Effects of traditional clam harvesting on the foraging ecology of migrating curlews (Numenius arquata)

One of the main conservation priorities in many geographical regions for intertidal estuaries is to protect and maintain internationally important numbers of migratory shorebirds (Charadrii). Traditional harvesting practices at low tide cause alterations to the surface of the intertidal habitat by trampling and digging the substrate to obtain benthic resources, which may have negative effects on shorebirds. Here, we conducted a BACI (Before-After Control-Impact) experiment to monitor changes in foraging ecology of Eurasian curlews Numenius arquata, before and after an experimental disturbance of the intertidal mud in a key stopover area for migratory shorebirds. Mean density of curlews, foraging activity, feeding rate, percentage of crabs in the diet, size of crabs, and foraging speed differed neither between treatment and control plots, nor between before and after mud disturbance. The feeding technique was similar between treatment and control plots, but differed significantly between periods. These findings suggest that mud disturbance by harvesters working by hand did not affect curlew fitness by altering the energy deposition necessary for long-distance migration. Although conclusions must be extrapolated with caution to other sites and/or species, the Santoña Marshes Natural Park may provide a potentially good example of compatibility between shorebird conservation and traditional low tide harvesting practices.     

Agroecosystems and conservation of migratory waterbirds: importance of coastal pastures and factors influencing their use by wintering shorebirds

Coastal pastures are common agroecosystems adjacent to estuarine areas that can provide valuable habitat for wildlife, particularly for migratory shorebirds. Disentangling the factors that influence coastal pasture use by wintering shorebirds will provide new insights into its role for buffering human disturbances and habitat loss in intertidal areas. We examined whether numbers of two shorebirds (Eurasian curlew and Black-tailed godwit) foraging actively on coastal pastures was affected by weather conditions, tidal stage (low/high tide) and number of harvesters at intertidal areas throughout winter. Both species frequently used coastal pastures and most individuals foraged actively there. The average percentage of the total wintering population of curlews and godwits foraging on coastal pastures was 27.4 and 7.8 %, respectively, and was significantly higher during high tide compared to low tide. The number of harvesters on mudflats also had a positive significant effect in explaining the presence of curlews, and to a lesser extent for godwits, on coastal pastures, and accumulated rainfall had a positive effect for both species too. These supratidal areas were consistently used as alternative foraging grounds during low tide by curlews, as well as supplementary foraging areas during high tide by wintering populations of both large shorebirds. By supplementary foraging, wintering curlews, and probably godwits, seemed to compensate for a negative effect of the presence of harvesters on their foraging activity. We recommend managing of those coastal agricultural fields adjacent to intertidal foraging grounds in order to increase the availability of supratidal foraging habitats for declining shorebird populations. These habitats may thus have a beneficial role in sustaining populations of wintering shorebirds, but further studies are needed to estimate if birds can compensate for any shortfall in daily energy budget by supplementary foraging on coastal pastures, thus providing insights into whether they are involved in large-scale population regulation of migratory birds.     

Humans alter habitat selection of birds on ocean‐exposed sandy beaches

Aim Resource‐selection functions (RSFs) can quantify and predict the density of animal populations across heterogeneous landscapes and are important conservation tools in areas subject to human disturbance. Sandy beach ecosystems have comparatively low habitat heterogeneity and structural relief in the intertidal zone, but intense human use. We aimed to develop predictive RSFs for birds on ocean‐exposed sandy beaches at two spatial scales, 25 ha (local scale) and 250 ha (landscape scale), and to test whether habitat selection of birds that commonly use the surf–beach–dune interface is influenced by the rates of human activities. Location Moreton and North Stradbroke Island, eastern Australia. Methods Avifauna and human activities were mapped on three sandy beaches covering 79 km of coastline for 15 months. Habitat characteristics of the surf–beach–dune interface were derived from remote sensing and ground surveys. RSFs were developed for 12 species of birds at two spatial scales: 25 ha (local scale) and 250 ha (landscape scale). Results  At local (25 ha) and landscape scales (250 ha), dune dimensions and the extent and type of vegetation structure were important predictors of bird density. Adding the frequency of human activities improved the predictive power of RSFs, suggesting that habitat selection of birds on beaches is modified by human use of these environments. Human activities occurred mostly in the mid‐ to lower intertidal zone of the beach, overlapping closely with the preferred habitats of Silver Gulls (Larus novaehollandiae), Pied Oystercatchers (Haematopus longirostris), Red‐capped Plovers (Charadrius ruficapillus) and endangered Little Terns (Sternula albifrons). Main conclusions In addition to demonstrating the appropriateness of RSFs to the surf–beach–dune interface, our results stress the need for systematic conservation planning for these ecosystems, where ecological values have traditionally been subsidiary to the maintenance of sand budgets and erosion control.     

Low tide distribution of wintering waders and shelduck on the Severn Estuary in relation to the proposed tidal barrage

Waders and shelduck were counted at low tide on 162 sectors comprising 85% of the intertidal area (21 467 ha) of the Severn Estuary on 12 occasions during winter 1987/88. On average, 50% of birds present at low tide utilized just 13 sectors (12% of the area); 90% of birds occurred on only 56 sectors, leaving large expanses of intertidal sand virtually devoid of birdlife. Dunlin, the numerically dominant species, occurred widely on the middle and outer estuary, whereas shelduck predominantly occurred on the outer estuary and redshank around many tributary river mouths. Curlew, the most ubiquitous species, was the only one concentrated on the inner estuary. Severe gales in both late December and mid-January concentrated all main species within fewer sectors, probably by the short-term removal of surface sediment from substantial areas. It is estimated that the proposed tidal barrage would eliminate intertidal areas accounting for between c. 40% (for shelduck and curlew) and 80% (for redshank) of current total low tide usage by the internationally important populations present.     

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

Abstract 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.     

Abundance and distribution of sturgeon feeding pits in a Washington estuary

Sturgeon diet and feeding habitats are notoriously difficult to document. We mapped the locations of feeding pits in Willapa Bay, Washington, to characterize estuarine habitats used by sub-adult and adult sturgeon for infaunal feeding. Monthly summer surveys of intertidal plots revealed that feeding pit density was highest in July and August, when sturgeon occupy Willapa Bay. The ephemeral nature of feeding pits and high daily densities (> 1000 pits/ha) indicated intensive sturgeon feeding over unvegetated littoral mud flats during high tide. Feeding pit density was lowest in subtidal areas, over sand (grain sizes primarily >63 μ), and at sites with dense stands of non-indigenous seagrass, Zostera japonica. Sub-adult and adult sturgeon apparently used these habitats significantly less than would be predicted based on their availability. Feeding pit formation was negatively correlated with Z. japonica shoot dry weight and positively correlated with the abundance of thalassinid shrimp burrows. Experimental removal of Z. japonica resulted in increased sturgeon feeding, but experimental removal of burrowing shrimp did not significantly affect feeding pit formation. Aquaculture activities that harden substrate and proliferation of invasive seagrass both appear to produce estuarine substrates that are unsuitable for benthic feeding by sturgeon.     

Predator presence may benefit: kestrels protect curlew nests against nest predators

We studied whether the presence of breeding kestrels (Falco tinnunculus) affected nest predation and breeding habitat selection of curlews (Numenius arquata) on an open flat farmland area in western Finland. We searched for nests of curlews from an area of 6 km² during 1985–1993. For each nest found, we recorded the fate of the nest, and the distance to the nearest kestrel nest and to the nearest perch. We measured the impact of breeding kestrels on nest predation by constructing artificial curlew nests in the vicinity of ten kestrel nests in 1993. Curlew nests were closer to kestrel nests than expected from random distribution, eventhough kestrels fed on average 5.5% of curlew chick production. Predation risk by kestrels was lower than predation risk by corvids and other generalist predators, which predated 9% of curlew nests surviving farming practices and an unknown proportion of chicks. Artificial nest experiment showed that nest predation was lower close to kestrel nests than further away suggesting that the breeding association of curlews and kestrels was a behavioural adaptation against nest predation. Thus, the presence of a predator may sometimes be beneficial to prey, and prey animals have behavioural adaptations to these situations.     

Experiments on epibenthic predation in the Wadden Sea

Field experiments were designed to evaluate the role of predators in the Wadden Sea, small predators like shore crabs, shrimps and gobies, and large ones like flatfish and birds. Exclosures, maintained in aCorophium volutator bed, an eelgrass bed, in a sandy and a muddy flat, protected the infauna from such epibenthic predation. The resulting changes in the macrofauna were recorded and compared with an unaffected control area. In sandy and muddy flats of the lower intertidal zone, cages (mesh size5 mm) altered abundance and composition of the infauna almost entirely. Nearly all species achieved higher population densities than in the control area, and in addition, the number of species increased as well. In the the scarcely populated mud flat a dense suspension-feeder assemblage emerged, associated with numerous tube-building polychaetes. The sand flat, normally dominated by deposit feeders, also became occupied by a dense suspension-feeder assemblage, mainly cockles. In contrast, narrowly meshed cages had only little effect in the beds of eelgrass and ofCorophium volutator. Both are positioned in the upper intertidal zone. Although a number of species still responded with significant increases in abundance, many remained indifferent or even tended to be less abundant within cages. Cages provided with a 20-mm mesh nylon net, excluding only birds, flatfish and the biggest crabs, increased significantly the survival of large-sized infaunal members. This was only apparent in the upper intertidal zone. It was concluded, that small sized epibenthic predators are the major determinants of the dynamic species abundance pattern of the lower intertidal flats. In the eelgrass bed, the meshwork of rootlets constitutes an important spatial refuge from these predators. The complex habitat structure causes a diversified faunal assemblage. TheCorophium bed in the uppermost intertidal zone is less accessable to predators like crabs, shrimp and gobies. The monotonous appearance of this faunal assemblage is assumed to be the outcome of competitive exclusion and of occasional harsh physical conditions.     

Estimating the Breeding Population of Long-Billed Curlew in the United States

ABSTRACT Determining population size and long-term trends in population size for species of high concern is a priority of international, national, and regional conservation plans. Long-billed curlews (Numenius americanus) are a species of special concern in North America due to apparent declines in their population. Because long-billed curlews are not adequately monitored by existing programs, we undertook a 2-year study with the goals of 1) determining present long-billed curlew distribution and breeding population size in the United States and 2) providing recommendations for a long-term long-billed curlew monitoring protocol. We selected a stratified random sample of survey routes in 16 western states for sampling in 2004 and 2005, and we analyzed count data from these routes to estimate detection probabilities and abundance. In addition, we evaluated habitat along roadsides to determine how well roadsides represented habitat throughout the sampling units. We estimated there were 164,515 (SE=42,047) breeding long-billed curlews in 2004, and 109,533 (SE=31,060) breeding individuals in 2005. These estimates far exceed currently accepted estimates based on expert opinion. We found that habitat along roadsides was representative of long-billed curlew habitat in general. We make recommendations for improving sampling methodology, and we present power curves to provide guidance on minimum sample sizes required to detect trends in abundance.     

Determination of substrate preferences of tench, Tinca tinca (L.), under controlled experimental conditions

The substrate preference of tench Tinca tinca (L.), 33 2‐year‐old individuals, mean size 10 ± 0.36 cm total length, was determined using four different substrates (concrete, artificial vegetation, sand and mud) in a 26 m long maquette. The species showed a high preference (100%) for the mud. Six more trials were performed combining the substrates by pairs (concrete–vegetation, concrete–mud, concrete–sand, mud–vegetation, mud–sand, sand–vegetation). Selection order was: mud (100% of the individuals when compared with concrete and sand and 98.26 ± 2.68% when compared with artificial vegetation), artificial vegetation (100% of the individuals when compared with concrete and 99.81 ± 0.76% when compared with sand), sand (98.29 ± 3.66% when compared with concrete) and concrete.     

Middle Triassic horseshoe crab reproduction areas on intertidal flats of Europe with evidence of predation by archosaurs

A systematically excavated track site in a 243.5 Myr old Middle Triassic (Karlstadt Formation, Pelsonian, middle Anisian) intertidal carbonate mud‐flat palaeoenvironment at Bernburg (Saxony‐Anhalt, central Germany) has revealed extensive horseshoe crab trackways attributable to the Kouphichnium Nopsca, 1923 ichnogenus. The exposed track bed of a Germanic Basin‐wide spanned intertidal megatrack site is a mud‐cracked biolaminate surface on which detailed tracks have been preserved because of rapid drying and cementation as a result of high temperatures, followed by rapid covering with a protective layer of arenitic storm or tsunami sediments. The different trackway types and their orientations have allowed a tidal sequence to be reconstructed, with the initial appearance of swimming horseshoe crabs followed by half‐swimming/half‐hopping limulids under the shallowest water conditions. The Bernburg trackways, which have mapped lengths of up to 40 m, were all produced by adult animals and exhibit a variety of shapes and patterns that reflect a range of subaquatic locomotion behaviour more typical of mating than of feeding activities. The closest match to the proportions and dimensions of the horseshoe crab tracks at Bernburg is provided by the largest known Middle Triassic limulid Tachypleus gadeai, which is known from the north‐western Tethys in Spain. The horseshoe crab body fossils recognized in the German Mesozoic intertidal zones, instead, are from juveniles. The uniformly adult size indicated by the trackways therefore suggests that they may record the oldest intertidal reproductive zones of horseshoe crabs known from anywhere in the world, with the track‐makers having possibly migrated thousands of kilometres from shallow marine areas of the north‐western Tethys to reproduce in the intertidal palaeoenvironments of the Germanic Basin. Chirotherium trackways of large thecodont archosaurs also appeared on these flats where they appear to have fed on the limulids. With the tidal ebb, smaller reptiles such as Macrocnemus (Rhynchosauroides trackways) appeared on the dry intertidal flats, probably feeding on marine organisms and possibly also on horseshoe crab eggs. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 76–105.     

Long-billed curlew nest site selection and success in the Intermountain West

Grassland birds have experienced steeper population declines between 1966 and 2015 than any other bird group on the North American continent, and migratory grassland birds may face threats in all stages of their annual cycle. The grassland-associated long-billed curlew (Numenius americanus) is experiencing population declines in regional and local portions of their North American breeding range. The nesting period is an important portion of the annual cycle when curlews may face demographic rate limitations from a suite of threats including predators and anthropogenic disturbance. We compared nest sites to random sites within breeding territories to examine nest site selection, and modeled correlates of nesting success for 128 curlew nests at 5 Intermountain West sites. Nest sites were 6 times more likely than random sites to be situated adjacent to existing cowpies. Additionally, curlews selected nest sites with shorter vegetation, and less bare ground, grass, and shrub cover than at random sites within their territories. Nest success varied widely among sites and ranged from 12% to 40% in a season with a mean of 27% for all nests during the 2015 and 2016 seasons. Higher nest success probability was associated with higher curlew densities in the area, greater percent cover of conspicuous objects (cowpies, rocks) near the nest, and higher densities of black-billed magpies (Pica hudsonia) and American crows (Corvus brachyrhynchos) at the site. We also found increased probability of nesting success with increased distance from a nest to the nearest potential perch in that territory. Given the central role of working lands to curlews in much of the Intermountain West, understanding limitations to nesting success in these diverse landscapes is necessary to guide adaptive management strategies in increasingly human-modified habitats. We suggest some grazing and irrigation practices already provide suitable nesting conditions for curlews, and others may require only minor temporal shifts to improve compatibility. © 2019 The Wildlife Society.     

Dynamics of sediment carbon stocks across intertidal wetland habitats of Moreton Bay,Australia

Coastal wetlands are known for high carbon storage within their sediments, but our understanding of the variation in carbon storage among intertidal habitats, particularly over geomorphological settings and along elevation gradients, is limited. Here, we collected 352 cores from 18 sites across Moreton Bay, Australia. We assessed variation in sediment organic carbon (OC) stocks among different geomorphological settings (wetlands within riverine settings along with those with reduced riverine influence located on tide‐dominated sand islands), across elevation gradients, with distance from shore and among habitat and vegetation types. We used mid‐infrared (MIR) spectroscopy combined with analytical data and partial least squares regression to quantify the carbon content of ~2500 sediment samples and provide fine‐scale spatial coverage of sediment OC stocks to 150 cm depth. We found sites in river deltas had larger OC stocks (175–504 Mg/ha) than those in nonriverine settings (44–271 Mg/ha). Variation in OC stocks among nonriverine sites was high in comparison with riverine and mixed geomorphic settings, with sites closer to riverine outflow from the east and south of Moreton Bay having higher stocks than those located on the sand islands in the northwest of the bay. Sediment OC stocks increased with elevation within nonriverine settings, but not in riverine geomorphic settings. Sediment OC stocks did not differ between mangrove and saltmarsh habitats. OC stocks did, however, differ between dominant species across the research area and within geomorphic settings. At the landscape scale, the coastal wetlands of the South East Queensland catchments (17,792 ha) are comprised of approximately 4,100,000–5,200,000 Mg of sediment OC. Comparatively high variation in OC storage between riverine and nonriverine geomorphic settings indicates that the availability of mineral sediments and terrestrial derived OC may exert a strong influence over OC storage potential across intertidal wetland systems.     

Measuring potential negative effects of traditional harvesting practices on waterbirds: a case study with migrating curlews

Because of potential conflicts between commercial and conservation interests, one challenge in coastal-area management is how to regulate harvesting practices in coastal areas without adversely affecting the survival of migratory waterbirds. In most Spanish intertidal areas of importance for shorebirds, managers have concentrated only on stock management of the shellfish species. Here, we studied aspects of the foraging behaviour of Eurasian curlews Numenius arquata migrating through a Ramsar area in north Spain in the presence and absence of hand harvesters. We aimed to assess potential negative effects on curlews with a view to making coastal management recommendations that would help reduce conflict between local people and waterbirds. The average density of hand harvesters and foraging curlews at low tide was 0.56±0.09 persons per 10 ha and 16.47±0.73 birds per 10 ha, respectively. The presence of harvesters had a significant effect on foraging activity (no harvesting: 86.47±1.01%; harvesting: 82.70±1.00%). However, the absence of significant differences in all other foraging variables, between days with and without harvesting, indicated that curlews were able to compensate for the impact of harvesters on their foraging activity. We recommend, as a point of departure, that intertidal coastal managers of this Spanish site and similar areas of importance for shorebirds limit the harvesting load to <0.56 persons per 10 ha⁻¹ at least during autumn migration.     

Eelgrass,Zostera marina,as essential fish habitat for young‐of‐the‐year winter flounder,Pseudopleuronectes americanus (Walbaum, 1792) in Maine estuaries

Distribution and density by habitat for age‐0, young‐of‐the‐year (YOY) winter flounder, Pseudopleuronectes americanus (Walbaum, 1792), were compared for two Maine estuaries to help define essential fish habitat for this life history stage. Two estuaries (Weskeag River and Penobscot Bay) along Mid‐coast Maine were sampled monthly with daytime 1.0 m² fixed‐frame throw traps around neap low tide, May–December over two consecutive years (2003–2004). Both eelgrass and adjacent sand/mud (20–60 cm deep) were randomly sampled with equal effort (4–12 samples per month) at two sites in both the Weskeag River and Penobscot Bay. Significantly higher densities of YOY winter flounder (2–9 cm TL) occurred in eelgrass relative to sand/mud. Density increased significantly in both habitats in 2004, and was higher in Penobscot Bay relative to the Weskeag River. YOY densities compared by eelgrass coverage within throw traps were found to be significantly higher in eelgrass that exceeded 30% coverage when compared with adjacent sand/mud areas and eelgrass coverage of 10–20%. YOY occurred in all months sampled (May–December); no density differences existed by month. These results indicate that very shallow (<0.6 m) eelgrass habitat is of key importance to YOY winter flounder in Maine estuaries and should be viewed as essential fish habitat (EFH) for this species and life stage.     

Modelling nearshore intertidal habitat use of young-of-the-year Pacific sand lance (Ammodytes hexapterus) in Barkley Sound, British Columbia, Canada

The Pacific sand lance (Ammodytes hexapterus) is an ecologically important prey species for many vertebrate marine predators in the Pacific Northwest. In this study, we examined the use of intertidal habitat by young of the year (YOY) sand lance in Barkley Sound, British Columbia. We selected 14 of the original 29 independent environmental variables for modelling based on univariate analysis for variable selection. These were then used to model sand lance presence-absence using a classification tree approach. Based on our models we found that sand lance avoided mud and intertidal eelgrass. For sites that had very little mud and no intertidal eelgrass, sand lance preferred sediment size mean ≥1,918 μm or sites with sediment size mean ≤1,918 μm but with relatively well sorted sediment (sorting values ≤2.56 SDs, used as a heterogeneity index of the substrate grain size). Adjacent subtidal characteristics were not found to be important at this scale of study. This suggests intertidal substrate characteristics and presence-absence of intertidal eelgrass are the main influences on occurrence of YOY in the intertidal. Our results support the hypothesis that sand lance are associated with particular sediment types, however intertidal sediment types used by sand lance in our study differ from known preferences of sand lance for subtidal sediment types. This difference may be due to unique habitat constraints for intertidal versus subtidal regions. Although it is unknown if the results of this study are widely applicable, the results begin to identify intertidal habitat features that are important for Pacific sand lance.     

Intertidal Benthic Invertebrates and Sediment Stability at the Mouth of Delaware Bay

From December 1970 to December 1972, the monthly species composition, density, species richness, and biomass of marine macroscopic invertebrates were quantitatively studied from a wave-protected, intertidal sand flat at the mouth of Delaware Bay. Considerable erosion, transportation, and deposition of sand occurred throughout the study period. The fauna was extremely flexible and resilient in its response to a gradient of considerable sediment mobility. This response was similar in scope to one recognized along the coast of California. Comparison of the local biota with other sand flats and beaches from Maine to South Carolina showed that (1). haustoriids were among the dominant species, (2). densities were roughly comparable, and (3). secondary species were similar. Based on the biological index value, three species contained 87.8% of the fauna from the Cape Henlopen tide flat. These species were the polychaete Scoloplos fragilis and the amphipod Neohaustorius biarticulatus, which were characteristic of sand bars, and the polychaete Heteromastus filiformis, which was more characteristic of troughs or depressions. Mean density varied considerably both monthly and annually with almost no cyclic repetition for any of the three species. There was evidence to indicate that specific habitat requirements of a few species can strongly affect the structure which emerges for a total fauna. Morphology, behavior, physiology, and reproductive strategy are aspects of biology which link dominant benthic species of intertidal sand beaches together.     

Food Resources for Wintering and Spring Staging Black Ducks

Abstract: A bioenergetic approach has been adopted as a planning tool to set habitat management objectives by several United States Fish and Wildlife Service North American Waterfowl Management Plan Joint Ventures. A bioenergetics model can be simplified into 2 major components, energetic demand and energetic supply. Our goal was to estimate habitat-specific food availability, information necessary for estimating energy supply for black ducks (Anas rubripes) wintering on Long Island, New York, USA. We collected both nektonic and benthic samples from 85 wetland sites dispersed among 5 habitat types (salt marsh, mud flat, submersed aquatic vegetation, brackish bay, and freshwater) commonly used by black ducks in proportion to expected use. Biomass varied among habitats (F_4,5 > 7.46, P < 0.03) in 2004–2005, but there was only marginal variation in 2005–2006 (F_3,4 = 5.75, P = 0.06). Mud flats had the greatest biomass (1,204 kg/ha, SE = 532), followed by submersed aquatic vegetation (61 kg/ha, SE = 18), and salt marsh (34 kg/ha, SE = 6). In the second year of the study, freshwater had the greatest biomass (306 kg/ha, SE = 286), followed by mud flats (85 kg/ha, SE = 63), and salt marsh (35 kg/ha, SE = 4). Our results suggest food density on wintering grounds of black ducks on coastal Long Island is considerably lower than for dabbling ducks using inland freshwater habitats, indicating black duck populations are more likely than other species of dabbling ducks to be limited by winter habitat. We recommend targeting preservation, restoration, and enhancement efforts on salt marsh habitat.     

Holocene reef evolution in a macrotidal setting: Buccaneer Archipelago,Kimberley Bioregion,Northwest Australia

This study uses information derived from cores to describe the Holocene accretion history of coral reefs in the macrotidal (up to 11 m tidal range) Buccaneer Archipelago of the southern Kimberley coast, Western Australia. The internal architecture of all cored reefs is broadly similar, constituting well-preserved detrital coral fragments, predominantly branching Acropora, in a poorly sorted sandy mud matrix. However, once the reefs reach sea level, they diverge into two types: low intertidal reefs that maintain their detrital character and develop relatively narrow, horizontal or gently sloping reef flats at approximately mean low water spring, and high intertidal reefs that develop broad coralline algal-dominated reef flats at elevations between mean low water neap and mean high water neap. The high intertidal reefs develop where strong, ebb-dominated, tidal asymmetry retains seawater over the low tide and allows continued accretion. Both reef types are ultimately constrained by sea level but differ in elevation by 3–4 m.     

The relative importance of geography and ecology in species diversification: evidence from a tropical marine intertidal snail (Nerita)

Aim In this study, I examined the relative contributions of geography and ecology to species diversification within the genus Nerita, a prominent clade of marine snails that is widely distributed across the tropics and intertidal habitats. Specifically, I tested whether geographical patterns of speciation correspond primarily to allopatric or sympatric models, and whether habitat transitions have played a major role in species diversification. Location Indo‐West Pacific, eastern Pacific, Atlantic, tropical marine intertidal. Methods I used a previously reconstructed molecular phylogeny of Nerita as a framework to assess the relative importance of geographical and ecological factors in species diversification. To evaluate whether recently diverged clades exhibit patterns consistent with allopatric or sympatric speciation, I mapped the geo‐graphical distribution of each species onto the species‐level phylogeny, and examined the relationship between range overlap and time since divergence using age–range correlation analyses. To determine the relative contribution of habitat transitions to divergence, I traced shifts in intertidal substrate affinity and vertical zonation across the phylogeny using parsimony, and implemented randomization tests to evaluate the resulting patterns of ecological change. Results Within the majority of Nerita clades examined, age–range correlation analysis yielded a low intercept and a positive slope, similar to that expected under allopatric speciation. Approximately 75% of sister species pairs have maintained allopatric distributions; whereas more distantly related sister taxa often exhibited complete or nearly complete geographical overlap. In contrast, only 19% of sister species occupy distinct habitats. For both substrate and zonation, habitat transitions failed to concentrate towards either the tips or the root of the phylogeny. Instead, habitat shifts have occurred throughout the history of Nerita, with a general transition from the lower and mid‐littoral towards the upper and supra‐littoral zones, and multiple independent shifts from hard (rock) to softer substrates (mangrove, mud and sand). Main conclusions Both geography and ecology appear to have influenced diversification in Nerita, but to different extents. Geography seems to play a principal role, with allopatric speciation driving the majority of Nerita divergences. Habitat transitions appear insignificant in shaping the early and recent history of speciation, and promoting successive diversification in Nerita; however, shifts may have been important for respective divergences (i.e. those that correspond to the transitions) and enhancing diversity throughout the clade.