Unravelling the migration and moult strategies of a long-distance migrant using stable isotopes: Red Knot Calidris canutus movements in the Americas

For long‐distance migrants, such as many of the shorebirds, understanding the demographic implications of behavioural strategies adopted by individuals is key to understanding how environmental change will affect populations. Stable isotopes have been used in the terrestrial environment to infer migratory strategies of birds but rarely in marine or estuarine systems. Here, we show that the stable isotope ratios of carbon and nitrogen in flight feathers can be used to identify at least three discrete wintering areas of the Red Knot Calidris canutus on the eastern seaboard of the Americas, ranging from southeastern USA to Patagonia and Tierra del Fuego. In spring, birds migrate northwards via Delaware Bay, in the northeastern USA, the last stopping point before arrival in Arctic breeding areas, where they fatten up on eggs of spawning Horseshoe Crabs Limulus polyphemus. The isotope ratios of feather samples taken from birds caught in the Bay during May 2003 were compared with feathers obtained from known wintering areas in Florida (USA), Bahia Lomas (Chile) and Rio Grande (Argentina). In May 2003, 30% of birds passing through the Bay had Florida‐type ‘signatures’, 58% were Bahia Lomas‐type, 6% were Rio Grande‐type and 7% were unclassified. Some of the southern wintering birds had started moulting flight feathers in northern areas, suspended this, and then finished their moult in the wintering areas, whereas others flew straight to the wintering areas before commencing moult. This study shows that stable isotopes can be used to infer migratory strategies of coastal‐feeding shorebirds and provides the basis for identifying the moult strategy and wintering areas of birds passing through Delaware Bay. Coupled with banding and marking birds as individuals, stable isotopes provide a powerful tool for estimating population‐specific demographic parameters and, in this case, further our understanding of the migration systems of the declining Nearctic populations of Red Knot.     

Seasonality in basal metabolic rate and thermal conductance in a long-distance migrant shorebird,the knot (Calidris canutus)

Knots Calidris canutus live highly seasonal lives, breeding solitarily on high arctic tundra and spending the non-breeding season in large social flocks in temperate to tropical estuaries. Their reproductive activities and physiological preparations for long flights are reflected in pronounced plumage and body mass changes, even in long-term captives of the islandica subspecies (breeding in north Greenland and northeast Canada and wintering in western Europe) studied in outdoor aviaries. The three to four fattening episodes in April-July in connection with the flights to and from the high arctic breeding grounds by free-living birds, are represented by a single period of high body mass, peaking between late May and early July in a sample of ten captive islandica knots studied over four years. There are consistent and synchronized annual variations in basal metabolic rate and thermal conductance in three islandica knots. Basal metabolic rate was highest during the summer body mass peak. Within the examined individuals, basal metabolic rate scales on body mass with an exponent of about 1.4, probably reflecting a general hypertrophy of metabolically expensive muscles and organs. Any potential effect of moult on basal metabolic rate was obscured by the large seasonal mass-associated variations. In breeding plumage, insulation (the inverse of thermal conductance) was a factor of 1.35 lower than in winter plumage. This was paralleled by the dry mass of contour feathers being a factor of 1.17 lower. In this subspecies the breeding season is indeed the period during which the costs of thermoregulation are lowest. In captive knots seasonal changes in basal metabolic rate and thermal conductance likely reflect an anticipatory programme adaptive to the variable demands made by the environment at different times of the year.     

Twofold seasonal variation in the supposedly constant, species-specific, ratio of upstroke to downstroke flight muscles in red knots Calidris canutus

We show that in a long-distance migrant shorebird species with outspoken seasonal changes in body mass and composition, the red knot Calidris canutus , the ratio between the masses of the small flight muscle ( musculus supracoracoideus , powering twists and active upstrokes of the wings) and the larger flight muscle ( musculus pectoralis , for the downstrokes) is far from constant. During an annual cycle the supracoracoideus / pectoralis ratio varied more than twofold between values of 0.058 (±0.005 SE) in early winter period and of 0.124 (±0.05 SE) on the High Arctic tundra breeding grounds. The ratios thus spanned a range from those typical of soaring raptors and seabirds to those of fast and agile fliers and birds with rapid take-offs. The overall average ratio was 0.102 (±0.001 SE, for non-starved knots, and 0.103±0.001 including starved knots) and did not differ between males and females. As predicted from the known functions of supracoracoideus and pectoralis , the ratio was a negative function of body mass. However, after arrival on the breeding grounds (0.124) and during winter starvation (0.135) particularly high ratios were reached: these may be times when wing-manoeuvrability (in flight display and during the evasive 'rodent run' away from predators at the nest) and an ability for rapid take-off and active up-strokes (from –near– the nest, and in times of depletion of flight muscle mass during winter starvation) may be at premium. The particularly low ratio of 0.06 in early winter is puzzling. Many aspects of avian phenotypes have recently been shown to be intraindividually variable. To a twofold seasonal variation in flight muscle mass ( Dietz et al. 2007 ), we can now add the twofold variation in the ratio between the muscles for the upstroke and the downstroke.     

Prebreeding moult, plumage and evidence for a presupplemental moult in the Great Knot Calidris tenuirostris

We studied the prebreeding moult and resulting plumage in a long-distance migrant sandpiper (Scolopacidae), the Great Knot Calidris tenuirostris , on the non-breeding grounds (northwest Australia), on arrival at the staging grounds after the first migratory flight (eastern China) and on or near the Russian breeding grounds (Russian data from museum specimens). We show that breeding plumage scores and breast blackness were affected not only by the increase in moulted feathers but also in the wearing down of overlaying pale tips of fresh feathers. Birds migrating from Australia and arriving in China had completed or suspended moult, but more moult must occur in Asia as Russian specimens had moulted more of their mantle and scapular feathers. Russian birds had significantly more red feathering on their upperparts than had birds in Australia or those arriving in China. The increase in reddish feathers cannot by accounted for simply by continuation of the prealternate moult. Instead, a third, presupplemental moult must occur, in which red-marked feathers replace some scapular and especially mantle feathers that were acquired in a prealternate moult only 1–3 months earlier. Great Knot sexes show little size and plumage dimorphism, whereas two other sandpipers that have supplemental plumages (Ruff Philomachus pugnax and Bar-tailed Godwit Limosa lapponica ) are thought to be highly sexually selected. Bidirectional sexual selection may therefore be involved in the evolution of a supplemental plumage in Great Knots.     

A rare case of Plasmodium (Haemamoeba) relictum infection in a free-living Red Knot (Calidris canutus rufa, Scolopacidae)

The Red Knot (Calidris canutus rufa) is a Nearctic migrant shorebird that breeds in the Canadian Arctic and spends the winter season in coastal sites in South America. A rare case of a blood protozoan was found by molecular analyses from an adult bird captured during spring migration at the last refuelling stopover in Delaware Bay USA in 2006. The parasite was identified as Plasmodium relictum belonging to subgenus Haemamoeba based on the shape of meronts, roundish gametocytes, and its position in the erythrocytes from the blood smears examination. A partial cytochrome b sequence was a 100% match to a sequence of Plasmodium relictum, sequence Genbank accession number: id DQ659543.1 (lineage code haplotype P5). This is the first report of avian malaria in a wild individual of C. c. rufa.     

Breeding phased dependent oxidative balance in a small High Arctic seabird,the little auk

Reproduction is a demanding phase of bird's life cycle and imposes various physiological challenges. Increasing oxidative stress (OS) has been proposed as one of the costs associated with reproduction. In this study, we investigate the level of OS in a small Arctic seabird, the little auk Alle alle in relation to sex and phase of breeding (incubation, chick‐rearing). We also examine whether OS is related to the birds leucocyte profile. We expected increase in OS with the progress of breeding period (due to increasing energetic demands) and higher values in females (due to high initial investments in production of a large egg). Surprisingly, we found higher OS during incubation compared to chick‐rearing period, suggesting that incubation is a highly demanding reproductive phase in terms of oxidative balance. We suggests that those changes may be attributed to changes in hormone levels affecting oxidative status. Also, in contrasts to our expectations, we did not find sex differences in OS throughout the studied periods of breeding. Finally, we found positive relationships between OS level and haematological parameters: heterophils/lymphocytes ratio, number of leucocytes and lymphocytes per 10 000 erythrocytes, suggesting the effect of the oxidative stress on the immunological system.     

Skipping‐type migration in a small Arctic wader,the Temminck's stint Calidris temminckii

By using morphometric data and geolocator tracking we investigated fuel loads and spatio‐temporal patterns of migration and non‐breeding in Temminck's stints Calidris temminckii. Body masses in stints captured at autumn stopover sites from Scandinavia to northern Africa were generally not much higher than during breeding and did not vary geographically. Thus, we expected migrating stints to make several stopovers and either circumventing the Sahara desert with low fuel loads or fuelling at north African stopover sites before desert crossing. Geolocation revealed that birds (n = 6) departed their Norwegian breeding site in the last part of July and all but one migrated south‐west over continental western Europe. A single bird headed south‐east to the Balkan Peninsula where the geolocator died. As predicted, southbound migration proceeded in a typical skipping manner with 1–4 relatively short stopovers (median 4 d) during 10–27 d of migration before reaching north‐west Africa. Here birds spent 11–20 d before crossing the Sahara. The non‐breeding sites were located at or near the Niger River in Mali and were occupied continuously for more than 215 d with no indications of itinerancy. Spring migration commenced in late April/early May when birds crossed the desert and used stopover sites in the western Mediterranean basin in a similar manner as during autumn. The lowest body masses were recorded in spring at islands in the central Mediterranean basin, indicating that crossing the Sahara and Mediterranean barriers is exhausting to these birds. Hence, the skipping‐type pattern of migration revealed by geolocators is likely to be natural in this species and not an effect of instrumentation.     

Combined Genetic and Telemetry Data Reveal High Rates of Gene Flow,Migration, and Long-Distance Dispersal Potential in Arctic Ringed Seals (Pusa hispida)

Ringed seals (Pusa hispida) are broadly distributed in seasonally ice covered seas, and their survival and reproductive success is intricately linked to sea ice and snow. Climatic warming is diminishing Arctic snow and sea ice and threatens to endanger ringed seals in the foreseeable future. We investigated the population structure and connectedness within and among three subspecies: Arctic (P. hispida hispida), Baltic (P. hispida botnica), and Lake Saimaa (P. hispida saimensis) ringed seals to assess their capacity to respond to rapid environmental changes. We consider (a) the geographical scale of migration, (b) use of sea ice, and (c) the amount of gene flow between subspecies. Seasonal movements and use of sea ice were determined for 27 seals tracked via satellite telemetry. Additionally, population genetic analyses were conducted using 354 seals representative of each subspecies and 11 breeding sites. Genetic analyses included sequences from two mitochondrial regions and genotypes of 9 microsatellite loci. We found that ringed seals disperse on a pan-Arctic scale and both males and females may migrate long distances during the summer months when sea ice extent is minimal. Gene flow among Arctic breeding sites and between the Arctic and the Baltic Sea subspecies was high; these two subspecies are interconnected as are breeding sites within the Arctic subspecies.     

Population structure,biomass, and diet of landlocked Arctic charr (Salvelinus alpinus) in a small,shallow High Arctic lake

Polar Biology - Although landlocked Arctic charr, Salvelinus alpinus, occur in a large number of High Arctic lakes and often as the only fish species, knowledge of densities and resource use is...     

Darting behavior of a sandburrower fish,Limnichthys nitidus (Creediidae), in the Red Sea

Synopsis Creediid fishes usually live burrowed in sand. When disturbed they make quick movements out and back into the sand. The darting behavior of 35 individualLimnichthys nitidus was recorded on videotape with the aid of SCUBA in the shallow bay, Marsa Muqabelah, Egypt, on the NE coast of Sinai in the Gulf of Aqaba, Red Sea. Thirty-nine sequences show that these fish travel distances of 26.7 to 110 mm in a single dart (1.7 to 5.5 times their total length), can reach speeds of nearly 0.5 m per sec, and execute turns up to 185° to avoid objects in their path. The darting movements, size and cryptic coloration ofL. nitidus make them difficult to see yet they may be the most numerous fish in shallow sandy areas of the Red Sea reaching concentrations of over 50 m^-2, a density exceeded in chordates only by the lancelet,Branchiostoma caribaeum. Their population in Marsa Muqabelah was drastically reduced, along with that of other sand fishes, between 1987 and 1989 when the sand became mostly covered with an overgrowth of algal mats and sea grasses.     

Social Behaviour and Cooperative Breeding in Arctic Foxes,Alopex lagopus (L.), in a Semi-natural Environment

Most canid species show cooperative breeding at least occasionally. The helper-at-the-den system, when extra adults serve as helpers by feeding and guarding the cubs of an alpha pair, has been observed but not studied in any detail in wild Arctic foxes (Alopex lagopus). During a 3-months study of arctic foxes in two enclosures of 4 ha each, we measured the social behaviour during the reproductive season. Older foxes dominated younger ones and males dominated females of the same age. A litter with one surviving cub was born in one enclosure. The alpha male increased his rate of urine marking and barking and fed the alpha female both before and after the birth of the litter. However, about 10 days after the birth, the alpha female died. The cub was fed by his putative father, his sister and his brother (both one year old). The one year old female increased her rate of territorial defence, measured as urine marking and barking, when the mother died. The subordinate females were probably suppressed from breeding by the high aggression levels and territorial defence of the dominant females in each enclosure. The dominant female in the second enclosure came into heat after the death of the alpha female (her mother) in the first enclosure. These changes in behaviour can probably be explained by sexual inhibition by the alpha female while she was present. The significance of territorial defence and dominance, inbreeding avoidance, sexual suppression and evolution of helping behaviour are discussed.     

Plural Breeding in the Splendid Fairy-wren,Malurus splendens (Aves: Maluridae), a Cooperative Breeder

Groups of the cooperatively breeding splendid fairy-wren Malurus splendens may include more than one female. Previously this species has been described as singular breeding (only one female breeds). This paper describes the occurrence of plural breeding (PB) groups in 10% of group years, in which two females had separate nests. In all cases, the secondary female (Y) was related to the primary breeding female (X) and was generally a 2-year old female which had helped in the group during the previous breeding season. Plural breeding was correlated with an increase in population density and in the number of female helpers; PB groups were larger than singular-breeding groups. In most cases, the X female was occupied with her own nest or offspring when the Y female began to nest, and there was no aggression between them. Which birds helped the Y female to feed at her nest depended on the time between the hatching of the two nests. If the interval was small, some group members helped at each nest; with longer intervals, the group members began to feed at the earlier nest, and the other female was left to raise her brood alone. Female helpers were very active in feeding at single nestings, and the cost to an X female of a Y female breeding was mainly a loss of this assistance. The success of individual X nests was not affected. Effects on productivity were slight, but fewer X females in PB groups raised second broods than did experienced singular breeding females. Y females were less productive than X females, but no less productive than singular breeding novice females without helpers. It is not known whether Y females copulated with primary or secondary males within their group, or with males from outside the group. Certainly, they did not form an observable pairing with any male in the group. Plural breeding occurred in a minority of group years in response to extrinsic conditions and the current demographic situation, and shows the extreme plasticity of the mating system in M. splendens.     

Identification of Beijerinckia in the High Arctic (Devon Island, Northwest Territories)

Although generally considered, with few exceptions, to be restricted to the acidic, tropical soils of the southern hemisphere, Beijerinckia species, resembling B. indica, were found at three sites on Devon Island (75°33′N, 84°40′W) in the Canadian Arctic.     

Population structure, ecological segregation and reproduction in non-anadromous Arctic charr, Salvelinus alpinus (L), in four unexploited lakes in the Canadian high Arctic

Land-locked populations of Arctic charr in four lakes on Northern Ellesmere Island (80° N) were found to consist of two distinct sizes: 'dwarf' and 'normal' charr. Both groups attained sexual maturity but differed in appearance and habitat. The smaller fish, occupying the more marginal habitats, retained their parr-markings; the larger group had the general characteristics of smolts, being more silvery and without parr-marks. In their juvenile stages, the charr destined to attain the larger group were indistinguishable from members of the smaller group. Although fish in the larger group were capable of cannibalism, this was rarely observed. In general, the fish in the larger group were older than the smaller ones but great variation in size at a given age resulted in certain age classes containing representatives of both groups. The population structure varied considerably between lakes; a high proportion of 'normal' charr correlated well with a high growth rate in the first few years of development. It is postulated that the two groups live in dynamic equilibrium where the advantages of progenesis (retention of juvenile characters by adults) in the smaller type are traded against the larger proportion of the energy resources available to the larger type. The concept of heterochrony in an ecological setting is introduced.     

Tetrameres (Tetrameres) megaphasmidiata n. sp. (Nematoda: Tetrameridae), a parasite of the two-banded plover, Charadrius falklandicus, and white-rumped sandpiper, Calidris fuscicollis, from Patagonia, Argentina

Tetrameres (Tetrameres) megaphasmidiata n. sp. is described from the proventriculus of the two-banded plover, Charadrius falklandicus, and the white-rumped sandpiper, Calidris fuscicollis, from Patagonia, Argentina. The new species shares with T. (T.) nouveli, T. (T.) paradisea, T. (T.) prozeskyi, T. paraaraliensis, T. (T.) cladorhynchi, and T. lobybicis the absence of the right spicule and the presence of 4 rows of somatic spines. Tetrameres (T.) megaphasmidiata n. sp. differs from the first 4 species mainly by its longer left spicule. The new species can be distinguished from T. (T.) cladorhynchi by the extension of the lateral alae, the number and arrangement of the caudal papillae, and the absence of polar filaments in the eggs. Tetrameres lobybicis differs from the new species by having shorter rows of dorsal spines and a different number and arrangement of the caudal papillae. This report is the first record of a species of Tetrameres in C. falklandicus and C. fuscicollis.     

Diet and population size structure of the Arctic shanny,Stichaeus punctatus (Pisces: Stichaeidae), at sites in eastern Newfoundland and the eastern Arctic

Synopsis The size structure and diet of the Arctic shanny,Stichaeus punctatus, were studied at two sites, one at Nuvuk Islands in the Arctic, and one at Newman Sound, in eastern Newfoundland. The Newfoundland population showed peaks corresponding to 0+,1+, and an accumulation of older individuals, while the Nuvuk population was dominated by 2+ and 3+ combined, and individuals aged 6+ and older. Epiphytic, harpacticoid copepods were the most important food items for the smaller Arctic shanny, but the importance of harpacticoid copepods decreased with increasing fish size. Harpacticoid copepods are thus a link between the macro-algae and the young stages of the Arctic shanny. Since Arctic shanny are the main food used by black guillemots to feed chicks, these copepods are also an indirect link between benthic algal production and these seabirds.     

The Breeding Ecology of a Critically Endangered Salamander,Hynobius amjiensis(Caudata:Hynobiidae),Endemic to Eastern China

Hynobius amjiensis is a critically endangered salamander species(IUCN Red List) endemic to eastern China. It currently has three known populations: one in Longwangshan, Zhejiang Province(type locality), and two in Qingliangfeng between Anhui and Zhejiang Provinces. We examined the relatively unstudied breeding ecology of this species in the field and at laboratory from March 2007 to May 2014. Adult males and females were year-round terrestrial, except for the February–April breeding season. During this period, we captured only a total of 16 breeding adults(11 males and 5 females). As few as 100 breeding females were estimated based on the number of egg sacs observed since 2007. This number was significantly reduced from the estimated number between 1992 and 1998. Males(mean total length = 16.21 cm, mean body mass = 18.8 g) were slightly smaller than females(16.51 cm, 19.2 g). Size of breeding pools ranged from 0.2 m~2 to 1.2 m~2(0.1–1.2 m depths). Each female deposits a pair of egg sacs by attaching the adhesive tips of the sacs to aquatic plants or dead twigs. Fifteen pairs of egg sacs had an average length of 28.6 cm and a diameter of 3.3 cm. On average, each egg sac contained 75 eggs with a diameter of 0.3 cm. Our field survey revealed that H. amjiensis used oviposition sites in small, cool, and weakly acidic pools at high elevations(1 300–1 600 m) where peat moss was abundant. Reduction in wetland size and disappearance of suitable breeding pools suggest that this salamander species is under threat of extinction, particularly at Longwangshan, where 5 of the 9 breeding pools have either dried up or disappeared. Combined size of the remaining 4 pools is less than 2 m~2. We urge immediate implementation of more effective conservation measures and suggest that preservation priority should be given to habitat that contains suitable breeding pools.     

Apogon smithi (Kotthaus, 1970) (Teleostei: Apogonidae), a Red Sea cardinalfish colonizing the Mediterranean Sea

Smith's cardinalfish Apogon smithi is recorded for the first time in the Mediterranean Sea. Six specimens were caught by commercial trawler north of Jaffa, Israel, at depths of 40 m and the relatively high number of specimens suggests that A. smithi has established a sustainable population. This is the third Indo-Pacific apogonid species documented in the Mediterranean Sea, and is assumed to be a migrant originating from the Red Sea.