Flexibility in the bimodal foraging strategy of a high Arctic alcid, the little auk Alle alle

A bimodal foraging strategy has previously been described for procellariiform seabird species and is thought to have evolved in response to local resource availability being too low for adult birds to meet chick requirements and simultaneously maintain their own body condition. Here, we examine the dual foraging trip pattern of an alcid, the little auk Alle alle , at five colonies with contrasting oceanographic conditions. In spite of large variation in local conditions, little auks at all colonies showed the same general pattern of alternating a single long-trip with several consecutive short-trips. However, we found that the foraging pattern was flexible and could be adjusted at three levels: (1) the length of long-trips, (2) the frequency of short-trips, and (3) the total time spent foraging. Birds facing unfavorable conditions increased the duration of long-trips and reduced the number of short-trips. These adjustments resulted in reduced provisioning rates of chicks despite the fact that birds also increased the time allocated to foraging. Travel times during foraging trips were positively correlated to the total duration of the trip suggesting that differences in trip length among colonies were partly driven by variation in the distance to foraging areas. Most birds spent substantially more time traveling during long compared to short-trips, indicating that they accessed distant foraging areas during long-trips but remained close to the colony during short-trips. However, the difference in travel times was small at the site with the most favorable conditions suggesting that bimodal foraging in the little auk may be independent of the existence of high-quality areas at distance from the breeding ground.     

No evidence of extra-pair paternity in the little auk Alle alle

Extra-pair paternity is typically rather infrequent in seabirds, as in most other long-lived and socially monogamous birds. Here we report the first paternity study of the little auk Alle alle , a high arctic seabird which raises only a single chick per year. Parentage was determined using three highly polymorphic microsatellite markers. We found that all 26 chicks in our sample were true genetic offspring of their social parents, with an upper 95% confidence limit of 10.88% for the frequency of extra-pair paternity. This level of extra-pair paternity is not significantly different from frequencies reported from the closely related common Uria aalge and Brünnich's guillemots U. lomvia .     

Divergent diving behavior during short and long trips of a bimodal forager,the little auk Alle alle

The purpose of this study was to characterize for the first time seabird diving behavior during bimodal foraging. Little auks Alle alle, small zooplanktivorous Alcids of the High Arctic, have recently been shown to make foraging trips of short and long duration. Because short (ST) and long trips (LT) are thought to occur in different locations and serve different purposes (chick‐ and self‐feeding, respectively) we hypothesized that foraging differences would be apparent, both in terms of water temperature and diving characteristics. Using Time Depth Recorders (TDRs), we tested this hypothesis at three colonies along the Greenland Sea with contrasting oceanographic conditions. We found that diving behavior generally differed between ST and LT. However, the magnitude of the disparity in diving characteristics depended on local foraging conditions. At the study site where conditions were favorable, diving behavior differed only to a small degree between LT and ST. Together with a lack of difference in diving depth and ocean temperature, this indicates that these birds did not increase their foraging effort during ST nor did they travel long distances to seek out more profitable prey. In contrast, where local foraging conditions were poor, birds increased their diving effort substantially to collect a chick meal during ST as indicated by longer, more U‐shaped dives with slower ascent rates and shorter resting times (post‐dive intervals and extended surface pauses). In addition, large differences in diving depth and ocean temperature indicate that birds forage on different prey species and utilize different foraging areas during LT, which may be up to 200 km away from the colony. Continued warming and deteriorating near‐colony foraging conditions may have energetic consequences for little auks breeding in the eastern Greenland Sea.     

Distribution of little auk (Alle alle) breeding colonies in Thule District, northwest Greenland

Large areas with hydrocarbon prospects offshore northwest Greenland necessitate seabird studies in this region. The little auk population breeding in Thule District is the largest seabird population in northwest Greenland, as well as the largest known population of this species. In 1994 and 1995 we mapped the distribution of little auk colonies in Thule District during aerial surveys. Most colonies are situated in sloping screes facing the sea, some as far as 11 km from the coast in valleys or facing glaciers. The total horizontal extent of the colonies is about 400 km. Our surveys generally confirm previous and local knowledge, although the information obtained is more detailed. Received: 11 November 1996 / Accepted: 21 September 1997     

Composition of chick meals from one of the main little auk (Alle alle) breeding colonies in Northwest Greenland

During the last decade, increasing information on little auk (Alle alle) biology, ecology and behaviour has been reported. However, only a few of these studies have focused on the breeding population in the Avanersuaq (Thule) district of Northwest Greenland, where 80 % of the global little auk population is estimated to breed. This study reports on the chick diet composition from one of the largest colonies, the Paakitsoq colony, located on the south-eastern margin of the North Water (NOW) Polynya. Results revealed the highest proportion of Calanus hyperboreus, a large lipid-rich copepod, in chick diet reported for any little auk colony. Results confirmed that the cold, highly productive waters of the NOW Polynya are favourable foraging grounds for the little auks during the breeding season. Species diversity within and between the chick meals was low, which probably reflects a high availability of a few preferred prey species. Individual chick meals were generally low in number of prey items and total energy content compared with other published results. This may be explained by a higher feeding frequency or by the samples being collected late in the breeding period (during late chick rearing), when chicks have a reduced growth rate and may require less energy than at earlier developmental stages.     

Winter diet of the little auk (Alle alle) in the Northwest Atlantic

The little auk (Alle alle) is one of the most numerous seabirds in the world, but their winter prey selection has never been thoroughly studied. In the present study, we analyzed the proventricular contents of 205 little auks caught in coastal areas off southwest Greenland during December–February and off Newfoundland in March. Large Calanoid copepods are known to be the main prey during summer. We found krill to become the crucial winter prey in both areas, followed by Themisto spp. and young capelin (Mallotus villosus). No difference was found between the diet of juvenile and adult birds. Copepods constituted around 1 % of the diet and close to all copepods were in birds caught near Cape Farwell in December. These findings provide new and important insight into the forage ecology of the little auk, and they support other studies showing that large Calanoid copepods in the Arctic decent for winter hibernation at depths that are below the diving range of the little auks. More studies to determine offshore diet and annual variation are, however, needed in order to get a more complete picture.     

The finding of a cestode in the little auk Alle alle on Franz Josef Land

One of 14 dovekeys (Alle alle) investigated in August 1993 on Hooker Island (Franz Joseph Land) was found to harbour a single specimen of an early immature dilepidid cestode attributed as Alcataenia sp. The rostellum bears a double crown of 22 hooks, 0.025-0.027 mm in length. Except Threlfall (1971) there was no information concerning dovekey's parasites.     

On the importance of fat reserves for the little auk (Alle alle) chicks

Summary The pattern of fat deposition and inter-year variation in the amount of deposited fat, lean dry mass (LDM) and water were studied in the chicks of little auk (Alle alle), an arctic alcid, at Spitsbergen in the 1984, 1986 and 1987 breeding seasons. The lipid index (lipid mass/LDM) rose from 0.29 at hatching to about 0.70 at the age of maximum body mass attained in the nest, the highest yet recorded for seabirds except procellariiforms. Just before fledging the index dropped to 0.38. Fat mass in 1984 chicks and also fresh body mass of older 1984 chicks was significantly lower than in the other two seasons, but there were no statistical year-to-year differences in LDM's of chicks. The amount of fat in chicks was significantly correlated with certain weather parameters. Fasting capability (FC) of the chicks (the time that chicks could survive when using only their lipid stores) was calculated from fat mass and previous data on metabolic rates of chicks. The FC's of all 108 chicks collected in three seasons were greater than maximum interval between two consecutive feeds to chicks reported in the study colony. Also the estimated fat reserves of 99.7% of chicks weighed in 1984 were sufficient to support them for longer than the maximum recorded intervals between feeds. In relation to death by starvation, little auk chicks, at least up to the age of maximum body mass, have a substantial surplus of fat. This is contrary to the hypothesis that fat depots in seabird chicks assist survival during the periodic fasts resulting from unpredictable feeding conditions.     

Climate warming decreases the survival of the little auk (Alle alle), a high Arctic avian predator

Delayed maturity, low fecundity, and high adult survival are traits typical for species with a long‐life expectancy. For such species, even a small change in adult survival can strongly affect the population dynamics and viability. We examined the effects of both regional and local climatic variability on adult survival of the little auk, a long‐lived and numerous Arctic seabird species. We conducted a mark‐resighting study for a period of 8 years (2006‐2013) simultaneously at three little auk breeding sites that are influenced by the West Spitsbergen Current, which is the main carrier of warm, Atlantic water into the Arctic. We found that the survival of adult little auks was negatively correlated with both the North Atlantic Oscillation (NAO) index and local summer sea surface temperature (SST), with a time lag of 2 and 1 year, respectively. The effects of NAO and SST were likely mediated through a change in food quality and/or availability: (1) reproduction, growth, and development of Arctic Calanus copepods, the main prey of little auks, are negatively influenced by a reduction in sea ice, reduced ice algal production, and an earlier but shorter lasting spring bloom, all of which result from an increased NAO; (2) a high sea surface temperature shortens the reproductive period of Arctic Calanus, decreasing the number of eggs produced. A synchronous variation in survival rates at the different colonies indicates that climatic forcing was similar throughout the study area. Our findings suggest that a predicted warmer climate in the Arctic will negatively affect the population dynamics of the little auk, a high Arctic avian predator.     

Potentially pathogenic yeast isolated from the throat and cloaca of an Arctic colonial seabird: the little auk (Alle alle)

Yeasts are a distinctive group of microfungi, but compared to other microorganisms, their ecological function and biodiversity are poorly known. This is especially so where polar ecosystems are concerned. With climate changes and increasing pollution levels in the Arctic, it can be anticipated that there will be an increase in the prevalence and diversity of fungi colonizing live organisms. With these changes, it is crucial to investigate and monitor species diversity and prevalence of fungi in this fragile environment. In this study, yeasts were examined from throat and cloaca of a small colonial seabird, the little auk (Alle alle), a keystone species in the Arctic ecosystem. Samples were collected from 94 adults and 17 nestlings in breeding colony in Magdalenefjorden (NW Spitsbergen) in 2009. In total, twelve species of yeast from eight genera were found in 12 % of the samples, with the Dipodascus genus being the most prevalent. All yeast species were found in the adults, but only one species, Cryptococcus macerans, was found in a single nestling. In individuals where fungus was isolated, it was only isolated from either the throat or the cloaca, except for two cases, where fungus was found in both throat and cloaca. The presence of yeast was not related to sex but age of the birds, with adults being more prone to colonization by yeasts than the nestlings. The relatively low prevalence and diversity of yeast in little auks suggest that these birds are random carriers of fungi, with minor health impacts.     

Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time‐constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early‐laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post‐fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first‐laid eggs.     

Spatial ecology of a root parasite – from pattern to process

Abstract Occurrence patterns of parasitic plants are constrained by the distribution of suitable hosts and movement patterns of seed vectors and, accordingly, represent a simplified system to study many aspects of spatial ecology and determinants of distribution. Previous work has focused on the aerially hemiparasitic mistletoes, and it is unclear whether root parasites are affected by similar factors. Here, we evaluate spatial patterns in the root parasitic Santalum lanceolatum in an arid shrubland in north‐western New South Wales, central Australia. In this region, the principal host is a long‐lived nitrogen fixing shrub Acacia tetragonophylla closely associated with ephemeral creek‐lines. The location of 765 individuals of both species was mapped along a 250‐m section of creek‐line using a total survey station, and occurrence patterns of the root parasite related to host distribution and landscape context. We used Ripley's K‐function and the O‐ring statistic to determine whether the distribution of S. lanceolatum was random, aggregated or regular; the spatial scales at which these patterns occurred; and to quantify any spatial associations between the parasite and its host, A. tetragonophylla. While acacias were closely associated with the creek‐line, S. lanceolatum plants were more tightly clustered, displaying significant clustering at two spatial scales (1.2 m and 8.8 m). We suggest that host quality may act as an important constraint, with only those acacias growing in or near the creek‐line being physiologically capable of supporting a parasite to maturity. Insights gained from spatial analysis are used to guide ongoing research in this system, and highlight the utility of the O‐ring statistic for understanding patterns of distribution affected by multiple processes operating at critical scales.     

Estimating prey capture rates of a planktivorous seabird,the little auk (<Emphasis Type="Italic">Alle alle</Emphasis>), using diet,diving behaviour,and energy consumption

Interpreting the impact of environmental change on food webs requires a clear understanding of predator–prey interactions. Such knowledge is often lacking in the marine environment where the foraging behaviour and prey requirements of some of the major top-predators remains mysterious. For example, very little is known about the underwater foraging behaviour of the little auk, the most numerous seabird in the North Atlantic. In 2004, we used time–depth-recorders at two breeding colonies in East Greenland to examine the diving behaviour of this small, planktivorous seabird during the chick-rearing period. Due to technical difficulties data were only collected for four individuals, but recordings showed that birds dive up to 240 times a day to maximum depths of 27 m (average 10 m), with maximum dive durations of 90 s (average 52 s). In addition, we collected the chick meals from 35 individuals, which were dominated by Calanus copepods (95%), and also determined the field metabolic rates (FMR) of 14 individuals using the doubly labelled water technique, which averaged 609.9 kJ day⁻¹. We integrated information on diving duration with chick diet and FMR to estimate the prey requirements and underwater capture rates of little auks using a Monte Carlo simulation. Chick-rearing little auks needed to catch about 59,800 copepods day⁻¹, which is equivalent to about six copepods caught per second spent underwater. These astonishing results strongly suggest that little auks are, at least partly, filter-feeding, and underline the importance of highly productive, cool marine areas that harbour dense patches of large, energy-rich copepods.     

Factors influencing the evolution of moult in the non‐breeding season: insights from the family Motacillidae

The number of moults per annual cycle and their final spatial pattern (i.e. topography) show high interspecific variation in the order Passeriformes. Factors behind this variability remain obscure, especially for variability in spatial pattern among species. Here, we explored the relative influence of ten ecological, ontogenetic, social and sexual factors on the evolution of autumn moult (feather replacement largely undertaken by migratory species, which is not necessarily an independent episode within their moult cycle) and prealternate moult among Northern Hemisphere species of the family Motacillidae using phylogenetically controlled analyses, ancestral state reconstruction and analyses of correlated evolution. The results strongly support the presence of prealternate moult and absence of autumn moult as ancestral states in this family. A high rate of change between related species indicates phylogenetic independence among prealternate moult patterns and examined factors. Migration distance and gregariousness are the most important factors influencing prealternate moult evolution, and point toward natural selection and sociality as the most important evolutionary drivers of prealternate moult in Motacillidae. Breeding latitude, seasonal plumage change, winter plumage conspicuousness, sexual dichromatism, plumage maturation and extent of preformative moult show a minor influence, and suggest that ontogeny and sexual selection may have played a limited role in shaping prealternate moult in Motacillidae.     

Extension of the growing season due to delayed autumn over mid and high latitudes in North America during 1982–2006

Aim We intend to characterize and understand the spatial and temporal patterns of vegetation phenology shifts in North America during the period 1982–2006. Location North America. Methods A piecewise logistic model is used to extract phenological metrics from a time‐series data set of the normalized difference vegetation index (NDVI). An extensive comparison between satellite‐derived phenological metrics and ground‐based phenology observations for 14,179 records of 73 plant species at 802 sites across North America is made to evaluate the information about phenology shifts obtained in this study. Results The spatial pattern of vegetation phenology shows a strong dependence on latitude but a substantial variation along the longitudinal gradient. A delayed dormancy onset date (0.551 days year^?1, P= 0.013) and an extended growing season length (0.683 days year^?1, P= 0.011) are found over the mid and high latitudes in North America during 1982–2006, while no significant trends in greenup onset are observed. The delayed dormancy onset date and extended growing season length are mainly found in the shrubland biome. An extensive validation indicates a strong robustness of the satellite‐derived phenology information. Main conclusions It is the delayed dormancy onset date, rather than an advanced greenup onset date, that has contributed to the prolonged length of the growing season over the mid and high latitudes in North America during recent decades. Shrublands contribute the most to the delayed dormancy onset date and the extended growing season length. This shift of vegetation phenology implies that vegetation activity in North America has been altered by climatic change, which may further affect ecosystem structure and function in the continent.     

Nutrient Fluxes from Land to Sea: Consequences of Future Scenarios on the Oder River Basin – Lagoon – Coastal Sea System

Eutrophication management is still one of the major challenges in the Baltic Sea region. Intense transformation processes in several Baltic Sea states have led to drastic changes in e.g., landuse and thereby nutrient emissions and water quality. Several future development directions are possible. The Oder catchment – lagoon – coastal water system serves as a pilot study area, since it has a major influence on the nutrient loads into the Baltic Sea and about 90% of the catchment is located in Poland, a state with transitional economy. Different scenarios for landuse changes in the Oder catchment are developed and their consequences on nutrient emissions simulated. Next to politically induced changes of agricultural landuse in general, specific aspects such as cultivation of energy maize and increased animal stocks are considered. Nitrogen emissions are likely to increase due to agricultural landuse changes whereas phosphorus emissions will not change or even decrease according to the application of the EC‐Urban Waste Water Treatment Directive. Resulting nitrogen loads to the Oder Lagoon could increase up to 23%, phosphorus loads could decrease by 11% compared to 2005. These trends may lead to higher nitrogen availability compared to phosphorus at least in the Oder lagoon. Interannual differences in discharge also have profound effects on nutrient emissions. A good status of the Oder river basin – lagoon – coastal sea system according to EC‐Directives is not very likely to be achieved under the investigated circumstances. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Process and pattern in the biogeography of New Zealand – a global microcosm?

Aim  To describe New Zealand's historical terrestrial biogeography and place this history in a wider Southern Hemisphere context.
Location  New Zealand.
Methods  The analysis is based primarily on literature on the distributions and relationships of New Zealand's terrestrial flora and fauna.
Results  New Zealand is shown to have a biota that has broad relationships, primarily around the cool Southern Hemisphere, as well as with New Caledonia to the north. There are hints of ancient Gondwanan taxa, although the long-argued predominance of taxa derived by vicariant processes, driven by plate tectonics and the fragmentation of Gondwana, is no longer accepted as a principal explanation of the biota's origins and relationships.
Main conclusions  Most of the terrestrial New Zealand flora and fauna has clearly arrived in New Zealand much more recently than the postulated separation of New Zealand from Gondwana, dated at c. 80 Ma. There is a view that New Zealand may have disappeared completely beneath the sea in the early Cenozoic, and acceptance of this would mean derivation of the entire biota by transoceanic dispersal. However, there are elements in the biota that seem to have broad distributions that date back to Gondwanan times, and also some that are thought unlikely to have been able to disperse to New Zealand across ocean gaps, especially freshwater organisms. Very strong connections to the biota of Australia, rather than to South America, are inconsistent with the timing of New Zealand's ancient and early separation from Gondwana and seem likely to have resulted from dispersal.     

Leptodora kindti and Flexible Foraging Behaviour of Fish – Factors behind the Delayed Biomass Peak of Cladocerans in Lake Hiidenvesi

In the eutrophic L. Hiidenvesi, the spring biomass maximum of cladoceran zooplankton is missing and the highest biomass takes place in July–August. The factors behind the delayed biomass peak were studied in four different basins of the lake with concomitant data on cladocerans assemblages, density of the predatory cladoceran Leptodora kindti and food composition of fish. In all the basins, the abundance of Leptodora peaked in June, being highest (up to 800 ind. m^–3) in the two most shallow basins (max depth < 4 m). The duration of the high population density was short and in July‐August Leptodora density stayed below 200 ind. m^–3, although the water temperature was still favourable. The collapse of the Leptodora population coincided with the change in the feeding habits of fish. In early summer, fish predation was targeted mainly on copepods and zoobenthos, while in high summer Leptodora was one of the main preys of perch, white bream and bleak. The biomass of herbivorous cladocerans was below 10 μg C l^–1 in June, and climbed to a maximum in August in the two most shallow basins (34 and 76 μg C l^–1), in July in the deepest basin (27 μg C l^–1), and in September in the intermediate basin (55 μg C l^–1). In the two most shallow basins, the death rate of the dominating cladoceran, Daphnia cristata, closely followed the food consumption rate by the Leptodora population. In the deeper basins, the agreement was not so close, smelts (Osmerus eperlanus) and chaoborids being important predators of herbivores. The duration of the period of high Leptodora density thus depended on the predation pressure by fish, while the increased fish predation on Leptodora in July–August allowed the elevation of the biomass of herbivorous cladocerans.