Post-fledging body mass as a determinant of subadult survival in Common Terns Sterna hirundo

Once a bird has fledged it becomes hardly accessible for researchers and consequently knowledge about post-fledging ontogeny is scarce. In this study on juvenile Common Terns (Sterna hirundo) we used an automated transponder-based detection and weighing system at Banter See colony, Northern Germany, which enabled us to investigate body mass growth of post-fledglings and its consequences for their survival until first return to the natal colony when 2 years old. We analysed data from two contrasting breeding seasons, 2000 and 2001, in order to determine inter-year and inter-sex variation of post-fledging parameters assumed to potentially affect subadult survival, such as the period a juvenile is still present at colony surroundings (departure age), its fledging mass and last recorded post-fledging body mass, and hatch date. Using an information-theoretic model selection approach, neither the date of hatching nor the departure age was found to affect survival. The only predictor of survival was last post-fledging body mass whereas fledging mass itself was of minor importance. Although there was weak evidence for an interaction with year, individuals of the cohort 2000, which left the colony area on average 5 g lighter than those reared under the more favourable conditions in 2001, did not exhibit lower return probability. We suggest that under unfavourable conditions selection had eliminated weak individuals prior to fledging or during the post-fledging period. This study underlines the importance of the post-fledging period and its consequences for survival, especially in species with prolonged parental care post-fledging.     

Spatial Heterogeneity as a Genetic Mixing Mechanism in Highly Philopatric Colonial Seabirds

How genetic diversity is maintained in philopatric colonial systems remains unclear, and understanding the dynamic balance of philopatry and dispersal at all spatial scales is essential to the study of the evolution of coloniality. In the King penguin, Aptenodytes patagonicus, return rates of post-fledging chicks to their natal sub-colony are remarkably high. Empirical studies have shown that adults return year after year to their previous breeding territories within a radius of a few meters. Yet, little reliable data are available on intra- and inter-colonial dispersal in this species. Here, we present the first fine-scale study of the genetic structure in a king penguin colony in the Crozet Archipelago. Samples were collected from individual chicks and analysed at 8 microsatellite loci. Precise geolocation data of hatching sites and selective pressures associated with habitat features were recorded for all sampling locations. We found that despite strong natal and breeding site fidelity, king penguins retain a high degree of panmixia and genetic diversity. Yet, genetic structure appears markedly heterogeneous across the colony, with higher-than-expected inbreeding levels, and local inbreeding and relatedness hotspots that overlap predicted higher-quality nesting locations. This points towards heterogeneous population structure at the sub-colony level, in which fine-scale environmental features drive local philopatric behaviour, while lower-quality patches may act as genetic mixing mechanisms at the colony level. These findings show how a lack of global genetic structuring can emerge from small-scale heterogeneity in ecological parameters, as opposed to the classical model of homogeneous dispersal. Our results also emphasize the importance of sampling design for estimation of population parameters in colonial seabirds, as at high spatial resolution, basic genetic features are shown to be location-dependent. Finally, this study stresses the importance of understanding intra-colonial dispersal and genetic mixing mechanisms in order to better estimate species-wide gene flows and population dynamics.     

The post-fledging survival of young Guillemots Uria aalge in relation to hatching date and growth

The post-fledging survival of a total of 1277 young Guillemots Uria aalge ringed in 6 years was assessed using sightings of 267 individuals back at the natal colony and recoveries of 46 ringed birds. In two years there was a significant decline in survival prospects with estimated hatching date, the first time such a trend has been demonstrated in the Alcidae. In these years pairs breeding early had a 2–3 times greater chance of having a young survive to return to the colony than pairs breeding three weeks later. Hatching date had no effect on survival prospects in the other four seasons and in none of the years did chick body condition have a demonstrable effect on post-fledging survival.     

Effect of fog on the arrival time of little penguins Eudyptula minor: a clue for visual orientation?

We report a dramatic change in the arrival time of little penguins Eudyptula minor at their breeding colony when a heavy low-level fog was present on the southern beaches of Phillip Island, Australia. On the evening of 21 December 2002, only 5% of penguins arrived at the expected time, and the peak time at arrival that night was 2 h behind the peak arrival times of previous and subsequent days. It appears that little penguins were unable or unwilling to return to the colony under the low visibility caused by the fog. Only half (52%) of the average number of penguins returned to the colony on the evening of the fog. We propose two explanations for our observation. First, penguins could have been close to the colony but did not come ashore in conditions of poor visibility to avoid predation. Secondly, the extensive fog may have affected the orientation of the penguins and they were unable to find their way back home. Our results could serve as indirect evidence for both hypotheses.     

The long engagement of the emperor penguin

In birds, courtship is generally short relative to the whole breeding cycle. Emperor penguins (Aptenodytes forsteri), however, are an exception as their courtship period is much longer (ca. 6 weeks) than the courtship of other penguin species. This strategy may appear surprising, as it is especially costly to fast and endure drastic climatic conditions for long periods at the colony (1.5 and up to 4 months for females and males, respectively). We examined here the reasons of this extended courtship period and found that emperor penguins returned earlier to the colony when primary oceanic production before breeding was high. This suggests that emperor penguins return to the colony as soon as primary oceanic production in summer allows them to replenish their body reserves. The extended period of time spent at the colony during courtship may therefore result from an evolutionary process that confers advantages to emperor penguins that arrive earlier at the colony by reducing predation risks and offering better chances of securing a partner.     

Breeding site fidelity and natal philopatry in the Redshank Tringa totanus

This paper presents the results of a study carried out on breeding Redshank in the Ribble Marshes, Lancashire, England.
Redshank tend to return to the same breeding area year after year. There was no detectable sex bias in return rates. Experienced birds were more site faithful than inexperienced birds, with previously successful birds exhibiting the highest degree of breeding site fidelity. Older, more experienced birds were more successful at hatching eggs than inexperienced birds.
Breeding dispersal was the same both within and between years. Faithful pairs and males nesting with a new mate dispersed significantly shorter distances than females nesting with a new partner. Dispersal distances in female Redshank were affected by breeding success: unsuccessful females, nesting with a new mate, dispersed significantly farther than successful females. A pair's breeding success influenced the following year's mate fidelity. However, other factors such as overwintering survival and date of return may also have influenced mate fidelity.
Redshank were highly faithful to their natal area; a high proportion of birds that survived post-fledging mortality returned to breed in their area of birth. No sex bias in natal dispersal was detected. Approximately 50% of Redshank breed in their first year of life.     

Post-fledging body mass increase in Common Terns Sterna hirundo: influence of age, sex and year

We studied the inter-year and inter-sex variation of the post-fledging body mass development of Common Terns Sterna hirundo in 2000 and 2001 at the Banter See colony, northern Germany. Here, post-fledglings can be identified and weighed remotely and automatically by a transponder system that makes use of automated balances installed at the colony. Individuals were sexed with PCR amplification methods. After fledging, young generally continued to increase their mass. However, in 2000, the young did not significantly increase their mass during the post-fledging period. In 2001, conditions were more favourable and body mass increased continuously. Further, in 2001, male post-fledglings were significantly heavier than female post-fledglings. Once having left the colony area (on average 18–23 days after fledging in 2000, and 14–16 days after fledging in 2001), post-fledgling body mass had still not reached adult body mass. The longer a juvenile stayed at the colony, the higher was its final body mass, which if acting as a threshold level may control departure time. Neither brood size nor hatching order affected post-fledging mass or period. In the unfavourable year 2000, when many individuals were found dead after fledging, fledging age but not fledging mass was found to be a predictor of post-fledging survival before departure: individuals fledging when older had a lower survival probability. Our results stress the importance of the post-fledging period for body mass increase and survival prior to departure. The variation in post-fledging mass growth between years and between the sexes is discussed with respect to parental effort and a possible selective provisioning of sons over daughters.     

Estimating the critical body mass of king penguins

King penguins (Aptenodytes patagonicus) can fast for over a month. However, they return to sea to forage before their body mass reaches a critical value (cMb), beyond which there is an increase in rate of mass loss and in protein catabolism, termed phase III of fasting. Thus when studying king penguins onshore, accurate estimation of their cMb and, in turn, the date at which that body mass would be reached, will be informative to behavioural and physiological data being collected. For penguins being studied during fasts in captivity, knowing cMb is particularly important because of the need to release the birds back into their colony while they are still in good nutritional condition. The present study investigates the validity of using measures of beak, flipper and foot length together to estimate cMb in king penguins and provides a simple and effective prediction equation for researchers. The three morphometric measurements, along with body mass just prior to going to sea after the moult fast (taken to represent cMb), were obtained for nine king penguins in a colony at the Crozet Archipelago. A multiple linear regression of the three morphometric measurements against cMb provided an R ² of 71.2%. Mean absolute percentage error of the estimate of cMb over the nine birds was 8.82 ± 1.20%. The described technique could probably be employed for estimating cMb in other long-fasting seabirds.     

Hints on Research for Bird-watchers

Capsule Fledgling Golden Eagles in northern Sweden preferred clearcuts and other open forest habitats, as well as steep slopes.

Aims To study the post-fledging habitat use and ranging behaviour of juvenile Golden Eagles on their natal territories.

Methods Fourteen juvenile Golden Eagles in northern Sweden were marked with GPS transmitters and tracked until they left their natal territory.

Results Eagles fledged at the end of July–beginning of August and remained on their natal territories until October–early November. Fledged eagles' home range size before flying south was on average 41?km². Juvenile eagles showed a preference for clearcuts, coniferous forest on lichen-covered bedrock and edges between clearcuts and forest, whilst all other habitat types were used less than expected. The eagles showed a preference for steep slopes, in particular south-facing ones, whilst north-facing slopes were used less than expected.

Conclusion Golden Eagles' preference for clearcuts and steep slopes can be used in the planning and management of ‘eagle friendly’ wind farms.     

Responses of king penguin Aptenodytes patagonicus adults and chicks to two food‐related odours

Increasing evidence suggests that penguins are sensitive to dimethyl sulphide (DMS), a scented airborne compound that a variety of marine animals use to find productive areas of the ocean where prey is likely to be found. Here we present data showing that king penguins Aptenodytes patagonicus are also sensitive to DMS. We deployed DMS on a lake near a king penguin colony at Ratmanoff beach in the Kerguelen archipelago. We also presented DMS to ‘sleeping’ adults on the beach. On the lake, penguins responded to the DMS deployments by swimming more, while on the beach, penguins twitched their heads and woke up more for the DMS than for the control presentations. Interestingly, penguins did not respond to cod liver oil deployments on the lake; mirroring at‐sea studies of other penguins. Although at‐sea studies are needed to confirm that king penguins use DMS as a surface cue that informs them of productivity under the water, this study is an important first step in understanding how these birds locate prey over significant distances.     

High juvenile annual survival probabilities in Southern Rockhopper Penguins Eudyptes chrysocome are independent of individual fledging traits

Juvenile survival is an important demographic parameter. Southern Rockhopper Penguins Eudyptes chrysocome have undergone a dramatic population decline in the past century across their distribution, but the demographic processes are poorly understood. To estimate juvenile annual survival probabilities, Rockhopper Penguin chicks from two cohorts on New Island, Falkland Islands, were marked with transponders and recorded in subsequent years using an automated gateway. We first estimated annual survival and detection probabilities using a Cormack‐Jolly‐Seber (CJS) model, and found that both probabilities were extremely high (81% in the first and 98% in the second, third and fourth years of life), even in comparison with adult birds. Because detection probability after 3 years was effectively 1, and our sample size (n = 114) was too small to explore the effects of individual traits on survival in a CJS model, we assessed whether sex, cohort, body mass and laying sequence affected whether juveniles returned to the colony during their first 3 years of life using a simple generalized linear model that assumed perfect detection. Juveniles from the first cohort and males showed a higher return probability than juveniles from the second cohort and females. There was no clear effect of fledging body mass on return rate, probably related to the favourable environmental conditions during the study period. The laying sequence did not markedly affect the return probability of chicks, indicating that, once fledged, first‐laid A‐chicks have the same probability to return as second‐laid B‐chicks despite a much larger initial maternal investment in B‐eggs in this species. This study demonstrates extraordinarily high juvenile survival probabilities and will help to understand the recent changes in the population dynamics of the Falkland Islands Southern Rockhopper Penguins.     

Do fledgling and pre‐breeding Common Swifts Apus apus take part in aerial roosting? An answer from a radiotracking experiment

This study has shown that fledgling Common Swifts Apus apus spend the first post-fledging night of their life on the wing and that pre-breeders also spend the full night on the wing. Even though this work was conducted during an unusually cold, wet period, the results show that fledglings do not return to their natal colony in the week after fledging. It also demonstrates that yearlings are only slightly more likely than fledglings to spend any time at a particular colony, but are more likely to move from colony to colony. Older pre-breeders are more likely to spend more time at and revisit a particular colony more often than yearlings. It is our observation that only right at the end of the breeding duties will the parents participate in an evening ascent, and even then many of them return to their nest for the evening. Breeding adults displayed the greatest devotion to a particular colony. But even among such adults we detected some that ceased caring for their young a few days prior to their fledging. Adults roost in their nestbox if they meet with bad weather.     

At‐sea distribution and habitat use in king penguins at sub‐Antarctic Marion Island

King penguins make up the bulk of avian biomass on a number of sub‐Antarctic islands where they have a large functional effect on terrestrial and marine ecosystems. The same applies at Marion Island where a substantial proportion of the world population breeds. In spite of their obvious ecological importance, the at‐sea distribution and behavior of this population has until recently remained entirely unknown. In addressing this information deficiency, we deployed satellite‐linked tracking instruments on 15 adult king penguins over 2 years, April 2008 and 2013, to study their post‐guard foraging distribution and habitat preferences. Uniquely among adult king penguins, individuals by and large headed out against the prevailing Antarctic Circumpolar Current, foraging to the west and southwest of the island. On average, individuals ventured a maximum distance of 1,600 km from the colony, with three individuals foraging close to, or beyond, 3,500 km west of the colony. Birds were mostly foraging south of the Antarctic Polar Front and north of the southern boundary of the Antarctic Circumpolar Current. Habitat preferences were assessed using boosted regression tree models which indicated sea surface temperate, depth, and chorophyll a concentration to be the most important predictors of habitat selection. Interestingly, king penguins rapidly transited the eddy‐rich area to the west of Marion Island, associated with the Southwest Indian Ocean Ridge, which has been shown to be important for foraging in other marine top predators. In accordance with this, the king penguins generally avoided areas with high eddy kinetic energy. The results from this first study into the behavioral ecology and at‐sea distribution of king penguins at Marion Island contribute to our broader understanding of this species.     

Early-life behaviour predicts first-year survival in a long-distance avian migrant

Early-life conditions have critical, long-lasting effects on the fate of individuals, yet early-life activity has rarely been linked to subsequent survival of animals in the wild. Using high-resolution GPS and body-acceleration data of 93 juvenile white storks (Ciconia ciconia), we examined the links between behaviour during both pre-fledging and post-fledging (fledging-to-migration) periods and subsequent first-year survival. Juvenile daily activity (based on overall dynamic body acceleration) showed repeatable between-individual variation, the juveniles'' pre- and post-fledging activity levels were correlated and both were positively associated with subsequent survival. Daily activity increased gradually throughout the post-fledging period, and the relationship between post-fledging activity and survival was stronger in individuals who increased their daily activity level faster (an interaction effect). We suggest that high activity profiles signified individuals with increased pre-migratory experience, higher individual quality and perhaps more proactive personality, which could underlie their superior survival rates. The duration of individuals’ fledging-to-migration periods had a hump-shaped relationship with survival: higher survival was associated with intermediate rather than short or long durations. Short durations reflect lower pre-migratory experience, whereas very long ones were associated with slower increases in daily activity level which possibly reflects slow behavioural development. In accordance with previous studies, heavier nestlings and those that hatched and migrated earlier had increased survival. Using extensive tracking data, our study exposed new links between early-life attributes and survival, suggesting that early activity profiles in migrating birds can explain variation in first-year survival.     

A method to assess population changes in king penguins: the use of a Geographical Information System to estimate area-population relationships

During the last decades, king penguin (Aptenodytes patagonicus) populations have been reported to increase throughout most of their breeding range. In this study, we compared the results obtained from direct counts of incubating king penguins with the results yielded by the estimation of the change in area occupied by breeding birds at the Ratmanoff king penguin colony at the Kerguelen Islands. The area of the colony was determined using a Geographical Information System with a georeferencing extension on aerial pictures taken in 1963, 1985 and 1998. Individual king penguin were counted on the same pictures or pictures taken on the same day. The overall population increase between 1963 and 1998 was 733% while the colony area increased by 677%. This study indicates that monitoring change in colony size is a good indicator for detecting and monitoring large population changes in king penguins, in particular for remote colonies. The discrepancy between the two results may be from two different kinds of bias. Firstly, there could be a possible error in the estimation of the area occupied by the colony resulting from the georeferencing of oblique pictures, and secondly, the density of king penguins may also change with population number. This method, which only requires high-altitude pictures, also reduces the possible disturbance to breeding made by low- to medium-altitude flights. Accepted: 7 February 2000     

Post-Fledging Dispersal of King Penguins (Aptenodytes patagonicus) from Two Breeding Sites in the South Atlantic

Most studies concerning the foraging ecology of marine vertebrates are limited to breeding adults, although other life history stages might comprise half the total population. For penguins, little is known about juvenile dispersal, a period when individuals may be susceptible to increased mortality given their naïve foraging behaviour. Therefore, we used satellite telemetry to study king penguin fledglings (n = 18) from two sites in the Southwest Atlantic in December 2007. The two sites differed with respect to climate and proximity to the Antarctic Polar Front (APF), a key oceanographic feature generally thought to be important for king penguin foraging success. Accordingly, birds from both sites foraged predominantly in the vicinity of the APF. Eight king penguins were tracked for periods greater than 120 days; seven of these (three from the Falkland Islands and four from South Georgia) migrated into the Pacific. Only one bird from the Falkland Islands moved into the Indian Ocean, visiting the northern limit of the winter pack-ice. Three others from the Falkland Islands migrated to the eastern coast of Tierra del Fuego before travelling south. Derived tracking parameters describing their migratory behaviour showed no significant differences between sites. Nevertheless, generalized linear habitat modelling revealed that juveniles from the Falkland Islands spent more time in comparatively shallow waters with low sea surface temperature, sea surface height and chlorophyll variability. Birds from South Georgia spent more time in deeper waters with low sea surface temperature and sea surface height, but high concentrations of chlorophyll. Our results indicate that inexperienced king penguins, irrespective of the location of their natal site in relation to the position of the APF, develop their foraging skills progressively over time, including specific adaptations to the environment around their prospective breeding site.     

Is Penguin banding harmless?

Thousands of penguins are banded annually world-wide, even though little is known about the potential impact of these flipper bands. In this paper, the possible effect of banding on the survivorship, breeding frequency and other ecological factors on king penguins was investigated. The extended laying period (3–4 months) of the king penguin allows the observation of non-lethal effects that could influence the time of laying and thus the reproductive success. Three hundred and eighty-three breeding king penguins on the colony of “la Baie du Marin” on Possession Island, Crozet Archipelago were either single- or double-flipper banded. The results show that the returning birds were laying late the following breeding season, and that double-banded birds lay significantly later than single-banded birds. Furthermore, our data suggest a lower return rate for double-banded birds than for single-banded birds (45% as opposed to 76%). The low return rate of single-banded birds, when compared to an interannual survivorship estimated to be 96.5%, also raises questions concerning the potential impact of single banding. Received: 2 January 1998 / Accepted: 23 May 1998     

Juvenile helping in the moorhen,Gallinula chloropus

In a 3-year study of the moorhen, Gallinula chloropus, some chicks from first broods stayed on their natal territory once they had reached independence, and helped to rear their younger, second brood, siblings. Juvenile dispersal was constrained by habitat saturation, and first brood young were forced to stay on their natal territory. Juveniles that hatched early in the year were forced to stay longer, and helped more than those that hatched late. The total feeding rates to broods with and without juvenile helpers were the same, but parents with helpers reduced their feeding rates relative to parents without helpers. Pairs with helpers (=pairs attempting second broods) reared more chicks per nesting, attempt than pairs rearing chicks at the same time of year without helpers (=pairs attempting first clutch renests). This was true both for all clutches, and for hatched clutches only, even when controlling for parental quality, territory size and scasonal effects.     

Behavioral and physiological significance of minimum resting metabolic rate in king penguins

Because fasting king penguins (Aptenodytes patagonicus) need to conserve energy, it is possible that they exhibit particularly low metabolic rates during periods of rest. We investigated the behavioral and physiological aspects of periods of minimum metabolic rate in king penguins under different circumstances. Heart rate (f(H)) measurements were recorded to estimate rate of oxygen consumption during periods of rest. Furthermore, apparent respiratory sinus arrhythmia (RSA) was calculated from the f(H) data to determine probable breathing frequency in resting penguins. The most pertinent results were that minimum f(H) achieved (over 5 min) was higher during respirometry experiments in air than during periods ashore in the field; that minimum f(H) during respirometry experiments on water was similar to that while at sea; and that RSA was apparent in many of the f(H) traces during periods of minimum f(H) and provides accurate estimates of breathing rates of king penguins resting in specific situations in the field. Inferences made from the results include that king penguins do not have the capacity to reduce their metabolism to a particularly low level on land; that they can, however, achieve surprisingly low metabolic rates at sea while resting in cold water; and that during respirometry experiments king penguins are stressed to some degree, exhibiting an elevated metabolism even when resting.     

Age-dependent prospecting and recruitment to a breeding colony of Short-tailed Shearwaters Puffinus tenuirostris

In a small colony of Short-tailed Shearwaters Puffinus tenuirostris , monitored annually since 1947, many of the young ringed as nestlings returned to their natal colony for one or more years before breeding for the first time. These 'prospectors' were first recorded at 4.1 ± 0.1 (range 2–7) years old; they started breeding at 7.0 ± 0.2 (4–14) years old. Overall, 31% of all prospectors bred on their natal island with no significant annual variation in this proportion. However, 8% of all natal recruits started breeding without having been recorded prospecting and bred first when significantly younger (5.6 ± 0.4 years old) than those first recorded prospecting. Shearwaters which subsequently bred at their natal colony started prospecting when significantly older (4.3 ± 0.1) than those that did not breed at their natal colony (4.0 ± 0.1 years). Non-natal recruits spent significantly less time prospecting (1.0 ± 0.1 years) than prospectors born on the island (1.9 ± 0.1 years). The shearwaters appeared to become progressively more faithful to the colony during their prospecting period.