Mercury exposure in an endangered seabird: long-term changes and relationships with trophic ecology and breeding success

Mercury (Hg) is an environmental contaminant which, at high concentrations, can negatively influence avian physiology and demography. Albatrosses (Diomedeidae) have higher Hg burdens than all other avian families. Here, we measure total Hg (THg) concentrations of body feathers from adult grey-headed albatrosses (Thalassarche chrysostoma) at South Georgia. Specifically, we (i) analyse temporal trends at South Georgia (1989–2013) and make comparisons with other breeding populations; (ii) identify factors driving variation in THg concentrations and (iii) examine relationships with breeding success. Mean ± s.d. feather THg concentrations were 13.0 ± 8.0 µg g⁻¹ dw, which represents a threefold increase over the past 25 years at South Georgia and is the highest recorded in the Thalassarche genus. Foraging habitat, inferred from stable isotope ratios of carbon (δ¹³C), significantly influenced THg concentrations—feathers moulted in Antarctic waters had far lower THg concentrations than those moulted in subantarctic or subtropical waters. THg concentrations also increased with trophic level (δ¹⁵N), reflecting the biomagnification process. There was limited support for the influence of sex, age and previous breeding outcome on feather THg concentrations. However, in males, Hg exposure was correlated with breeding outcome—failed birds had significantly higher feather THg concentrations than successful birds. These results provide key insights into the drivers and consequences of Hg exposure in this globally important albatross population.     

Hidden keys to survival: the type,density, pattern and functional role of emperor penguin body feathers

Antarctic penguins survive some of the harshest conditions on the planet. Emperor penguins breed on the sea ice where temperatures drop below −40°C and forage in −1.8°C waters. Their ability to maintain 38°C body temperature in these conditions is due in large part to their feathered coat. Penguins have been reported to have the highest contour feather density of any bird, and both filoplumes and plumules (downy feathers) are reported absent in penguins. In studies modelling the heat transfer properties and the potential biomimetic applications of penguin plumage design, the insulative properties of penguin plumage have been attributed to the single afterfeather attached to contour feathers. This attribution of the afterfeather as the sole insulation component has been repeated in subsequent studies. Our results demonstrate the presence of both plumules and filoplumes in the penguin body plumage. The downy plumules are four times denser than afterfeathers and play a key, previously overlooked role in penguin survival. Our study also does not support the report that emperor penguins have the highest contour feather density.     

Where do penguins go during the inter-breeding period? Using geolocation to track the winter dispersion of the macaroni penguin

Although penguins are key marine predators from the Southern Ocean, their migratory behaviour during the inter-nesting period remains widely unknown. Here, we report for the first time, to our knowledge, the winter foraging movements and feeding habits of a penguin species by using geolocation sensors fitted on penguins with a new attachment method. We focused on the macaroni penguin Eudyptes chrysolophus at Kerguelen, the single largest consumer of marine prey among all seabirds. Overall, macaroni penguins performed very long winter trips, remaining at sea during approximately six months within the limits of the Southern Ocean. They departed from Kerguelen in an eastward direction and distributed widely, over more than 3.10⁶ km². The penguins spent most of their time in a previously unrecognized foraging area, i.e. a narrow latitudinal band (47–49° S) within the central Indian Ocean (70–110° E), corresponding oceanographically to the Polar Frontal Zone. There, their blood isotopic niche indicated that macaroni penguins preyed mainly upon crustaceans, but not on Antarctic krill Euphausia superba, which does not occur at these northern latitudes. Such winter information is a crucial step for a better integrative approach for the conservation of this species whose world population is known to be declining.     

Rapidly increasing methyl mercury in endangered ivory gull (Pagophila eburnea) feathers over a 130 year record

Mercury (Hg) is increasing in marine food webs, especially at high latitudes. The bioaccumulation and biomagnification of methyl mercury (MeHg) has serious effects on wildlife, and is most evident in apex predators. The MeHg body burden in birds is the balance of ingestion and excretion, and MeHg in feathers is an effective indicator of overall MeHg burden. Ivory gulls (Pagophila eburnea), which consume ice-associated prey and scavenge marine mammal carcasses, have the highest egg Hg concentrations of any Arctic bird, and the species has declined by more than 80% since the 1980s in Canada. We used feathers from museum specimens from the Canadian Arctic and western Greenland to assess whether exposure to MeHg by ivory gulls increased from 1877 to 2007. Based on constant feather stable-isotope (δ¹³C, δ¹⁵N) values, there was no significant change in ivory gulls'' diet over this period, but feather MeHg concentrations increased 45× (from 0.09 to 4.11 µg g⁻¹ in adults). This dramatic change in the absence of a dietary shift is clear evidence of the impact of anthropogenic Hg on this high-latitude threatened species. Bioavailable Hg is expected to increase in the Arctic, raising concern for continued population declines in high-latitude species that are far from sources of environmental contaminants.     

Activity Time Budget during Foraging Trips of Emperor Penguins

We developed an automated method using depth and one axis of body acceleration data recorded by animal-borne data loggers to identify activities of penguins over long-term deployments. Using this technique, we evaluated the activity time budget of emperor penguins (n = 10) both in water and on sea ice during foraging trips in chick-rearing season. During the foraging trips, emperor penguins alternated dive bouts (4.8±4.5 h) and rest periods on sea ice (2.5±2.3 h). After recorder deployment and release near the colony, the birds spent 17.9±8.4% of their time traveling until they reached the ice edge. Once at the ice edge, they stayed there more than 4 hours before the first dive. After the first dive, the mean proportions of time spent on the ice and in water were 30.8±7.4% and 69.2±7.4%, respectively. When in the water, they spent 67.9±3.1% of time making dives deeper than 5 m. Dive activity had no typical diurnal pattern for individual birds. While in the water between dives, the birds had short resting periods (1.2±1.7 min) and periods of swimming at depths shallower than 5 m (0.25±0.38 min). When the birds were on the ice, they primarily used time for resting (90.3±4.1% of time) and spent only 9.7±4.1% of time traveling. Thus, it appears that, during foraging trips at sea, emperor penguins traveled during dives >5 m depth, and that sea ice was primarily used for resting. Sea ice probably provides refuge from natural predators such as leopard seals. We also suggest that 24 hours of sunlight and the cycling of dive bouts with short rest periods on sea ice allow emperor penguins to dive continuously throughout the day during foraging trips to sea.     

Making the most of the old age: Autumn breeding as an extra reproductive investment in older seabirds

The extrinsic and intrinsic factors affecting differing reproductive strategies among populations are central to understanding population and evolutionary ecology. To evaluate whether individual reproductive strategies responded to annual patterns in marine productivity and age‐related processes in a seabird we used a long term (2003–2013), a continuous dataset on nest occupancy and attendance at the colony by little penguins (Eudyptula minor) at Phillip Island (Victoria, Australia). We found that concurrent with a secondary annual peak of marine productivity, a secondary peak in colony attendance and nest occupancy was observed in Autumn (out of the regular breeding season in spring/summer) with individuals showing mating‐like behavior. Individuals attending this autumn peak averaged 2.5 years older than those individuals that exclusively bred during spring/summer. Rather than being a naïve response by younger and inexperienced birds misreading environmental cues, our data indicate that the autumn peak attendance is an earlier attempt to breed by older and more experienced penguins. Therefore, we provide strong support for the fundamental prediction of the life‐history theory of increasing investment in reproduction with age to maximize lifetime fitness as future survival prospects diminish and experience increases.     

Transition between moult and migration in a long-distance migratory passerine: organ flexibility in the African wintering area

Phenotypic flexibility of organs in migratory birds has been documented for a variety of species of different genera during the migratory period. However, very little is known about phenotypic mass changes of organs with respect to other events within the annual cycle. This seems particularly interesting when birds face different physiological challenges in quick succession. We investigated mass changes of 13 organs from garden warblers (Sylvia borin) during the transition from moult to migration. These long-distance migratory birds perform a complete moult within their wintering area just shortly before the onset of spring migration. Birds were sampled in three successive stages according to their moult status: group I consisted of birds with growing primary or secondary wing feathers, group II consisted of birds with completed wing moult but with still moulting body feathers, and group III consisted of birds that had completed wing moult and body moult. Size-corrected flight muscle, kidney mass, and pancreas mass differed significantly among the three groups. Flight muscle was heaviest in birds that were about to leave their wintering area (group III) compared with birds still in body moult (group II). Kidney and pancreas showed a pattern similar to each other, with the heaviest mass occurring in birds with moulting wing feathers (group I) and significantly reduced mass in birds that had completed wing moult (group II) or both wing and body moult (group III). Mass reductions of kidney and pancreas during the transition from moult to migration are considered to be related to the demands of moult, while increased flight muscle may be due to moult, migration, or both. Phenotypic mass changes of organs in birds occur during their migration, but they also occur during the transition between other phases of the annual cycle such as moult and migration and are not restricted to the flight muscle.     

The Role of Canine Distemper Virus and Persistent Organic Pollutants in Mortality Patterns of Caspian Seals (Pusa caspica)

Persistent organic pollutants are a concern for species occupying high trophic levels since they can cause immunosuppression and impair reproduction. Mass mortalities due to canine distemper virus (CDV) occurred in Caspian seals (Pusa caspica), in spring of 1997, 2000 and 2001, but the potential role of organochlorine exposure in these epizootics remains undetermined. Here we integrate Caspian seal mortality data spanning 1971–2008, with data on age, body condition, pathology and blubber organochlorine concentration for carcases stranded between 1997 and 2002. We test the hypothesis that summed PCB and DDT concentrations contributed to CDV associated mortality during epizootics. We show that age is the primary factor explaining variation in blubber organochlorine concentrations, and that organochlorine burden, age, sex, and body condition do not account for CDV infection status (positive/negative) of animals dying in epizootics. Most animals (57%, n = 67) had PCB concentrations below proposed thresholds for toxic effects in marine mammals (17 µg/g lipid weight), and only 3 of 67 animals had predicted TEQ values exceeding levels seen to be associated with immune suppression in harbour seals (200 pg/g lipid weight). Mean organonchlorine levels were higher in CDV-negative animals indicating that organochlorines did not contribute significantly to CDV mortality in epizootics. Mortality monitoring in Azerbaijan 1971–2008 revealed bi-annual stranding peaks in late spring, following the annual moult and during autumn migrations northwards. Mortality peaks comparable to epizootic years were also recorded in the 1970s–1980s, consistent with previous undocumented CDV outbreaks. Gompertz growth curves show that Caspian seals achieve an asymptotic standard body length of 126–129 cm (n = 111). Males may continue to grow slowly throughout life. Mortality during epizootics may exceed the potential biological removal level (PBR) for the population, but the low frequency of epizootics suggest they are of secondary importance compared to anthropogenic sources of mortality such as fishing by-catch.     

How Much Is Too Much? Assessment of Prey Consumption by Magellanic Penguins in Patagonian Colonies

Penguins are major consumers in the southern oceans although quantification of this has been problematic. One suggestion proposes the use of points of inflection in diving profiles (‘wiggles’) for this, a method that has been validated for the estimation of prey consumption by Magellanic penguins (Spheniscus magellanicus) by Simeone and Wilson (2003). Following them, we used wiggles from 31 depth logger-equipped Magellanic penguins foraging from four Patagonian colonies; Punta Norte (PN), Bahía Bustamente (BB), Puerto Deseado (PD) and Puerto San Julián (PSJ), all located in Argentina between 42–49° S, to estimate the prey captured and calculate the catch per unit time (CPUT) for birds foraging during the early chick-rearing period. Numbers of prey caught and CPUT were significantly different between colonies. Birds from PD caught the highest number of prey per foraging trip, with CPUT values of 68±19 prey per hour underwater (almost two times greater than for the three remaining colonies). We modeled consumption from these data and calculate that the world Magellanic penguin population consumes about 2 million tons of prey per year. Possible errors in this calculation are discussed. Despite this, the analysis of wiggles seems a powerful and simple tool to begin to quantify prey consumption by Magellanic penguins, allowing comparison between different breeding sites. The total number of wiggles and/or CPUT do not reflect, by themselves, the availability of food for each colony, as the number of prey consumed by foraging trip is strongly associated with the energy content and wet mass of each colony-specific ‘prey type’. Individuals consuming more profitable prey could be optimizing the time spent underwater, thereby optimizing the energy expenditure associated with the dives.     

Contrasting strategies for wing‐moult and pre‐migratory fuelling in western and eastern populations of Common Whitethroat Sylvia communis

Trade‐offs between moult and fuelling in migrant birds vary with migration distance and the environmental conditions they encounter. We compared wing moult and fuelling at the northern and southern ends of migration in two populations of adult Common Whitethroats Sylvia communis. The western population moults most remiges at the breeding grounds in Europe (e.g. Poland) and migrates 4000–5000 km to western Africa (e.g. Nigeria). The eastern population moults all remiges at the non‐breeding grounds and migrates 7000–10 000 km from western Asia (e.g. southwestern Siberia) to eastern and southern Africa. We tested the hypotheses that: (1) Whitethroats moult their wing feathers slowly in South Africa, where they face fewer time constraints than in Poland, and (2) fuelling is slower when it coincides with moulting (Poland, South Africa) than when it occurs alone (Siberia, Nigeria). We estimated moult timing of primaries, secondaries and tertials from moult records of Polish and South African Whitethroats ringed in 1987–2017 and determined fuelling patterns from the body mass of Whitethroats ringed in all four regions. The western population moulted wing feathers in Poland over 55 days (2 July–26 August) at a varying rate, up to 13 feathers simultaneously, but fuelled slowly until departure in August–mid‐September. In Nigeria, during the drier period of mid‐February–March they fuelled slowly, but the fuelling rate increased three‐fold in April–May after the rains before mid‐April–May departure. The eastern population did not moult in Siberia but fuelled three times faster before mid‐July–early August departure than did the western birds moulting in Poland. In South Africa, the Whitethroats moulted over 57 days (2 January–28 February) at a constant rate of up to nine feathers simultaneously and fuelled slowly from mid‐December until mid‐April–May departure. These results suggest the two populations use contrasting strategies to capitalize on food supplies before departure from breeding and non‐breeding grounds.     

Feather corticosterone reveals effect of moulting conditions in the autumn on subsequent reproductive output and survival in an Arctic migratory bird

For birds, unpredictable environments during the energetically stressful times of moulting and breeding are expected to have negative fitness effects. Detecting those effects however, might be difficult if individuals modulate their physiology and/or behaviours in ways to minimize short-term fitness costs. Corticosterone in feathers (CORTf) is thought to provide information on total baseline and stress-induced CORT levels at moulting and is an integrated measure of hypothalamic–pituitary–adrenal activity during the time feathers are grown. We predicted that CORTf levels in northern common eider females would relate to subsequent body condition, reproductive success and survival, in a population of eiders nesting in the eastern Canadian Arctic during a capricious period marked by annual avian cholera outbreaks. We collected CORTf data from feathers grown during previous moult in autumn and data on phenology of subsequent reproduction and survival for 242 eider females over 5 years. Using path analyses, we detected a direct relationship between CORTf and arrival date and body condition the following year. CORTf also had negative indirect relationships with both eider reproductive success and survival of eiders during an avian cholera outbreak. This indirect effect was dramatic with a reduction of approximately 30% in subsequent survival of eiders during an avian cholera outbreak when mean CORTf increased by 1 standard deviation. This study highlights the importance of events or processes occurring during moult on subsequent expression of life-history traits and relation to individual fitness, and shows that information from non-destructive sampling of individuals can track carry-over effects across seasons.     

Evaluating otter reintroduction outcomes using genetic spatial capture–recapture modified for dendritic networks

Monitoring the demographics and genetics of reintroduced populations is critical to evaluating reintroduction success, but species ecology and the landscapes that they inhabit often present challenges for accurate assessments. If suitable habitats are restricted to hierarchical dendritic networks, such as river systems, animal movements are typically constrained and may violate assumptions of methods commonly used to estimate demographic parameters. Using genetic detection data collected via fecal sampling at latrines, we demonstrate applicability of the spatial capture–recapture (SCR) network distance function for estimating the size and density of a recently reintroduced North American river otter (Lontra canadensis) population in the Upper Rio Grande River dendritic network in the southwestern United States, and we also evaluated the genetic outcomes of using a small founder group (n = 33 otters) for reintroduction. Estimated population density was 0.23–0.28 otter/km, or 1 otter/3.57–4.35 km, with weak evidence of density increasing with northerly latitude (β = 0.33). Estimated population size was 83–104 total otters in 359 km of riverine dendritic network, which corresponded to average annual exponential population growth of 1.12–1.15/year since reintroduction. Growth was ≥40% lower than most reintroduced river otter populations and strong evidence of a founder effect existed 8–10 years post‐reintroduction, including 13–21% genetic diversity loss, 84%–87% genetic effective population size decline, and rapid divergence from the source population (F _ST accumulation = 0.06/generation). Consequently, genetic restoration via translocation of additional otters from other populations may be necessary to mitigate deleterious genetic effects in this small, isolated population. Combined with non‐invasive genetic sampling, the SCR network distance approach is likely widely applicable to demogenetic assessments of both reintroduced and established populations of multiple mustelid species that inhabit aquatic dendritic networks, many of which are regionally or globally imperiled and may warrant reintroduction or augmentation efforts.     

Temporal Variation in Fish Mercury Concentrations within Lakes from the Western Aleutian Archipelago,Alaska

We assessed temporal variation in mercury (Hg) concentrations of threespine stickleback (Gasterosteus aculeatus) from Agattu Island, Aleutian Archipelago, Alaska. Total Hg concentrations in whole-bodied stickleback were measured at two-week intervals from two sites in each of two lakes from June 1 to August 10, 2011 during the time period when lakes were ice-free. Across all sites and sampling events, stickleback Hg concentrations ranged from 0.37–1.07 µg/g dry weight (dw), with a mean (± SE) of 0.55±0.01 µg/g dw. Mean fish Hg concentrations declined by 9% during the study period, from 0.57±0.01 µg/g dw in early June to 0.52±0.01 µg/g dw in mid-August. Mean fish Hg concentrations were 6% higher in Loon Lake (0.56±0.01 µg/g dw) than in Lake 696 (0.53±0.01 µg/g dw), and 4% higher in males (0.56±0.01 µg/g dw) than in females (0.54±0.01 µg/g dw). Loon Lake was distinguished from Lake 696 by the presence of piscivorous waterbirds during the breeding season. Mercury concentrations in stickleback from Agattu Island were higher than would be expected for an area without known point sources of Hg pollution, and high enough to be of concern to the health of piscivorous wildlife.     

Plumage development and moult in the European Quail Coturnix c. coturnix: criteria for age determination

Sequence, rate and duration of moult were studied in captive bred European Quail Coturnix coturnix coturnix. The founder population originated from southwest France. The study was conducted between 1986 and 1989 on birds aged from 1 day to 2 years, exposed to a seasonal photoperiod corresponding to latitude 16°N during autumn and winter and latitude 48°N during the remainder of the year. Under these conditions, adult quail showed two annual moults with only the post-breeding one being complete. The pre-breeding moult essentially involved the throat feathers. Large interindividual variation was observed in the duration, timing and development of the post-breeding moult: 60% of the studied birds suspended moult when they developed migratory restlessness and then finished renewing their feathers during the winter. The post-juvenile moult was also suspended when 7–9 weeks old (3–6 primaries and 1–10 secondaries renewed). After this suspension, the length of which was related to the hatching date, the moult continued up to p7. The three outer primaries were kept for the first year and were replaced only during the post-breeding moult. Based on the examination of wing patterns, our study provides reliable criteria for discriminating between age classes. The numbers of primaries and secondaries simultaneously in growth or renewed were different between the age classes. The secondaries of adults were renewed later in the moult stage than were the secondaries of juveniles. These criteria provide field researchers with a guide that enables them to age quail with reasonable accuracy.     

Mercury in Nelson's Sparrow subspecies at breeding sites

Background

Mercury is a persistent, biomagnifying contaminant that can cause negative effects on ecosystems. Marshes are often areas of relatively high mercury methylation and bioaccumulation. Nelson''s Sparrows (Ammodramus nelsoni) use marsh habitats year-round and have been documented to exhibit tissue mercury concentrations that exceed negative effects thresholds. We sought to further characterize the potential risk of Nelson''s Sparrows to mercury exposure by sampling individuals from sites within the range of each of its subspecies.

Methodology/Principal Findings

From 2009 to 2011, we captured adult Nelson''s Sparrows at sites within the breeding range of each subspecies (A. n. nelsoni: Grand Forks and Upham, North Dakota; A. n. alterus: Moosonee, Ontario; and A. n. subvirgatus: Grand Manan Island, New Brunswick) and sampled breast feathers, the first primary feather (P1), and blood for total mercury analysis. Mean blood mercury in nelsoni individuals captured near Grand Forks ranged from 0.84±0.37 to 1.65±1.02 SD ppm among years, between 2.0 and 4.9 times as high as concentrations at the other sites (P<0.01). Breast feather mercury did not vary among sites within a given sampling year (site means ranged from 0.98±0.69 to 2.71±2.93 ppm). Mean P1 mercury in alterus (2.96±1.84 ppm fw) was significantly lower than in any other sampled population (5.25±2.24–6.77±3.51 ppm; P≤0.03).

Conclusions/Significance

Our study further characterized mercury in Nelson''s Sparrows near Grand Forks; we documented localized and potentially harmful mercury concentrations, indicating that this area may represent a biological mercury hotspot. This finding warrants further research to determine if wildlife populations of conservation or recreational interest in this area may be experiencing negative effects due to mercury exposure. We present preliminary conclusions about the risk of each sampled population to mercury exposure.     

Condition-dependent variation in the blue-ultraviolet coloration of a structurally based plumage ornament

After years of investigation into the function of sexually dimorphic ornamental traits, researchers are beginning to understand how bright plumage colour in birds acts as an intraspecific signal. This work has focused primarily on pigment-based ornaments because they are highly variable in patch size, hue and brightness for some species. In contrast, structurally based ornaments have been little studied, in part because they do not appear to be as variable as pigment-based ornaments. We investigated a structurally based plumage ornament in a wild population of blue grosbeaks (Guiraca caerulea), a sexually dimorphic passerine. We report plumage variation that extends into the ultraviolet region of the spectrum. The pattern of covariation between four out of five elements of plumage variation suggests that structurally based ornamentation is pushed towards extreme expression of the trait as predicted by the sexual selection theory. The ''bluest'' birds have the highest percentage of blue feathers on the body. These ornamental feathers reflect light maximally at the shortest wavelengths (ultraviolet), with the greatest intensity and the greatest contrast. Age may have some effect on expression of blueness. In addition, plumage variables are correlated with growth bars in tail feathers (a record of nutritional condition during moult in a non-ornamental trait). This suggests that the ornament is partially condition dependent. Thus, blue plumage in male grosbeaks may serve as an honest indicator of age and quality.     

A Method for Investigating Population Declines of Migratory Birds Using Stable Isotopes: Origins of Harvested Lesser Scaup in North America

Background

Elucidating geographic locations from where migratory birds are recruited into adult breeding populations is a fundamental but largely elusive goal in conservation biology. This is especially true for species that breed in remote northern areas where field-based demographic assessments are logistically challenging.

Methodology/Findings

Here we used hydrogen isotopes (δD) to determine natal origins of migrating hatch-year lesser scaup (Aythya affinis) harvested by hunters in the United States from all North American flyways during the hunting seasons of 1999–2000 (n = 412) and 2000–2001 (n = 455). We combined geospatial, observational, and analytical data sources, including known scaup breeding range, δD values of feathers from juveniles at natal sites, models of δD for growing-season precipitation, and scaup band-recovery data to generate probabilistic natal origin landscapes for individual scaup. We then used Monte Carlo integration to model assignment uncertainty from among individual δD variance estimates from birds of known molt origin and also from band-return data summarized at the flyway level. We compared the distribution of scaup natal origin with the distribution of breeding population counts obtained from systematic long-term surveys.

Conclusions/Significance

Our analysis revealed that the proportion of young scaup produced in the northern (above 60°N) versus the southern boreal and Prairie-Parkland region was inversely related to the proportions of breeding adults using these regions, suggesting that despite having a higher relative abundance of breeding adults, the northern boreal region was less productive for scaup recruitment into the harvest than more southern biomes. Our approach for evaluating population declines of migratory birds (particularly game birds) synthesizes all available distributional data and exploits the advantages of intrinsic isotopic markers that link individuals to geography.     

Intraspecific Relationships among Wood Density,Leaf Structural Traits and Environment in Four Co-Occurring Species of Nothofagus in New Zealand

Plant functional traits capture important variation in plant strategy and function. Recent literature has revealed that within-species variation in traits is greater than previously supposed. However, we still have a poor understanding of how intraspecific variation is coordinated among different traits, and how it is driven by environment. We quantified intraspecific variation in wood density and five leaf traits underpinning the leaf economics spectrum (leaf dry matter content, leaf mass per unit area, size, thickness and density) within and among four widespread Nothofagus tree species in southern New Zealand. We tested whether intraspecific relationships between wood density and leaf traits followed widely reported interspecific relationships, and whether variation in these traits was coordinated through shared responses to environmental factors. Sample sites varied widely in environmental variables, including soil fertility (25–900 mg kg^–1 total P), precipitation (668–4875 mm yr^–1), temperature (5.2–12.4 °C mean annual temperature) and latitude (41–46 °S). Leaf traits were strongly correlated with one another within species, but not with wood density. There was some evidence for a positive relationship between wood density and leaf tissue density and dry matter content, but no evidence that leaf mass or leaf size were correlated with wood density; this highlights that leaf mass per unit area cannot be used as a surrogate for component leaf traits such as tissue density. Trait variation was predicted by environmental factors, but not consistently among different traits; e.g., only leaf thickness and leaf density responded to the same environmental cues as wood density. We conclude that although intraspecific variation in wood density and leaf traits is strongly driven by environmental factors, these responses are not strongly coordinated among functional traits even across co-occurring, closely-related plant species.     

Post-breeding dispersal of Ad��lie penguins (Pygoscelis adeliae) nesting at Signy Island, South Orkney Islands

ARGOS satellite telemetry and Global Location Sensors (geolocators) were used to identify the moult locations and the winter foraging dispersal of Adélie penguins after they left their breeding colonies on Signy Island in the South Orkney Islands. Animals were tracked during the period December 2004 to October 2005. All birds displayed a similar pattern of migratory behaviour, remaining away from colonies for approximately 9 months, at distances of up to 2,235 km. Moult locations were within the pack ice. Mean daily travel speeds to the moult locations were significantly faster when moving through open water than through pack ice. Moult occurred during February/March within a narrow latitudinal range (65–71°S), at a mean distance of 126 km from the ice edge; the mean duration of individual moult was c. 18.6 days. After moult, penguins spent the subsequent winter months moving north or north-eastward within the expanding winter pack ice, at a mean distance of 216 km from the ice edge, and in areas with ice cover >80%. The penguins returned to the vicinity of their colony between September 26 and October 22, 2005. This dependence of Adélie penguins on sea ice habitat suggests that any further reductions in sea ice extent in the Weddell Sea region would potentially have important impacts on the population processes of this pagophilic species.