Correlates of distribution and nesting success in a Welsh upland Eurasian Curlew Numenius arquata population

Capsule: Habitat structure and composition explained spatial variation in breeding distribution and nesting success in a declining upland Eurasian Curlew Numenius arquata population in North Wales.

Aims: To identify environmental correlates of Curlew breeding distribution, nesting success and change in distribution.

Methods: Thirty random 1?km squares stratified by historical population trend were surveyed for Curlew density and nesting success, and habitat- and predation-related variables in a landscape containing agriculturally improved farmland, and moorland that was partly protected and subject to grazing reductions for nature conservation. Analyses tested for associations between Curlew measures and environmental variables.

Results: Curlew breeding density declined by 29% between 1994 and 2008, and was highest in squares comprising a mixture of moorland and agriculturally improved farmland, and in squares with lower vegetation density and higher cover of Nardus stricta (characteristic of rough grazing). Nesting success was positively associated with cover of Trichophorum germanicum (characteristic of mire). Vegetation density was lower than average in squares with the highest Curlew densities, while in the protected area vegetation density was higher than average.

Conclusion: Habitat and vegetation variables influenced Curlew distribution and nesting success in North Wales, largely in line with previous results but with no evidence for predator-related influences. Habitat condition for Curlew in the protected area could probably be increased through targeted increases in grazing alongside the protection of priority habitats.     

Placement,survival and predator identity of Eurasian Curlew Numenius arquata nests on lowland grass-heath

ABSTRACT

Capsule

Within the UK’s largest lowland Eurasian Curlew Numenius arquata population, curlew preferentially nested on physically disturbed (treated) than undisturbed (control) grassland, and low nest survival rates were primarily attributable to predation by Red Fox Vulpes vulpes.     

Environmental correlates of breeding abundance and population change of Eurasian Curlew Numenius arquata in Britain

Capsule: Across Britain, breeding Eurasian Curlew Numenius arquata are less numerous and have shown greater population declines in areas with more arable farming, woodland cover and higher generalist predator abundance.

Aims: We present the first national-scale analysis of the potential drivers of Curlew population change in Britain, which is needed to guide conservation action for this globally near-threatened, declining species.

Methods: Breeding Bird Survey data and environmental predictors were used to model variation in Curlew abundance in 1995–99 and 2007–11, and population change between these periods.

Results: Arable farming and woodland cover were negatively associated with Curlew abundance and population declines. Curlew abundance was positively associated with extent of protected area coverage and gamebird numbers. Abundance and population change were positively associated with cooler temperatures and higher summer rainfall, but negatively associated with numbers of generalist predators.

Conclusions: We found support for the negative effects of intensive agriculture, forestry, increases in generalist predator populations and climate warming on Curlew abundance and population change. Effective site protection and measures to reduce generalist predator abundance may be important conservation measures, together with improving breeding habitat quality in the wider countryside.     

Population density and the intensity of paternity assurance behaviour in a monogamous wader: the Curlew Numenius arquata

We compared paternity assurance behaviour and related displays in high (1.6 pairs perkm²) and low (6.7 pairs per km²) density populations of the Curlew Numenius arquata breeding on arable farmland in western Finland. There was little evidence of individuals pursuing extra-pair copulations or males exhibiting paternity assurance behaviour. Furthermore, there was no variation in the frequency of intrusions or in the intensity of paternity assurance behaviour relative to lay date. However, intrusions and approaches to the pair female by extra-pair males were more frequent at high breeding density, and pair males remained closer to their female, followed her more and exhibited a higher frequency of copulatory behaviours in the high-density population. Therefore, high breeding density appeared to increase opportunities for individuals to copulate outside the pair-bond and resulted in more intense male paternity guards. Male song flight displays were also more frequent in the high-density population. Territorial absences by males were infrequent in both areas. The increased frequency of intrusions and interactions between non-pair individuals (male-male and male-female) at high density were probably the major factors in explaining area differences in the intensity of paternity assurance and territorial behaviour.     

Values and perceptions of landowners within remaining breeding territories of Eurasian Curlew Numenius arquata in Ireland

Habitat loss and degradation have been identified as some of the main threats to breeding Curlew (Numenius arquata) across much of Europe. In Ireland, marginal habitats such as rough or wet grasslands and peatlands have been fragmented or degraded by activities including afforestation, drainage and intensification. The management implemented by landowners directly affects Curlew breeding territories. However, the values and perceptions held by landowners whose lands contain Curlew breeding territories, or the factors driving the decisions behind farming practices in these areas are rarely considered when looking at the causes of changes in these bird populations. This study, as part of the Curlew Conservation Programme established in 2017, gathered data through the distribution of questionnaires to landowners found within three kilometres of Curlew breeding territories in Ireland. In this study, we identify the current land uses being employed in Curlew breeding territories, and query future projections of land use in these areas. We investigate landowners’ perceptions of the requirements to sustain favourable environments for breeding Curlew. We also explore landowner values with respect to farming. The landowners in this study identified habitat loss and predation as the main drivers for Curlew declines. The majority of farming systems in this study were cattle rearing, the sustainability of which is under threat across Ireland. The results indicate that these landowners are not financially motivated, however, the availability of financial aid and expert advice are listed by landowners as requirements for traditional farming practices to continue. These results give an insight to the lifestyle, values and perceptions owners of land adjacent or within Curlew breeding territories. This information can be used to design Curlew conservation programmes that align with these values.     

Effects of radiotagging on the weight gain and survival of Curlew Numenius arquata chicks

CapsuleNo detrimental effects on weight gain or survival to fledging stage were found.

Aims To assess the effects of radiotagging on the weight gain and survival of Curlew chicks.

Methods Small (≤ 1.5 g) radiotags were attached to Curlew chicks at hatching, during studies in four different areas of the UK. Tagged chicks were recaptured regularly and their weight gain and survival compared to non-tagged chicks using within and between-brood comparisons.

Results Radiotagging did not affect weight gain. While there was no evidence of reduced survival of tagged chicks, comparisons were based on small samples.

Conclusions Small radiotags can be invaluable in determining the survival and causes of mortality of the chicks of waders and other precocial birds.     

Estimating the abundance and hatching success of breeding Curlew Numenius arquata using survey data

Counts of individual Curlew and estimates of breeding pairs from ‘field-by-field’ surveys were highly correlated with the numbers of nesting pairs, as determined by intensive studies on 27 sites located on four grassland-dominated study areas. The mean count of individual Curlew over three standard survey visits to each site was used to estimate numbers of nesting pairs. This estimate exceeded the number of nesting pairs on all study areas (as assessed by intensive studies) by 12% but was more accurate than the maximum number of pairs estimated from the three survey visits on each site (the previous convention for estimating breeding pairs). Estimates of the number of pairs hatching chicks were assessed by recording alarm-calling Curlew during late survey visits. The maximum number of alarm-calling pairs was used to estimate the number of pairs hatching chicks, overestimating this by 54%. Three of the study sites were adjacent to extensive moorland which produced overestimates during surveys because moorland nesting birds fed and led broods onto these sites. Omitting these sites from consideration reduced the degree of overestimation to 1% for the number of pairs and 7% for the number of pairs hatching chicks.     

Population estimates,trends and habitat associations of breeding Lapwing Vanellus vanellus,Curlew Numenius arquata and Snipe Gallinago gallinago in Northern Ireland in 1999

Capsule Declines in the breeding populations of Snipe, Lapwing and Curlew were recorded between April and June 1999 and compared with previous estimates in 1987.

Aims To compare populations of non-coastal breeding waders between 1987 and 1999.

Methods In 106 2 km × 2 km square tetrads, observers recorded the number of breeding pairs of waders and habitat details on 1:10 000 scale maps.

Results A significant decline of c. 60% for Lapwing and Curlew, and a non-significant decline of 30% for Snipe was recorded over 12 years. Concentrations of these species were found in County Tyrone, but Counties Antrim, Down and Armagh supported few breeding pairs of any species. Very few pairs of any species were recorded on improved grassland despite its widespread availability.

Conclusion A successful conservation strategy for these species must address the wider countryside and not just key sites. Intensive pastoral farming in upland and lowland areas and activities such as drainage and peat extraction will further reduce the suitability of open habitats for these wader species.     

Brood amalgamation in the Bristle-thighed Curlew Numenius tahitiensis: process and function

Alloparental care in birds generally involves nonbreeding adults that help at nests or breeding adults that help raise young in communal nests. A less often reported form involves the amalgamation of broods, where one or more adults care for young that are not their own. We observed this phenomenon among Bristle-thighed Curlew Numenius tahitiensis broods in western Alaska during 1990–1992. Amalgamation of broods generally involved the formation of temporary and extended associations. Temporary associations were formed by the incidental convergence of broods soon after they left their nests. During this period, parents defended distinct brood-rearing areas, were antagonistic to conspecifics and remained together for less than 3 days. Extended associations formed when chicks were 1–2 weeks old. Here, parents and their broods occupied distinct, but adjacent, brood-rearing areas and moved around as a unit. Whether a brood participated in either temporary or extended associations or remained solitary appeared to depend on brood density in the immediate area and on hatching date. When chicks were 3–4 weeks old, aggregations of up to ten broods formed wherein young mixed and parents defended a common brood-rearing area. All broods (n = 48) that survived to fledging joined such aggregations. Alloparental care involved only antipredator defence and was not associated with activities such as feeding and brooding. Most female parents abandoned their broods shortly after the young could fly and when aggregations were forming. The female parent of a pair always deserted its young before or on the same day as the male parent and, in every aggregation, one or two males continued to tend young for about 5 days longer than other male parents. In most cases, adults deserted the young 2–6 days before the young departed the area when about 38 days old. Bristle-thighed Curlews also formed temporary associations with American and Pacific Golden Plover Pluvialis dominica and Pluvialis fulva, Whimbrel Numenius phaeopus, Bar-tailed Godwit Limosa lapponica, Western Sandpiper Cal-idris mauri and Long-tailed Skua Stercorarius longicaudus. Curlews and other larger bodied species commonly attack-mobbed predators together, whereas smaller bodied species generally gave alarm calls and circled the predators. For all species, the intensity of antipredator defence by attending adults gradually decreased as young became older and aggregations formed. We suggest that amalgamation of broods among Bristle-thighed Curlew enhances predator defence, aids in the process of flock formation for migrating young, and allows females and some males to desert their young earlier.     

Sex roles in the monogamous Purple Sandpiper Calidris maritima in Svalbard

The Purple Sandpiper Calidris maritima, an Arctic shorebird, displays unusual sex roles during breeding. In a 5-year study in Svalbard, in the Norwegian high Arctic, data were collected on the roles of the sexes in this species. Purple Sandpipers were similar to most shorebirds in that males actively courted females, established territories and vigorously defended their territories from intruders. Both sexes shared incubation duties approximately equally over the entire 21-day incubation period. Males incubated very little initially, but increased their effort significantly during the first 11 days of incubation to over 50% of the time from days 11–21. However, Purple Sandpipers contrasted with most other shorebird species in that females discontinued their breeding efforts at hatching. In nearly all cases, broods were attended solely by the male until the chicks reached fledging age (or even longer). Nonetheless, the pair bond must be described as monogamous because neither males nor females were found to re-mate or lay a second clutch during a season. This pattern of parental care, which is found only in very few other shorebird species, is discussed in an evolutionary context.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.     

Double brooding and offspring desertion in the barn owl Tyto alba

Many bird species produce two annual broods during a single breeding season. However, not all individuals reproduce twice in the same year suggesting that double brooding is condition‐dependent. In contrast to most raptors and owls, the barn owl Tyto alba produces two annual clutches in most worldwide distributed populations. Nevertheless, the determinants of double brooding are still poorly studied. We performed such a study in a Swiss barn owl population monitored between 1990 and 2014. The annual frequency of double brooding varied from 0 to 14% for males and 0 to 59% for females. The likelihood of double brooding was higher when individuals initiated their first clutch early rather than late in the season and when males had few rather than many offspring at the first nest. Despite the reproductive benefits of double brooding (single‐ and double‐brooded individuals produced 3.97 ± 0.11 and 7.07 ± 0.24 fledglings, respectively), double brooding appears to be traded off against offspring quality because at the first nest double‐brooded males produced poorer quality offspring than single‐brooded males. This might explain why females desert their first mate to produce a second brood with another male without jeopardizing reproductive success at the first nest. Furthermore, the reproductive cycle being very long in the barn owl (120 d from start of laying to offspring independence), selection may have favoured behaviours that accelerate the initiation of a second annual brood. Accordingly, half of the double‐brooded females abandoned their young offspring to look for a new partner in order to initiate the second breeding attempt, 9.48 d earlier than when producing the second brood with the same partner. We conclude that male and female barn owls adopt different reproductive strategies. Females have more opportunities to reproduce twice in a single season than males because mothers are not strictly required during the entire rearing period in contrast to fathers. A high proportion of male floaters may also encourage females to desert their first brood to re‐nest with a new male who is free of parental care duties.     

Offspring desertion and parental care in the Whiskered Tern Chlidonias hybrida

In species with biparental care, a conflict of interest can arise if one mate tries to maximize its own reproductive success at the expense of the other's. One of the mates can desert the brood to accrue a number of benefits to enhance its own fitness, leaving parental care to the remaining parent. This study is the first to describe the desertion pattern in a tern species (Sternidae). We investigated offspring desertion in the Whiskered Tern Chlidonias hybrida, a species with semi‐precocial chicks. Offspring desertion was recorded in 52% of nests prior to fledging (n = 131 nests). Females also deserted during the post‐fledging period. Of the deserters, 97% were females. Desertions started when chicks were 5 days old and no longer required intense brooding. Desertions before fledging did not affect fledging success. Provisioning rates between pair members differed, and females supplied much less food than males. Female provisioning rate affected the chances of nest desertion significantly: daily desertion rates were lower when females supplied more food. After females had deserted, males increased their provisioning rates but compensated for the loss of female care only partly in two‐ and three‐chick broods. Only in small (one‐chick) broods was compensation full. We conclude that male and female Whiskered Terns adopt different reproductive strategies in the population studied here. Females invest much less in parental care than males, providing less food and deserting more frequently. Given the ready availability of food and low predation pressure, benefits appear to accrue to females that desert; selection forces may therefore not be acting against female desertion.     

Factors affecting female parental investment in the monogamous goby, Valenciennea longipinnis

Female Valenciennea longipinnis construct a conspicuous rubble mound on a burrow after spawning while the paired male tends eggs in the burrow until hatching occurs. The mound has a function of promoting water-exchange in the burrow through hydrodynamic effects, contributing to prevention of the male egg-desertion by reducing his parental costs. Although higher mounds are more effective in water exchange, they cost females much work after spawning. In this study, I investigated effects of six ecological and environmental factors on the mound height, i.e., female parental investment. Multiple regression analysis indicated that only female body size could explain the female parental investment: larger females tended to construct higher mounds. The size-assortative mound building suggests that the females strive to construct mounds as high as they can irrespective of the other ecological and environmental factors. Because current strength and oozing of underground water fluctuated even in a day, females may be obliged to construct high mounds on the basis of the worst condition.     

Brood desertion in Kentish plover: the value of parental care

To understand the evolution of parental care, one needs to estimatethe payoffs from providing care for the offspring and from terminatingcare and deserting them. In this study we estimated the payofffrom care provision, and in a companion paper we analyze thepayoff from offspring desertion. In the current study we experimentallyinvestigated the influence of the number and sex of attendingparents on growth and survival of offspring in the Kentish ploverCharadrius alexandrinus, in two sites (A and B). Either the maleor the female parent was removed from some broods at hatchingof the chicks (female-only and male-only broods, respectively),whereas in control broods both parents were allowed to attendtheir young. At site A survival of the chicks was lower in uniparental(male-only and female-only) broods than in control broods, whereaswe found no difference in brood survival at site B. Brood survivaldecreased over the season. Removal of either parent did not influencethe growth of the young, although growth varied over the breeding season,and it was significantly different between the sites. Theseresults suggest that the payoff from parental care decreasesover the breeding season and that the value of parental care(i.e., the contribution of parents to the survival of theiryoung) may depend on the environment.     

A case of offspring desertion by a female chimpanzee and the behavioral changes of the abandoned offspring

In the Mahale Mountains National Park, Tanzania, a chimpanzee mother moved to a neighboring unit-group leaving her 5-year-old son behind in the original group. The mother remained in the new group for ten months and then after conceiving, returned to the original group. The mother and son were reunited again but the son died of illness two months later. The mother gave birth to a baby in the original group but eventually transferred to the neighboring group after that baby disappeared. This is the first record of an immature chimpanzee abandoned by its emigrating mother, although a similar phenomenon has been reported for gorillas and Hanuman langurs. The abandoned offspring followed an adult male of the unit-group and the male was the individual who gave most of the care to the offspring. The offspring's time spent on grooming others increased, the time spent on being groomed decreased, and the rate of aggression toward him increased after his mother's desertion. This report will occasion to reconsider the previous notion of all-powerful mother-offspring bond in chimpanzees.     

Sexual differences in reproductive effort: time-activity budgets of monogamous killdeer,Charadrius vociferus

By measuring the time allocated to various reproductive and somatic activities it was possible to look at differences in the strategies of the sexes for killdeer, a monogamous species. Killdeer time-activity budgets supported the general hypothesis that males and females maximize fitness in different ways. Males allocated more time than females to mating activities: mate attraction and territorial defence. However, the parental roles of the sexes were not those predicted for a monogamous species; males allocated more time than females to parental activities throughout a nesting attempt. Males appeared to be less time-constrained, and better able to perform parental activities. Females allocated more time to foraging than did males. It is suggested that the amount of time females allocated to parental activities is limited by the high energy demands of egg production. Further, high nest-failure rates may select for females maintaining good condition for re-nesting. The costs of desertion for killdeer were considerable. Adult removal experiments supported the hypothesis that two parents were necessary for successful reproduction. No individuals managed to raise offspring alone successfully. Lone males, however, were able to hatch chicks while females were not. The need for continuous parental attentiveness may results in selection for biparental care for killdeer.