Effects of flipper bands and injected transponders on the survival of adult Little Penguins Eudyptula minor

Tagging is essential for many types of ecological and behavioural studies, and it is generally assumed that it does not affect the fitness of the individuals being examined. However, the tagging of birds has been shown to have negative effects on some aspects of their lives. Here we investigate the influence of tagging on apparent survival. We examined the effects of flipper bands and injected transponders on the apparent survival of adult Little Penguins by comparing the survival probabilities of 2483 Little Penguins marked at Phillip Island, Australia, between 1995 and 2001 in one of three ways: with bands, with transponders or with both. The design of the study and our method of analysis allowed us to estimate tag loss and ensured that tag loss did not bias the survival estimates. Birds marked with flipper bands had lower survival probabilities than those marked with transponders (with apparent survival probabilities in the first year after tagging of 75% for banded birds and 80% for birds fitted with transponders, and in subsequent years of 87% for banded birds and 91% for birds fitted with transponders). We estimated both band and transponder loss probabilities for the first time, and found that transponder loss probabilities were substantially higher than band loss probabilities, particularly in the first year after marking when the tag loss probability was 5% for transponders and 0.7% for bands. Survival probabilities were lower in the first year after marking than in subsequent years for all birds. Studies of penguins that have used flipper bands to identify individuals may have underestimated annual adult survival probabilities, as banded penguins were likely to have lower than average survival probabilities than those of unbanded birds. The higher annual survival probabilities of individuals marked with transponders indicate that this should be the preferred marking technique for Little Penguins. However, future studies will, like ours, need to consider the higher rates of transponder loss when estimating survival, possibly by double‐tagging some birds.     

Flipper bands do not affect foraging‐trip duration of Magellanic Penguins

ABSTRACT Flipper bands are used to mark penguins because leg bands can injure their legs. However, concerns remain over the possible effects of flipper bands on penguins. We examined the effects of stainless‐steel flipper bands on the duration of foraging trips by Magellanic Penguins (Spheniscus magellanicus) at Punta Tombo, Argentina, using an automated detection system. We predicted that, if bands were costly and increased drag, flipper‐banded penguins would make longer foraging trips than those with small or no external markings. We tagged 121 penguins with radio‐frequency identification (RFID) tags and an additional external mark. We placed either a stainless‐steel band on the left flipper (N= 62) or a 2×10‐mm small‐animal ear tag in the outside web of the left foot (N= 59). We measured foraging‐trip durations (N= 376 trips) for 68 adult penguins with chicks from 15 December 2007 to 28 February 2008. Contrary to predictions, trip duration was similar for banded and web‐tagged penguins (P= 0.22) and for males and females (P= 0.52), with no interaction between tag type and sex (P= 0.52). No penguins marked in the 2007 breeding season and recaptured between 30 September and 30 November 2008 (N= 113) lost flipper bands or web tags, but three RFID tags failed between March and September 2008. Properly designed and applied flipper bands were a reliable marking method for Magellanic Penguins, had a lower failure rate than RFIDs, and did not affect foraging‐trip duration.     

Effects of double bands on Magellanic Penguins

ABSTRACT Banding penguins is controversial because bands can alter the survival, reproduction, and behavior of marked individuals. The effects of bands are not consistent among band types and, although stainless steel is thought to be better than other materials, tests of the long‐term impact of bands on tag‐loss rates and the reproduction and survival of individuals are needed. We tested three types of external tags on Magellanic Penguins (Spheniscus magellanicus) to measure band effects and tag‐loss rates. In 1993, we double‐tagged 300 penguins with aluminum flipper bands, stainless‐steel flipper bands, or small (2 mm × 10 mm) metal tags attached to foot webbing. We searched for double‐tagged birds for 13 of 15 yrs (1994–2008). Aluminum bands deformed, caused feather wear, injured and killed some penguins, and were lost more often than stainless‐steel bands or web tags. During the first 2 yrs of our study, at least nine penguins lost one aluminum band (N= 71 penguins resighted), but no penguins lost a stainless‐steel band (N= 84) or a web tag (N= 88). During the next 13 yrs, five penguins lost one of their two web tags (N= 89), but none lost a stainless‐steel band (N= 84). Females laid eggs of similar size before they carried a band and in the year following tagging (P= 0.09). The type of tags a female carried did not significantly change egg size (P > 0.22). During the first breeding season after tagging, penguins with aluminum bands had lower reproductive success than penguins with stainless‐steel bands or web‐tags (P= 0.04). The annual survival of females with two stainless‐steel bands was lower (0.79) than that of males with two stainless‐steel bands or males and females with two web‐tags (0.87). Aluminum bands injured Magellanic Penguins, were lost at high rates, and should not be used. Double stainless‐steel bands had no apparent effects on adult male Magellanic Penguins, but reduced survival rates of adult females. A single stainless‐steel band would likely have less impact than two bands, and our results suggest that the impact of a single band would be difficult to measure.     

Flipper Bands Modify the Short-Term Diving Behavior of Little Penguins

ABSTRACT Flipper banding has long been the primary method to identify individual penguins, despite studies indicating that it may be detrimental to breeding success and survival. Our objectives were to measure the effects that flipper bands may have on diving performance of little penguins to determine whether the bands may be detrimental. We studied short- and long-term direct effects of flipper banding on diving behavior of free-ranging little penguins (Eudyptula minor) by comparing diving behavior before and after banding and by comparing diving performance of unbanded birds to those that had carried flipper bands for several years, respectively. Recently banded birds displayed increases in multiple variables following banding. Long-term banded penguins did not exhibit differences to their unbanded counterparts in most variables examined. Our findings are useful to those considering or reviewing the use of bands in penguin study and management.     

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     

The starvation–predation risk trade-off, body mass and population status in the Common Starling Sturnus vulgaris

It is theoretically and empirically well established that body mass variation in small birds reflects a trade-off between starvation risk and predation risk. This occurs because carrying increased fat reserves reduces starvation risk but also results in a higher predation risk due to reduced escape flight performance and/or the increased foraging exposure needed to maintain a higher body mass. In principle, therefore, the theory of mass-dependent predation risk could be used to understand how a bird perceives and responds to the risks in its environment, because its mass will reflect the predictability of foraging opportunities and predation risk. Mass in birds may then provide a relatively straightforward way of assessing the foraging environment of birds and so the potential conservation problems a species faces. This study tests, for the first time for any species, how body mass changes in response to changing starvation risk, changing predation risk and changing population status. Common Starling Sturnus vulgaris mass varies as predicted by starvation–predation risk trade-off theory: mass is lower when foraging conditions are more favourable and when predation risk is increased. The populations that are declining the most strongly have higher mass, which is most likely indicative of a poor foraging environment, leading to lower relative survival. The results suggest that increased mass in Starlings, and possibly in other species, may provide an indication of the poor quality of the foraging environment and/or rapidly declining populations.     

Social learning in birds and its role in shaping a foraging niche

We briefly review the literature on social learning in birds, concluding that strong evidence exists mainly for predator recognition, song, mate choice and foraging. The mechanism of local enhancement may be more important than imitation for birds learning to forage, but the former mechanism may be sufficient for faithful transmission depending on the ecological circumstances. To date, most insights have been gained from birds in captivity. We present a study of social learning of foraging in two passerine birds in the wild, where we cross-fostered eggs between nests of blue tits, Cyanistes caeruleus and great tits, Parus major. Early learning causes a shift in the foraging sites used by the tits in the direction of the foster species. The shift in foraging niches was consistent across seasons, as showed by an analysis of prey items, and the effect lasted for life. The fact that young birds learn from their foster parents, and use this experience later when subsequently feeding their own offspring, suggests that foraging behaviour can be culturally transmitted over generations in the wild. It may therefore have both ecological and evolutionary consequences, some of which are discussed.     

长距离迁徙鸟类对应于能量积累状态的取食行为调整

迁徙鸟类能够预计到迁徙过程中对能量需求的增加和迁徙途中获得能量的不确定性。最佳迁徙理论指出：迁徙停留期的一系列决策受到体内能量状态、取食机会和迁徙时间的影响。利用刚完成春季跨越墨西哥湾迁徙的鸫类,我们研究了取食行为、体内能量状态和能量积累速度的相互关系。我们用雾网捕获了停留的鸫,然后给每只鸟进行了环志和称重,并估测了表皮下积累的脂肪。为了检测体内能量积累大小对取食行为的影响,我们把在野外观察到的鸟分成肥、瘦两组。当一天里被捕获鸟的平均体重低于相关种的瘦体重时,这一天被观察到的取食鸟被归到瘦组,反之就属于肥组。我们同时对一部分鸟用有色环进行了标记,以便能在野外观察到它们时能准确地知道每一只鸟的初始能量积累状态。在春天完成跨越墨西哥湾迁徙以后的鸫大约有50％在到达停留地时就已经消耗了所有的表皮下脂肪。与肥组鸟相比,瘦组的个体在停留期扩展了取食方法和取食基底,增加了取食速度。与此相关的是,瘦鸟表现出体重积累更多、速度更快。我们的数据表明迁徙鸟在到达迁徙中途停留地时的能量状态会影响到它们停留期间的取食行为和继续迁徙时的生理机能,从而影响停留期的长短和在迁徙途中停留与否的决定[动物学报51(1)：12—23,2005]。     

Foraging methods can affect patch choice: an experimental study in Mallard (Anas platyrhynchos)

Animals can adapt to changes in feeding conditions by switching between foraging methods. Dabbling ducks use different foraging methods, including dabbling in deep water with the head and neck submerged, and grubbing in the mud (or shallow water) where the eyes are above the surface, so the bird can visually monitor its environment while foraging. Deep foraging is considered to provide lower intake rates and to have high associated costs, such as predation risk, compared to shallow foraging. Ducks should thus prefer shallow foraging and switch to deeper methods when feeding conditions deteriorate. We conducted a set of experiments with Mallard to assess the importance of intake rate as a cue to choose between patches associated with different foraging methods, and evaluate the influence of food depletion on the decision to switch between methods. When 50 g of wheat were presented in two patches, one at a depth of 5 cm and one at 35 cm, most of the foraging was in the shallow area. Reducing food abundance to 10 g in the shallow area led to an increase in deep foraging, although the birds still preferred the shallow area at the beginning of the tests despite the fact that it did not provide a higher intake rate. This area was used until complete depletion, and birds did not turn to deep foraging before ensuring that the shallow patch was empty. These results show that food depletion affects the choice between feeding patches hence foraging method. However the value of intake rate is not the main cue for decision, rather the birds appear to choose between patches with different methods on account of their respective costs.     

Ecology and sociality in a multilevel society: ecological determinants of spatial cohesion in hamadryas baboons

The multilevel society of hamadryas baboons, consisting of troops, bands, clans, and one-male units (OMUs), is commonly perceived to be an effective means of adapting to variable food availability while allowing spatial cohesion in response to predator pressure. The relationship between these variables, however, has never been tested quantitatively. The Filoha site in Awash National Park, Ethiopia is ideally suited to such an investigation as it contains nutrient-dense palm forests in addition to the Acacia scrublands typical of hamadryas distribution elsewhere, allowing comparisons of spatial cohesion across habitat types. Here, we use observations over a 1-year period to examine the relationship between resource availability, perceived predator pressure, and spatial cohesion in a band of wild hamadryas baboons at Filoha. Our results demonstrate that the band was more likely to break into OMUs when foraging in habitats with lower food availability, and that the band fissioned into independent clans more often when preferred resources were not available. Furthermore, the baboons remained in larger aggregations for longer periods of time (i.e., prior to embarking on their daily foraging route) on mornings after predators were heard in the vicinity, and increased cohesion in response to encounters with people who may have been perceived as predators. These results support the notion that hamadryas baboons change their social groupings in response to both food availability and predation risk and that the ability of hamadryas bands to cleave and coalesce in response to changes in these factors underlies the evolution of the hamadryas modular social structure.     

Wintering birds avoid warm sunshine: predation and the costs of foraging in sunlight

Wintering birds can gain significant thermal benefits by foraging in direct sunlight. However, exposure to bright sunlight might make birds easier to detect by predators and may also cause visual glare that can reduce a bird’s ability to monitor the environment. Thus, birds likely experience a trade-off between the thermal benefits and predation-related costs of foraging in direct sunlight. To examine this possible thermoregulation-predation trade-off, we monitored the behavior of mixed-species flocks of wintering emberizid sparrows foraging in alternating strips of sunlight and shade. On average, these sparrows routinely preferred to forage in the shade, despite midday air temperatures as much as 30 °C below their thermoneutral zone. This preference for shade was strongest at relatively high temperatures when the thermal benefits of foraging in sunlight were reduced, suggesting a thermoregulation-predation trade-off. Glare could be reduced if birds faced away from the sun while feeding in direct sunlight, but we found that foraging birds tended to face southward (the direction of the sun). We speculate that other factors, such as the likely direction of predator approach, may explain this southerly orientation, particularly if predators use solar glare to their advantage during an attack. This interpretation is supported by the fact that birds had the weakest southerly orientation on cloudy days. Wintering birds may generally avoid foraging in direct sunlight to minimize their risk of predation. However, given the thermal benefits of sunshine, such birds may benefit from foraging in habitats that provide a mosaic of sunlit and shaded microhabitats.     

Bird mixed‐species flock formation is driven by low temperatures between and within seasons in a Subtropical Andean‐foothill forest

According to both the predation avoidance and foraging efficiency hypotheses, birds within mixed flocks increase their foraging efficiency and/or can spend more time feeding and less time looking out for predators. These hypotheses predict that birds in mixed flocks obtain benefits. Thus, mixed flock formation could serve as a strategy to cope with difficult conditions imposed on birds such as climatic conditions that ultimately result in a change in predation pressure or food resources. We evaluate the hypotheses that forming part of a flock confers benefits to its members and the associated prediction that birds will take advantage of these benefits and flock more often under cold and dry weather conditions between and within seasons to cope with such conditions. We surveyed the presence of mixed flocks, flocking propensity, number of species and individuals in mixed flocks in the Subtropical Yungas foothill of Argentina, to examine seasonality, flocking behavior of birds and their responses to two climatic variables: temperature and humidity. Bird species presented a higher flocking propensity and mixed flocks occurred more frequently during the dry and cold seasons than during the more benign seasons, and lower values of temperature within seasons triggered the flocking behavior. Although effects between seasons were expected, birds also showed a short‐term response to small changes in temperature within seasons. These results strengthen the ideas proposed by the foraging hypothesis. Although benefits derived from flocking have yet to be determined, whatever they are should be understood in the context of seasonal variation in life‐history traits.     

The social organization of non-breeding Magpies Pica pica

The social behaviour of non-breeding individuals in a colour-marked population of Magpies was studied. In early autumn most non-breeders began to forage in a common area, the 'Non-breeding Flock Area'. A few individuals remained on their natal territories away from the flock area as solitary non-breeders. Most non-breeders were first-year birds, but some were second years or adults which no longer held a territory. Birds foraged in groups, their food intake rate differing with both group size and location. It appeared that the area in which birds foraged had a significant effect on food intake rate; foraging groups tended to form at sites rich in food. There was a dominance hierarchy amongst non-breeders; an individual's foraging behaviour, survivorship and chances of breeding were status dependent, with subordinates feeding less in groups and being less likely to survive and breed. Solitary individuals' chances of breeding were similar to those of high status birds, although their survivorship to breeding age may have been lower. Non-breeding Magpies are compared with non-breeders of other species and the factors which may influence their social behaviour are discussed. It is suggested that remaining as a solitary non-breeder is a viable alternative to becoming a low-status flock member for some birds.     

GPS tracking devices reveal foraging strategies of Black-legged Kittiwakes

The Black-legged Kittiwake Rissa tridactyla is the most abundant gull species in the world, but some populations have declined in recent years, apparently due to food shortage. Kittiwakes are surface feeders and thus can compensate for low food availability only by increasing their foraging range and/or devoting more time to foraging. The species is widely studied in many respects, but long-distance foraging and the limitations of conventional radio telemetry have kept its foraging behavior largely out of view. The development of Global Positioning System (GPS) loggers is advancing rapidly. With devices as small as 8 g now available, it is possible to use this technology for tracking relatively small species of oceanic birds like kittiwakes. Here we present the first results of GPS telemetry applied to Black-legged Kittiwakes in 2007 in the North Pacific. All but one individual foraged in the neritic zone north of the island. Three birds performed foraging trips only close to the colony (within 13 km), while six birds had foraging ranges averaging about 40 km. The maximum foraging range was 59 km, and the maximum distance traveled was 165 km. Maximum trip duration was 17 h (mean 8 h). An apparently bimodal distribution of foraging ranges affords new insight on the variable foraging behaviour of Black-legged Kittiwakes. Our successful deployment of GPS loggers on kittiwakes holds much promise for telemetry studies on many other bird species of similar size and provides an incentive for applying this new approach in future studies.     

Do hunters target auxiliary markers? An example using black brant

Auxiliary markers play an essential role in understanding migration, movement, demography, and behavior of migratory birds. Use of such markers relies on the assumption that the markers do not affect the traits of interest. Neck collars, among the most conspicuous of markers, substantially affect risk of harvest, and survival even in the absence of harvest. Effects of less-conspicuous markers, such as colored plastic tarsal bands, are not well understood. We used 30 years (1986–2015) of banding, recovery, and recapture data from the Yukon-Kuskokwim Delta in Alaska, USA, to assess differences in direct band recovery rates (DRRs) between black plastic and brightly colored plastic bands applied to black brant (Branta bernicla nigricans). We also assessed the effect of the color of plastic tarsal bands on annual survival, risks of natural mortality harvest, and fidelity to the breeding colony of adult female black brant. When assessing only DRRs we found that brightly colored bands were recovered at higher rates than black plastic bands in the early 2000s, but DRRs for black bands increased more rapidly through time, resulting in similar DRRs for the 2 band colors at the end of the study. Using a Burnham model structure, our results demonstrated that individuals fitted with colored bands had slightly lower hazards of dying from natural causes or hunting than individuals carrying less-conspicuous black tarsal bands. Differences on annual probability scales were small and credible intervals broadly overlapped between band types, indicating minimal differences between individuals with different band types; however, we could not resolve all confounding in our study design and we suggest that specific studies directed at assessing marker effects are warranted. We encourage education of hunters about their roles as citizen scientists and the potentially detrimental effect of targeting birds with auxiliary markers.     

Ultimate and proximate factors affecting the breeding performance of a marine top-predator

Variability in ecosystems affects the life history of organisms. In marine ecosystems where interannual variability is high, relationships between fluctuations in oceanographic parameters and top-predator breeding performance are increasingly documented but it is less clear why such relationships exist. In this study, we examined the connections between marine environment fluctuations and breeding performance of a long-lived top-predator, the black-browed albatross Diomedea melanophris at Kerguelen, through study of resource acquisition and allocation processes. Our results show that this population used the same foraging zones and spent similar time foraging year after year, but adult body condition varied between years. Foraging trips are regulated mainly by changes in body condition. During years of low resource availability, birds return to their nest with lower body condition and adults in low body condition were more frequent and therefore were more likely to stop breeding. Poor breeding success was related to the presence of colder waters in the foraging zones of breeding albatrosses as measured by the positive correlation between sea surface temperatures and breeding success measured over 18 years. Lower breeding success was mainly due to failure by inexperienced birds. The results of this study demonstrate how oceanographic conditions affect breeding performance through allocation processes. We compared these results to those at South Georgia where the breeding success is lower and more variable. This population relies mainly on krill, a resource that shows a very variable year-to-year availability compared to fish prey consumed by Kerguelen birds. This study shows that, in the same species, differences in resource variability and availability affect the demographic strategies probably through differences in allocation strategies.     

Mallards Feed Longer to Maintain Intake Rate under Competition on a Natural Food Distribution

Animals foraging in groups may benefit from a faster detection of food and predators, but competition by conspecifics may reduce intake rate. Competition may also alter the foraging behaviour of individuals, which can be influenced by dominance status and the way food is distributed over the environment. Many studies measuring the effects of competition and dominance status have been conducted on a uniform or highly clumped food distribution, while in reality prey distributions are often in‐between these two extremes. The few studies that used a more natural food distribution only detected subtle effects of interference and dominance. We therefore conducted an experiment on a natural food distribution with focal mallards Anas platyrhynchos foraging alone and in a group of three, having a dominant, intermediate or subordinate dominance status. In this way, the foraging behaviour of the same individual in different treatments could be compared, and the effect of dominance was tested independently of individual identity. The experiment was balanced using a 4 × 4 Latin square design, with four focal and six non‐focal birds. Individuals in a group achieved a similar intake rate (i.e. number of consumed seeds divided by trial length) as when foraging alone, because of an increase in the proportion of time feeding (albeit not significant for subordinate birds). Patch residence time and the number of different patches visited did not differ when birds were foraging alone or in a group. Besides some agonistic interactions, no differences in foraging behaviour between dominant, intermediate and subordinate birds were measured in group trials. Possibly group‐foraging birds increased their feeding time because there was less need for vigilance or because they increased foraging intensity to compensate for competition. This study underlines that a higher competitor density does not necessarily lead to a lower intake rate, irrespective of dominance status.     

The effect of social dominance on foraging by the Three-toed Woodpecker Picoides tridactylus

During the winter, female Three-toed Woodpeckers Picoides tridactylus , when unaccompanied by the male, foraged at a lower height above the ground and on tree trunks of greater diameter than when foraging together with a male. Among males, however, no such differences were found between those birds foraging in the absence of, or in company with, a female. The niche breadth of the males was less than that of the females, indicating a higher degree of specialization by the males. The niche overlap between the sexes was less when the birds foraged together in pairs, especially as regards foraging height and foraging site diameter. The intersexual segregation of foraging niche recorded for the Three-toed Woodpecker is therefore probably not determined genetically, but is due to social dominance by males which obliges the females to occupy the less preferred niche. Both sexes spent less time being vigilant when foraging together, than when alone.     

Experimental increase of flying costs in a pelagic seabird: effects on foraging strategies,nutritional state and chick condition

A central point in life history theory is that parental investment in current reproduction should be balanced by the costs in terms of residual reproductive value. Long-lived seabirds are considered fixed investors, that is, parents fix a specific level of investment in their current reproduction independent to the breeding requirements. We tested this hypothesis analysing the consequences of an experimental increase in flying costs on the foraging ecology, body condition and chick condition in Cory’s shearwaters Calonectris diomedea. We treated 28 pairs by reducing the wing surface in one partner and compared them with 14 control pairs. We monitored mass changes and incubation shifts and tracked 19 foraging trips per group using geolocators. Furthermore, we took blood samples at laying, hatching and chick-rearing to analyse the nutritional condition, haematology, muscle damage and stable isotopes. Eighty-day-old chicks were measured, blood sampled and challenged with PHA immune assay. In addition, we analysed the effects of handicap on the adults at the subsequent breeding season. During incubation, handicapped birds showed a greater foraging effort than control birds, as indicated by greater foraging distances and longer periods of foraging, covering larger areas. Eighty-day-old chicks reared by treated pairs were smaller and lighter and showed a lower immunity than those reared by control pairs. However, oxygen demands, nutritional condition and stable isotopes did not differ between control and handicapped birds. Although handicapped birds had to increase their foraging effort, they maintained physical condition by reducing parental investment and transferred the experimentally increased costs to their partners and the chick. This result supports the fixed investment hypothesis and is consistent with life history theory.