Food Dispersion and Foraging Energetics: a Mechanistic Synthesis for Field Studies of Avian Benthivores

Much effort has focused on modeling and measuring the energy costs of free existence and the foraging strategies of animals. However, few studies have quantitatively linked these approaches to the patch structure of foods in the field. We developed an individual—based model that relates field measurements of the dispersion of benthic foods to search costs and foraging profitability of diving ducks. On Lake Mattamuskeet, North Carolina, Canvasback ducks (Aythya valisineria) eat only the belowground winter buds of the submerged plant Vallisneria americana. We measured and modeled the patch structure of winter buds at the level of potential foraging loci, defined as contiguous circles 1 m in diameter. In the field and in the model, Canvasbacks make repeated vertical dives in such loci, foraging in the sediments by touch, before surface—swimming to another locus. We quantified first—order patchiness by fitting a negative binomial distribution to core samples taken at 50-m intervals along transects, to yield the frequencies of loci with different bud densities. Second-order patchiness was measured by taking cores at 1-m increments radiating from each sampling point, and regressing bud density at each sampling point on densities at these increments. No significant correlations were found, indicating that Canvasbacks could not predict food densities based on densities in nearby foraging loci. For the model, we generated food grids from the negative binomial distributions of core samples. Energy costs of diving were calculated by applying aerobic efficiencies (mechanical power output / aerobic power input) to biomechanical models. Unlike respirometry alone, this method accounts for effects on dive costs of varying water depth and dive duration. We used measurements of Canvasback intake rates at different bud densities to calculate profitability (energy intake minus expenditure) for each dive. Multivariate uncertainty analyses (Latin hypercube) indicated that profitability for Canvasbacks foraging on Vallisneria buds is determined mainly by food—item size and locomotor costs of descent. Bud metabolizable energy, water temperature, bud dispersion, and search and handling time coefficients of the functional response for intake rate have relatively minor influence. Individual-parameter perturbations indicated that to maintain the same foraging benefits, the total area of Vallisneria habitat would have to increase by 1.4-fold if dry mass per bud decreased from 0.10 to 0.03 g, and by 2.1-fold if water depth increased from 0.5 to 2 m. Our method allows study of interactions between patch structure and foraging energetics without detailed spatial mapping of foods, which is not feasible at appropriate scales for highly mobile benthivores. The model yields estimates of energy balance, contaminant intake, and amount and quality of foraging habitat required to sustain diving duck populations under varying environmental conditions. More accurate prediction of giving-up times and giving-up food densities will require better understanding of the time scale over which ducks balance their energy budgets

Relative Foraging Value to Lesser Scaup Ducks of Native and Exotic Clams from San Francisco Bay

Invasions of exotic invertebrates have greatly altered many aquatic communities, but impacts on the foraging energetics of predators seldom have been assessed. In San Francisco Bay, California (USA), a major community change occurred with introduction of the Asian clam (Potamocorbula amurensis) in 1986. This species now greatly outnumbers the previous clam prey of a variety of sharks, rays, sturgeon, flatfish, and crabs, as well as several diving duck species for which the bay is the most important wintering area on the U.S. Pacific Coast. P. amurensis also accumulates much higher levels of some contaminants than the formerly dominant prey. Because alteration of the food base or contaminated foods on wintering areas may be factors in the population decline of scaup ducks, effects of this exotic invasion are important to assess. For Lesser Scaup (Aythya affinis), we studied effects of differences in nutrient content, digestibility, crushing resistance of shells, areal density, size, and depth in the sediments on the relative foraging value of exotic P. amurensis vs. the formerly dominant native clam Macoma balthica. P. amurensis, including shells, had higher nitrogen and energy content per clam of the same length class, and higher digestibility of energy, than M. balthica. Gut retention time did not differ between clam species, so their relative profitability for scaup was determined mainly by the intake rate of digestible nutrients during short, costly dives. For scaup foraging in an aquarium 1.8 m deep, intake rates (number of prey per second) of food items buried in sand-filled trays increased with increasing prey density up to at least 4000 prey/m2. For items buried 3 cm deep, intake rates did not differ for prey \u3c6 mm long vs. prey\u3e6–12 mm long; however, intake rates were much lower when prey were deeper in the sediments (6 cm vs. 3 cm). In the field, a much higher percentage of P. amurensis were in the length range most commonly eaten by Lesser Scaup (\u3c12 \u3emm), and unlike M. balthica, almost all P. amurensis were in the top 5 cm of sediments where scaup intake rates are highest. In tensometer measurements, shells of P. amurensis were much harder to crush than shells of M. balthica, which might partly offset the apparent energetic advantages of P. amurensis. In many respects, the exotic P. amurensis appears to be a more valuable food than the native M. balthica for Lesser Scaup. However, because P. amurensis accumulates much higher levels of some contaminants, this exotic invasion increases the risk of toxicity to scaup and a range of other benthic predators

Algal vs. Macrophyte Inputs to Food Webs of Inland Saline Wetlands

Invertebrate food webs in wetlands were traditionally thought to be fueled mainly by decaying macrophytes, but recently it has been recognized that microalgae may be more important. In particular, the paradigm that shredders of vascular plant litter dominate food web processes may not apply to many wetlands where shredders are rare and microalgae more abundant. This issue is complicated by potential consumption of flocs of dissolved organic matter (DOM) released from living plants, and of exopolymer secretions (EPS) from both autotrophic and heterotrophic microbes. In Wyoming, we used gut contents and stable isotopes to investigate organic matter sources for the dominant invertebrates in oligosaline (0.5–5 g/L total dissolved solids) and mesosaline (5–18 g/L) wetlands. We examined the trophic importance of microalgae vs. macrophytes in wetlands with and without emergent vegetation (Scirpus acutus), with different growth forms and species of submersed plants (Chara spp. vs. Potamogeton pectinatus), with dominance by different microalgal types (phytoplankton, epiphyton, epipelon), and with different primary consumers (mainly amphipods vs. chironomid larvae). In all wetlands studied, guts of the major primary consumers contained little or no macrophyte tissue, but rather mostly amorphous detritus (organic particles with no recognizable cellular structure). Values of δ13C indicated that organic matter entering foodwebs was not from submersed macrophytes, but that emergent plants might be a source of DOM or EPS in amorphous detritus. However, in some wetlands, amphipods eating mainly amorphous detritus had the same δ13C values as chironomids eating a much higher fraction of diatoms, indicating that amorphous detritus was derived mainly from diatoms. Patterns of temporal change of δ13C in consumers, seston, and emergent plants supported this interpretation. We conclude that microalgae rather than macrophytes provided most organic matter for these food webs via amorphous detritus. Amorphous detritus is often thought to have poor nutrient quality and low assimilation efficiency, but this idea may not be true if amorphous detritus is largely flocs of labile DOM/EPS. Our results suggest that characterizing the origin and nature of amorphous detritus is key to understanding variations in macroinvertebrate production among saline wetlands and a broad range of wetland types

OPPORTUNITY COSTS OF WATER LEASING: IRRIGATION, INSTREAM FLOW, AND WETLAND CONSIDERATIONS IN THE LARAMIE BASIN, WYOMING

Flood irrigation in the Laramie Basin of southeast Wyoming has created many wetlands that rely directly on irrigation inputs for water. The Laramie Basin is a proposed water source for enhancing Platte River instream flows, to the benefit of endangered cranes, terns, plovers, and sturgeons. Increasing irrigation efficiency, or retiring irrigated lands would transform Laramie Basin agriculture and cause a high fraction of the Basins wetlands to be lost. This study explores the limitations of traditional water transfer tools when regional instream-flow requirements compete for water with local irrigation-dependent wetlands. A rotating short-term water lease program is proposed. The program would allow Laramie Basin producers to contribute to instream flow without causing permanent wetland damage or loss. Short-term water leasing programs could allow agricultural communities to contribute to regional environmental water needs without sacrificing local, agriculturally-based ecological resources. An estimate of minimum water costs, advantages and disadvantages of short-term water leasing are discussed.Resource /Energy Economics and Policy,

Modeling Underwater Visual and Filter Feeding by Planktivorous Shearwaters in Unusual Sea Conditions

Short-tailed Shearwaters (Puffinus tenuirostris) migrate between breeding areas in Australia and wintering areas in the Bering Sea. These extreme movements allow them to feed on swarms of euphausiids (krill) that occur seasonally in different regions, but they occasionally experience die-offs when availability of euphausiids or other prey is inadequate. During a coccolithophore bloom in the Bering Sea in 1997, hundreds of thousands of Short-tailed Shearwaters starved to death. One proposed explanation was that the calcareous shells of phytoplanktonic coccolithophores reduced light transmission, thus impairing visual foraging underwater. This hypothesis assumes that shearwaters feed entirely by vision (bite-feeding), but their unique bill and tongue morphology might allow nonvisual filter-feeding within euphausiid swarms. To investigate these issues, we developed simulation models of Short-tailed Shearwaters bite-feeding and filter-feeding underwater on the euphausiid Thysanoessa raschii. The visual (bite-feeding) model considered profiles of diffuse and beam attenuation of light in the Bering Sea among seasons, sites, and years with varying influence by diatom and coccolithophore blooms. The visual model indicated that over the huge range of densities in euphausiid swarms (tens to tens of thousands per cubic meter), neither light level nor prey density had appreciable effects on intake rate; instead, intake was severely limited by capture time and capture probability after prey were detected. Thus, for shearwaters there are strong advantages of feeding on dense swarms near the surface, where dive costs are low relative to fixed intake rate, and intake might be increased by filter-feeding. With minimal effects of light conditions, starvation of shearwaters during the coccolithophore bloom probably did not result from reduced visibility underwater after prey patches were found. Alternatively, turbidity from coccolithophores might have hindered detection of euphausiid swarms from the air

Modeling Marine Protected Areas for Threatened Eiders in a Climatically Changing Bering Sea

Delineating protected areas for sensitive species is a growing challenge as changing climate alters the geographic pattern of habitats as well as human responses to those shifts. When human impacts are expected within projected ranges of threatened species, there is often demand to demarcate the minimum habitat required to ensure the species\u27 persistence. Because diminished or wide-ranging populations may not occupy all viable (and needed) habitat at once, one must identify thresholds of resources that will support the species even in unoccupied areas. Long-term data on the shifting mosaic of critical resources may indicate ranges of future variability. We addressed these issues for the Spectacled Eider (Somateria fischeri), a federally threatened species that winters in pack ice of the Bering Sea. Changing climate has decreased ice cover and severely reduced the eiders\u27 benthic prey and has increased prospects for expansion of bottom trawling that may further affect prey communities. To assess long-term changes in habitats that will support eiders, we linked data on benthic prey, sea ice, and weather from 1970 to 2001 with a spatially explicit simulation model of eider energy balance that integrated field, laboratory, and remote-sensing studies. Areas estimated to have prey densities adequate for eiders in 1970–1974 did not include most areas that were viable 20 years later (1993–1994). Unless the entire area with adequate prey in 1993–1994 had been protected, the much reduced viable area in 1999–2001 might well have been excluded. During long non-foraging periods (as at night), eiders can save much energy by resting on ice vs. floating on water; thus, loss of ice cover in the future might substantially decrease the area in which prey densities are adequate to offset the eiders\u27 energy needs. For wide-ranging benthivores such as eiders, our results emphasize that fixed protected areas based on current conditions can be too small or inflexible to subsume long-term shifts in habitat conditions. Better knowledge of patterns of natural disturbance experienced by prey communities, and appropriate allocation of human disturbance over seasons or years, may yield alternative strategies to large-scale closures that may be politically and economically problemati

Walruses Attack Spectacled Eiders Wintering in Pack Ice of the Bering Sea

We observed walruses (Odobenus rosmarus) pursuing spectacled eiders (Somateria fischeri) within pack ice of the Bering Sea, 70–90 km from the nearest land. We used both direct observations from a helicopter and a heligimbal camera system that can film animals from a helicopter at high altitudes. The eiders were in monospecific flocks of thousands of birds within large leads. The walruses apparently tried to catch the eiders from below; the eiders responded with a “flash expansion” (explosive radial movement), wing-flapping and running along the water surface to escape. Disturbance by individual walruses could restrict flocks of thousands of birds to small portions of the open water. In eight such events that we witnessed over 75 min of observations, we were unable to confirm that walruses captured any of these full-grown, flight-capable eiders. However, the high rate of attacks and the eiders’ dramatic escape response suggest that walruses can at times be effective predators on them, and may affect the eiders’ dispersion and energy balance.Nous avons observé des morses (Odobenus rosmarus) en train de pourchasser des eiders à lunettes (Somateria fischeri) sur la banquise de la mer de Béring, soit à une distance de 70 à 90 kilomètres de la terre ferme la plus près. Nous nous sommes servis à la fois d’observations directes faites à partir d’un hélicoptère et d’un appareil héligimbal capable de filmer les animaux à partir d’un hélicoptère à haute altitude. Les eiders se tenaient en bandes monospécifiques constituées de milliers d’oiseaux faisant partie de gros groupements. Il semblait que les morses essayaient d’attraper les eiders par en-dessous; les eiders réagissaient en faisant une « expansion éclair » (un mouvement radial explosif), en battant des ailes et en courant le long de la surface de l’eau afin de s’échapper. Les perturbations exercées par les morses individuels pouvaient avoir pour effet de restreindre les bandes de milliers d’oiseaux à de petites nappes d’eau libre. Dans huit cas de telle nature dont nous avons été témoins pendant plus de 75 minutes d’observation, nous n’avons pas été en mesure de confirmer si les morses avaient réussi à capturer des eiders adultes en état de voler. Cependant, le taux élevé d’attaques de même que la réaction dramatique des eiders qui tentaient de s’échapper laissent entrevoir que les morses pourraient être des prédateurs efficaces en ce qui les concerne, ce qui pourrait exercer une influence sur l’expansion des eiders et sur leur bilan énergétique

Wintering Eiders Acquire Exceptional Se and Cd Burdens in the Bering Sea: Physiological and Oceanographic Factors

During late winter (March) in the Bering Sea, levels of Se in livers and Cd in kidneys of spectacled eiders Somateria fischeri were exceptionally high (up to 489 and 312 µg g−1 dry mass, respectively). Comparison of organ and blood samples during late winter, early spring migration, and breeding suggests that the eiders’ high Se and Cd burdens were accumulated at sea, with highest exposure during winter. High exposure may have resulted from high metabolic demands and food intake, as well as concentrations in food. In the eiders’ remote wintering area, their bivalve prey contained comparable Se levels and much higher Cd levels than in industrialized areas. Patterns of chlorophyll a in water and sediments indicated that phytoplankton detritus settling over a large area was advected into a persistent regional eddy, where benthic prey densities were higher than elsewhere and most eider foraging occurred. Se and Cd assimilated or adsorbed by bloom materials apparently also accumulated in the eddy, and were incorporated into the bivalve prey of eiders. Atmospheric deposition of dust-borne trace elements from Asia, which peaks during the ice-edge phytoplankton bloom from March to May, may augment processes that concentrate Se and Cd in eider prey. Compared with freshwater birds, some sea ducks (Mergini) accumulate much higher concentrations of trace elements, even with the same levels in food, with no apparent ill effects. Nevertheless, the absolute and relative burdens of different elements in sea ducks vary greatly among areas. Our results suggest these patterns can result from (1) exceptional accumulation and tolerance of trace elements when exposure is elevated by high food intake or levels in food, and (2) atmospheric and oceanographic processes that concentrate trace elements in local benthic food webs