On the migration of the Whimbrel,Numenius phaeopus,in Turkey

Abstract

The Whimbrel (Numenius phaeopus) is a scarce but regular passage visitor in Turkey. Spring migration has its peak in April, autumn migration in August/September (medians are April 23th and August 29th). Detailed information on flock size, wintering, main resting sites etc. is given and compared with other Middle East countries.     

Adaptive variation in the foraging behaviour of Grey Plover Pluvialis squatarola and Whimbrel Numenius phaeopus

Between-site variation in the diet and foraging behaviour of a fixed-method forager, the Grey Plover Pluvialis squatarola , and a versatile forager, the Whimbrel Numenius phaeopus , was examined and compared at nine tropical and south temperate sites. Grey Plover always foraged in a run-stop-search manner but changed the emphasis of components of this behaviour in response to prey type. Whimbrel foraged tactilely or visually and readily changed foraging speed and habitat. Both species ate a variety of prey species depending on their availability. The diet of Grey Plover was dominated by small prey, polychaetes and crabs, and Whimbrel ate mostly crabs, with Whimbrel being the more specialized in prey choice. The broader diversity of prey types consumed by Grey Plover was attributed to limitations imposed by obligate visual foraging, whereas the flexible foraging behaviour of Whimbrel allowed the latter to concentrate their efforts on the most profitable prey. Grey Plover appeared to have a density upper limit, determined by their stereotyped foraging behaviour. Whimbrel densities varied greatly in response to prey type and foraging method. Foraging effort could not be predicted from measurements of instantaneous daytime energy intake rates, and reasons for the lack of this relationship are discussed.     

Nesting densities,productivity and survival of breeding Whimbrel Numenius phaeopus in Shetland

Between 1986 and 1988 data on the nesting densities, productivity, and survival of breeding Whimbrel Numenius phaeopus were collected from 5 study sites in the Northern Isles of Shetland. Nesting densities on sites varied between 11 and 21 pairs per 100 ha. Few studies elsewhere have recorded such high densities of nesting Whimbrel as those on certain sites used in the present study. Each breeding pair produced an average of between 0.75 and 0.91 fledglings per year. Productivity varied considerably between different study sites however, primarily because of differences in the survival rate of chicks between hatching and fledging (ranging from 3% to 55% in one year). The annual survival rate of breeding adults was at least 89%. Considering adult survival in relation to overall productivity, it appears likely that the production of fledglings from study sites is in excess of that required to balance adult mortality. This result is consistent with the current increase of the Whimbrel population in Shetland.     

Predation risk and resource availability explain roost locations of Whimbrel Numenius phaeopus in a tropical mangrove delta

Roosts are important sites for shorebirds in non‐breeding areas at night and during high tides. How the spatial configuration of food and risk of predation and disturbance influence roost site use in tropical locations remains poorly known. We analysed the locations of nocturnal roosts of Whimbrel Numenius phaeopus in mangroves of Sanquianga National Park, Colombia, with respect to variation in spatial variables related to food resources and risk of predation and disturbance. We contrasted characteristics of all 13 known nocturnal roost locations with those of all other mangrove islands (n = 209) within the limits of the park. We estimated the distance from roosts and other mangrove islands to foraging sites, and sources of predators and human disturbance. Larger areas of feeding habitat surrounded nocturnal roosts than other mangrove islands, and the average distance to individual feeding patches was shorter. Roosts were also more isolated than other islands, but proximity to sources of human disturbance did not differ. We conclude that Whimbrel roost site use in Sanquianga was best explained by a combination of access to feeding territories and isolation from potential sources of mainland predators, but not by avoidance of human disturbance. Beyond identifying factors influencing roost site selection, the large aggregations of individuals in single locations may suggest that presence of conspecifics itself also plays a role in the formation of Whimbrel roosts. We highlight the interaction of food and risk landscapes with intraspecific attraction on the roost site selection by Whimbrels and the importance of mangroves as roosting sites in tropical regions.     

Relationships between egg size, chick size at hatching, and chick survival in the Whimbrel Numenius phaeopus

The relationships between egg size, chick size at hatching and chick survival in Whimbrels Numenius phaeopus were studied over a three-year period in the Shetland Isles. Three measurements of chick size at hatching were all positively correlated with egg volume, though the relationship was strongest with hatchling body-weight. In two of the three years the proportion of chicks from a brood which survived to fledging increased significantly with the mean hatching weight of chicks in the brood. Within broods, a significant effect of hatching weight on survival was detectable only up to 7 days after hatching. Between years the egg volumes and hatchling weights of individual female Whimbrels showed relatively little variability, indicating that these attributes could be controlled to a large extent by inheritance.     

Territorial aggression and hormones during the non-breeding season in a tropical bird

The hormonal control of territorial aggression in male and female vertebrates outside the breeding season is still unresolved. Most vertebrates have regressed gonads when not breeding and do not secrete high levels of sex steroids. However, recent studies implicate estrogens in the regulation of non-breeding territoriality in some bird species. One possible source of steroids during the non-breeding season could be the adrenal glands that are known to produce sex steroid precursors such as dehydroepiandrosterone (DHEA). We studied tropical, year-round territorial spotted antbirds (Hylophylax n. naevioides) and asked (1). whether both males and females are aggressive in the non-breeding season and (2). whether DHEA is detectable in the plasma at that time. We conducted simulated territorial intrusions (STIs) with live decoys to male and female free-living spotted antbirds in central Panama. Non-breeding males and females displayed robust aggressive responses to STIs, and responded more intensely to decoys of their own sex. In both sexes, plasma DHEA concentrations were detectable and higher than levels of testosterone (T) and 17beta-estradiol (E(2)). In males, plasma DHEA concentrations were positively correlated with STI duration. Next, we conducted STIs in captive non-breeding birds. Captive males and females displayed robust aggressive behavior. Plasma DHEA concentrations were detectable in both sexes, whereas T was non-detectable (E(2) was not measured). Plasma DHEA concentrations of males were positively correlated with aggressive vocalizations and appeared to increase with longer STI durations. We conclude that male and female spotted antbirds can produce DHEA during the non-breeding season and DHEA may serve as a precursor of sex steroids for the regulation of year-round territorial behavior in both sexes.     

Sex promotes spatial and dietary segregation in a migratory shorebird during the non-breeding season

Several expressions of sexual segregation have been described in animals, especially in those exhibiting conspicuous dimorphism. Outside the breeding season, segregation has been mostly attributed to size or age-mediated dominance or to trophic niche divergence. Regardless of the recognized implications for population dynamics, the ecological causes and consequences of sexual segregation are still poorly understood. We investigate the foraging habits of a shorebird showing reversed sexual dimorphism, the black-tailed godwit Limosa limosa, during the winter season, and found extensive segregation between sexes in spatial distribution, microhabitat use and dietary composition. Males and females exhibited high site-fidelity but differed in their distributions at estuary-scale. Male godwits (shorter-billed) foraged more frequently in exposed mudflats than in patches with higher water levels, and consumed more bivalves and gastropods and fewer polychaetes than females. Females tended to be more frequently involved and to win more aggressive interactions than males. However, the number of aggressions recorded was low, suggesting that sexual dominance plays a lesser role in segregation, although its importance cannot be ruled out. Dimorphism in the feeding apparatus has been used to explain sex differences in foraging ecology and behaviour of many avian species, but few studies confirmed that morphologic characteristics drive individual differences within each sex. We found a relationship between resource use and bill size when pooling data from males and females. However, this relationship did not hold for either sex separately, suggesting that differences in foraging habits of godwits are primarily a function of sex, rather than bill size. Hence, the exact mechanisms through which this segregation operates are still unknown. The recorded differences in spatial distribution and resource use might expose male and female to distinct threats, thus affecting population dynamics through differential mortality. Therefore, population models and effective conservation strategies should increasingly take sex-specific requirements into consideration.     

Oestrogen regulates male aggression in the non-breeding season

Extensive research has focused on territorial aggression during the breeding season and the roles of circulating testosterone (T) and its conversion to 17beta-oestradiol (E2) in the brain. However, many species also defend territories in the non-breeding season, when circulating T-levels are low. The endocrine control of non-breeding territoriality is poorly understood. The male song sparrow of Washington State is highly territorial year-round, but plasma T is basal in the non-breeding season (autumn and winter). Castration has no effect on aggression in autumn, suggesting that autumnal territoriality is independent of gonadal hormones. However, non-gonadal sex steroids may regulate winter territoriality (e.g. oestrogen synthesis by brain aromatase). In this field experiment, we treated wild non-breeding male song sparrows with a specific aromatase inhibitor (fadrozole, FAD) using micro-osmotic pumps. FAD greatly reduced several aggressive behaviours. The effects of FAD were reversed by E2 replacement. Treatment did not affect body condition or plasma corticosterone, suggesting that all subjects were healthy These data indicate that E2 regulates male aggression in the non-breeding season and challenge the common belief that aggression in the non-breeding season is independent of sex steroids. More generally, these results raise fundamental questions about how sexual and/or aggressive behaviours are maintained in a variety of model vertebrate species despite low circulating levels of sex steroids or despite castration. Such non-classical endocrine mechanisms may be common among vertebrates and play an important role in the regulation of behaviour.     

Territorial aggression does not feed back on testosterone in a multiple-brooded songbird species with breeding and non-breeding season territoriality,the European stonechat

Testosterone mediates reproductive behaviours in male vertebrates. For example, breeding season territoriality depends on testosterone in many species of birds and in some, territorial interactions feed back on testosterone concentrations. However, the degree to which territorial behaviour and testosterone are associated differs even between species with seemingly similar life histories, especially between species that also defend territories outside the breeding season. Here, we investigate the link between territorial behaviour and testosterone in European stonechats. Previous studies found that territorial aggression in stonechats depends on testosterone in a breeding, but not in a non-breeding context. We investigated whether stonechats show a rise in testosterone during simulated territorial intrusions (STI) during the breeding season. Post-capture testosterone concentrations of males caught after an STI were not higher than those of males caught in a control situation regardless of breeding stage. However, most of the males would have been able to mount a testosterone response because the same individuals that did not increase testosterone during the STI showed a substantial increase in testosterone after injections of gonadotropin-releasing hormone (GnRH). GnRH-induced and post-capture testosterone concentrations were positively correlated and both decreased with successive breeding stages. Further, territory owners with a short latency to attack the decoy expressed higher post-capture testosterone concentrations than males with a longer latency to attack the decoy. Thus, there is no evidence for behavioural feedback on testosterone concentrations during male-male interactions in stonechats. In combination with previous studies our data suggest that testosterone functions as an on/off switch of high intensity territorial aggression during the breeding season in stonechats. The among-species variation in the androgen control of territorial behaviour may be only partly a result of environmental differences. Instead, potential differences in how territoriality evolved in different species may have influenced whether and how a reproductive hormone such as testosterone was co-opted into the mechanistic control of territorial behaviour.     

Coal Tit fat-storing patterns during the non-breeding season: the role of residence status

According to the optimal body mass hypothesis, resident individuals, having priority of resource access and better knowledge of food availability in an area, should carry smaller fat reserves than transients whose resource predictability is lower. We tested this prediction in a free-living population of Coal Tits Parus ater in sub-alpine coniferous forest during three winters. We examined the role of residence status in determining the daily patterns of fat accumulation of individuals using time of day, temperature, season, year and body size as covariates. Only time of day, residence status and temperature significantly affected daily variation of fat score in Coal Tits. Fat scores were higher in transients than residents, although within residents, juveniles did not carry significantly more fat than adults. Our results show that fattening strategies are associated with residence status in the Coal Tit, in support of the hypothesis that transient individuals facing lower food predictability carry larger body fat reserves. The finding that within residents, patterns of fat accumulation did not differ between adults and juveniles, suggests that residence per se , involving higher food predictability, rather than age-related dominance or hoarding behaviour, determines fattening strategies in wild Coal Tits.     

Time budgets of Tibetan eared pheasants during the non-breeding season in an alpine scrub habitat

Time allocations of the group-living Tibetan eared pheasants Crossoptilon harmani with and without supplementary food were investigated by full-day sampling from winter through spring in an alpine scrub zone,south Tibet.At a flock scale,both the different food-supply flocks displayed similar daily patterns of activity,foraging in the morning and evening,and resting around midday.In terms of individual activity,either foraging or resting was highly synchronous with the flock's.Non-provisioned birds spent mor...     

Morphological and functional changes of stallion spermatozoa after cryopreservation during breeding and non-breeding season

The study compared quality and freezability of stallion semen during breeding and non-breeding seasons. Ejaculates were collected twice per week from four stallions during May (n = 24) and December (n = 24). The semen was mixed with skim milk extender, centrifuged and resuspended in fresh extender. Aliquots of this sperm suspension were separated from extender and diluted in TALP medium for sperm evaluation or with cryoextender (type "Gent" or a combination of Triladyl and skim milk). Samples of 0.5ml were cryopreserved in straws using a programmed freezer. Parameters of sperm quality were evaluated before and after freezing/thawing. These included percentages of motile spermatozoa and of morphological intact sperm. Typical injuries were demonstrated by scanning electron microscopy (S.E.M.). The acrosomal status was visualised using FITC-conjugated peanut agglutinin, and the acrosome reaction was induced by calcium ionophore A 23187. The chromatin stability was estimated by acridine orange test.In winter, the average percentages of motile and morphologically normal sperm (67 and 74.3%, respectively) were higher than during the breeding season in May (59 and 65.9%; P < 0.05). After freezing/thawing the proportions of vital and intact sperm decreased significantly. The number of motile sperm declined to 15 and 18% in May and December (range 5-40%), and of morphologically intact sperm to 51% in both seasons. Results of S.E.M. showed typical membrane ruptures in the acrosomal region and some sperm with abnormal necks. The proportion of frozen sperm with spontaneous acrosome reaction was higher during winter (86.5 versus 77.0%), suggesting a higher degree of membrane reactivity. Percentages of spermatozoa with denaturated chromatin were minimal and showed minimal differences between fresh and frozen state, stallions or seasons. An additional decondensation treatment with papain and DTE revealed a slightly enhanced number of spermatozoa with denaturable DNA after cryopreservation, especially in December (5.4 +/- 1.3%). The influence of cryoextenders was not significant for most sperm parameters, but there was a high variability between the stallions. Altogether, the influence of factors on the quality of spermatozoa has the following rank order: cryopreservation > stallion > season. Different cellular structures seem to have different susceptibilities to physicochemical stress. The cryopreservation of sperm during December results in survival rates similar to those measured during the breeding season, even more important for successful preservation is the selection of suitable semen donors.     

House Finch Carpodacus mexicanus roosting behaviour during the non-breeding season and possible effects of mycoplasmal conjunctivitis

We studied House Finch Carpodacus mexicanus roosting behaviour during the non-breeding season using radiotracking and direct observations in Ithaca, NY, USA. In contrast to what has been reported in the literature and in contrast to what should be expected from Newton's European studies of cardueline finches, House Finches roost in small groups (mean 3.5; maximum 11) and do not display at roost sites. Seventy-seven per cent of the birds re-used the same tree on successive observations. In winter, birds re-used the same roost tree more often than in the autumn, and birds with mycoplasmal conjunctivitis tended to move more between roost trees than did birds without conjunctivitis. A small number of radiotagged birds that roosted in the same tree were observed together in the daytime more often than by chance, suggesting the existence of social bonds between birds (some same sex) during the non-breeding season. In the autumn the birds often roosted in leafed deciduous trees and closer to their daytime feeding locations than they did in winter. In winter all birds roosted in evergreen trees. It is possible that the reliable and predictable food sources at feeding sites offered by the public might have changed House Finch roosting behaviour.     

The induction of oestrus in ewes during the non-breeding season using pre-used CIDRs and oestradiol-17β treatment

The fertility obtained in sheep after the use of intravaginal progesterone devices is related to the content of progesterone of the device. The hypothesis of this study was that the reproductive response of anoestrous ewes to the ram-effect could be improved by the administration of oestradiol-17β in conjunction with CIDRs treatment—using previously used CIDRs in a 5-day progestagen priming. Therefore, the objective was to determine if oestradiol-17β treatment increases fertility of anoestrous ewes primed with used CIDRs and stimulated by the ram-effect. The hypothesis was tested with CIDRs that had been previously used for 12 or 18 days. The trial was performed during the non-breeding season using 158 Corriedale ewes. Ewes had been isolated from rams since Day −35 (Day 0 = introduction of the rams). A CIDR (0.3 g progesterone, InterAg, Hamilton, New Zealand) was inserted on Day −5 in all ewes with CIDR that had been previously used for 12 days (n = 62) or 18 days (n = 96). Also on Day −5, 29 and 53 ewes that had received CIDRs of 12 or 18 days, respectively, received an intra-muscular treatment of 50 μg of oestradiol-17β (E groups). The ewes that did not receive the oestradiol-17β treatment remained as the control group (C group). Overall the treatment groups were thus: C12 (n = 33), C18 (n = 43), E12 (n = 29), and E18 (n = 53). On Day 0 all CIDRs were withdrawn, and ewes were placed with 18 rams and 20 ewes hormonally induced to exhibit oestrus. Sexual receptivity of ewes treated with CIDRs was estimated from marks on the rumps of the ewes daily from Day 0 to Day 5, and the pregnancy status diagnosed with transrectal ultrasonography on Day 40. The percentage of ewes exhibiting oestrus and pregnancy rates were lower in ewes synchronized with previously used CIDRs for 18 days, compared to those used for 12 days. The responses of ewes in oestrus were 39.4, 14.0, 65.5, and 32.1% for the C12, C18, E12, and E18 groups respectively, with pregnancy rates of 30.3, 14.0, 34.5, and 17.0%. Administration of oestradiol-17β increased the frequency of oestrous response in ewes that were treated with CIDRs previously used for 12 days (P < 0.05), but not in those treated with CIDRs used for 18 days. It could be concluded that the administration of oestradiol-17β only improved the percentage of ewes responding to oestrus when CIDRs previously used for 12 days were used for 5 days before the introduction of rams. No positive effect on fertility was observed irrespective of the period during which CIDR had been previously used.     

Sex differences in cortisol secretion after administration of an ACTH analogue in sheep during the breeding and non-breeding season

The aim of this study was to compare the response of cortisol in sheep of different sex and gonadal status to adrenal cortex stimulation by an ACTH analogue in the breeding and non-breeding season. Twenty-four adult Corriedale sheep were used in the non-breeding season, and 19 in the breeding season. Three weeks prior to the first trial (non-breeding season), six rams and six ewes were gonadectomised. In each trial, blood was obtained every 15min for 9h and the animals received 0.5mg of ACTH (Tetracosactid, Synacthen Depot i.m., after 1.5h of sampling. Sampling began at 10:00a.m. in the non-breeding season and at 9:00a.m. in the breeding season. Three main effects (sex, gonadal status and season) were evaluated, each with two levels (male and female, intact and gonadectomised, breeding and non-breeding season, respectively). In both seasons, the females showed higher cortisol levels after ACTH than males (P<0.001), though the difference seemed less marked in the non-breeding season. The cortisol response in the ewes was not affected by season. The rams, however, showed a lower response in the breeding season (P<0.03). Gonadectomy reduced the response in the ewes (P<0.001) but had no effect in the rams. Nevertheless, gonadectomy also eliminated the differences between the ewes and the rams, such that the intact rams had lower levels of cortisol compared to the intact females, with those of the gonadectomised animals of both sexes being intermediate between the gonad-intact groups. The results of this study confirm sex differences in ACTH induced cortisol secretion in intact sheep in vivo. Furthermore, by applying exogenous ACTH we have directly stimulated the adrenal cortex, indicating the existence of sex differences also at this level. The circulating gonadal steroids, which are responsible at least in part for the sex differences in the responses to stress, may influence cortisol secretion from the adrenal gland by direct action at the cortex.