Sex‐specific winter distribution in a sexually dimorphic shorebird is explained by resource partitioning

Sexual size dimorphism (SSD) implies correlated differences in energetic requirements and feeding opportunities, such that sexes will face different trade‐offs in habitat selection. In seasonal migrants, this could result in a differential spatial distribution across the wintering range. To identify the ecological causes of sexual spatial segregation, we studied a sexually dimorphic shorebird, the bar‐tailed godwit Limosa lapponica, in which females have a larger body and a longer bill than males. With respect to the trade‐offs that these migratory shorebirds experience in their choice of wintering area, northern and colder wintering sites have the benefit of being closer to the Arctic breeding grounds. According to Bergmann's rule, the larger females should incur lower energetic costs per unit of body mass over males, helping them to winter in the cold. However, as the sexes have rather different bill lengths, differences in sex‐specific wintering sites could also be due to the vertical distribution of their buried prey, that is, resource partitioning. Here, in a comparison between six main intertidal wintering areas across the entire winter range of the lapponica subspecies in northwest Europe, we show that the percentage of females between sites was not correlated with the cost of wintering, but was positively correlated with the biomass in the bottom layer and negatively with the biomass in the top layer. We conclude that resource partitioning, rather than relative expenditure advantages, best explains the differential spatial distribution of male and female bar‐tailed godwits across northwest Europe.     

Spatially Extended Habitat Modification by Intertidal Reef-Building Bivalves has Implications for Consumer-Resource Interactions

Ecosystem engineers can strongly modify habitat structure and resource availability across space. In theory, this should alter the spatial distributions of trophically interacting species. In this article, we empirically investigated the importance of spatially extended habitat modification by reef-building bivalves in explaining the distribution of four avian predators and their benthic prey in the Wadden Sea—one of the world’s largest intertidal soft-sediment ecosystems. We applied Structural Equation Modeling to identify important direct and indirect interactions between the different components of the system. We found strong spatial gradients in sediment properties into the surrounding area of mixed blue mussel (Mytilus edulis) and Pacific oyster (Crassostrea gigas) reefs, indicating large-scale (100s of m) engineering effects. The benthic community was significantly affected by these gradients, with the abundance of several important invertebrate prey species increasing with sediment organic matter and decreasing with distance to the reefs. Distance from the reef, sediment properties, and benthic food abundance simultaneously explained significant parts of the distribution of oystercatchers (Haematopus ostralegus), Eurasian curlews (Numenius arquata), and bar-tailed godwits (Limosa lapponica). The distribution of black-headed gulls (Chroicocephalus ridibundus)—a versatile species with many diet options—appeared unaffected by the reefs. These results suggest that intertidal reef builders can affect consumer-resource dynamics far beyond their own boundaries, emphasizing their importance in intertidal soft-bottom ecosystems like the Wadden Sea.     

How do red knots Calidris canutus leave Northwest Australia in May and reach the breeding grounds in June? Predictions of stopover times,fuelling rates and prey quality in the Yellow Sea

In general, Arctic-breeding waders leave non-breeding grounds in Australasia from March (New Zealand) to mid-April (Northwest Australia). Here we provide evidence from radio-tracking and visual observations that many red knots Calidris canutus do not leave Roebuck Bay, Northwest Australia, until early or mid-May. Late-departing red knots probably belong to the subspecies piersmai , which breeds on the New Siberian Islands, 10,400 km from Northwest Australia. Based on comparisons of temperatures on the breeding grounds of different knot subspecies, we predict that piersmai knots would not arrive on the breeding grounds until early June, leaving at most 3–4 weeks refuelling in Asia. Using a model of fuelling capacity in relation to prey quality and gizzard mass, we show that these knots must fuel very differently in Australia and Asia. In Australia, knots have seemingly suboptimal gizzard sizes and deposit fuel slowly. In the Yellow Sea, birds could only fuel up within the available time if they either enlarged their gizzards substantially or encountered prey qualities much higher than in Australia, for which we provide quantitative predictions.     

Scale dependence of diversity measures in a leaf-litter ant assemblage

A reliable characterization of community diversity and composition, necessary to allow inter‐site comparisons and to monitor changes, is especially difficult to reach in speciose invertebrate communities. Spatial components of the sampling design (sampling interval, extent and grain) as well as temporal variations of species density affect the measures of diversity (species richness S, Buzas and Gibson's evenness E and Shannon's heterogeneity H). Our aim was to document the small‐scale spatial distribution of leaf litter ants in a subtropical dry forest of the Argentinian Chaco and analyze how the community characterization was best achieved with a minimal sampling effort. The work was based on the recent standardized protocol for collecting ants of the leaf litter (“A.L.L.”: 20 samples at intervals of 10 m). To evaluate the consistency of the sampling method in time and space, the selected site was first subject to a preliminary transect, then submitted after a 9‐month interval to an 8‐fold oversampling campaign (160 samples at interval of 1.25 m). Leaf litter ants were extracted from elementary 1 m² quadrats with Winkler apparatus. An increase in the number of samples collected increased S and decreased E but did not affect much H. The sampling interval and extent did not affect S and H beyond a distance of 10 m between samples. An increase of the sampling grain had a similar effect on S than a corresponding increase of the number of samples collected, but caused a proportionaly greater increase of H. The density of species m^?2 varied twofold after a 9‐month interval; the effect on S could only be partially corrected by rarefaction. The measure of species numerical dominance was little affected by the season. A single standardized A.L.L. transect with Winkler samples collected <45% of the species present in the assemblage. All frequent species were included but their relative frequency was not always representative. A log series distribution of species occurrences was oberved. Fisher's α and Shannon's H were the most appropriate diversity indexes. The former was useful to rarefy or abundify S and the latter was robust against sample size effects. Both parametric and Soberón and Llorente extrapolation methods outperformed non‐parametric methods and yielded a fair estimate of total species richness along the transect, a minimum value of S for the habitat sampled.     

Detection of earthworm prey by Ruff Philomachus pugnax

Ruff Philomachus pugnax staging in the Netherlands forage in agricultural grasslands, where they mainly eat earthworms (Lumbricidae). Food intake and the surface availability of earthworms were studied in dairy farmland of southwest Friesland in March–April 2011. Daily changes in earthworm availability were quantified by counting visible earthworms. No earthworms were seen on the surface during daytime, but their numbers sharply increased after sunset and remained high during the night. Nevertheless, intake rates of individual Ruff in different grasslands measured during daytime showed the typical Holling type II functional response relationship with the surfacing earthworm densities measured at night. Radiotagging of Ruff in spring 2007 revealed that most, if not all, feeding occurs during the day, with the Ruff assembling at shoreline roosts at night. This raises the question of why Ruff do not feed at night, if prey can be caught more easily than during daytime. In March–May 2013 we experimentally examined the visual and auditory sensory modalities used by Ruff to find and capture earthworms. Five males were kept in an indoor aviary and we recorded them individually foraging on trays with 10 earthworms mixed with soil under various standardized light and white noise conditions. The number of earthworms discovered and eaten by Ruff increased with light level, but only when white noise was played, suggesting that although they can detect earthworms by sight, Ruff also use auditory cues. We suggest that although surfacing numbers of earthworms are highest during the night, diurnal intake rates are probably sufficient to avoid nocturnal foraging on a resource that is more available but perhaps less detectable at that time.     

Does wintering north or south of the Sahara correlate with timing and breeding performance in black‐tailed godwits?

Migrating long distances requires time and energy, and may interact with an individual's performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.     

Carrying capacity models should not use fixed prey density thresholds: a plea for using more tools of behavioural ecology

Earlier studies have developed models of carrying capacity to predict the number of animals a certain area can support. These models assume that resources are not renewed after consumption ('standing stock' models), and that the initial number of prey and the rate of prey consumption determine the time a population of foragers can live in an area. Within such areas, foragers give up feeding at a sub-site or patch when intake rates no longer cover energy expenditure. To improve the success rate of the models' predictions, we here change the existing rate-maximising models into fitness-maximising models, and include dynamics in the availability of patches. These new (conceptual) models show that the approaches used so far may over- as well as underestimate carrying capacity. We review empirical studies that have aimed to estimate carrying capacity, and discuss how concepts have been confused. We make explicit suggestions on how to proceed in predicting carrying capacities in future studies.     

Do body condition and plumage during fuelling predict northwards departure dates of Great Knots Calidris tenuirostris from north‐west Australia?

It is often assumed that strong selection pressures give rise to trade-offs between body condition and time in long-distance migrating birds. Birds that are 'behind schedule' in fuel deposition or moult should delay departure, and this should result in a negative correlation between initial condition and departure date. We tested this hypothesis in the Great Knot Calidris tenuirostris migrating from north-west Australia to eastern Asia en route to Siberia. Great Knot gain mass and moult into breeding plumage before leaving northern Australia in late March and early April, and fly 5400–6000 km to eastern China and Korea. We radiotracked 27 individuals (17 males and ten females) to determine departure dates; 23 migrated and four remained in Australia. We characterized body condition at capture using body mass, predicted pectoral muscle mass (based on ultrasound estimates of the size of the pectoral muscles) and breeding plumage scores. Residual condition indices were uncorrelated, indicating that at the individual level, variation in one fuelling component was not strongly associated with variation in the other components. Birds that did not depart had lower residual body mass and breeding plumage indices than those that did migrate; these four birds may have been subadults. Neither sex, size nor the condition indices explained variation in departure date of migrants. Reasons for this are explored. Departure dates for northward migrating waders indicate that the migration window (span over which birds depart) decreases with proximity to the northern breeding grounds. We suggest that migration schedules become tighter as birds get nearer to the breeding grounds. Thus the lack of a relationship between condition and departure date in Great Knots may reflect the fact that the departure episode under study is the first one in sequence and is still 4–8 weeks before breeding.     

Competitive food exploitation of smelt Osmerus eperlanus by great crested grebes Podiceps cristatus and perch Perca fluviatilis at Lake IJsselmeer,The Netherlands

The impact of predation by an avian predator (great crested grebe Podiceps cristatus) and a predatory fish (perch Perca fluviatilis) on a common resource, the small planktivorous fish smelt (Osmerus eperlanus) was studied in Lake IJsselmeer, The Netherlands, in July–October 1985–1988. At this time of the year the grebes are in simultaneous wing-moult and therefore temporarily flightless. Four factors limit the predation pressure by grebes: (1) an underwater visibility threshold of c. 40 cm Secchi depth is the lower limit at which large numbers attend the moulting site, (2) a lower density threshold of exploitable smelt biomass of c. 30 kg/ha determines the actual fishing areas, (3) a fish size threshold of 6.5 cm total length further limits prey availability, and (4) the vertical movements of the prey impose a strong (diel) time constraint on the birds, allowing only crepuscular foraging. Perch, the other main predator in this system, is less restricted in its foraging. Perch were found to be in direct competition with the grebes, preying heavily on the juvenile smelt. Over the 4 years of study 90% of all predation from 15 July to 15 October (72–233 kg/ha) was due to perch. The high level of predation by birds and fish was only possible due to a continuous immigration of smelt into the area (1.45 kg/ha per day). In three of the four years, however, the combined predation by fish and birds surpassed the immigration rate of the prey, which led to a strong reduction in smelt stocks in the study area. A conceptual model is developed to describe the different sets of constraints on the grebes' foraging. First, the state of eutrophication in relation to the weather condition determines the degree of algal blooms, and thus underwater visibility, in late summer. This is the major factor governing the numbers of grebes on the moulting area. Second, the size of the population of predatory fish determines the overall food availability (biomass and size distribution of smelt). These factors are partly interconnected and related to human action (pollution, fisheries). It is suggested that, despite deteriorating visibility conditions, the largest moulting site for grebes in Europe at Lake IJsselmeer exists because the stocks of predatory fish are kept low by overfishing.     

Metabolic profile of long-distance migratory flight and stopover in a shorebird

Migrating birds often complete long non-stop flights during which body energy stores exclusively support energetic demands. The metabolic correlates of such long-distance travel in free-living migrants are as yet poorly studied. Bar-tailed godwits, Limosa lapponica taymyrensis, undertake a 4500 km flight to their single spring stopover site and thus provide an excellent model in which to determine the energy fuels associated with endurance travel. To this end, we evaluated plasma concentrations of six key metabolites in arriving godwits caught immediately upon landing near their stopover site. Initial metabolite levels were compared with levels after 5 h of inactive rest to determine how flight per se affects energy metabolism. Birds refuelling on the stopover site were also examined. Arriving godwits displayed elevated plasma free fatty acids, glycerol and butyrate, confirming the importance of lipid fuel in the support of extended migratory activity. Further-more, elevated plasma triglycerides in these birds suggest that fatty acid provisioning is facilitated through hepatic synthesis and release of neutral lipids, as previously hypothesized for small migrants with high mass-specific metabolic rates. Finally, elevations in plasma uric acid suggest that protein breakdown contributes to the support of long-distance movement, to possibly maintain citric acid cycle intermediates, gluconeogenesis and/or water balance.