THE LEVEL OF RESERVE PROTEIN AS THE PROXIMATE FACTOR CONTROLLING THE TIMING OF BREEDING AND CLUTCH-SIZE IN THE RED-BILLED QUELEA QUELEA QUELEA

Changes in the fat and flight muscle protein reserves of adult Red-billed Queleas Quelea quelea were followed in two colonies in Tanzania and Botswana. At the start of a breeding attempt the protein reserves were higher that at any other time of the year, particularly in females which had heavier flight muscles (non-fat dry weight) than did the (larger) males at that time. The pre-breeding increase in the labile component of the muscle protein (the actual protein reserve) is calculated at 80% for females, but only 14% for males. The fat reserve was only increased slightly at the beginning of the breeding attempt.
In both sexes, though for different reasons, the protein and fat reserves fell rapidly during the first few days of the attempt, in some individuals to dangerously low levels. During the incubation period there was rapid recovery while the situation during the rearing period appeared to vary between colonies.
It is proposed that the proximate control of breeding is provided by the individual's own body condition, and particularly the state of its protein reserves. No environmental releasers are required for the birds to breed at the appropriate time of the year.
Individual females producing two, three and four egg clutches differed in the rate at which their reserves fell during egg formation. The results are used to support the view that in this quelea the actual clutch-size produced on any occasion is the largest the female can produce before becoming too enfeebled. Thus, clutch-size appears to be determined phenotypically downward from a maximum which is indirectly under genetic control.     

Isotopic (δ2Hf) evidence of “loop migration” and use of the Gulf of Maine Flyway by both western and eastern breeding populations of Blackpoll Warblers

Declining numbers of Blackpoll Warblers (Setophaga striata) have been documented at long‐term migration monitoring sites as well as in breeding areas. However, the “loop migration” of Blackpoll Warblers makes it difficult to ascribe population change at migration monitoring sites to specific breeding populations. Individuals from all populations across the breeding range of Blackpoll Warblers concentrate in fall along the Atlantic coastline of eastern North America prior to initiating a transoceanic flight to wintering areas. In spring, Blackpoll Warblers return along a different route, moving north into the southeastern United States where birds from eastern and western breeding populations then diverge during migration to reach their respective breeding areas. To monitor breeding populations outside of breeding areas and identify factors potentially affecting those populations, we must be able to identify where birds captured during migration breed and map seasonal variation in population‐specific flyways. To “map” population‐specific migration movements of Blackpoll Warblers, we used feather deuterium (δ²H_f) values and a spatially explicit model to assign molt origins of 289 Blackpoll Warblers moving through sites in the Gulf of Maine (GOM) region and at three locations further west and south (northern Great Lakes area, Pennsylvania, and Florida). The assignment method was validated with feather samples from 35 birds captured during the breeding season at Churchill, Manitoba, Canada. As predicted, the spatial pattern of movement within and between seasons reflected “loop migration.” Blackpoll Warblers captured during fall migration in the GOM region included birds from across their breeding range, whereas birds captured during the spring were exclusively from northeastern populations. During fall migration, Blackpoll Warblers captured at two sites west of the GOM were from breeding areas further northwest than those from western Canada that were captured in the GOM. Blackpoll Warblers captured in eastern Florida during spring migration were assigned exclusively to breeding areas in the northeast, suggesting that eastern and western populations diverge soon after entering the United States. Finally, most Blackpoll Warblers sampled at Manomet Bird Observatory originated from breeding populations in Alaska and western Canada that have shown a similar (70–90%) decline over the same period. Our results, therefore, not only document the “loop migration” pattern of Blackpoll Warblers, but, by mapping patterns of connectivity between breeding and non‐breeding areas, may help target conservation efforts for breeding populations of Blackpoll Warblers where most needed.     

Evidence of itinerant breeding of the Red-billed Quelea Quelea quelea in the Ethiopian Rift Valley

This paper provides evidence for itinerant breeding by Red-billed Quelea Quelea quelea in the Ethiopian Rift Valley. Queleas were mass-marked with aerially applied fluorescent particles in two separate nesting areas in southwestern Ethiopia during June 1981. Marked adults from both areas were recovered from nesting colonies in the Awash River Valley during August and September, up to 100 days post-spray and between 500 and 700 km to the north of the spray sites. In the Awash colonies, the presence of marked adults in breeding plumage with interrupted primary wing-moult, together with two age classes of juveniles, suggests that this was the second nesting. The progress of both the post-breeding and post-juvenal primary moults was consistent with an earlier breeding in May and June. The timing of the arrival and departure of birds from these nesting areas also supported the occurrence of double breeding by the same birds. Furthermore, the composition of black facial mask types of adult males was more similar between samples from the southern Ethiopian Rift and Awash Valley than between samples from either of these areas and samples from outside the Rift, providing additional evidence that the same population of birds bred in these two areas of the Rift Valley. Nesting colonies in both areas were scattered in time and space. Colonies in the Awash were distributed for more than 300 km and were established over a two-month period, which coincided with local differences in the timing of the seeding in grasses. This wide distribution probably increases nesting success in such areas of locally variable rainfall. This contrasts with a stategy of mass migration, where concentrated breeding occurs where and when suitable conditions are first encountered. Knowledge of the location and timing of previously successful nesting areas may increase the success of itinerant breeding, as Queleas seem to use the same areas in successive years if conditions are favourable. Recoveries of marked birds in the Awash were segregated according to both spray site and sex, suggesting some degree of group cohesion by sex from the first to the second nesting. We speculate that post-nesting group cohesion of adults may provide a means to maintain the integrity of collective group information on seasonal movements. Group cohesion may be facilitated by the high degree of nesting synchrony within colonies.     

DISPERSION, POPULATION ECOLOGY AND MIGRATION OF EASTERN GREAT REED WARBLERS ACROCEPHALUS ORIENTALIS WINTERING IN MALAYSIA

A population of 400–600 Acrocephalus orientalis wintering in a Phragmites habitat at 3°N in West Malaysia was studied during four northern hemisphere winters, by means of systematic mist-netting. Data from other study-areas, other habitats and other winters are also used. Intensive mist-netting appears to have made birds move over longer distances than they did in the absence of disturbance, and to have led to the emigration of marked birds from the study-area. Trapping also affected feeding behaviour, resulting in weight-loss; repeated trapping may have increased mortality. Males and females could be separated by means of wing-length in fresh plumage. Females were largely confined to Phragmites; males were more numerous on the edge of reed-beds and in scrub vegetation. Males suffered greater feather-wear than females. As measured by the trapping rate, birds were uniformly distributed throughout the Phragmites habitat, at the same density in different winters. Undisturbed birds used a “home-range” of 1–4 ha, overlapping with 15–50 other individuals. Disturbed birds overlapped with 100–200 others. Individual birds returned to exactly the same “home-range” in successive winters. After correcting for the effects of disturbance and incomplete sampling, the proportion of adults ringed in one winter which returned in the next is estimated as 65% in each of two study-areas. This is a minimum estimate of the annual survival rate for adults. Mean total body-weights were at a minimum in midwinter (November-February). Fat-free weights were also lower in midwinter than in autumn and spring. Body-moult was observed in March and April. Moult of the flight-feathers takes place between July and September, on the breeding grounds or slightly to the south. Females departed on spring migration between 10 and 25 May; males some 11–14 days earlier. Adults arrived in autumn between 8 September and 7 October; males and females often came in in separate “waves”. Females were absent for only about 127 days, about the minimum required for migration, breeding and moult. Dates of migration match those of the more northern breeding populations. Spring departure is later than dates of passage recorded in south China; hence birds of this population appear to make long nights. On average, birds departing in spring carried about 9 g of fat, roughly 40% of total fat-free body-weight. This is about half the energy reserve required for the entire journey. Dates of passage in central China are consistent with a hypothesis that they make the journey (4,500-5,000 km) in two “hops”. A few birds which remained light until very late in the spring showed a significantly lower return rate in the next year. Most birds arriving in autumn appear to have carried 1–2 g of fat, but some were at or below the normal fat-free weight. Many birds appear to have lost weight soon after arrival. Returning ringed adults were amongst the very first birds trapped in September. Individual birds appear to have migrated on very similar dates in different years: many of the dates of trapping differed by 2 days or less in successive years. Trapping rates reached a peak in early October and then declined rapidly, reaching the midwinter level by 21 October. The decline coincided with the differential disappearance of juvenile birds. However, birds collected at this time had adequate fat reserves, and the disappearance appears to have preceded the period of food-shortage. It is suggested that the loss of juvenile birds resulted from behavioural interactions favouring the more dominant individuals, as has been described for several temperate zone residents. The first few weeks in the wintering area may thus be the critical period of mortality during the year. Because birds from different breeding areas are expected to be mixed in the winter-quarters, and vice versa, local mortality factors in winter may affect a number of breeding populations. High adult survival rates have been recorded in several other birds which breed in the temperate zones and winter in the tropics. In general their breeding success appears to be high, so the first-year mortality must be high. The closely related A. arundinaceus, which winters in Africa, differs from A. orientalis in size, wing-shape, timing of spring migration and timing of moult. These differences can be interpreted as adaptations to different environmental (primarily climatic) factors experienced during migration and on the breeding grounds. The segregation of males and females into different habitats probably reduces inter-sexual competition in winter, but this is not necessarily its primary function. Males collected in the evening in Phragmites had smaller fat reserves than females, suggesting that the females are better adapted to this habitat. The large size of the males is probably maintained in part by sexual selection in the breeding season. On the other hand, the size of females and their habitat is probably limited by the specialisation of their nest. These factors would suffice to explain the sexual dimorphism in size and habitat.     

Year‐round spatial movements and trophic ecology of Trindade Petrels (Pterodroma arminjoniana)

Trindade Petrels (Pterodroma arminjoniana) are vulnerable gadfly petrels that breed on the remote Trindade Island, located ~1100 km off the Brazilian coast. Little is known about their spatial ecology, and their trophic ecology has only been described for the breeding season. We tagged four Trindade Petrels with global location sensing loggers (GLS) from October 2013 to November 2014 and sampled the blood and feathers (innermost primary and the eighth secondary) of 14 individuals to evaluate their year‐round spatial and isotopic ecology. We examined individual distributions, habitat use and suitability, activity, and isotopic values during the breeding, migration, and non‐breeding periods. Trindade Petrels used areas in the southwest Atlantic Ocean (between 10°N and 50°S in latitude) during the breeding season. They migrated through pelagic waters of the tropical Atlantic to the northwest Atlantic, where they spent the non‐breeding season. Trindade Petrels used mostly tropical to subtropical waters in areas of intermediate to high wind speeds and low marine productivity. Individuals spent more time foraging at night than during the day. During the breeding season, birds in northerly areas had higher carbon‐13 values, and birds that used more pelagic areas foraged on prey at a higher trophic level (higher nitrogen‐15 values) than those in more southern and coastal areas. Isotopic values during the breeding, migration, and non‐breeding periods differed, possibly due to differences among individuals in their at‐sea distribution throughout the year. We confirmed the non‐breeding distribution of Trindade Petrels, which was previously known only from vessel sightings and stranded birds. Our results also suggest a strong temporal segregation in the at‐sea distribution and trophic ecology between two groups of individuals, which might indicate the existence of two separate breeding populations.     

FEEDING ECOLOGY OF THE BLACK-FACED DIOCH QUELEA QUELEA IN NIGERIA

The results of a three-year study of the diet of Quelea quelea in the Lake Chad region of Nigeria indicate why this bird periodically does extensive damage to crops of dry-season guinea-corn.
The normal food of the birds in the early dry-season consists of small grass seeds collected off the ground. As the supply of these diminishes there is a gradual change-over to larger seeds.
In years when this change begins early, it is liable to coincide with the ripening of the cereal crop and severe damage may be caused.
At the beginning of the rains the dry-season food supply is lost owing to the simultaneous germination of the seeds over large areas.
The birds lay down fat reserves in preparation for this lean period.
After a short time the birds migrate south to regions where rain has been falling for some weeks.
In the short period spent in these parts, Queleas feed on ripening grass seeds—including those of cultivated millet.
There is evidence for considerable mortality during this time of food scarcity and it is suggested that the numbers of Queleas are limited by the food supply.
The significance of communal roosting is discussed. It is proposed that the phenomenon is related to food finding, the roost being considered as an "information centre" for the large area prospected by the birds constituting the roosting community. A method is outlined by which information on food availability within the feeding area may feasibly be transmitted within the community.     

THE BREEDING SEASON IN S.W. ECUADOR

The breeding season on the Santa Elena peninsula in S.W. Ecuador is described on the basis of 1761 nests found in the four years 1955 to 1958. The environment and climate are outlined—a cool, dry season from about May to November and a warm, variable wet season from December to April, when alone rain may fall. It is shown that each year the general breeding season is closely correlated with the rainfall. The peak of breeding varies from year to year by at least a month and its length from about six weeks to three and a half months.
Although all species for which there are enough data, are stimulated to breed after important falls of rain, most of those species of small land-birds which are completely resident, attempt to nest before the rain and often continue for some time afterwards. Some specific differences, between seed- and insect-eating finches, seem to be adaptations to food supply and the availability of nest sites and building material. Such evidence as there is suggests that raptors, waders and waterfowl also nest in the wet season (often late), rather than in the dry season. (Unfortunately herons and sea-birds do not nest in the area.)
Annual differences in the amount of breeding by the same species are thought not to be entirely due to differences in weather between years.
It is suggested that the late (August-September) breeding in the Galápagos Islands described by Lack (1950) and the apparent dry-season nesting of raptors, waders and waterfowl there might be found to be linked with variable or abnormally wet weather, if records of rainfall from the islands were available.     

Hatching date influences age at first reproduction in the black-headed gull

In long-lived colonial birds, age at recruitment is an important life-history character. Variation in this parameter may reflect differences in several factors, including competitive ability and breeding strategies. Further, these differences may be due to timing of hatching (for instance through differences in competitive ability). We investigated the age of first-time breeders in relation to hatching date in a black-headed gull Larus ridibundus colony situated in central France, from 1979 to 1993. Age at first breeding was estimated for four groups of individuals (total n=550) according to their hatching date, using a recent capture-recapture methodology which allowed us to estimate recruitment rate without the limiting assumptions of methods relying on simple return rates. The age at first breeding was negatively correlated with the hatching date of individuals: individuals hatched earlier in the season started breeding at a younger age than individuals born later. Proportionally more 2-year-old late-hatched individuals were seen breeding on small peripheral colonies than young early-hatched individuals. This difference disappeared after age 3 years. These results strongly suggest that individuals hatched late in the season start to breed on peripheral colonies before recruiting to their natal colony. A difference of few weeks in hatching date has consequences which can last for several years.     

Delayed female reproduction in equilibrium and chaotic populations

Behavioural and life history polymorphisms are often observed in animal populations. We analyse the timing of maturation and reproduction in risky and resource-limited environments. Field and laboratory evidence suggests that female voles and mice, for example, can adjust their breeding according to the level of risk to their own survival and to survival probabilities and recruitment of young produced under different environmental conditions. Under risky or harsh conditions breeding can be postponed until later in the current breeding season or even to the next breeding season. We develop a population dynamics and life history model for polymorphism in reproduction (co-existence of breeding and non-breeding behaviours) of females in an age-structured population, with two temporally distinct mating events within the breeding season. We assume that, after overwintering, the females can breed in spring and again in summer or they can delay breeding in spring and breed in summer only. Young females born in spring can either mature and breed in summer or stay immature and postpone breeding over the winter to the next breeding season. We show that an evolutionarily stable breeding strategy is either an age-structured combination of pure breeding behaviours (old females breed and young delay maturity) or a mixed breeding behaviour within age-classes (a fraction of females breed and the rest of the age class postpones breeding). Co-occurrence of mixed reproductive behaviour in spring and summer within a single breeding season is observed in fluctuating populations only. The reproductive patterns depend on intraspecific, possibly interspecific, and ecological factors. The density dependence (e.g. social suppression) and predation risk are shown to be possible evolutionary mechanisms in adjusting the relative proportions of the different but co-existing reproductive behaviours.     

‘Same procedure as last year?‘ Repeatedly tracked swifts show individual consistency in migration pattern in successive years

Individual migration pattern during non‐breeding season is still a black box in many migratory birds. However, knowledge on both individual level and population level in migration and overwintering is fundamental to understand the life cycle of these birds and the constraints affecting them. We showed in a highly aerial migrant, the common swift Apus apus, that repeatedly tracked birds breeding at one site in Germany used the same individual‐specific migration routes and wintering areas in subsequent years. In contrast, different individuals from the same breeding colony showed diverse movement patterns during non‐breeding season suggesting that several suitable areas for overwintering coexist. We found lower variation in timing of autumn and spring migration within than between individuals. Our findings provide first indication of individual consistency but between‐individual variation in migration pattern in a small non‐passerine bird revealed by geolocators. This supports that swifts have diverse but individual‐specific ‘step‐by‐step’ migration patterns revealing high flexibility through individual strategies.     

Different means to the same end: long-distance migrant seabirds from two colonies differ in behaviour, despite common wintering grounds

Although seabirds that are trans-equatorial migrants show apparently broad overlap among populations in the non-breeding season, such large-scale pattern may conceal subtle but nevertheless key differences in migratory behaviour. These specializations could reflect adaptation to different environments during the breeding season, carry-over effects from the breeding to the nonbreeding period, or asymmetries in competitive ability of birds of different origin. We compared the migratory and wintering behaviour of Cory's shearwaters Calonectris diomedea nesting in Berlengas and in the Selvagens, two colonies in contrasting oceanographic environments, separated by ca. 1200 km. Although no differences were found in winter distribution, there was a marked divergence in timing, route and the use of staging areas during the postbreeding (autumn) migration. Birds from Berlengas typically travelled to oceanic waters in the North Atlantic for an extended stopover, whereas those from Selvagens rarely did so. In the South Atlantic, birds from Selvagens spent more time in flight, perhaps because they had higher energy and nutrient requirements for feather replacement compared to birds from Berlengas, which moult more flight feathers during breeding. Stable isotope analyses of feathers suggested that this variation in activity patterns was unrelated to trophic ecology. Differences in migration routes and stopovers may expose populations to distinct threats, and should be taken into consideration when defining units for conservation purposes and developing appropriate management strategies.     

Consistent annual schedules in a migratory shorebird

Many migratory birds start prebreeding moult and premigratory fuelling some months before the breeding season and face severe time constraints, while travelling up to 15,000 km between non-breeding and breeding grounds. Shorebirds typically leave Southern Hemisphere non-breeding areas over a 3-4 week period, but whether they benefit from interannually consistent timing of departure is unknown. Here, I show that individual bar-tailed godwits (Limosa limosa baueri) from New Zealand are highly consistent in their migratory scheduling. Most birds left within the same week each year (between-year repeatability, r, of 0.83) and adult males, which moult into a bright breeding plumage, were also highly repeatable in the extent of their prebreeding moult (r=0.86). This is consistent with the hypothesis that birds have individually optimized migration schedules. Within adult males, but not females, smaller birds tended to migrate earlier than large birds. Whether this reflects differences in size-related migration speed, optimal breeding time at different sites or size-related natural or sexual selection pressures, remains unknown.     

Resource use for reproduction depends on spring arrival time and wintering area in an arctic breeding shorebird

Resources for egg production may come from body reserves stored before breeding (“capital breeders”) or from food acquired at the breeding site (“income breeders”). Arctic migrants were long thought to be capital breeders, because they often arrive at a time when local food availability is still limited. However, later evidence suggested that arctic breeding shorebirds are primarily income breeders, or that they use a mixed strategy depending on laying date. We explored the relationship between laying date and resource use for reproduction in the pectoral sandpiper Calidris melanotos breeding in the Alaskan arctic by contrasting carbon isotope (δ¹³C) values of the local diet and of maternal plasma, cellular blood, feather and claw with those of the eggs produced. Our results revealed that early breeding females utilize resources for egg production that were acquired recently at staging areas, whereas later breeding females mostly relied on nutrients derived from local food sources. These findings suggest that the resource allocation strategy used for reproduction differs among females, and varies depending on the timing of arrival and the start of reproduction. The arrival date at the breeding ground and laying date may critically depend on non‐breeding season events such as winter habitat choice, staging areas or migration routes. By comparing maternal feather δ¹³C, claw δ¹³C and feather δD, we examined whether non‐breeding season events influenced the use of resources for egg production through variation in capture date or clutch initiation date. Female pectoral sandpipers originating from moulting areas characterized by higher (more positive) δD signatures were caught earlier and started laying earlier, and they used stored resources for reproduction. Using regional maps of δD values for precipitation in the wintering sites in South America, we compared the spatial variation in the observed feather δD signatures. This analysis indicated that female pectoral sandpipers with higher δD signatures, presumably coming from more north‐easterly wintering sites in southern America, started laying earlier and used mostly stored resources for egg production, compared to females that wintered (or at least moulted) further south. Our results thus show that winter moulting habitat is linked to breeding resource allocation strategy in this high‐arctic breeding shorebird.     

Moult Strategies Affect Age Differences in Autumn Migration Timing in East Mediterranean Migratory Passerines

Adult passerines renew their flight feathers at least once every year. This complete moult occurs either in the breeding areas, just after breeding (summer moult), or, in some long-distance migratory species, at the non-breeding areas, after arrival to the southern wintering area at the end of autumn migration (winter moult). The aim of this study was to relate moult strategies with the DMD, the difference in median migration date, through Israel, between juveniles and adults. Our data on autumn migration timing in juveniles and adults was based on ringing data of 49,125 individuals belonging to 23 passerine species that breed in Europe and Western Asia and migrate through Israel. We found that DMD was associated with moult timing. In all species that perform a winter moult, adults preceded juveniles during autumn. Among migrants who perform a summer moult, we found evidence of both migration timing patterns: juveniles preceding adults or adults preceding juveniles. In addition, in summer moulters, we found a significant, positive correlation between mean breeding latitude and DMD. Although previous studies described that moult duration and extent can be affected by migration, we suggest that moult strategies affect both migration timing and migration strategy. These two moult strategies (summer or winter moult) also represent two unique migration strategies. Our findings highlight the evolutionary interplay between moult and migration strategies.     

Interrelationships between breeding frequency,timing and outcome in King Penguins Aptenodytes patagonicus: are King Penguins biennial breeders?

King Penguin Aptenodytes patagonicus chick growth is interrupted by a winter fast which extends the length of the breeding cycle (laying to chick independence) to 14–16 months, so that continuous annual successful breeding appears to be impossible. The 3–month laying period imposes further constraints with respect to timing of breeding attempts in relation to the onset of fasting. By the frequent resighting of individually marked birds at Marion Island, we examined the frequency of breeding and the relationships between timing, outcome and frequency in the same and in consecutive years. A total of 3101 adult King Penguins were banded between 1984 and 1991, yielding continuous breeding histories spanning a maximum of 5 years. Most penguins attempted to breed in consecutive years, although the likelihood of taking a year off increased with the number of consecutive attempts. In any one season, about 19% of potentially breeding adults did not breed. Early breeders were more likely to succeed than late breeders and comprised 84% of breeding attempts of known timing. Successful birds in one season usually bred late in the following season, whereas failed attempts were usually followed by an early attempt. Non-breeding was more likely to occur after a successful outcome than a failed one, and after a year off 93% of birds bred early. It is incorrect to refer to the King Penguin as a biennial breeder.     

Flexibility in the timing of post-nuptial moult among Red-billed Queleas Quelea quelea in Botswana in relation to the timing of breeding

The timing of primary moult of adult Red-billed Queleas Quelea quelea, captured as they were completing an unusually late breeding attempt at Francistown, northern Botswana, in June 2004, was compared with the timing of moult of birds breeding earlier in the season in north-west Botswana during two earlier years, 1971 and 1972. Differences between years in the dates when local colonies finished breeding (mid-March to late June) and between two localities in the same year (mid-March and late May) were matched by corresponding differences in the estimated dates of moult onset, ranging from mid-April to mid-June. Flexibility in the timing of moult among Red-billed Queleas in southern Africa evidently enables birds to take advantage of unusually late breeding opportunities by delaying moult onset and overlapping moult and breeding at the end of the nesting cycle. Such flexibility may also include moult interruption to permit late breeding, although its incidence in southern Africa is apparently low.     

Annual cycles in Jamaican forest birds

The annual cycles of forest birds in Jamaica were found to be very similar to those at higher latitudes. Most species bred between March and September, though a few possibly breed throughout the year, especially in cultivated areas. Primary moult followed immediately after breeding, and in some species was apparently arrested to allow a further breeding attempt. Several species were fatter outside the breeding season than during it, and this is interpreted as "winter fattening" comparable to that found in many birds at higher latitudes. Weights varied little but individuals retrapped were usually heavier outside the breeding season. In some species the first complete moult took place at the end of the first year, implying that the birds do not breed until at least two years old.     

Testing the reproductive and somatic trade‐off in female Columbian ground squirrels

Energetic trade‐offs in resource allocation form the basis of life‐history theory, which predicts that reproductive allocation in a given season should negatively affect future reproduction or individual survival. We examined how allocation of resources differed between successful and unsuccessful breeding female Columbian ground squirrels to discern any effects of resource allocation on reproductive and somatic efforts. We compared the survival rates, subsequent reprodction, and mass gain of successful breeders (females that successfully weaned young) and unsuccessful breeders (females that failed to give birth or wean young) and investigated “carryover” effects to the next year. Starting capital was an important factor influencing whether successful reproduction was initiated or not, as females with the lowest spring emergence masses did not give birth to a litter in that year. Females that were successful and unsuccessful at breeding in one year, however, were equally likely to be successful breeders in the next year and at very similar litter sizes. Although successful and unsuccessful breeding females showed no difference in over winter survival, females that failed to wean a litter gained additional mass during the season when they failed. The next year, those females had increased energy “capital” in the spring, leading to larger litter sizes. Columbian ground squirrels appear to act as income breeders that also rely on stored capital to increase their propensity for future reproduction. Failed breeders in one year “prepare” for future reproduction by accumulating additional mass, which is “carried over” to the subsequent reproductive season.     

Comparative analysis of factors associated with first‐year survival in two species of migratory songbirds

Our understanding of the full life cycle of most migratory birds remains limited. Estimates of survival rates, particularly for first‐year birds are notably lacking. This knowledge gap results in imprecise parameters in population models and limits our ability to fully understand life history trade‐offs. We used eleven years of field data to estimate first‐year apparent survival (φ_1st) for two species of migratory grassland songbirds that breed in the same managed habitats but have substantially different migration distances. We used a suite of life‐history, habitat and individually‐based covariates to explore causes of variation in φ_1st. The interaction between fledge date and body mass was the best supported model of apparent survival. We found differential effects of fledging date based on nestling body mass. Overall, lighter nestlings had greater apparent survival than heavier nestlings; average or heavy nestlings within‐brood had greater apparent survival when they fledged earlier in the summer. We hypothesize that heavier birds that fledge earlier in the season have a longer window of opportunity to evaluate potential breeding sites and are more likely to disperse greater distances from the natal region, thus confounding survival with permanent emigration. Lighter birds, particularly those fledged late in the breeding season may spend more time on self‐maintenance and consequently have less time to evaluate potential future breeding sites, showing greater fidelity to their natal region. We found no support for management treatment (timing of mowing), sex, brood size, or species as important covariates in explaining apparent survival. Our results suggest that differential migration distances may not have a strong effect on first‐year apparent survival.     

DISTRIBUTION, SYSTEMATICS, AND POLYMORPHISM OF THE AFRICAN WEAVER-BIRD QUELEA QUELEA

Quelea quelea is distributed throughout Africa south of the Sahara in the Grass Steppe, Dry Savanna, and Montane Grassland biomes. It is most numerous where its staple food–the seeds of annual grasses–is most plentiful. In all parts of the range, the males in nuptial plumage are dimorphic. Everywhere the “black—faced” morph is commoner, but with up to 25% of “white-faced”. Proportions of the morphs are given for samples from various parts of Africa. Only three races are accepted: quelea of western West Africa, aethiopica of the Sudan area, and lathami of southern Africa. In the Lake Chad basin there are hybrid swarms breeding (queleaj aethiopica), as there are in East Africa where aethiopica/lathami swarms occur.