decade^?1 earlier and delayed departure by 8.1 days decade^?1, thus extending the time spent in their breeding grounds by ～11 days decade^?1. The average advance in arrival at the nonbreeding grounds of long‐distance migrants is 6.8 days decade^?1. These species, however, have also advanced departure by an average of 6.9 days decade^?1. Hence, the length of stay has not changed but rather, the timing of events has advanced. The patterns of change in FAD and LDD of Australian migratory birds are of a similar magnitude to changes undergone by Northern Hemisphere species, and add further evidence that the modest warming experienced over the past few decades has already had significant biological impacts on a global scale.     

Starting over: experimental effects of breeding delay on reproductive success in early‐arriving male American redstarts

Birds that arrive and breed early often have higher reproductive success than late individuals, either as a consequence of timing‐specific advantages (the timing hypothesis) or because these individuals and/or their resources are of higher quality (the quality hypothesis). In this study, we examined the potential influence of several factors affecting reproductive success by experimentally delaying breeding of early‐arriving male American redstarts Setophaga ruticilla, a species for which early male arrival is strongly related to increased reproductive success. Our manipulation involved the capture, holding, and release of males following pairing and territory establishment, resulting in the majority of subjects (67%) losing their initial mate (47%) or mate and territory (20%) and forcing them to start over approximately 12 d after their initial arrival. Males forced to start over (i.e. those losing their first territory and/or mate) did not experience any decrease in body condition, nor did their reproductive behaviour differ from that of early‐arriving control males. We found that naturally early‐arriving but experimentally manipulated males suffered reduced fledging success in comparison to early‐arriving males that bred early or late, but equivalent success in comparison to males that arrived and bred late. Based on our results, we propose that the relationship between early arrival and higher reproductive success in this species is mediated not simply by individual male quality or absolute arrival timing alone, but rather some other aspect of resource quality is likely important. We discuss and present evidence for two alternative explanations under the quality hypothesis: female quality and territory quality. To our knowledge, our study is the first to investigate the effects of experimentally delaying male breeding time, strengthening previous correlational evidence for resource quality as a potentially important selective agent driving early arrival in migratory birds.     

Early arrival, initiation of nesting, and social status: an experimental study of breeding female red-winged blackbirds

Red-winged blackbirds (Agelaius phoeniceus) arrive at breedingsites several months before nests are constructed. Males inthis highly polygynous species presumably return early to defenddesirable territories. Females, however, also begin to arrivealmost two months before nesting is attempted. Early returnto breeding sites could enhance reproductive success by increasinga female's social status and thereby allowing earlier nesting.I measured the effect of experimentally delayed arrival on thetiming of nesting, reproductive success, and social status offemales. Birds were captured as they arrived in early springand detained in an aviary. These experimental females were laterreleased at their capture sites before control females beganconstruction of nests; controls had arrived during the sameperiod as experimentals but were not detained. Experimentalfemales nested, on average, more than a week later than controls,although I could detect no effect of timing on reproductivesuccess during this study. By manipulating arrival date, butnot covariates of arrival time such as age or experience, thisresult indicates that timing of arrival directly influencedlaying date. Delayed females were subordinate to control femaleswith the same mate. This decline in social status may have beenresponsible for the delay in nesting and could have fitnessconsequences for females in some years, because the offspringof earlier nesting, dominant birds are more likely to receivemale parental care in this population.     

Relationship between arrival date, hatching date and breeding success of the white stork (Ciconia ciconia) in Slovakia

Changes in the spring arrival dates of migrant birds have been reported from a range of locations and many authors have focused on long-term trends and their relationship to temperature and other climatic events. Perhaps more importantly, changed arrival dates may have consequences for the breeding dates of birds which strongly influence breeding success. In this paper we take the opportunity provided by a monitoring scheme of the white stork (Ciconia ciconia) to examine several features of the timing of arrival and breeding in relation to chick production in Slovakia during the period 1978–2002. First arrival dates ranged from 5^th March to 30^th April, and hatching dates varied between 26^th April and 8^th July. Generally, early arriving pairs started breeding earlier and a shorter interval between the arrival of the first partner and breeding, expressed here as hatching date, resulted in higher breeding success.     

Spring migration phenology of birds in the Northern Prairie region is correlated with local climate change

ABSTRACT In apparent response to recent periods of global warming, some migratory birds now arrive earlier at stopover sites and breeding grounds. However, the effects of this warming on arrival times vary among locations and species. Migration timing is generally correlated with temperature, with earlier arrival during warm years than during cold years, so local variation in climate change might produce different effects on migration phenology in different geographic regions. We examined trends in first spring arrival dates (FADs) for 44 species of common migrant birds in South Dakota (1971–2006) and Minnesota (1964–2005) using observations compiled by South Dakota and Minnesota Ornithologists’ Unions. We found significant trends in FAD over time for 20 species (18 arriving earlier and two later) in South Dakota and 16 species (all earlier) in Minnesota. Of these species, 10 showed similar significant trends for both states. All 10 of these species exhibited significantly earlier arrival, and all were early spring migrants, with median FADs before 10 April in both states. Eighteen of the 44 species showed significant negative correlations of FADs with either winter (December–February) or spring (arrival month plus previous month) temperatures in one or both states. Interestingly, spring temperatures in both South Dakota and Minnesota did not warm significantly from 1971–2006, but winter temperatures in both states warmed significantly over the same time period. This suggests that the warmer winters disproportionately affected early spring migrants, especially those associated with aquatic habitats (seven of the 10 species showing significantly earlier spring arrival in both states). The stronger response to climate change by early spring migrants in our study is consistent with the results of several other studies, and suggests that migrants, especially early migrants, are capable of responding to local temperature conditions experienced on wintering grounds or along the migration route.     

A SUMMARY OF THE BIRDS OF WESTMINSTER O.F.S. AND SURROUNDINGS (1953-1955)

Results are presented from point-counts at six sites in the Kalahari in Botswana. Counts were repeated three times: during a dry season following good rains (1991), during the next wet season when rains were far below average, and the following dry season (1992) when the area became drought-stricken. Compared to the wet season, bird numbers decreased during the drought by 37–81% and species by 8–52%; compared to the previous dry season, birds decreased by 5–71% and species by 2–47%. Bird diversity (relative to numbers) tended to increase during the wet season but was little affected by drought, except in the northern Kalahari, where a greater proportion of birds moved out in response to drought. This gave the northern Kalahari the most distinct bird community during a wet cycle, but it became again typically Kalahari during the drought. Thus, the typical Kalahari bird communities expanded their range during drought into the moister periphery. Changes in numbers, most probably resulting from (local) movements were found in many species. Most confirmed earlier reports on their nomadic nature but some, like Red-crested Lophotis ruficrista and Northern Black Afrotis afraoides Korhaan, Chestnut-vented Tit-Babbler Parisoma subcaeruleum, Ant-eating Chat Myrmecocichla formicivora, Tinkling Cisticola Cisticola rufilatus, Black-chested Prinia Prinia flavicans, Marico Flycatcher Bradornis mariquensis and Brown-crowned Tchagra Tchagra australis have not been or are not widely recognised as mobile species.     

Climate and the complexity of migratory phenology: sexes,migratory distance,and arrival distributions

The intra- and inter-season complexity of bird migration has received limited attention in climatic change research. Our phenological analysis of 22 species collected in Chicago, USA, (1979–2002) evaluates the relationship between multi-scalar climate variables and differences (1) in arrival timing between sexes, (2) in arrival distributions among species, and (3) between spring and fall migration. The early migratory period for earliest arriving species (i.e., short-distance migrants) and earliest arriving individuals of a species (i.e., males) most frequently correlate with climate variables. Compared to long-distance migrant species, four times as many short-distance migrants correlate with spring temperature, while 8 of 11 (73%) of long-distance migrant species’ arrival is correlated with the North Atlantic Oscillation (NAO). While migratory phenology has been correlated with NAO in Europe, we believe that this is the first documentation of a significant association in North America. Geographically proximate conditions apparently influence migratory timing for short-distance migrants while continental-scale climate (e.g., NAO) seemingly influences the phenology of Neotropical migrants. The preponderance of climate correlations is with the early migratory period, not the median of arrival, suggesting that early spring conditions constrain the onset or rate of migration for some species. The seasonal arrival distribution provides considerable information about migratory passage beyond what is apparent from statistical analyses of phenology. A relationship between climate and fall phenology is not detected at this location. Analysis of the within-season complexity of migration, including multiple metrics of arrival, is essential to detect species’ responses to changing climate as well as evaluate the underlying biological mechanisms.     

Finish with a sprint: Evidence for time‐selected last leg of migration in a long‐distance migratory songbird

Under time‐selected migration, birds should choose a strategy for outcompeting rivals over securing access to prime resources at the final destination. Thus, migration can be viewed as a race among individuals where winners are arriving first when conditions are suitable. The sprint migration hypothesis predicts that individuals shift from maximum sustained speed to a final burst of sprint to shorten the transition from migration to breeding (Alerstam, 2006). In this study, we test the hypothesis of a final sprint migration in a long‐distance Afro‐Palearctic migrant, the collared flycatcher Ficedula albicollis, during autumn and spring, and compare migration strategies between the seasons. In both seasons, collared flycatchers evidently exhibited sprint migration by increasing their overall speed over the last leg of migration after the Sahara crossing. This phenomenon was more pronounced in spring, contributing to overall faster spring migration and possibly highlighting higher importance for early arrival at the breeding grounds. In both seasons and particularly in spring, late departing individuals flew at a faster rate, partially being able to catch up with their early departing conspecifics. Differential fueling strategies may play an important role in determining migration speed, especially during the early stages of the migration, and might explain the observed differences in migration speeds between late and early departing individuals. Our findings suggest competition for early arrival at the breeding and at the nonbreeding destinations alike. Sprint migration might be an appropriate strategy to gain advantage over conspecifics and settle in prime territories as well as to cope with the increasingly earlier springs at high latitudes.     

Arrival date and microorganisms in barn swallows

Migration between breeding sites and winter quarters constitute a major life history strategy in birds. The benefits of such migrations must at least equal the costs for such behavior to evolve and be maintained. We tested whether there was a relationship between abundance and diversity of microorganisms on nest lining feathers and timing of arrival by barn swallows Hirundo rustica. Nest lining feathers are chosen and transported by adult barn swallows to their nests just before and during egg laying, at a time when barn swallows have arrived weeks earlier, implying that any heterogeneity in abundance and diversity of microorganisms on feathers in nests must arise from feather preferences. There was a negative relationship between arrival date and the total number of fledglings showing that early arrival is advantageous. The arrival date of adult barn swallows was significantly positively correlated with the abundance of specific bacteria (Bacillus licheniformis) and positively correlated with the abundance of the fungus Trichoderma reesei and negatively correlated with the abundance of the fungus Mucor circinelloides. Moreover, we found a significant positive relationship between arrival date and mean total number of bacterial colonies in TSA medium. There was a significant negative relationship between arrival date and mean total number of bacterial colonies in FMA medium, and Simpson's diversity index of the abundance of bacteria in FMA medium. Such heterogeneity may arise from some microorganisms being beneficial, others detrimental and yet others benign and of no significant importance. In contrast, there was no significant relationship between arrival date and age of individuals. These findings are consistent with the hypothesis that early arriving barn swallows differ in abundance and diversity of microorganisms from late arriving conspecifics, and that they choose feathers for their nests that differ in terms of microorganisms from those chosen by late arrival individuals.     

Does wintering north or south of the Sahara correlate with timing and breeding performance in black‐tailed godwits?

Migrating long distances requires time and energy, and may interact with an individual's performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.     

Absence of protandry in the spring migration of a population of Song Sparrows Melospiza melodia

Early male arrival at breeding sites, or protandry, is thought to have evolved from intrasexual competition among males for access to mates or breeding resources. Males of polygynous species tend to be larger than females and have exaggerated secondary sexual traits. Additionally, such species show a high degree of protandry, suggesting that timing of arrival is sexually selected. Species showing limited sexual dimorphism and showing sexual monochromatism may be expected to show limited early male arrival. However, there are very few studies of migration timing of the sexes in such species because individuals cannot be readily identified to sex in the hand. In this study, we genetically sexed birds and found no evidence for early male arrival, for a population of migratory Song Sparrows Melospiza melodia . For our study population, males and females display limited sexual size dimorphism and are sexually monochromatic which is characteristic of the species. Fat scores for males were inversely associated with timing of arrival, whereas for females, larger-winged birds arrived sooner – suggesting that early migration timing may be selected for in both sexes.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

Temporal patterns of arrival from migration as a response to competition for breeding space

Birds arriving from spring migration have higher chances of securing a territory if they reach the breeding area before their companions. However, an early arrival may compromise their chances of surviving until reproduction begins. Here, we study how evolutionarily stable responses to these opposite constraints may shape patterns of arrival in a population of migratory birds, with the help of a simple evolutionary algorithm. We show that, initially, an increasing level of competition for breeding territories generates a massive peak of early arrivals. However, when the level of competition becomes very high, the temporal distribution of arrivals may take different shapes, depending on the modalities of competition. If individuals trying to usurp territories have relatively high chances of success, a second peak of arrival emerges at the very end of the arrival period. Birds arrive then in two successive waves. By contrast, if these chances are low, a single early peak of arrival is still present. Many individuals do however continuously arrive afterwards, at a regular and increasing tempo, until reproduction begins. The prior residence effect, that rules competition between resident birds and their challengers, is thus expected to influence the whole distribution of arrival dates on breeding grounds. This distribution will, in turn, affect the ability of the population to respond to changing environmental conditions.     

Fat stores of American redstarts Setophaga ruticilla arriving at northerly breeding grounds

Many long-distance passerine migrants arrive with more fat stores than necessary to have reached their northerly breeding grounds. Researchers have argued for adaptive advantages associated with arriving with 'surplus' migratory fat, including increased breeding performance and insurance against adverse weather, and reduced food resources during the days following arrival. The objectives of this work were to document fat stores in American redstarts Setophaga ruticilla arriving to breed in northern Michigan and to test predictions associated with the hypothesis that arrival fat serves an insurance role. Results suggest that redstarts arrived in northern Michigan with fat stores sufficient to have continued migrating an additional distance greater than 1000 km. Significant yearly variation in arrival fat corresponded in part with environmental conditions measured at the breeding grounds. Birds arrived with the most fat in the year with the coolest temperatures and the lowest food abundance at the time of arrival to the site. Further, an inverse relationship between arrival fat and arrival day in males indicated that early arrivals carried more fat than later arrivals. Birds that arrived early faced cooler temperatures and lower densities of terrestrial invertebrates, and arrival fat may have provided a mechanism to overcome poor early season foraging conditions. However, our results are not entirely consistent with the hypothesis that arrival fat serves only as insurance. Arrival fat appeared important even during the most benign year of this study. Further, evidence suggests that females may have derived more benefit from arrival fat than males. These results highlight the connection between phases of a bird's annual cycle. Migrants that do well en route , arrive at breeding grounds in better condition, which may contribute to survival and reproductive success.     

Leaf flushing phenology and herbivory in a tropical dry deciduous forest,southern India

Patterns of leaf-flushing phenology of trees in relation to insect herbivore damage were studied at two sites in a seasonal tropical dry forest in Mudumalai, southern India, from April 1988 to August 1990. At both sites the trees began to flush leaves during the dry season, reaching a peak leaf-flushing phase before the onset of rains. Herbivorous insects emerged with the rains and attained a peak biomass during the wet months. Trees that flushed leaves later in the season suffered significantly higher damage by insects compared to those that flushed early or in synchrony during the peak flushing phase. Species whose leaves were endowed with physical defenses such as waxes suffered less damage than those not possessing such defenses. There was a positive association between the abundance of a species and leaf damage levels. These observations indicate that herbivory may have played a major role in moulding leaf flushing phenology in trees of the seasonal tropics.