Influence of growth rate retardation on time budgets and energetics of Arctic Tern Sterna paradisaea and Common Tern S. hirundo chicks

Time budgets of free-living chicks of Arctic Terns Sterna paradisaea and Common Terns S. hirundo throughout development are presented with special reference to changes in time allocation when growth rate varies. Chicks of both species were inactive most of the time observed (87%). Time allocated to the different behaviours changed during development and was generally better correlated with body mass than age. Slower growing nestlings were brooded more and allocated more time to quiescence and less time to locomotion, preening, begging and attacking (the latter two significant only for the Arctic Tern). The energetic implications of variation in time budgets with age and growth rate were considered. Parental brooding resulted in an average energy saving of nearly 40% of an individual nestling's thermoregulatory costs. Whereas thermoregulatory costs remained nearly unchanged in Arctic Tern chicks, these were negatively correlated with growth rate in Common Terns. Tentatively, we estimated a 30% reduction in a nestling's total energy requirement for a 50% reduction in average growth rate for both species.     

Effects of egg size, parental quality and hatch-date on growth and survival of Common Tern Sterna hirundo chicks

We examined the relative contributions of egg size, parental quality and hatch-date to growth and survival of second-hatched chicks (those chicks making the greatest contribution to differences in productivity among pairs) by exchanging clutches among nests of Common Terns Sterna hirundo matched for lay-date (range 13 May to 9 June). The mass of a second-laid egg in an exchanged clutch ranged from 17.70 to 23.80 g. Growth and survival were studied during three periods: early (days 0–3), middle (days 3–12) and late (days 12–25). Both egg mass and hatch-date were important predictors of hatchling mass (positive relationships), although there was no seasonal trend in egg mass. During the middle period, hatch-date was a significant predictor of mass gain and survival (inverse relationships). After controlling for hatch-date, other indices of parental quality made only small contributions to chick mass gain and survival. Our results suggest that although breeding early generally leads to greater overall survival of chicks, several important interactions among egg 'quality', parental quality and early laying may affect breeding success under specific conditions.     

The Function of Embryonic Vocalization in the Little Tern (Sterna albifrons)

Various functional explanations can be proposed for the evolution of bird embryonic vocalizations during the pre-hatching period, namely: 1. To elicit switching of parents from incubation to parental behaviour typical of the chick period; 2. To allow thermoregulation of embryos by soliciting parents to incubate; or 3. To establish parent—offspring individual recognition. In this paper, we present the results of field experiments designed to test hypotheses 1 and 3 in the colonial, ground-nesting little tern. Parents that had their hatching eggs cross-fostered with foreign eggs at the same hatching stage exhibited a parental behaviour similar to unmanipulated controls. Parents that incubated foreign eggs up to a stage in which embryos were not yet vocalizing, and were challenged with their own hatching eggs that had been incubated in foster nests, performed less efficient parental cares than unmanipulated control pairs and pairs that had cross-fostered hatching eggs. The results do not support the hypothesis of early individual recognition and suggest that embryonic vocalizations in little terns have the function of promoting switching of parents from incubation to accepting and feeding hatchlings. Similar to other tern and gull species, the duration of incubation period in the little tern varies markedly among pairs and years. In these species, embryonic vocalizations can be adaptive since they provide parents a cue to switch at a proper time from incubation to parental cares typical of the chick period.     

Colony size and not nest density drives reproductive output in the Common Tern Sterna hirundo

Density is known to be an important factor in population size regulation. Several mechanisms of density limitation have been identified in colonial birds. We studied competition in Common Terns Sterna hirundo to assess whether the factor limiting reproductive output was competition for nest‐sites, which is dependent on local nest density, or density‐dependent competition for food resources, which is dependent on overall colony size using the same foraging area. We found strong associations of both colony size and nest density with reproductive output in five colonies of Common Terns in three different habitats (one marine, two freshwater). Based on detailed long‐term datasets of six separate sub‐colonies of the Banter See colony that differed in nest density, we found that reproductive success was not related to nest density but to overall colony size, possibly a result of resource depletion and food competition. We also found carry‐over effects of colony size during rearing on post‐fledging return rate. These results have important implications for the conservation management plans aimed at recovering declining populations of Common Terns.     

Feeding ecology of the Common Tern Sterna hirundo in a wintering area in southern Brazil

The wintering diet of Common Terns Sterna hirundo was studied by using 714 pellets collected on roosting sites at the mouth of the Lagoa dos Patos and on adjacent coastal beaches, in Rio Grande do Sul, southern Brazil, from March 1999 to February 2000. A total of 12 340 individual prey items of 35 different food types was found. Fish was the most important food type in the diet, constituting 32% by number and 93% by mass. Insects contributed 67% by number but only 3% by mass. The main food types were sciaenid fishes Paralonchurus brasiliensis, Micropogonias furnieri, Cynoscion guatucupa and Macrodon ancylodon. Several of these are important commercial species; fisheries potentially impact food availability to the terns, and terns may contribute significantly to the natural mortality of these fishes. Clupeiform fishes, the urophycid fish Urophycis brasiliensis and flying ants (Camponotus sp.) were also important. Species composition of the diet (food types), both by number and by mass, differed significantly between months. Prey sizes ranged in length from 12.7 mm to 217.4 mm. The average estimated total length of fish taken was 77.7 mm, but the mean differed significantly among prey species. The importance of demersal sciaenids to the diet of the Common Tern, a surface predator, may be explained by their association with aquatic predators, especially adult Bluefish Pomatomus saltatrix and Striped Weakfish Cynoscion guatucupa, and the Franciscana Dolphin Pontoporia blainvillei, which drive these fish to the surface. The occurrence of flying ants in the diet was related to offshore winds, which carried these insects out to sea. The occasional high availability of insects possibly changed the cost/benefit relationship of several food types, causing diet changes. The high number of prey species, the temporal variations in the composition of the diet and the wide range of prey sizes are evidence of the high dietary plasticity of the Common Tern, at wintering areas in southern Brazil.     

Variation in the growth and survival of Golden Plover Pluvialis apricaria chicks

The growth of Golden Plover Pluvialis apricaria chicks is modelled in detail for the first time. The pattern of growth is typical of postnatal development in waders, although the mean fledging time of 37 days is slower than would be expected from adult body weight. Bill length and weight at hatching had significant effects on the rate of weight gain shown by 2-day-old chicks, although this effect was not noticeable at 4 days of age. Chick survival was significantly affected by bill length, as a result of the variation in weight gain. The growth of older chicks was positively correlated with mean minimum temperature. The slow rate of growth exhibited by Golden Plover chicks is discussed in relation to breeding habitat and the effects of weather and hatchling biometrics.     

Offspring sex ratio in the Common Tern Sterna hirundo, a species with negligible sexual size dimorphism

In many vertebrates, male offspring are affected more than female offspring by adverse conditions during growth, resulting in facultative adjustment of offspring sex ratio by parents in response to social and environmental conditions during breeding. The greater vulnerability of male offspring is generally attributed to their higher energy requirements associated with their larger size, although greater sensitivity to adverse conditions could be related to other factors such as negative effects of androgens on male physiology. To control for sexual differences in body size, we examined variation in offspring sex ratio in the Common Tern Sterna hirundo , a species with negligible sexual size dimorphism. In this species, the last-laid egg (termed the c-egg) is smaller than the first two and hatches last, so that the chick obtains relatively little food and hence has a low probability of survival to fledging. This species thus provides a powerful model for examining sex-linked mortality and sex ratio variation under natural conditions. We found that the sex ratio of c-eggs, but not of earlier laid eggs, was significantly biased in favour of females. Chicks hatched from c-eggs (termed c-chicks) had low survival but female c-chicks had significantly higher survival than male c-chicks. These data provide strong evidence that factors other than sexual size dimorphism are responsible for producing greater vulnerability of male offspring to adverse conditions during growth.     

Effects of food availability on fledging condition and post-fledging survival in king penguin chicks

Effects of summer food shortage on king penguin Aptenodytes patagonicus chicks were studied at South Georgia. Two cohorts were compared, fledging in the austral summers of 1992 (n = 32) and 1994 (n = 33) when availability of food was judged good and poor, respectively. The former cohort had a higher pre-fledging mean mass (12.78 kg vs ≤ 10.03 kg), fledged earlier (median 5 January vs 21 January), and a higher proportion was re-sighted within 2 years of fledging (28% vs 0%). Within 4 years, 47% of the former cohort had been re-sighted (i.e. post-fledge survival); in addition, one was observed at the Falkland Islands, and 22% had bred (i.e. recruitment) in their colony of origin. The re-sighted chicks of the 1992 cohort fledged earlier than those not re-sighted (median 24 December vs 10 January), but it remain unclear if they were heavier at fledging. All chicks in this study (n = 65) were marked with both transponders (subcutaneously implanted) and flipper bands (on one flipper), and no losses of any markings were found (controlled up to 4 years afterwards). Therefore, data on chick post-fledging survival and recruitment were not adjusted for losses of markings, as has been done in other studies. Received: 21 October 1996 / Accepted: 2 February 1997     

红嘴巨燕鸥云南鸟类种的新记录

在最近整理标本的过程中,作者发现1981年4月13日采自云南昆明滇池草海的一只红嘴巨燕鸥(Sterna caspia),为云南鸥科(Laridae)鸟类种的新记录。据约翰.马敬能等(2000)记录,红嘴巨燕鸥(Sterna caspiaPallas,1770,Caspian Tern)中文名为红嘴巨鸥。作者考虑该种鸟类隶属于鸥科的燕鸥属(Sterna),故建议将中文名修订为“红嘴巨燕鸥”。鉴别特征:体形较大,体重610g,体长470mm,嘴峰长60mm,翅长340mm,尾长160mm,跗长40mm。前额、头顶至后枕白色,布有黑色纵纹,枕部和眼下的纵纹较为密集而粗著,眼先和额部的纵纹较纤细而稀疏。背部和两翅表面银灰…     

Latitudinal trends, population size and habitat preferences of the Damara Tern Sterna balaenarum on Namibia's desert coast

The Damara Tern Sterna balaenarum is a rare, near-endemic breeder along the southwestern coast of Africa. Found predominantly in Namibia, its world population was once believed to be 4000 birds. Population surveys in the northern Namib Desert have recently estimated that 12,000 adults exist. The aims of this study were to (1) determine the population status along the southern coast, thereby completing Namibian surveys, (2) assess latitudinal trends (over 10̀) in relation to ocean upwellings for the entire coast and (3) determine breeding habitat preferences. This was completed in 1994 and 1996 using random sampling techniques. Survey squares of 1 km², randomly plotted on 1:50,000 maps, were located in the desert with a Global Positioning System (GPS) unit and searched for breeding terns. A total of 1780 km² was sampled in the southern desert and extrapolations indicated that about 1450 adult terns occur in this region. This gives a global population estimate of 13,500 adult Damara Terns. Significant habitat preferences were apparent with terns nesting most often on gravel plains in the north but switching to salt pans in the south. Lower salt pan temperatures in the south may account for these trends. The peak density was in central areas (23|dGS) with a decreasing density north and south along Namibia's 1470 km-coast. This is parallel to recent findings for linear shorebird densities and macro-invertebrate density. This suggests that primary productivity of the Benguela is highest in the central regions and not where the upwellings occur 400 km south. We propose a delayed blooming effect to account for these trends and conclude that this diminutive tern is influenced in many aspects of its life history by Namibia's cold offshore upwellings.     

Consequences of a female-biased sex-ratio in a socially monogamous bird: female-female pairs in the Roseate Tern Sterna dougallii

In the socially monogamous gulls and terns, female-biased sex ratios are sometimes revealed by the occurrence of ‘supernormal clutches’, which are usually attended by female-female pairs or other multi-female associations. We studied these phenomena in the endangered Roseate Tern Sterna dougallii at Bird Island, USA, from 1970 to 1995. DNA-techniques were used to sex breeding adults in 1992–94. Supernormal clutches (with three or four eggs) have comprised 1–7% of all Roseate Tern clutches at Bird Island since at least 1970, probably increasing in frequency since 1980. Supernormal clutches were spatially clustered; most were laid late in the peak period of nesting during each season. More than 80% of supernormal clutches and at least 7% of normal clutches were attended by multi-female associations; most of these were female-female pairs, with a few trios (male + two females, or three females) and one quartet (four females). More than half of the multi-female associations attended normal clutches. Some female-female pairs were maintained for up to five years. The age-distribution of females mated to females did not differ significantly from that of females mated to males. Females mated together usually laid eggs synchronously (±2 days). Such females laid fewer eggs than females mated to males (means 1.20 versus 1.73), and had lower fertility and hatching success (about 46% versus 98%); they were less successful in raising young from eggs that did hatch (means 58% versus 73%), but this difference was not significant. Their overall breeding success was much lower (about 0.34 fledglings per female versus 1.35). The sex-ratio of breeders was about 127 females to 100 males; about 20% of breeding females did not have male mates. Female Roseate Terns that do not obtain male mates appear to be of low phenotypic ‘quality’ - based on late laying, small clutches and small eggs. Our data support the hypothesis that such females have a higher fitness if they mate with each other and raise a few young than if they do not breed at all.     

Abundance and breeding of the Antarctic Tern Sterna vittata at the James Ross and Seymour Islands, NE Antarctic Peninsula

The Antarctic Peninsula (AP) is experiencing rapid environmental change associated with warming and sea ice retreat, which is likely to affect locally breeding birds. Yet, contrary to the knowledge of bird biology along the maritime West coasts of the AP, there is a remarkable lack of data from the more continental East coast. We report on the distribution, abundance, and breeding of the Antarctic Tern at the James Ross and Seymour islands, the two largest snow-free areas in the NE part of the AP, where this species breeds under harsh climate conditions probably close to its limits. Terns were found breeding in most ice-free areas, with nests located up to 2.9 km from coastlines at altitudes up to 180 m a.s.l. While the large-scale density was relatively low (c. 450 pairs per 127 km² of surveyed ice-free area), the local density (total colony area: 3 nests per ha; nest clusters: 100–140 nests per ha) was as high as elsewhere. Mean clutch size (1.21, n = 196) was smaller than in the west AP or in the maritime Antarctic. Daily nest survival rate during incubation varied between years and locations (mean = 0.977; 95 % CI: 0.966–0.985). While both predation and weather-caused mortality were locally important, the impact of skua predation might be lower in areas with alternative prey (penguin colonies). We suggest that the Antarctic Tern deserves attention as a species potentially suitable for monitoring of environmental impacts upon bird populations around the AP.     

Conservation genetics of the Common Tern (Sterna hirundo) in the North Atlantic region; implications for the critically endangered population at Bermuda

Common Terns nesting at Bermuda are isolated by 1,000–4,000?km from other populations of the species around the North Atlantic Ocean. This population experienced a severe demographic bottleneck as a result of a hurricane in 2003 and was subsequently re-established by four males and four females. Using seven microsatellite loci, we compared the genetic diversity of the pre- and post-bottleneck populations, compared the genetic profile of the Bermuda population with those of other populations around the North Atlantic Ocean and mainland Europe, and assessed the potential contribution of immigration to genetic diversity. We found a loss of genetic diversity (number of alleles and heterozygosity) in the post-bottleneck Bermudian population (4.6–2.9 and 0.56–0.52, respectively). We also report significant differentiation among all sampled locations (global F_ST?=?0.16) with no evidence for immigration into Bermuda. Common Terns from the Azores were genetically more similar to those from mainland North America than to those from Bermuda or mainland Europe. Our results suggest that the critically endangered population in Bermuda is genetically distinct and requires continued and enhanced conservation priority.     

Variation in early neonatal mortality for different types of fetal growth retardation

Birth and first-48-hr death records were analyzed for 10,024 liveborn infants in Mexico City and 12,786 liveborn infants in Santa Cruz, Bolivia. The objective of the analysis was to characterize the early postnatal mortality rates for different types of fetal growth retardation and prematurity. Infants who were delivered prior to 37 weeks of gestation had 23-100 times the mortality risk of infants born at full term and normal weight. Light-for-gestational-age infants (birth weight less than 2,900 g) were further divided into proportionately growth-retarded with normal Rohrer's index (weight/height) and disproportionately growth-retarded with low Rohrer's index. The proportionately growth-retarded infant had nearly twice the mortality of the full-term, appropriate-weight infants, whereas the disproportionately growth-retarded infants had 2.9-5.7 times the mortality rate of the full-term, appropriate-weight infants. There were some differences between samples in mortality rates and prevalence of the different classes of small infants, but the pattern of mortality within samples was consistent between samples.     

Effects of clutch size and egg-laying order on the breeding success in the Little Tern Sterna albifrons on the Nakdong Estuary, Republic of Korea

Survivorship of Little Tern Sterna albifrons eggs and chicks was followed on an islet in the Nakdong Estuary, Republic of Korea, in 1995 and 1996. Mean egg size and incubation period were significantly different between the 2 years. The maximum clutch size was three eggs, and the second egg in the clutch often hatched earlier than the first, while most of the third eggs hatched last. In 1996, when the fate of 249 eggs from 106 nests was followed for 40 days, hatching success, fledging success and breeding success were 77%, 40% and 31%, respectively. High mortality occurred in the early chick stage, mostly because of rain and predation by Weasels Mustela sibirica. The breeding success per egg was 14% in one-egg clutches, 28% in two-egg clutches and 34% in three-egg clutches. This difference was mainly attributed to the lower hatching success in the smaller clutches. In three-egg clutches, the third egg showed significantly lower breeding success than siblings. The main foods of the Little Tern were Tridentiger obscurus, Engraulis japonicus, Hyporhamphus intermedius, Acanthogobius flavimanus (all fish), Palaemon sp. and Crangon affinis (shrimps). The feeding frequency was, apparently, not affected by time of day and age of chicks but was probably influenced by weather conditions. Newly hatched chicks failed to eat 25% of the prey brought to them, although this decreased with the age of the chicks.     

Effects of testosterone on growth, plumage pigmentation, and mortality in Black-headed Gull chicks

In the Black-headed Gull Larus ridibundus, sibling chicks defend small territories against conspecifics with testosterone-dependent aggressive behaviour. The energetic requirements for the performance of this behaviour may trade off against the energetic requirements for growth. There are indications that testosterone suppresses growth in birds and, therefore, regulate this trade-off. In this study, the effect of testosterone on growth and plumage pigmentation of Black-headed Gull chicks was analysed. Young chicks in small groups were treated for ten days with testosterone or sham treated. Testosterone-treated birds showed decreased growth rate (daily increase in body mass, head-bill length and tarsus-length) and a marked decrease in juvenile pigmentation of the plumage (tail-bar, back, and secondary coverts). Field measurements revealed a negative correlation between nest density, which correlates positively with aggressive behaviour of adults, and plumage coloration. Furthermore, these measurements showed an increase in mortality of chicks that had low levels of pigmentation early in life. The data suggest that chicks face a testosterone-regulated trade-off between growth and territory defence.     

Variation in milk production and lactation performance in grey seals and consequences for pup growth and weaning characteristics

Phocid seals are one of the few groups of mammals capable of sustaining the energetic demands of lactation entirely through body nutrient stores while fasting. Lactation performance of the female in turn influences the rate and pattern of pup growth. We examined variation in and patterns of milk composition and production, maternal energy output, and pup growth and energy deposition over the entire lactation period in 18 grey seal mother-pup pairs using hydrogen isotope (3H2O and D2O) dilution. Milk composition was independent of maternal mass and nutrient stores, indicating dependence on other physiological and genetic factors. Heavier females lactated longer (r2=0.653, P<0.001), had higher total milk outputs (r2=0.652, P<0.001), and produced larger pups at weaning (r2=0.417, P=0.005). While fatter females lactated for longer periods of time (r2=0.595, P<0.001), females with a larger lean body mass at parturition produced more milk (r2=0.579, P<0.001). Total milk energy output was the strongest predictor of pup weaning mass, which, along with the pup's efficiency of energy storage, accounted for 91% of the variation in weaning mass. Nevertheless, there was sufficient plasticity in milk composition and energy output that some smaller females produced relatively large pups. Few females appeared to deplete body nutrients to the point where it might limit the duration of lactation.     

Body condition,hormonal correlates and consequences for survival in common tern chicks

Although sibling competition in avian species has been extensively studied, the proximate mechanisms remain largely unknown. Recent research proposed that steroid hormones, in particular testosterone (T) and corticosterone (CORT), might play a role either in promoting competitive behavioral displays or in response to chronic nutritional stress accompanied by a sustained competitive situation. Here, we examine body condition, endogenous T levels and fecal glucocorticoid metabolites (FGM) as non-invasive measures of CORT in sibling broods of wild common tern chicks (Sterna hirundo) during three post-natal developmental stages. In this species, distinct within-brood size asymmetries are imposed by an asynchronous hatching interval, and sexes show slightly different growth patterns. First-hatched (a-)chicks were in better condition than their later-hatched sibling (b-chick). FGM levels inversely covaried with condition and were elevated at the end of pre-fledging development. T levels of a- and b-chicks changed with age, although the direction of the changes differed, with b-chicks eventually having higher levels than their older siblings. Survival to fledging was not associated with FGM but with T levels, which tended to be higher in surviving chicks. Our results are discussed with regard to how plasticity in steroid hormones could be involved in mediating sibling competition in common terns.