Potential interactions between humans and non-breeding shorebirds on a subtropical intertidal flat

The primary aim of this study was to investigate whether bait harvesting, with all its inherent effects, occurring in the intertidal zone of a subtropical estuary, had an impact on a migratory shorebird, the eastern curlew Numenius madagascariensis. In a large-scale manipulative study (units of experiment were 1 ha plots), callianassid shrimp Trypaea australiensis populations were harvested simulating the technique (manual pumping) and the levels of harvesting intensity per unit area (347 shrimp per hectare per harvesting event) exhibited by bait-collectors in SE Australia and South Africa. It was found that at present levels of harvesting intensity per unit area (approximately 1% of standing stock removed per harvesting event) there is no threat to the stocks of Trypaea exploited by the curlews in Moreton Bay, Australia. However, the results show that the curlews themselves apply a considerable predation pressure on Trypaea. Based on the birds' foraging rates and densities, it was estimated that they would consume up to 100% of the initial Trypaea stock over the course of a non-breeding season (October to March). However, the stable seasonal trend in the density of the size-cohort of Trypaea preyed upon by the curlews indicates that the existing rates of predation are easily counterbalanced, e.g. through continuous density-dependent recruitment of these crustaceans. We suggest that this mechanism will provide for a stable foraging environment for both the shorebirds and bait collectors.

Why do eastern curlews Numenius madagascariensis feed on prey that lowers intake rate before migration?

Eastern curlews Numenius madagascariensis spending the nonbreeding season in eastern Australia foraged on three intertidal decapods: soldier crab Mictyris longicarpus, sentinel crab Macrophthalmus crassipes and ghost-shrimp Trypaea australiensis. Due to their ecology, these crustaceans were spatially segregated (=distributed in 'patches') and the curlews intermittently consumed more than one prey type. It was predicted that if the curlews behaved as intake rate maximizers, the time spent foraging on a particular prey (patch) would reflect relative availabilities of the prey types and thus prey-specific intake rates would be equal. During the mid-nonbreeding period (November-December), Mictyris and Macrophthalmus were primarily consumed and prey-specific intake rates were statistically indistinguishable (8.8 versus 10.1 kJ x min(-1)). Prior to migration (February), Mictyris and Trypaea were hunted and the respective intake rates were significantly different (8.9 versus 2.3 kJ x min(-1)). Time allocation to Trypaea-hunting was independent of the availability of Mictyris. Thus, consumption of Trypaea depressed the overall intake rate. Six hypotheses for consuming Trypaea before migration were examined. Five hypotheses: the possible error by the predator, prey specialization, observer overestimation of time spent hunting Trypaea, supplementary prey and the choice of higher quality prey due to a digestive bottleneck, were deemed unsatisfactory. The explanation for consumption of a low intake-rate but high quality prey (Trypaea) deemed plausible was diet optimisation by the Curlews in response to the pre-migratory modulation (decrease in size/processing capacity) of their digestive system. With a seasonal decrease in the average intake rate, the estimated intake per low tide increased from 1233 to 1508 kJ between the mid-nonbreeding and pre-migratory periods by increasing the overall time spent on the sandflats and the proportion of time spent foraging.

Intake rates and the functional response in shorebirds (Charadriiformes) eating macro-invertebrates

As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type 11 ('disc equation') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching. A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (< 150/m(-2)). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote. A multivariate analysis of 468 'spot' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81 % of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3 %), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6%, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93 % of cases. We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested. As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.

Estimating survival and movements using both live and dead recoveries : a case study of oystercatchers confronted with habitat change

1. Animals facing partial habitat loss can try to survive in the remaining habitat or emigrate. Effects on survival and movements should be studied simultaneously since survival rates may be underestimated if emigrants are not considered, and since emigrants may experience reduced survival.

2. We analysed movements and survival of adult wintering oystercatcher Haematopus ostralegus in response to the 1986–1987 partial closure of the Oosterschelde in the Dutch Delta. This reduced by one-third the tidal area of this major European wintering area for waders.

3. We developed a novel variant of a multistate capture–recapture model allowing simultaneous estimation of survival and movement between sites using a mixture of data (live recaptures and dead recoveries). We used a two-step process, first estimating movements between sites followed by site-specific survival rates.

4. Most birds were faithful to their ringing site. Winter survival was negatively affected by winter severity and was lowest among birds changing wintering site (i.e. moving outside of the Oosterschelde).

5. During mild winters, survival rates were very high, and similar to before the closure in both changed and unchanged sectors of the Oosterschelde. However, the combined effect of habitat loss with severe winters decreased the survival of birds from changed sectors and induced emigration.

6. The coastal engineering project coincided with three severe winters and high food stock, making assessment of its effects difficult. However, the habitat loss seems to have had less impact on adult survival and movements than did winter severity.

7. Synthesis and applications. Human-induced habitat change may result in population decline through costly emigration or reduced survival or reproduction of individuals that stay. Long-term monitoring of marked individuals helps to understand how populations respond to environmental change, but site-specific survival and movement rates should be integrated in the same model in order to maximize the information yield. Our modelling approach facilitates this because it allows the inclusion of recoveries from outside the study area.

Comparing the seasonal survival of resident and migratory oystercatchers : carry-over effects of habitat quality and weather conditions

Events happening in one season can affect life-history traits at (the) subsequent season(s) by carry-over effects. Wintering conditions are known to affect breeding success, but few studies have investigated carry-over effects on survival. The Eurasian oystercatcher Haematopus ostralegus is a coastal wader with sedentary populations at temperate sites and migratory populations in northern breeding grounds of Europe. We pooled continental European ringing-recovery datasets from 1975 to 2000 to estimate winter and summer survival rates of migrant and resident populations and to investigate long-term effects of winter habitat changes. During mild climatic periods, adults of both migratory and resident populations exhibited survival rates 2% lower in summer than in winter. Severe winters reduced survival rates (down to 25% reduction) and were often followed by a decline in survival during the following summer, via short-term carry-over effects. Habitat changes in the Dutch wintering grounds caused a reduction in food stocks, leading to reduced survival rates, particularly in young birds. Therefore, wintering habitat changes resulted in long-term (>10 years) 8.7 and 9.4% decrease in adult annual survival of migrant and resident populations respectively. Studying the impact of carry-over effects is crucial for understanding the life history of migratory birds and the development of conservation measures.

Nonbreeding Eastern Curlews Numenius madagascariensis Do Not Increase the Rate of Intake or Digestive Efficiency before Long-Distance Migration because of an Apparent Digestive Constraint

The possibility of premigratory modulation in gastric digestive performance was investigated in a long-distance migrant, the eastern curlew (Numenius madagascariensis), in eastern Australia. The rate of intake in the curlews was limited by the rate of digestion but not by food availability. It was hypothesized that before migration, eastern curlews would meet the increased energy demand by increasing energy consumption. It was predicted that (1) an increase in the rate of intake and the corresponding rate of gastric throughput would occur or (2) the gastric digestive efficiency would increase between the mid-nonbreeding and premigratory periods. Neither crude intake rate (the rate of intake calculated including inactive pauses; 0.22 g DM [grams dry mass] or 3.09 kJ min(-1)) nor the rate of gastric throughput (0.15 g DM or 2.85 kJ min(-1)) changed over time. Gastric digestive efficiency did not improve between the periods (91%) nor did the estimated overall energy assimilation efficiency (63% and 58%, respectively). It was concluded that the crustacean-dominated diet of the birds is processed at its highest rate and efficiency throughout a season. It appears that without a qualitative shift in diet, no increase in intake rate is possible. Accepting these findings at their face value poses the question of how and over what time period the eastern curlews store the nutrients necessary for the ensuing long, northward nonstop flight.

Selecting umbrella species for conservation: a test of habitat models and niche overlap for beach-nesting birds

Umbrella species are rarely selected systematically from a range of candidate species. On sandy beaches, birds that nest on the upper beach or in dunes are threatened globally and hence are prime candidates for conservation intervention and putative umbrella species status. Here we use a maximum-likelihood, multi-species distribution modeling approach to select an appropriate conservation umbrella from a group of candidate species occupying similar habitats. We identify overlap in spatial extent and niche characteristics among four beach-nesting bird species of conservation concern, American oystercatchers (Haematopus palliatus), black skimmers (Rynchops niger), least terns (Sterna antillarum) and piping plovers (Charadrius melodus), across their entire breeding range in New Jersey, USA. We quantify the benefit and efficiency of using each species as a candidate umbrella on the remaining group. Piping plover nesting habitat encompassed 86% of the least tern habitat but only 15% and 13% of the black skimmer and American oystercatcher habitat, respectively. However, plovers co-occur with all three species across 66% of their total nesting habitat extent (~ 649 ha), suggesting their value as an umbrella at the local scale. American oystercatcher nesting habitat covers 100%, 99% and 47% of piping plover, least tern and black skimmer habitat, making this species more appropriate conservation umbrellas at a regional scale. Our results demonstrate that the choice of umbrella species requires explicit consideration of spatial scale and an understanding of the habitat attributes that an umbrella species represents and to which extent it encompasses other species of conservation interest. Notwithstanding the attractiveness of the umbrella species concept, local conservation interventions especially for breeding individuals in small populations may still be needed.