Kerry J. Barton

Diet and foraging effort of Adélie penguins in relation to pack-ice conditions in the southern Ross Sea

We investigated the diet and aspects of foraging effort among Adélie penguins (Pygoscelis adeliae) breeding at three colonies on Ross Island, in the southwestern Ross Sea – Capes Royds, Bird and Crozier – during the chick-provisioning period of three austral summers, 1994–1995, 1995–1996 and 1996–1997. During the study period, pack-ice cover differed in waters offshore of these colonies, by colony, seasons and year. Diet differed among colonies only slightly. The fish Pleuragramma antarcticum was the most important prey, especially during years or periods within years when little pack ice was present. With respect to krill, which composed the remainder of diet, juvenile Euphausia crystallorophias were consumed predominantly in a year of heavy pack-ice cover; more adult krill were consumed in 2 years when pack ice was sparse. Foraging trip duration differed by colony, season and year and was related directly to distance from the colony to the nearest pack ice. The amount of food brought to chicks increased as trip duration increased, to a point (2 days), but then decreased as duration increased further (up to 4 days). On the basis of data on mass of parents and of meal sizes to chicks, it appeared that on the longest trips more of the food gathered by parents was used for self maintenance; on the longest trips, parents lost body mass. Successful foraging during chick rearing, the period when adult foraging is most intense, appears to depend on the proximity of pack ice to nesting colonies for this penguin species.

GEOGRAPHIC STRUCTURE OF ADÉLIE PENGUIN POPULATIONS: OVERLAP IN COLONY‐SPECIFIC FORAGING AREAS

In an investigation of the factors leading to geographic structuring among Adelie Penguin (Pygoscelis adeliae) populations, we studied the size and overlap of colony- specific foraging areas within an isolated cluster of colonies. The study area, in the south- western Ross Sea, included one large and three smaller colonies, ranging in size from 3900 to 135 000 nesting pairs, clustered on Ross and Beaufort Islands. We used triangulation of radio signals from transmitters attached to breeding penguins to determine foraging loca- tions and to define colony-specific foraging areas during the chick-provisioning period of four breeding seasons, 1997-2000. Colony populations (nesting pairs) were determined using aerial photography just after egg-laying; reproductive success was estimated by com- paring ground counts of chicks fledged to the number of breeding pairs apparent in aerial photos. Foraging-trip duration, meal size, and adult body mass were estimated using RFID (radio frequency identification) tags and an automated reader and weighbridge. Chick growth was assessed by weekly weighing. We related the following variables to colony size: foraging distance, area, and duration; reproductive success; chick meal size and growth rate; and seasonal variation in adult body mass. We found that penguins foraged closest to their respective colonies, particularly at the smaller colonies. However, as the season pro- gressed, foraging distance, duration, and area increased noticeably, especially at the largest colony. The foraging areas of the smaller colonies overlapped broadly, but very little foraging area overlap existed between the large colony and the smaller colonies, even though the foraging area of the large colony was well within range of the smaller colonies. Instead, the foraging areas of the smaller colonies shifted as that of the large colony grew. Colony size was not related to chick meal size, chick growth, or parental body mass. This differed from the year previous to the study, when foraging trips of the large colony were very long, parents lost mass, and chick meals were smaller. In light of existing data on prey abundance in neritic waters in Antarctica suggesting that krill are relatively evenly distributed and in high abundance in the Southern Ross Sea, we conclude that penguins depleted or changed the availability of their prey, that the degree of alteration was a function of colony size, and that the large colony affected the location (and perhaps ultimately the size) of foraging areas for the smaller colonies. It appears, therefore, that foraging dynamics play a role in the geographic structuring of colonies in this species.

SPATIAL AND TEMPORAL VARIATION OF DIET WITHIN A PRESUMED METAPOPULATION OF ADÉLIE PENGUINS

Abstract We investigated temporal and spatial variability in the diet of chick-provisioning Adélie Penguins (Pygoscelis adeliae) breeding at all colonies within one isolated cluster in the southwestern Ross Sea, Antarctica, 1994–2000. We wished to determine if prey quality explained different population growth and emigration rates among colonies. Diet composition was described both by conventional means (stomach samples) and by analysis of stable isotopes in chick tissues (toenails of individuals killed by skuas [Stercorarius maccormicki]). Diets were similar among the four study colonies compared to the disparity apparent among 14 widely spaced sites around the continent. Calorimetry indicated that fish are more energetically valuable than krill, implying that if diet varied by colony, diet quality could attract recruits and help to explain differential rates of colony growth. However, a multiple-regression analysis indicated that diet varied as a function of year, time within the year, and percent of foraging area covered by sea ice, but not by colony location. Stable isotopes revealed similarity of diet at one colony where conventional sampling was not possible. We confirmed that sea ice importantly affects diet composition of this species in neritic waters, and found that (1) quality of summer diet cannot explain different population growth rates among colonies, and (2) stable isotope analysis of chick tissues (toenails) is a useful tool to synoptically describe diet in this species over a large area. Variación Espacial y Temporal de la Dieta en una Supuesta Metapoblación de Pygoscelis adeliae Resumen. Investigamos la variabilidad temporal y espacial en la dieta de Pygoscelis adeliae que se encontraban aprovisionando a sus polluelos en todas las colonias dentro de un grupo aislado en el mar de Ross sud-occidental, Antártica, entre 1994 y 2000. Deseábamos determinar si la calidad de la presa explicaba las diferentes tasas de crecimiento poblacional y emigración entre colonias. La composición de la dieta fue descrita por medios convencionales (muestras estomacales) y por el análisis de isótopos estables en tejidos de los polluelos (uñas de las patas de los individuos matados por Stercorarius maccormicki). Las dietas fueron similares entre las 4 colonias estudiadas en relación a la disparidad de la dieta evidente entre 14 sitios dispersos a través del continente. Las análisis de calorimetría indicaron que los peces tienen un mayor valor energético que el krill, sugiriendo que si la dieta varía entre colonias, la calidad de la dieta podría atraer a reclutas y ayudaría a explicar los diferentes índices de crecimiento entre colonias. Sin embargo, un análisis de regresión múltiple indicó que la dieta varió en función del año, de la época dentro de un año, y del porcentaje de área de forrajeo cubierta por el hielo del mar, pero no en función de la localización de la colonia. Los isótopos estables revelaron la semejanza de la dieta en una colonia en que no era posible el muestreo convencional. Confirmamos que el hielo en el mar afecta de manera importante la composición de la dieta de esta especie en aguas neríticas, y encontramos que (1) la calidad de la dieta de verano no puede explicar las diferentes tasas de crecimiento poblacional entre colonias, y que (2) el análisis de isótopos estables de los tejidos de polluelos (uñas de las patas) es una herramienta útil para describir, sinópticamente, la dieta en esta especie a través de un área extensa.

Survival differences and the effect of environmental instability on breeding dispersal in an Adélie penguin meta-population

High survival and breeding philopatry was previously confirmed for the Adélie penguin (Pygoscelis adeliae) during a period of stable environmental conditions. However, movements of breeding adults as a result of an unplanned natural experiment within a four-colony meta-population provided interesting insights into this species’ population dynamics. We used multistate mark-recapture models to investigate apparent survival and dispersal of breeding birds in the southwestern Ross Sea during 12 breeding seasons (1996–2007). The natural experiment was facilitated by the temporary grounding of two immense icebergs that (i) erected a veritable fence separating colonies and altering migration routes and (ii) added additional stress by trapping extensive sea ice in the region during 5 of 12 y. Colony size varied by orders of magnitude, allowing investigation of apparent survival and dispersal rates in relation to both environmental conditions and colony size within this meta-population. Apparent survival was lowest for the smallest colony (4,000 pairs) and similar for the medium (45,000 pairs) and large colonies (155,000 pairs), despite increased foraging effort expended by breeders at the largest colony. Dispersal of breeding birds was low (<1%), except during years of difficult environmental conditions when movements increased, especially away from the smallest colony (3.5%). Decreased apparent survival at the smallest colony could reflect differences in migration chronology and winter habitat use compared with the other colonies, or it may reflect increased permanent emigration to colonies outside this meta-population. Contrary to current thought, breeding penguins are not always philopatric. Rather, stressful conditions can significantly increase dispersal rates.

Trends in the Breeding Population of Adélie Penguins in the Ross Sea, 1981–2012: A Coincidence of Climate and Resource Extraction Effects

Measurements of the size of Adélie penguin (Pygoscelis adeliae) colonies of the southern Ross Sea are among the longest biologic time series in the Antarctic. We present an assessment of recent annual variation and trends in abundance and growth rates of these colonies, adding to the published record not updated for more than two decades. High angle oblique aerial photographic surveys of colonies were acquired and penguins counted for the breeding seasons 1981–2012. In the last four years the numbers of Adélie penguins in the Ross and Beaufort Island colonies (southern Ross Sea metapopulation) reached their highest levels since aerial counts began in 1981. Results indicated that 855,625 pairs of Adélie penguins established breeding territories in the western Ross Sea, with just over a quarter (28%) of those in the southern portion, constituting a semi-isolated metapopulation (three colonies on Ross Island, one on nearby Beaufort Island). The southern population had a negative per capita growth rate of −0.019 during 1981–2000, followed by a positive per capita growth rate of 0.067 for 2001–2012. Colony growth rates for this metapopulation showed striking synchrony through time, indicating that large-scale factors influenced their annual growth. In contrast to the increased colony sizes in the southern population, the patterns of change among colonies of the northern Ross Sea were difficult to characterize. Trends were similar to southern colonies until the mid-1990s, after which the signal was lost owing to significantly reduced frequency of surveys. Both climate factors and recovery of whale populations likely played roles in the trends among southern colonies until 2000, after which depletion of another trophic competitor, the Antarctic toothfish (Dissostichus mawsoni), may explain the sharp increasing trend evident since then.

Climate Change Winners: Receding Ice Fields Facilitate Colony Expansion and Altered Dynamics in an Adélie Penguin Metapopulation

There will be winners and losers as climate change alters the habitats of polar organisms. For an Adélie penguin (Pygoscelis adeliae) colony on Beaufort Island (Beaufort), part of a cluster of colonies in the southern Ross Sea, we report a recent population increase in response to increased nesting habitat as glaciers have receded. Emigration rates of birds banded as chicks on Beaufort to colonies on nearby Ross Island decreased after 2005 as available habitat on Beaufort increased, leading to altered dynamics of the metapopulation. Using aerial photography beginning in 1958 and modern satellite imagery, we measured change in area of available nesting habitat and population size of the Beaufort colony. Population size varied with available habitat, and both increased rapidly since the 1990s. In accord with glacial retreat, summer temperatures at nearby McMurdo Station increased by ∼0.50°C per decade since the mid-1980s. Although the Ross Sea is likely to be the last ocean with an intact ecosystem, the recent retreat of ice fields at Beaufort that resulted in increased breeding habitat exemplifies a process that has been underway in the Ross Sea during the entire Holocene. Furthermore, our results are in line with predictions that major ice shelves and glaciers will retreat rapidly elsewhere in the Antarctic, potentially leading to increased breeding habitat for Adélie penguins. Results further indicated that satellite imagery may be used to estimate large changes in Adélie penguin populations, facilitating our understanding of metapopulation dynamics and environmental factors that influence regional populations.

Population status and breeding of New Zealand fur seals (Arctocephalus forsteri) in the Nelson‐northern Marlborough region, 1991–94

Abstract The population size, distribution, breeding, and onshore habitats of New Zealand fur seals in the Nelson‐northern Marlborough region, New Zealand, were investigated. Breeding occurs at four relatively new rookeries along the southern shores of Cook Strait: at Stephens Island in the outer Marlborough Sounds, at Archway Islands and Pillar Point near Cape Farewell, and at Tonga Island in Tasman Bay. These rookeries were established in about 1970, 1980, 1988, and 1988, respectively. In 1993–94 about 500 pups were born and the extant population breeding or born at the four rookeries was assessed as 2410. There were also 17 hauling grounds, each with 25 or more seals ashore in winter, and about 50 other sites where lesser numbers hauled out regularly. Comparison with earlier estimates indicates that the summer population increased from about 70 in 1973 to about 1000 in 1994—a sevenfold increase of yearling and older seals. The number of pups produced increased at a mean annual rate of 23% since 1970–71...

Intra-seasonal variation in foraging behavior among Adélie penguins (Pygocelis adeliae) breeding at Cape Hallett, Ross Sea, Antarctica

We investigated intra-seasonal variation in foraging behavior of chick-rearing Adélie penguins, Pygoscelis adeliae, during two consecutive summers at Cape Hallett, northwestern Ross Sea. Although foraging behavior of this species has been extensively studied throughout the broad continental shelf region of the Ross Sea, this is the first study to report foraging behaviors and habitat affiliations among birds occupying continental slope waters. Continental slope habitat supports the greatest abundances of this species throughout its range, but we lack information about how intra-specific competition for prey might affect foraging and at-sea distribution and how these attributes compare with previous Ross Sea studies. Foraging trips increased in both distance and duration as breeding advanced from guard to crèche stage, but foraging dive depth, dive rates, and vertical dive distances travelled per hour decreased. Consistent with previous studies within slope habitats elsewhere in Antarctic waters, Antarctic krill (Euphausia superba) dominated chick meal composition, but fish increased four-fold from guard to crèche stages. Foraging-, focal-, and core areas all doubled during the crèche stage as individuals shifted distribution in a southeasterly direction away from the coast while simultaneously becoming more widely dispersed (i.e., less spatial overlap among individuals). Intra-specific competition for prey among Adélie penguins appears to influence foraging behavior of this species, even in food webs dominated by Antarctic krill.

Semi-automated penguin counting from digital aerial photographs

Semi-automated software has been written in Matlab to enable nesting Adélie penguins (Pygoscelis adeliae), on aerial photographs of the Ross Sea sector of Antarctica, to be counted. Previously, this task had been accomplished by manual marking of printed images; this is slow, and highly-dependent on the skill of the operator. The basis of the semiautomated counting procedure is to use linear discriminant analysis to separate the background (snow, water, rock, bare ground) from the guano-covered colony area, followed by morphological image processing operators to select the breeding penguins. Many interactive features are provided that allow an operator to delete penguins singly or in groups, selectively process a defined area, and record the running census counts.

Global phenological insensitivity to shifting ocean temperatures among seabirds

Reproductive timing in many taxa plays a key role in determining breeding productivity1, and is often sensitive to climatic conditions2. Current climate change may alter the timing of breeding at different rates across trophic levels, potentially resulting in temporal mismatch between the resource requirements of predators and their prey3. This is of particular concern for higher-trophic-level organisms, whose longer generation times confer a lower rate of evolutionary rescue than primary producers or consumers4. However, the disconnection between studies of ecological change in marine systems makes it difficult to detect general changes in the timing of reproduction5. Here, we use a comprehensive meta-analysis of 209 phenological time series from 145 breeding populations to show that, on average, seabird populations worldwide have not adjusted their breeding seasons over time (−0.020 days yr−1) or in response to sea surface temperature (SST) (−0.272 days °C−1) between 1952 and 2015. However, marked between-year variation in timing observed in resident species and some Pelecaniformes and Suliformes (cormorants, gannets and boobies) may imply that timing, in some cases, is affected by unmeasured environmental conditions. This limited temperature-mediated plasticity of reproductive timing in seabirds potentially makes these top predators highly vulnerable to future mismatch with lower-trophic-level resources2.Time of reproduction may be altered as the climate changes. For seabirds, it is shown that there has not been an adjustment in timing as the climate changes and the sea surface warms. This lack of plasticity could result in a mismatch with food resources.