Calidris Mauri Research Articles

Trends in timing of spring migration along the Pacific Flyway by Western Sandpipers and Dunlins

ABSTRACT Long-distant migrants timing their arrival on the breeding grounds face a tradeoff between optimal timing for breeding and optimal timing for survival. For many shorebird species, the flyway northward spans thousands of kilometers, and both conditions encountered en route and priorities of individuals can affect the timing of migration. We used data from spring migration surveys of Western Sandpipers (Calidris mauri) and Pacific Dunlins (Calidris alpina pacifica) along the Pacific Flyway of North America to determine whether the timing of their northward migration changed from 1985 to 2016. We compiled survey data for both species from 6 sites of varying size along the northern portion of the flyway from Washington, USA, through British Columbia, Canada, to Alaska, USA, and estimated interannual trends in the timing of passage through each site. Peak passage dates at the sites closest to the species' breeding grounds in Alaska became later by 1–2 days over the study period, while dates of peak pa...

Effects of environmental conditions on reproductive effort and nest success of Arctic‐breeding shorebirds

The Arctic is experiencing rapidly warming conditions, increasing predator abundance, and diminishing population cycles of keystone species such as lemmings. However, it is still not known how many Arctic animals will respond to a changing climate with altered trophic interactions. We studied clutch size, incubation duration and nest survival of 17 taxa of Arctic‐breeding shorebirds at 16 field sites over 7 years. We predicted that physiological benefits of higher temperatures and earlier snowmelt would increase reproductive effort and nest survival, and we expected increasing predator abundance and decreasing abundance of alternative prey (arvicoline rodents) to have a negative effect on reproduction. Although we observed wide ranges of conditions during our study, we found no effects of covariates on reproductive traits in 12 of 17 taxa. In the remaining taxa, most relationships agreed with our predictions. Earlier snowmelt increased the probability of laying a full clutch from 0.61 to 0.91 for Western Sandpipers, and shortened incubation by 1.42 days forarcticolaDunlin and 0.77 days for Red Phalaropes. Higher temperatures increased the probability of a full clutch from 0.60 to 0.93 for Western Sandpipers and from 0.76 to 0.97 for Red‐necked Phalaropes, and increased daily nest survival rates from 0.9634 to 0.9890 for Semipalmated Sandpipers and 0.9546 to 0.9880 for Western Sandpipers. Higher abundance of predators (foxes) reduced daily nest survival rates only in Western Sandpipers (0.9821–0.9031). In contrast to our predictions, the probability of a full clutch was lowest (0.83) for Semipalmated Sandpipers at moderate abundance of alternative prey, rather than low abundance (0.90). Our findings suggest that in the short‐term, climate warming may have neutral or positive effects on the nesting cycle of most Arctic‐breeding shorebirds.

Delayed egg‐laying and shortened incubation duration of Arctic‐breeding shorebirds coincide with climate cooling

Biological impacts of climate change are exemplified by shifts in phenology. As the timing of breeding advances, the within‐season relationships between timing of breeding and reproductive traits may change and cause long‐term changes in the population mean value of reproductive traits. We investigated long‐term changes in the timing of breeding and within‐season patterns of clutch size, egg volume, incubation duration, and daily nest survival of three shorebird species between two decades. Based on previously known within‐season patterns and assuming a warming trend, we hypothesized that the timing of clutch initiation would advance between decades and would be coupled with increases in mean clutch size, egg volume, and daily nest survival rate. We monitored 1,378 nests of western sandpipers, semipalmated sandpipers, and red‐necked phalaropes at a subarctic site during 1993–1996 and 2010–2014. Sandpipers have biparental incubation, whereas phalaropes have uniparental incubation. We found an unexpected long‐term cooling trend during the early part of the breeding season. Three species delayed clutch initiation by 5 days in the 2010s relative to the 1990s. Clutch size and daily nest survival showed strong within‐season declines in sandpipers, but not in phalaropes. Egg volume showed strong within‐season declines in one species of sandpiper, but increased in phalaropes. Despite the within‐season patterns in traits and shifts in phenology, clutch size, egg volume, and daily nest survival were similar between decades. In contrast, incubation duration did not show within‐season variation, but decreased by 2 days in sandpipers and increased by 2 days in phalaropes. Shorebirds demonstrated variable breeding phenology and incubation duration in relation to climate cooling, but little change in nonphenological components of traits. Our results indicate that the breeding phenology of shorebirds is closely associated with the temperature conditions on breeding ground, the effects of which can vary among reproductive traits and among sympatric species.

Effects of intertidal habitat availability on the use of anthropogenic habitats as foraging grounds by shorebirds: a case study on semi-intensive shrimp farms

Loss of wetlands may restrict foraging opportunities for many species of migratory shorebirds that depend on such habitats during the non-breeding periods. In addition, tidal cycles, both daily (high and low tides) and lunar (spring and neap tides), periodically modulate the availability of foraging areas within coastal wetlands, which can limit area and foraging time for shorebirds. We tested whether densities of shorebirds (marbled godwit, willet, western sandpiper, black-necked stilt, American avocet, two dowitcher species, and whimbrel) observed at a shrimp farm varied with predictable tidal cycles on the coast of northwestern Mexico. We found that most species occurred in higher densities during high- and neap-tides, when nearby intertidal areas were flooded and thus unavailable for foraging shorebirds. The majority of shorebirds at the shrimp farm were actively feeding, indicating that the shrimp farm can provide an alternate foraging habitat. As an exception, western sandpipers were found in lower densities during neap tides, and we suggest that predation risk from peregrine falcons at the shrimp farm may displace this species to other nearby safe habitats. Understanding the spatio-temporal variability on the use of shrimp farms will increase the general knowledge about the function of alternative anthropogenic habitats for migratory shorebirds.

Testing of an oral dosing technique for double-crested cormorants, Phalacocorax auritus, laughing gulls, Leucophaeus atricilla, homing pigeons, Columba livia, and western sandpipers, Calidris mauri, with artificially weather MC252 oil

Scoping studies were designed to determine if double-crested cormorants (Phalacocorax auritus), laughing gulls (Leucophaues atricilla), homing pigeons (Columba livia) and western sandpipers (Calidris mauri) that were gavaged with a mixture of artificially weathered MC252 oil and food for either a single day or 4–5 consecutive days showed signs of oil toxicity. Where volume allowed, samples were collected for hematology, plasma protein electrophoresis, clinical chemistry and electrolytes, oxidative stress and organ weigh changes. Double-crested cormorants, laughing gulls and western sandpipers all excreted oil within 30min of dose, while pigeons regurgitated within less than one hour of dosing. There were species differences in the effectiveness of the dosing technique, with double-crested cormorants having the greatest number of responsive endpoints at the completion of the trial. Statistically significant changes in packed cell volume, white cell counts, alkaline phosphatase, alanine aminotransferase, creatine phosphokinase, gamma glutamyl transferase, uric acid, chloride, sodium, potassium, calcium, total glutathione, glutathione disulfide, reduced glutathione, spleen and liver weights were measured in double-crested cormorants. Homing pigeons had statistically significant changes in creatine phosphokinase, total glutathione, glutathione disulfide, reduced glutathione and Trolox equivalents. Laughing gulls exhibited statistically significant decreases in spleen and kidney weight, and no changes were observed in any measurement endpoints tested in western sandpipers.

Light oiling of feathers increases flight energy expenditure in a migratory shorebird.

Flying birds depend on their feathers to undertake most activities, and maintain them in peak condition through periodic molt and frequent preening. Even small exposures to crude oil reduce the integrity of feathers, and could impair flight performance. We trained wild western sandpipers (Calidris mauri) to perform endurance flights in a wind tunnel, and used magnetic resonance body composition analysis to measure energy expenditure after birds were exposed to weathered MC252 crude oil from the Deepwater Horizon oil spill. The cost of transport was 0.26±0.04 kJ km-1 in controls, and increased by 22% when the trailing edges of the wing and tail were oiled (<20% of body surface; considered light oiling). Additional crude oil on breast and back feathers (∼30% total surface; moderate oiling) increased the cost of transport by 45% above controls. Oiling tended to decrease flight control, and only half of moderately oiled birds completed the flight test. We then flew birds at a range of speeds to estimate basic kinematic parameters. At low speeds, light and moderately oiled birds had larger wingbeat amplitudes than controls, while moderately oiled birds showed greater wingbeat frequencies across all speeds, and a shift in optimal flight speed towards higher wind speeds. We suggest these changes reflect poorer lift production and increased drag on the wings and body. Oiling will increase the difficulty and energy costs of locomotion for daily and seasonal activities such as foraging, predator evasion, territory defense, courtship, chick provisioning, commuting and long-distance migration. These sub-lethal effects must be considered in oil spill impact assessments.

Even light oiling from spills can be catastrophic for birds

![Figure][1] An unimpeded western sandpiper flying in the wind tunnel at Western University. Photo credit: Brock Fenton. It's a depressingly familiar sight when an oil well blows or a tanker runs aground: thousands of stranded, helpless animals wallowing in cloying crude oil. ‘Birds are

Reprint of: Trouble on takeoff: Crude oil on feathers reduces escape performance of shorebirds

The ability to takeoff quickly and accelerate away from predators is crucial to bird survival. Crude oil can disrupt the fine structure and function of feathers, and here we tested for the first time how small amounts of oil on the trailing edges of the wings and tail of Western sandpipers (Calidris mauri) affected takeoff flight performance. In oiled birds, the distance travelled during the first 0.4s after takeoff was reduced by 29%, and takeoff angle was decreased by 10° compared to unoiled birds. Three-axis accelerometry indicated that oiled sandpipers produced less mechanical power output per wingbeat during the initial phase of flight. Slower and lower takeoff would make oiled birds more likely to be targeted and captured by predators, reducing survival and facilitating the exposure of predators to oil. Whereas the direct mortality of heavily-oiled birds is often obvious and can be quantified, our results show that there are significant sub-lethal effects of small amounts crude oil on feathers, which must be considered in natural resource injury assessments for birds.

Toxicological and thermoregulatory effects of feather contamination with artificially weathered MC 252 oil in western sandpipers (Calidris mauri)

The external contamination of bird feathers with crude oil might have effects on feather structure and thus on thermoregulation. We tested the thermoregulatory ability of western sandpipers (Calidris mauri) in a respirometry chamber with oil applied either immediately prior, or three days before the experiment. The birds were then exposed to a sliding cold temperature challenge between 27°C and −3°C to calculate thermal conductance. After the experiment, a large blood sample was taken and the liver extracted to measure a range of parameters linked to toxicology and oxidative stress. No differences in thermal conductance were observed among groups, but birds exposed to oil for three days had reduced body temperatures and lost more body mass during that period. At necropsy, oiled birds showed a decrease in plasma albumin and sodium, and an increase in urea. This is reflective of dysfunction in the kidney at the loop of Henle. Birds, especially when exposed to the oil for three days, showed signs of oxidative stress and oxidative damage. These results show that the ingestion of externally applied oil through preening or drinking can cause toxic effects even in low doses, while we did not detect a direct effect of the external oil on thermoregulation over the temperature range tested.

Effect of oral exposure to artificially weathered Deepwater Horizon crude oil on blood chemistries, hepatic antioxidant enzyme activities, organ weights and histopathology in western sandpipers (Calidris mauri)

Shorebirds were among birds exposed to Mississippi Canyon 252 (MC252) crude oil during the 2010 Deep Water Horizon (DWH) oil spill in the Gulf of Mexico. The western sandpiper (Calidris mauri) was chosen as one of four species for initial oral dosing studies conducted under Phase 2 of the avian toxicity studies for the DWH Natural Resource Damage Assessment (NRDA). Thirty western sandpipers were assigned to one of three treatment groups, 10 birds per group. The control group was sham gavaged and the treatment groups were gavaged with 1 or 5mL oil kg bw−1 daily for 20 days. Periodic blood samples for hemoglobin measurements were collected during the trial. A final blood sample used to determine hemoglobin concentration in addition to complete blood counts, plasma clinical chemistries, haptoglobin concentration and plasma electrophoresis was collected when birds were euthanized and necropsied on day 21. Tissues were removed, weighed and processed for subsequent histopathological evaluation. There were numerical decreases in hemoglobin concentrations in oil-dosed birds over the 21-day trial, but values were not significantly different compared to controls on day 21. There were no significant differences between controls and oiled birds in complete blood counts, plasma chemistries, haptoglobin concentration, and plasma electrophoresis endpoints. Of the hepatic oxidative stress endpoints assessed, the total antioxidant capacity assessment (Trolox equivalents) for the control group was lower compared to the 1mL oil kg bw−1 group. Absolute liver weights in the 5mL oil kg bw−1 group were significantly greater compared to controls. While not conclusive, the numerical decrease in hemoglobin concentration and significant increase in absolute liver weight are consistent with exposure to oil. Histological changes in the adrenal gland could be considered a non-specific indicator of stress resulting from exposure to oil. It is possible that the quantity of oil absorbed was not sufficient to induce clearly evident hemolytic anemia or that the western sandpiper is relatively insensitive to ingested oil.

Trouble on takeoff: Crude oil on feathers reduces escape performance of shorebirds

The ability to takeoff quickly and accelerate away from predators is crucial to bird survival. Crude oil can disrupt the fine structure and function of feathers, and here we tested for the first time how small amounts of oil on the trailing edges of the wings and tail of Western sandpipers (Calidris mauri) affected takeoff flight performance. In oiled birds, the distance travelled during the first 0.4s after takeoff was reduced by 29%, and takeoff angle was decreased by 10° compared to unoiled birds. Three-axis accelerometry indicated that oiled sandpipers produced less mechanical power output per wingbeat during the initial phase of flight. Slower and lower takeoff would make oiled birds more likely to be targeted and captured by predators, reducing survival and facilitating the exposure of predators to oil. Whereas the direct mortality of heavily-oiled birds is often obvious and can be quantified, our results show that there are significant sub-lethal effects of small amounts crude oil on feathers, which must be considered in natural resource injury assessments for birds.

Migration as flow: using hydrological concepts to estimate the residence time of migrating birds from the daily counts

Summary Estimating the length‐of‐stay, the number of days a bird can be expected to stay at a site, at stopover sites is critical to understanding the migration ecology and estimating the population sizes of birds as they move between breeding and non‐breeding sites. Estimating the length‐of‐stay of migrating animals at stopover sites has an analogue in the hydrological concept of transit time, the amount of time that water spends in a reservoir, which can be calculated as a numerical integration of inflow and outflow rates with an underlying Storage Age Selection function. We used this approach to estimate the lengths‐of‐stay of migrating Western Sandpiper (Calidris mauri) and Dunlin (Calidris alpina) based on the time series of daily counts at two sites in British Columbia, Canada. The approach yielded mean transit times for Western Sandpiper during southward migration at Sidney Island that ranged between 9·6 days and 3·8 days, and showed a significant decline over time, 1992–2001, and is consistent with the estimates obtained from the capture‐mark‐resight studies. Transit times during northward migration at Roberts Bank, Fraser River Delta, based on the best available information ranged from 1·8 to 3·2 days for Western Sandpiper, and had a median value of 2·0 days for Dunlin, which is consistent with the estimates obtained from the radio‐telemetry studies. These results indicate that the hydrological flow models may offer a means to estimate the length‐of‐stay from the daily counts of birds during migration. The models present an opportunity for testing the alternate hypotheses concerning the roles of behavioural‐ vs. habitat‐related mechanisms driving shorebird population sizes.

Flow pattern similarities in the near wake of three bird species suggest a common role for unsteady aerodynamic effects in lift generation.

Analysis of the aerodynamics of flapping wings has yielded a general understanding of how birds generate lift and thrust during flight. However, the role of unsteady aerodynamics in avian flight due to the flapping motion still holds open questions in respect to performance and efficiency. We studied the flight of three distinctive bird species: western sandpiper (Calidris mauri), European starling (Sturnus vulgaris) and American robin (Turdus migratorius) using long-duration, time-resolved particle image velocimetry, to better characterize and advance our understanding of how birds use unsteady flow features to enhance their aerodynamic performances during flapping flight. We show that during transitions between downstroke and upstroke phases of the wing cycle, the near wake-flow structures vary and generate unique sets of vortices. These structures appear as quadruple layers of concentrated vorticity aligned at an angle with respect to the horizon (named 'double branch'). They occur where the circulation gradient changes sign, which implies that the forces exerted by the flapping wings of birds are modified during the transition phases. The flow patterns are similar in (non-dimensional) size and magnitude for the different birds suggesting that there are common mechanisms operating during flapping flight across species. These flow patterns occur at the same phase where drag reduction of about 5% per cycle and lift enhancement were observed in our prior studies. We propose that these flow structures should be considered in wake flow models that seek to account for the contribution of unsteady flow to lift and drag.

Effects of migration distance on life history strategies of Western and Semipalmated sandpipers in Perú

Migration distances of shorebird species correlate with life history strategies. To assess age-specific migratory preparation and adult wing-molt strategies, we studied Western Sandpipers (Calidris mauri) and Semipalmated Sandpipers (C. pusilla) with different migration routes at the Paracas National Reserve in Peru, one of the most austral non-breeding areas for these sandpipers, from 2012 to 2015. Western Sandpipers breed near the Bering Sea, ~11,000 km from Paracas. Semipalmated Sandpiper populations at Paracas are a mixture of short-billed birds from western Arctic breeding sites, plus long-billed birds from eastern sites, ~8000 km distant. Adults of both species arrive in October with primary feathers already partially renewed so wing molt starts at sites further north. Semipalmated Sandpipers with longer bills completed wing molt later than shorter billed birds. Adults of both species prepared for migration in February and March. No juvenile Western Sandpipers prepared for migration, confirming the “slow” over-summering life history strategy of more southerly non-breeding populations. Juvenile Semipalmated Sandpipers showed bimodality in strategies. Most showed no migratory preparation, but, during three non-breeding periods, from 27% to 31% fattened, molted, and partially replaced outer primaries during the pre-migratory period. Juveniles with longer culmens were heavier and tended to have more alternate plumage. Juveniles that were partially molting primaries had longer culmens and more alternate plumage. Juvenile Semipalmated Sandpipers from eastern-breeding populations thus have a higher propensity for a fast life history strategy, and western birds a slow one, at this non-breeding site in Peru. Western-breeding Semipalmated Sandpiper populations thus resemble Western Sandpipers, suggesting a common, possibly distance-related, effect on life history strategy.

Monitoring Migrating Shorebirds at the Tofino Mudflats in British Columbia, Canada: is Disturbance a Concern?

Abstract. The Tofino Wah-nah-jus Hilth-hoo-is Mudflats on Vancouver Island, British Columbia, Canada, include both pristine areas and sites with residential and resort development. Shorebird counts were conducted at six sites around these mudflats in 2011 and compared with historical counts from 1988, 1989, and 1995. Temporal trends of counts did not vary among sites with different levels of disturbance. Over the entire mudflats, counts of Dunlin (Calidris alpina) moving northward had a negative trend over time (βyr = -0.04), as did counts of Western Sandpiper (Calidris mauri; βyr = -0.04), Least Sandpiper (Calidris minutilla; βyr = -0.03) and Short-billed Dowitcher (Limnodromus griseus; βyr = -0.04) moving southward. In contrast, counts of Black-bellied Plover (Pluvialis squatarola; βyr = 0.03) and Semipalmated Plover (Charadrius semipalmatus; βyr = 0.02) moving northward increased over time. Counts of Dunlin and Short-billed Dowitcher were lower at disturbed sites relative to non-disturbed sites, indica...

Effects of geolocators on hatching success, return rates, breeding movements, and change in body mass in 16 species of Arctic-breeding shorebirds.

BackgroundGeolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8–2.0 g total, representing 0.1–3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2–4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26–1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables.ResultsWe detected negative effects of tags for three small-bodied species. Geolocators reduced annual return rates for two of 23 taxa: by 63 % for semipalmated sandpipers and by 43 % for the arcticola subspecies of dunlin. High resighting effort for geolocator birds could have masked additional negative effects. Geolocators were more likely to negatively affect return rates if the total mass of geolocators and color markers was 2.5–5.8 % of body mass than if tags were 0.3–2.3 % of body mass. Carrying a geolocator reduced nest success by 42 % for semipalmated sandpipers and tripled the probability of partial clutch failure in semipalmated and western sandpipers. Geolocators mounted perpendicular to the leg on a flag had stronger negative effects on nest success than geolocators mounted parallel to the leg on a band. However, parallel-band geolocators were more likely to reduce return rates and cause injuries to the leg. No effects of geolocators were found on breeding movements or changes in body mass. Among-site variation in geolocator effect size was high, suggesting that local factors were important.ConclusionsNegative effects of geolocators occurred only for three of the smallest species in our dataset, but were substantial when present. Future studies could mitigate impacts of tags by reducing protruding parts and minimizing use of additional markers. Investigators could maximize recovery of tags by strategically deploying geolocators on males, previously marked individuals, and successful breeders, though targeting subsets of a population could bias the resulting migratory movement data in some species.Electronic supplementary materialThe online version of this article (doi:10.1186/s40462-016-0077-6) contains supplementary material, which is available to authorized users.

Biofilm consumption and variable diet composition of western sandpipers (Calidris mauri) during migratory stopover.

Many shorebird species undertake long-distance migrations punctuated by brief stays at food-rich, estuarine stopover locations. Understanding use of these food resources helps guide conservation and responsible development decisions. We determined the extent and degree to which Western Sandpiper (Calidris mauri) utilized biofilm as a food resource across a large and variable stopover location during northward (spring) migration. We investigated the spatial heterogeneity in diet composition, to determine whether shorebirds were consistently feeding on biofilm or whether diet varied between naturally and anthropogenically delineated sites. We used stable isotope analysis to estimate that biofilm conservatively comprised 22% to 53% of Western Sandpiper droppings across all sampling sites and that prey composition differed significantly between areas within the stopover location. Widespread biofilm consumption demonstrates the importance of biofilm as a dietary component. Variable diet composition suggests that habitat heterogeneity may be an important component of high quality stopover locations in the context of “state-dependant trade-offs” of Western Sandpiper population sub-groups. Future management decisions must consider and address potential impacts on the biofilm community throughout a stopover location, as single site studies of diet composition may not be adequate to develop effective management strategies for entire stopover sites.

Sex Determination of Least Sandpiper(Calidris minutilla)and Western Sandpiper (Calidris mauri): Comparing Methodological Robustness of Two Morphometric Methods

Abstract Least (Calidris minutilla, n = 110) and Western (Calidris mauri, n = 37) sandpipers collected in Cuban wetlands were sexed by gonadal examination on dissection and used to assess the robustness of two morphometric methods commonly used to assign sex in shorebirds. Discriminant function analyses were performed and then tested by jackknife validation. The best discriminant function for Least Sandpiper included culmen and wing lengths and correctly classified 91% of the birds. Using culmen length, the discriminant function correctly sexed 97% of the Western Sandpiper. Sex ratio and morphometric estimates obtained through sexing Least and Western sandpipers by discriminant functions were not significantly different from the population sexed by gonadal examination on dissection. The range of the bill lengths used for sex determination of Least and Western sandpipers were then assessed by fitting the known-sex data. Classification accuracy of the bill length method was high for Western Sandpiper (95%),...

Intertidal biofilm distribution underpins differential tide-following behavior of two sandpiper species (Calidris mauri and Calidris alpina) during northward migration

The discovery that some shorebird species graze heavily on biofilm adds importance to elucidating coastal processes controlling biofilm, as well as impetus to better understand patterns of shorebird use of intertidal flats. Western sandpipers (Calidris mauri) and dunlin (Calidris alpina) stopover in the hundreds of thousands on the Fraser River estuary, British Columbia, Canada, during northward migration to breeding areas. Western sandpipers show greater modification of tongue and bill morphology for biofilm feeding than dunlin, and their diet includes more biofilm. Therefore, we hypothesized that these congeners differentially use the intertidal area. A tide following index (TFI) was used to describe their distributions in the upper intertidal during ebbing tides. Also, we assessed sediment grain size, biofilm (= microphytobenthic or MPB) biomass and invertebrate abundance. Foraging dunlin closely followed the ebbing tide line, exploiting the upper intertidal only as the tide retreated through this area. In contrast, western sandpipers were less prone to follow the tide, and spent more time in the upper intertidal. Microphytobenthic biomass and sediment water content were highest in the upper intertidal, indicating greater biofilm availability for shorebirds in the first 350 m from shore. Invertebrate density did not differ between sections of the upper intertidal. Overall, western sandpiper behaviour and distribution more closely matched MPB biofilm availability than invertebrate availability. Conservation of sandpipers should consider physical processes, such as tides and currents, which maintain the availability of biofilm, a critical food source during global migration.

Macroalgal Mats in a Eutrophic Estuary Obscure Visual Foraging Cues and Increase Variability in Prey Availability for Some Shorebirds

Conservation of habitat for flagship species, such as birds, aids in the protection of biodiversity in critical ecosystems. Extensive mats of macroalgae stimulated by nutrient input to estuaries threaten critical successes in conservation made by legislation that reduces habitat destruction. Macroalgae can cover intertidal mudflats that are key foraging grounds for obligate visual foragers such as black-bellied plovers (Pluvialis squatarola), as well as foragers that routinely switch from visual to tactile foraging such as least sandpipers (Calidris minutilla), western sandpipers (Calidris mauri), and willets (Tringa semipalmata), and predominately tactile foragers such as marbled godwits (Limosa fedoa). We hypothesized that macroalgae would affect shorebirds directly by altering foraging behavior and indirectly by reducing prey availability. Mats reduced visual foraging (pecking) for sandpipers and marbled godwits and caused them to probe more often. Willets spent overall less time foraging than sandpipers and marbled godwits and did not alter their foraging strategy due to macroalgae. While foraging, black-bellied plovers, marbled godwits, and willets avoided macroalgae while sandpipers did not select between habitat types. This suggests macroalgae may have negative effects on plovers, marbled godwits, and willets by reducing foraging area but that sandpipers may utilize both macroalgae and bare sediments. Macroalgal mats indirectly influenced shorebirds by enhancing variability in prey availability across the mudflat landscape. By quantifying avoidance of or preference for mats, foraging behavior, and variation in prey availability, we showed that macroalgae have differential effects across shorebird foraging guilds.