The scaling of diving time budgets: insights from an optimality approach

Simple scaling arguments suggest that, among air-breathing divers, dive duration should scale approximately with mass to the one-third power. Recent phylogenetic analyses appear to confirm this. The same analyses showed that duration of time spent at the surface between dives has scaling very similar to that of dive duration, with the result that the ratio of dive duration to surface pause duration is approximately mass invariant. This finding runs counter to other arguments found in the diving literature that suggest that surface pause duration should scale more positively with mass, leading to a negative scaling of the dive-pause ratio. We use a published model of optimal time allocation in the dive cycle to show that optimal decisions can predict approximate mass invariance in the dive-pause ratio, especially if metabolism scales approximately with mass to the two-thirds power (as indicated by some recent analyses) and oxygen uptake is assumed to have evolved to supply the body tissues at the required rate. However, emergent scaling rules are sensitive to input parameters, especially to the relationship between the scaling of metabolism and oxygen uptake rate at the surface. Our results illustrate the utility of an optimality approach for developing predictions and identifying key areas for empirical research on the allometry of diving behavior.     

Resource limitation of diving ducks at Myvatn: Food limits production

We monitored numbers and production of diving ducks on Lake Myvatn and its outlet river, the Laxá, over a period of 25 years, 1975–1999. Aquatic insects, a major food source, were monitored for 23 years, 1977–1999, using window traps. In this paper, we focus on the relationship between the demography of five species of diving ducks and the abundance of aquatic insect species, mainly Chironomidae and Simuliidae. Production of ducks was estimated when most young were 2–4 weeks old, either as young per female (Aythya marila and Aythya fuligula) or as total young found (Melanitta nigra, Bucephala islandica, Histrionicus histrionicus). Variation between years in production of young ducks was strongly correlated with variation in the abundance of aquatic insects. Procladius islandicus and Psectrocladius barbimanus, both of which are univoltine with peak emergence in late July (when many young ducks hatch), as well as the bivoltine Orthocladius consobrinus, showed particularly high correlations with duckling numbers. In one year, 1992, catastrophic weather depressed production. Food appeared to determine production, primarily through its availability to the young. Changes in spring density of breeding ducks of all five species were negatively associated with density in the year before. Resources, estimated as food availability or reproductive success in the year before, were positively correlated with change in breeding numbers in A. marila, A. fuligula and B. islandica. Production three years earlier had a weak but significant effect on change in numbers of M. nigra.     

Movements and diving behavior of pelagic spotted dolphins

We radio tracked 20 pantropical spotted dolphins ( Stenella attenuata ) in the pelagic waters of the eastern tropical Pacific Ocean during three research cruises in 1992, 1993, and 2001. Dolphins were tracked for up to 6 d from 20 to 1,000 km offshore. Seventeen of these dolphins also carried time–depth recorders, nine of which were recovered with a combined total of 477 h of dive data. The movement data suggested that the dolphins associate with areas of relatively high biological productivity. Dolphins close to shore moved along the continental slope, while some dolphins traveled farther offshore along thermocline "ridges." Tracking and tagging multiple dolphins demonstrated the dynamic nature of dolphin herds, which change size and membership over the course of a day. At night, the dolphins traveled more slowly but dove deeper and longer, with more rapid ascents and descents than during daylight hours. These nighttime dive characteristics support the hypothesis suggested by some food habit studies that spotted dolphins are nocturnal feeders. Comparison of dive data with acoustic backscatter data indicates that spotted dolphins typically begin and end nighttime feeding activities with dusk and dawn diving bouts that track the vertical migration of organisms associated with the deep scattering layer.     

Movements of wintering surf scoters: predator responses to different prey landscapes

The distribution of predators is widely recognized to be intimately linked to the distribution of their prey. Foraging theory suggests that predators will modify their behaviors, including movements, to optimize net energy intake when faced with variation in prey attributes or abundance. While many studies have documented changes in movement patterns of animals in response to temporal changes in food, very few have contrasted movements of a single predator species naturally occurring in dramatically different prey landscapes. We documented variation in the winter movements, foraging range size, site fidelity, and distribution patterns of a molluscivorous sea duck, the surf scoter (Melanitta perspicillata), in two areas of coastal British Columbia with very different shellfish prey features. Baynes Sound has extensive tidal flats with abundant clams, which are high-quality and temporally stable prey for scoters. Malaspina Inlet is a rocky fjord-like inlet where scoters consume mussels that are superabundant and easily accessible in some patches but are heavily depleted over the course of winter. We used radio telemetry to track surf scoter movements in both areas and found that in the clam habitats of Baynes Sound, surf scoters exhibited limited movement, small winter ranges, strong foraging site fidelity, and very consistent distribution patterns. By contrast, in mussel habitats in the Malaspina Inlet, surf scoters displayed more movement, larger ranges, little fidelity to specific foraging sites, and more variable distribution patterns. We conclude that features associated with the different prey types, particularly the higher depletion rates of mussels, strongly influenced seasonal space use patterns. These findings are consistent with foraging theory and confirm that predator behavior, specifically movements, is environmentally mediated.     

Variation in home-range size and movements of wintering dabbling ducks

Despite a long historical record of radio-tracking analyses, basic home-range information is still lacking for most common waterfowl species, especially during the winter. We investigated how dabbling duck home ranges and daily foraging movements are influenced by extrinsic (site, temperature, date) and intrinsic factors (species, sex, age). We radio-tagged and monitored 125 individuals of three duck species (mallard Anas platyrhynchos, Eurasian teal A. crecca crecca and northern pintail A. acuta) in three French wetlands over four winters. Home-range sizes for a given species varied greatly among our study sites. Moreover, species differed according to home-range structure and distance traveled to reach their foraging grounds (teal had a more patchy home range and traveled farther distances than mallards). Foraging distances increased with temperature and time (over the winter season), but this effect differed among species, suggesting that they behave differently in response to food depletion and/or cold weather. The commuting behavior (i.e., the decision to leave the roost at night for foraging) differed among species and season. Teals were more risk-prone because they were more likely to leave the roost at night. In our study, ducks foraged at distances of 1–2 km from roosts, whereas distances of 2–48 km have been recorded in North America. We suggest that food supply, hunting pressure or population density may account for these inter-continental differences.     

Sitting ducks: diet of wintering wildfowl in Lake Tonga,northeast Algeria

Dietary studies of five species of Anatidae (Gadwall Anas strepera, Shoveler Anas clypeata, Pochard Aythya ferina, Ferruginous Duck Aythya nyroca, and White-headed Duck Oxyura leucocephala) were carried out at Lake Tonga, northeast Algeria, from December 2011 to March 2012. Diet analysis of all five wintering species indicated a heavy reliance on seeds, which included a wide variety of aquatic plants, such as Myriophyllum spicatum, Ceratophyllum demersum, Sparganium erectum, Scirpus lacustris and Potamogeton pectinatus. Noteworthy is the paucity of ingested macroinvertebrates. Despite dietary overlap, the type, abundance and frequency of food consumed clearly indicated differences in resource use among the ducks. Incidentally, the study also highlighted the high vulnerability of threatened ducks to poaching (harvesting and disturbance) in a protected area. We discuss the implications of these findings for the management and conservation of wintering wildfowl and their North African habitats.     

The diving paradox: new insights into the role of the dive response in air-breathing vertebrates

When aquatic reptiles, birds and mammals submerge, they typically exhibit a dive response in which breathing ceases, heart rate slows, and blood flow to peripheral tissues is reduced. The profound dive response that occurs during forced submergence sequesters blood oxygen for the brain and heart while allowing peripheral tissues to become anaerobic, thus protecting the animal from immediate asphyxiation. However, the decrease in peripheral blood flow is in direct conflict with the exercise response necessary for supporting muscle metabolism during submerged swimming. In free diving animals, a dive response still occurs, but it is less intense than during forced submergence, and whole-body metabolism remains aerobic. If blood oxygen is not sequestered for brain and heart metabolism during normal diving, then what is the purpose of the dive response? Here, we show that its primary role may be to regulate the degree of hypoxia in skeletal muscle so that blood and muscle oxygen stores can be efficiently used. Paradoxically, the muscles of diving vertebrates must become hypoxic to maximize aerobic dive duration. At the same time, morphological and enzymatic adaptations enhance intracellular oxygen diffusion at low partial pressures of oxygen. Optimizing the use of blood and muscle oxygen stores allows aquatic, air-breathing vertebrates to exercise for prolonged periods while holding their breath.     

Modelling pink-footed goose (Anser brachyrhynchus) wintering distributions for the year 2050: potential effects of land-use change in Europe

Feeding on farmland by overwintering populations of pink-footed geese ( Anser brachyrhynchus ) conflicts with agricultural interests in Northern Europe. In order to forecast the potential future of this conflict, we used generalized linear models to relate the presence and absence of pink-footed geese to variables describing the contemporary landscape, and predicted their future distributions in relation to two land-use scenarios for the year 2050. One future scenario represented a global, economically orientated world (A1) and the other represented a regional, environmentally concerned world (B2). The probability of goose occurrence increased within cropland and grassland, and could be explained by their proximity to coast, elevation, and the degree of habitat closure. Predictions to future scenarios revealed noticeable shifts in the suitability of goose habitat evident at the local and regional scale in response to future shifts in land use. In particular, as grasslands and croplands give way to unsuitable land-use types (e.g. woody biofuel crops, increased urbanization, and forest) under both future scenarios, our models predicted a decrease in habitat suitability for geese. If coupled with continued goose population expansion, we expect that the agricultural conflict will intensify under the A1 and particularly the B2 scenarios.     

Characteristics of foraging sites and protein status in wintering muskoxen: insights from isotopes of nitrogen

Identifying links between nutritional condition of individuals and population trajectories greatly enhances our understanding of the ecology, conservation, and management of wildlife. For northern ungulates, the potential impacts of a changing climate to populations are predicted to be nutritionally mediated through an increase in the severity and variance in winter conditions. Foraging conditions and the availability of body protein as a store for reproduction in late winter may constrain productivity in northern ungulates, yet the link between characteristics of wintering habitats and protein status has not been established for a wild ungulate. We used a non‐invasive proxy of protein status derived from isotopes of N in excreta to evaluate the influence of winter habitats on the protein status of muskoxen in three populations in Alaska (2005–2008). Multiple regression and an information‐theoretic approach were used to compare models that evaluated the influence of population, year, and characteristics of foraging sites (components of diet and physiography) on protein status for groups of muskoxen. The observed variance in protein status among groups of muskoxen across populations and years was partially explained (45%) by local foraging conditions that affected forage availability. Protein status improved for groups of muskoxen as the amount of graminoids in the diet increased (?0.430 ± 0.31, β± 95% CI) and elevation of foraging sites decreased (0.824 ± 0.67). Resources available for reproduction in muskoxen are highly dependent upon demographic, environmental, and physiographic constraints that affect forage availability in winter. Due to their very sedentary nature in winter, muskoxen are highly susceptible to localized foraging conditions; therefore, the spatial variance in resource availability may exert a strong effect on productivity. Consequently, there is a clear need to account for climate–topography effects in winter at multiple scales when predicting the potential impacts of climatic shifts on population trajectories of muskoxen.     

Habitat selection by postbreeding female diving ducks: influence of habitat attributes and conspecifics

Habitat selection studies of postbreeding waterfowl have rarely focused on within‐wetland attributes such as water depth, escape cover, and food availability. Flightless waterfowl must balance habitat selection between avoiding predation risks and feeding. Reproductively successful female ducks face the greatest challenges because they begin the definitive prebasic molt at or near the end of brood rearing, when their body condition is at a low point. We assessed the relative importance of habitat attributes and group effects in habitat selection by postbreeding female lesser scaup Aythya affinis on a 2332‐ha montane wetland complex during the peak flightless period (August) over seven years. Hypothesis‐based habitat attributes included percent open water, open water:emergent edge density, water depth, percent flooded bare substrate, fetch (distance wind can travel unobstructed), group size, and several interactions representing functional responses to interannual variation in water levels. Surveys of uniquely marked females were conducted within randomly ordered survey blocks. We fitted two‐part generalized linear mixed‐effects models to counts of marked females within survey blocks, which allowed us to relate habitat attributes to relative probability of occurrence and, given the presence of a marked female, abundance of marked individuals. Postbreeding female scaup selected areas with water depths > 40 cm, large open areas, and intermediate edge densities but showed no relation to flooded bare substrate, suggesting their habitat preferences were more influenced by avoiding predation risks and disturbances than in meeting foraging needs. Grouping behavior by postbreeding scaup suggests habitat selection is influenced in part by behavioral components and/or social information, conferring energetic and survival benefits (predation and disturbance risks) but potentially also contributing to competition for food resources. This study demonstrates the importance of incorporating group effects and interannual variability in habitat conditions when investigating habitat selection, particularly for seasons when waterfowl are aggregated.     

西藏雅鲁藏布江中游斑头雁的越冬种群数量、分布和活动区

斑头雁(Anser indicus)是高致病性禽流感病毒的易感鸟类和潜在的传播源,在禽流感防控中占有重要地位。有关斑头雁的越冬种群现状缺乏研究。2009年1月,在雅鲁藏布江中游共统计到越冬斑头雁44657只,主要分布于一江两河地带的林周县、日喀则市、白朗县、拉孜县、江孜县和贡嘎县境内。这一统计数据远高于1990年代的统计数量,由此可将其全球种群数量估计值由5.2-6万只修正为至少7万只。越冬斑头雁的平均集群大小为(208±262)只(n=215),主要栖息环境包括冬小麦田、冬歇期农田、河流湖泊和沼泽湿地,其中在冬歇期农田中统计到的数量占72.1%。2006-2008年,对青海湖繁殖斑头雁的卫星跟踪表明,其越冬地点主要位于西藏雅鲁藏布江流域、拉萨河流域和印度,越冬期为11月至翌年的3月份,在越冬地平均停留(108±30)d,越冬活动区大小为(122.22±124.94)km2(n=3)。受西藏独特的宗教传统和农耕畜牧结合的生产方式的影响,越冬斑头雁经常与大量家畜和水鸟混杂在农田觅食,在禽流感疫情防控时应引起重视。     

GnRH deficiency: new insights from genetics

The acquisition of a sexually dimorphic phenotype is a critical event in mammalian development. Hypogonadotropic hypogonadism (HH) results from impaired secretion of GnRH. The patients display with delayed puberty, micropenis and cryptorchidism in the male reflecting gonadotropin insufficiency, and amenorrhea in the female. Kallmann's syndrome (KS) is defined by the association of HH and anosmia or hyposmia (absent smelling sense). Segregation analysis in familial cases has demonstrated diverse inheritance patterns, suggesting the existence of several genes regulating GnRH secretion. The X-linked form of the disease was associated with a genetic defect in the KALI gene located on the Xp22.3 region. KAL1 gene encodes an extracellular matrix glycoprotein anosmin-1, which facilitates neuronal growth and migration. Abnormalities in the migratory processes of the GnRH neurons with the olfactory neurons explain the association of HH with anosmia. Recently, mutations in the FGF recepteur 1 (FGFR1) gene were found in KS with autosomal dominant mode of inheritance. The role of FGFR1 in the function of reproduction requires further investigation. Besides HH with anosmia, there are isolated HH (IHH). No human GnRH mutations have been reported although hypogonadal mice due to a GnRH gene deletion exist. In patients with idiopathic HH and without anosmia an increasing number of GnRH receptor (GnRHR) mutations have been described which represent about 50% of familial cases. The clinical features are highly variable and there is a good relationship between genotype and phenotype. A complete loss of function is associated with the most severe phenotype with resistance to pulsatile GnRH treatment, absence of puberty and cryptorchidism in the male. In contrast, milder loss of function mutations causes incomplete failure of pubertal development. The preponderant role of GnRH in the secretion of LH by the gonadotrophs explains the difference of the phenotype between male and female with partial GnRH resistance. Affected females can have spontaneous telarche and normal breast development while affected males exhibit no pubertal development but normal testis volume, a feature described as "fertile-eunuch". High-dose pulsatile GnRH has been used to induce ovulation. Another gene, called GPR54, responsible for idiopathic HH has been recently described by segregation analysis in two different consanguineous families. The GPR54 gene is an orphan receptor, and its putative ligand is the product of the KISS-1 gene, called metastine. Their roles in the function of reproduction are still unknown.     

Geographical origin of dabbling ducks wintering in Iberia: sex differences and implications for pair formation

Natural and anthropogenic Iberian wetlands in southern Europe are well known for supporting large numbers of migratory Palaearctic waterbirds each winter. However, information on the geographical origin of dabbling ducks overwintering in these wetlands is scarce and mostly limited to data from ringing recoveries. Here, we used intrinsic isotopic markers to determine the geographical origin of male and female Northern Pintails Anas acuta and Eurasian Teal Anas crecca in Extremadura, inland Iberia, a key site for overwintering dabbling ducks. Additionally, we fitted six Northern Pintails with GPS‐GSM tags to complement the data derived from stable isotope analysis. Most (> 70%) first calendar‐year Northern Pintails were assigned to regions above 55°N, flying 2600–5600 km from their main natal regions to Extremadura. Mean values of δ²H_f varied significantly between male and female Northern Pintails, suggesting that the sexes had different geographical origins. Data from tagged adult Northern Pintails supported the isotopic data, one male flying more than 5000 km to the coast of the Pechora Sea (Russia). Most (> 70%) first calendar‐year Eurasian Teal were assigned to the region between 48° and 60°N, travelling 1500–4500 km to arrive in Extremadura. Male and female Eurasian Teal showed marginal differences in mean values of δ²H_f. In migratory dabbling ducks, pairing typically occurs on the wintering grounds, and ducks in their first winter can breed the following spring. For Northern Pintails, pair formation in Extremadura could occur between individuals with different geographical origins, which could contribute to the genetic variability of their offspring.     

Profilicollis botulus (Van Cleave, 1916) from diving ducks and shore crabs of British Columbia

Adults of Profilicollis botulus were found in 6 species of diving ducks in British Columbia including 3 new hosts: common goldeneye, Bucephala clangula (L.); Barrow's goldeneye, B. islandica (Gmelin); and greater scaup, Aythya marila (L.). The identification of the species was verified by the examination of co-types and specimens from eider ducks, Somateria mollissima (L.), from Scotland and oldsquaw, Clangula hyemalis (L.), from New Brunswick. Cystacanths from the hairy shore crab, Hemigrapsus oregonensis (Dana), were similar in morphology to those from Carcinus maenas (L.) from Scotland.