Shorebird abundance, migration phenology, and invertebrate prey availability at a mid-continental stopover site in Kentucky, USA

Shorebird use of an exposed sandy beach in southern California

Biguglia (Corse), quartier d'hiver d'importance internationale pour l'hivernage des canards plongeurs et des foulques macroules - fonctionnement et perspectives (Biguglia Corsica - an internationally important area for wintering coot and diving duck, functioning and outlook): G. Rocamora, Alauda, 60(4), 1992, pp 239–250

Multiple environmental factors and prey depletion determine declines in abundance and timing of departure in migratory shorebirds in the west coast of India

SummaryThe effects of agricultural intensification on biodiversity in arable systems of western Europe have received a great deal of attention. However, the recent transformation of grassland systems has been just as profound.In Britain, the management of grassland has changed substantially in the second half of the 20th century. A high proportion of lowland grassland is managed intensively. The major changes include a doubling in the use of inorganic nitrogen, a switch from hay to silage, and increased stocking densities, particularly of sheep. Structurally diverse and species‐rich swards have been largely replaced by relatively dense, fast‐growing and structurally uniform swards, dominated by competitive species.Most of these changes have reduced the suitability of grassland as feeding and breeding habitat for birds.The most important direct effects have been deterioration of the sward as nesting and wintering habitat, and loss of seed resources as food. Short uniform swards afford poor shelter and camouflage from predators, whereas increased mowing intensities and trampling by stock will destroy nests and young. Increased frequency of sward defoliation reduces flowering and seed set, and hence food availability for seed‐eating birds.The indirect effects of intensification of management on birds relate largely to changes in the abundance and availability of invertebrate prey. The effects of management vary with its type, timing and intensity, and with invertebrate ecology and phenology, but, in general, the abundance and diversity of invertebrates declines with reductions in sward diversity and structural complexity.Low input livestock systems are likely to be central to any future management strategies designed to maintain and restore the ecological diversity of semi‐natural lowland grasslands. Low additions of organic fertilizer benefit some invertebrate prey species, and moderate levels of grazing encourage sward heterogeneity.There is now a need to improve understanding of how grassland management affects bird population dynamics. Particularly important areas of research include: (i) the interaction between changes in food abundance, due to changes in fertilizer inputs, and food accessibility, due to changes in sward structure; (ii) the interaction between predation rates and management‐related changes in habitat; and (iii) the impact of alternative anti‐helminithic treatments for livestock on invertebrates and birds.

Many migratory shorebirds rely on estuaries as stop-over sites to refuel during migration, and the loss of stop-over sites is a primary threat to shorebird populations on the West Coast of the United States (e.g. Calidris alpina pacifica, C. mauri). Conservation and research has focused on the largest of these sites; however, smaller estuaries also host thousands of migratory shorebirds. Furthermore, the reasons for site selection are largely unknown. Estuarine inter-tidal microhabitats are non-uniform and both abiotic and biotic factors may serve as predictors of whether an abundance of shorebirds will use a site. I investigated shorebird site selection on broad and fine scales within Oregon estuaries. To identify factors that relate to shorebird abundance on large spatial scales, I compiled shorebird abundance data from estuaries throughout the Pacific Northwest as well as data on site quality factors. To investigate site selection on a finer scale I measured shorebird abundance, habitat characteristics, and food resources―invertebrates and a newly considered source, biofilm―within two Oregon estuaries during the fall migration period. Finally, I examined whether channels are preferentially used by foraging Calidrid shorebirds by conducting observations during the spring migration. I investigated whether channels may be superior foraging habitat possibly because prey are more abundant, are found at shallower depths, or because sediments are more penetrable (increasing the opportunity for shorebird probing) by taking infauna cores and measuring force required to probe in the sediment at channel and open mudflat sites. Among estuaries, shorebird densities in spring were best predicted by estuary size, as opposed to the amount of any one habitat. During fall migration, the amount of grassland in the surrounding watershed was also a good predictor, pointing to the probable importance of roost sites as well as feeding grounds. The amount of infauna also related to the density of shorebirds using a site. Within estuaries, shorebird distribution in the inter-tidal region was not generally predicted by prey abundance. Channels were used preferentially by shorebirds, and infauna abundance along channels was greater than in the surrounding mudflats. The more penetrable sediments of the channel also made it easier for shorebirds to probe and capture prey. Identification of these large-scale and fine-scale factors that influence site quality for migratory shorebirds will assist land and wildlife managers' efforts to protect these species.

When prey and predator are seasonal migrants, encounters depend on migration phenologies and environmental constraints on predation. Here we investigate the relative contribution of seasonality in irradiance and prey abundance in shaping the rapid seasonal body condition increase of a migrating predator searching visually for its prey: the Norwegian spring-spawning herring, Clupea harengus, feeding on the copepod Calanus finmarchicus. Two main seasonal pulses of prey are available to herring: (1) the parent generation of C. finmarchicus, with peak abundance in March-April, which appear too early to cause the main increase in herring condition; and (2) the abundant offspring generation of C. finmarchicus, with peak abundance in June-July, too late to explain the main increase in body condition. However, a mechanistic model of ingestion rate, including both solar irradiance and prey abundance, predicted seasonal food intake in good accordance with observed herring body condition. This suggests that the seasonality in herring foraging and energy storage is closely linked to the return of longer days in spring, and less dependent on a match or mismatch with seasonal peaks in abundance of their zooplankton prey. Consequently, light related constraints on foraging may make visually searching predators at high latitudes resilient to changes and fluctuations in prey phenology and abundance, but vulnerable to changes in the light regime, such as water clarity.

Estuaries provide crucial foraging resources and nursery habitat for threatened populations of anadromous salmon. As such, there has been a global undertaking to restore habitat and tidal processes in modified estuaries. The foraging capacity of these ecosystems to support various species of out‐migrating juvenile salmon can be quantified by monitoring benthic, terrestrial, and pelagic invertebrate prey communities. Here, we present notable trends in the availability of invertebrate prey at several sites within a restoring large river delta in Puget Sound, Washington, U.S.A. Three years after the system was returned to tidal influence, we observed substantial additions to amphipod, copepod, and cumacean abundances in newly accessible marsh channels (from 0 to roughly 5,000–75,000 individuals/m2). In the restoration area, terrestrial invertebrate colonization was dependent upon vegetative cover, with dipteran and hymenopteran biomass increasing 3‐fold between 1 and 3 years post‐restoration. While the overall biodiversity within the restoration area was lower than in the reference marsh, estimated biomass was comparable to or greater than that found within the other study sites. This additional prey biomass likely provided foraging benefits for juvenile Chinook, chum, and coho salmon. Primary physical drivers differed for benthic, terrestrial, and pelagic invertebrates, and these invertebrate communities are expected to respond differentially depending on organic matter exchange and vegetative colonization. Restoring estuaries may take decades to meet certain success criteria, but our study demonstrates rapid enhancements in foraging resources understood to be used for estuary‐dependent wildlife.

The diet of long-nosed bandicoots (Perameles nasuta) on the central coast of New South Wales, Australia, was examined over two summers and two winters using a combination of faecal scat analysis for food fragments and stable isotope analysis (ratios of 13C/12C and 15N/14N) of blood. Isotope ratios in blood overlapped most strongly with those in invertebrate prey, and varied much less between seasons than did those in most dietary items, suggesting that the assimilated diet of long-nosed bandicoots is dominated by invertebrates throughout the year. Invertebrate remains dominated collected faeces in both seasons, even though the availability of invertebrate prey was higher in summer. Thus both techniques indicated that long-nosed bandicoots were primarily insectivorous year-round. Faecal scat analysis indicated that invertebrate eggs were more abundant in summer than winter. At a finer scale, spiders, orthopterans, lepidopteran larvae, ants, leaf material (non-grass monocot) and seeds were more abundant in summer, while cicada larvae, roots, fungi, grass leaves and Acacia bract (small modified leaves appearing as scales) were more abundant in winter. Subterranean foods (cicada larvae, plant roots and hypogeous fungi) were more abundant in winter and more abundant in the diet of males than of either lactating or non-lactating females.

We analyzed the relationship between the abundance of shorebirds and that of their intertidal invertebrate prey, and attempted to determine if shorebird predation significantly affects prey density. The study was conducted at three sites in the Chacopata Lagoon complex in northeastern Venezuela between January 1985 and September 1986, using shorebird census data, monthly sampling of intertidal invertebrates, and exclosure experiments. The invertebrates collected were divided into three groups: (i) polychaetes (≤50 mm in length), (ii) small bivalves (2 – 3 mm in length), and (iii) other species. The overall density of shorebirds foraging in the Chacopata Lagoon complex was very high, exceeding the densities reported for most other staging and overwintering areas by a wide margin. The abundance of shorebirds in the study area was related primarily to the density of polychaetes, their main prey. At two of the study sites, the arrival of fall migrants followed a significant increase in polychaete numbers. The increase in shorebird numbers in autumn was negligible at the third site, where polychaete density was lower than at the other sites. The impact of shorebird predation on their invertebrate prey varied seasonally, as did the extent of the foraging area available to them. The exclosure experiment revealed significant differences in the density of polychaetes inside and outside the exclosures only during, or shortly after, the fall migration period. The variability in the conclusions drawn from other studies examining prey depletion by shorebirds in tropical environments may be explained by differences in the length of sampling periods and seasonal variations in the numbers of foraging shorebirds, the energy demands of moult, fat deposition, and the intertidal surface area available for feeding. The impact of seasonal variables such as these should be taken into consideration when designing studies to measure prey depletion by shorebirds; such objectives may necessitate long-term studies.

We compared prefledging growth, energy expenditure, and time budgets in the arctic-breeding red knot (Calidris canutus) to those in temperate shorebirds, to investigate how arctic chicks achieve a high growth rate despite energetic difficulties associated with precocial development in a cold climate. Growth rate of knot chicks was very high compared to other, mainly temperate, shorebirds of their size, but strongly correlated with weather-induced and seasonal variation in availability of invertebrate prey. Red knot chicks sought less parental brooding and foraged more at the same mass and temperature than chicks of three temperate shorebird species studied in The Netherlands. Fast growth and high muscular activity in the cold tundra environment led to high energy expenditure, as measured using doubly labelled water: total metabolised energy over the 18-day prefledging period was 89% above an allometric prediction, and among the highest values reported for birds. A comparative simulation model based on our observations and data for temperate shorebird chicks showed that several factors combine to enable red knots to meet these high energy requirements: (1) the greater cold-hardiness of red knot chicks increases time available for foraging; (2) their fast growth further shortens the period in which chicks depend on brooding; and (3) the 24-h daylight increases potential foraging time, though knots apparently did not make full use of this. These mechanisms buffer the loss of foraging time due to increased need for brooding at arctic temperatures, but not enough to satisfy the high energy requirements without invoking (4) a higher foraging intake rate as an explanation. Since surface-active arthropods were not more abundant in our arctic study site than in a temperate grassland, this may be due to easier detection or capture of prey in the tundra. The model also suggested that the cold-hardiness of red knot chicks is critical in allowing them sufficient feeding time during the first week of life. Chicks hatched just after the peak of prey abundance in mid-July, but their food requirements were maximal at older ages, when arthropods were already declining. Snow cover early in the season prevented a better temporal match between chick energy requirements and food availability, and this may enforce selection for rapid growth.

It is imperative to manage wetlands appropriately in the non‐breeding range of migratory shorebirds because their habitat quality impacts survival and reproductive performance. However, it is challenging to measure and relate these parameters to the habitat quality of individual sites because they are influenced by a composite contribution to fitness across all sites used in the annual cycle. Identifying important habitat quality features for shorebirds, and assessing whether management actions aimed at improving habitat quality are effective, poses a substantial challenge. Nonetheless, within a site, shorebird subsite selection and foraging behaviour could reveal variation in habitat quality proxies and suitability. We developed a field study to investigate habitat selection and habitat quality proxies for shorebirds in the Coorong, an internationally important non‐breeding site in Australia. We examined the abundance of shorebirds among multiple subsites in the wetland and their foraging behaviour in relation to multiple variables (potential habitat area, salinity, prey density, wind, temperature), the first three of which can be influenced by management. Larger potential habitat area, defined as the combined area of bare mud/sand and shallow water (<20 cm depth), was associated with higher shorebird abundance across subsites with the same shore length, suggesting this is the most important site feature for habitat selection. Shorebird step rate, which reflects the effort associated with catching prey, was higher at subsites with lower prey density. This suggests that step rate can be used as an indicator of prey availability. Potential habitat area is strongly positively correlated with shorebird abundance. At sites where water levels can be managed, flows should be regulated so that water levels are not too high (limiting habitat area) or too low (drying out mudflats). However, it is critical that large potential habitat areas that attract shorebirds are productive so that they do not become ecological traps. Management should aim to maintain healthy prey abundance on areas attractive for shorebirds, while also minimising predation and disturbance of shorebirds. Monitoring prey abundance directly can be challenging, but our study shows that shorebird step rate provides a low‐cost indicator of relative macroinvertebrate prey abundance.

Fishes use their mechanoreceptive lateral line system to sense nearby objects by detecting slight fluctuations in hydrodynamic motion within their immediate environment. Species of fish from different habitats often display specialisations of the lateral line system, in particular the distribution and abundance of neuromasts, but the lateral line can also exhibit considerable diversity within a species. Here, we provide the first investigation of the lateral line system of the Australian western rainbowfish (Melanotaenia australis), a species that occupies a diversity of freshwater habitats across semi-arid northwest Australia. We collected 155 individuals from eight populations and surveyed each habitat for environmental factors that may contribute to lateral line specialisation, including water flow, predation risk, habitat structure and prey availability. Scanning electron microscopy and fluorescent dye labelling were used to describe the lateral line system in M. australis, and to examine whether the abundance and arrangement of superficial neuromasts (SNs) varied within and among populations. We found that the SNs of M. australis were present in distinct body regions rather than lines. The abundance of SNs within each body region was highly variable, and also differed among populations and individuals. Variation in SN abundance among populations was best explained by habitat structure and the availability of invertebrate prey. Our finding that specific environmental factors explain among-population variation in a key sensory system suggests that the ability to acquire sensory information is specialised for the particular behavioural needs of the animal.

Suitability of habitat patches affects demographic processes and consequently influences the viability of populations. In order for managers to consider the potential of resources on a scale appropriate to their use, it is important to understand the processes that influence the ecology of threatened species. Differential growth rates of Litoria aurea (green and golden bell frog) at Sydney Olympic Park, Australia, may be explained by prey availability and diet. We tested: (1) whether food availability differed among precincts at Sydney Olympic Park and (2) whether the diet of L. aurea was influenced by availability of invertebrate prey. Diets were distinct among precincts and reflected the variation in biomass and richness of invertebrate assemblages. Precincts with greater biomass corresponded to areas with faster individual growth rates and greater habitat structure. The differences in diet and individual growth rates of L. aurea among precincts at Sydney Olympic Park demonstrate how caution must be applied to generalising population function, even within a small area.

Many Neartic shorebirds migrate to South America for the non-breeding season, and use several key stopover and wintering sites along the Atlantic coast. However, there is little information about annual shorebird use along the northeast coast of Brazil. This year-long study examined the seasonal occurrence and abundance of shorebirds at Atalaia Nova beach in Sergipe State, Brazil. Of the 13 species recorded, the most abundant were Semipalmated Plovers (Charadrius semipalmatus), Semipalmated Sandpipers (Calidris pusilla), and Sanderlings (Calidris alba). Highest numbers occurred between September and March, and the lowest numbers occurred between April and August. Some species were present throughout the year. Atalaia Nova beach is an important migration stopover and wintering site for shorebirds in northeastern Brazil.

The timing of annual events in migratory species is determined by the position of breeding and wintering grounds, length of the breeding season, the occurrence of their preferred diet or the timing of moult in birds. Previous studies found significant differences in the migration phenology of Asian Phylloscopus warbler species, a group of long-distance migratory passerines. However, the factors that influence the observed differences in migration phenology have not been studied yet. Here, we analyze the effect of the above-mentioned factors on the migration timing of seven Phylloscopus species at a stopover site in the Russian Far East. We found that spring migration phenology was best explained by moult strategy and the preferred prey size, while autumn migration phenology was linked to the southernmost wintering latitude of the species. Interspecific differences in migration timing were much higher in autumn than in spring. The duration of the species-specific migration periods was also longer in autumn than in spring, most likely caused by higher competition during spring to arrive early at the breeding grounds. Our results contribute to the understanding of migration ecology in songbirds moving along the little-studied East Asian flyway.