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At least 1,099 records · Page 61Linked to original sources

Detecting grizzly bear use of ungulate carcasses using global positioning system telemetry and activity data

Global positioning system (GPS) wildlife collars have revolutionized wildlife research. Studies of predation by free-ranging carnivores have particularly benefited from the application of location clustering algorithms to determine when and where predation events occur. These studies have changed our understanding of large carnivore behavior, but the gains have concentrated on obligate carnivores. Facultative carnivores, such as grizzly/brown bears ( Ursus arctos ), exhibit a variety of behaviors that can lead to the formation of GPS clusters. We combined clustering techniques with field site investigations of grizzly bear GPS locations ( n = 732 site investigations; 2004–2011) to produce 174 GPS clusters where documented behavior was partitioned into five classes (large-biomass carcass, small-biomass carcass, old carcass, non-carcass activity, and resting). We used multinomial logistic regression to predict the probability of clusters belonging to each class. Two cross-validation methods—leaving out individual clusters, or leaving out individual bears—showed that correct prediction of bear visitation to large-biomass carcasses was 78–88%, whereas the false-positive rate was 18–24%. As a case study, we applied our predictive model to a GPS data set of 266 bear-years in the Greater Yellowstone Ecosystem (2002–2011) and examined trends in carcass visitation during fall hyperphagia (September–October). We identified 1997 spatial GPS clusters, of which 347 were predicted to be large-biomass carcasses. We used the clustered data to develop a carcass visitation index, which varied annually, but more than doubled during the study period. Our study demonstrates the effectiveness and utility of identifying GPS clusters associated with carcass visitation by a facultative carnivore.

Greater Yellowstone Ecosystem↗

Carryover effects across juvenile periods vary by species, cause-specific mortality, and stage-specific behavior

Conditions during one life-stage can carry over to influence fitness outcomes in later stages. How carryover effects manifest, however, may vary depending on the type and magnitude of risks that animals face in subsequent life-stages. The influence of carryover effects on cause-specific mortality remains largely unclear, particularly during vulnerable life-stages, including juvenile periods. We proposed and investigated the Cause-specific carryover hypothesis, that the relative influence of traits developed during a previous stage on mortality during the next vary by cause of mortality, using juvenile sagebrush songbirds (Brewer’s sparrow, Spizella breweri ; sagebrush sparrow, Artemisiospiza nevadensis ; sage thrasher, Oreoscoptes montanus ) as a focal system. We tracked birds during the post-fledging period (a juvenile period following the nestling period), identified the rates and causes of mortality, and tested the relative influence of different nestling traits (wing length, tarsus length, feather development, and body condition) on post-fledging mortality risk. Carryover effects varied across co-occurring species and differed by cause of mortality, and tarsus length had an unexpected and outsized impact on predation risk in the two sparrows. Although wing morphology typically is considered the most important trait for mobility in birds, most mortalities occurred during the first 5 days of the post-fledging period when altricial fledglings were incapable of flight and likely relied on running or scrambling ability to escape predators. Therefore, carryover effects were driven by stage-specific behavior during the period of highest risk. Developing animals may face investment trade-offs between morphological traits that influence different types of risk during different subsequent stages.

Wyoming↗

A Geochemical Trophic Cascade in Yellowstone's Geothermal Environments

We contrast the geochemistry of the Madison drainage, which has high concentrations of geothermal features, with the Lamar drainage of Yellowstone National Park, USA, and trace the consequences of geochemical differences through abiotic and biotic linkages in the ecosystem. Waters in the geothermal-dominated drainage contained anomalously high levels of fluoride (F) and silica (SiO 2 ). Soils, stream sediments, and surface waters that interact or mix with geothermal waters, in turn, had elevated F and SiO 2 concentrations compared to similar samples from the Lamar drainage. The geochemical differences were reflected in the chemistry of forage plants, with some plants from geothermally influenced areas containing four- to eightfold higher concentrations of F and SiO 2 than similar plants in the Lamar drainage. Geothermal heat reduced snowpack, and we found that elk (Cervus elaphus) concentrated in these refugia as snowpack increased each winter. The consequent high dietary intake of F in animals associated with the geothermal areas was confirmed by the finding that bone samples from elk living in the Madison drainage contained sixfold higher concentrations of F than samples collected from animals wintering in the Lamar drainage. High F exposure resulted in compromised dentition due to fluoride toxicosis, which was undoubtedly exacerbated by the abrasive action of silica. The consequent accelerated and aberrant tooth wear resulted in early onset of senescence, reduced life span, and an abbreviated age structure. We speculate that these altered demographics, combined with spatial heterogeneity of snowpack, will result in increased vulnerability of this large herbivore population to wolf predation and less resiliency to compensate demographically for predation.

Wyoming↗

Spatial relationships between an introduced snapper and native goatfishes on Hawaiian reefs

It has been suggested that the introduced blueline snapper (Lutjanus kasmira, Family: Lutjanidae) may adversely affect populations of native fishery species in Hawai'i through competition for spatial or dietary resources, or through predation on young fish. We studied the habitat use patterns of L. kasmira and several native reef fish species using direct observation by SCUBA divers. Habitat use patterns of the yellowtail goatfish (Mulloidichthys vanicolensis, Family: Mullidae) were most similar to those of L. kasmira. Both species were primarily found low in the water column and were closely associated with areas of vertical relief. Individual M. vanicolensis were found higher in the water column when L. kasmirawere present, but L. kasmira were not similarly affected by M. vanicolensis. This finding suggests asymmetrical competition for shelter, in which the dominant L. kasmira displaces M. vanicolensis farther into the water column. This displacement from the protection of the reef could increase the vulnerability of M. vanicolensisto predators and fishers. ?? Springer 2005.

Biological Invasions↗

Diet and conservation implications of an invasive chameleon, Chamaeleo jacksonii (Squamata: Chamaeleonidae) in Hawaii

We summarize information on current distribution of the invasive lizard Chamaeleo jacksonii and predict its potential distribution in the Hawaiian Islands. Potential distribution maps are based on climate models developed from known localities in its native range and its Hawaiian range. We also present results of analysis of stomach contents of a sample of 34 chameleons collected from native, predominantly dryland, forest on Maui. These data are the first summarizing prey range of this non-native species in an invaded native-forest setting. Potential distribution models predict that the species can occur throughout most of Hawaii from sea level to >2,100 m elevation. Important features of this data set are that approximately one-third of the diet of these lizards is native insects, and the lizards are consuming large numbers of arthropods each day. Prey sizes span virtually the entire gamut of native Hawaiian arthropod diversity, thereby placing a large number of native species at risk of predation. Our dietary results contrast with expectations for most iguanian lizards and support suggestions that chameleons comprise a third distinct foraging-mode category among saurians. The combination of expanding distribution, large potential range size, broad diet, high predation rates, and high densities of these chameleons imply that they may well become a serious threat to some of the Hawaiian fauna.

Hawai'i↗

Free-ranging domestic cats ( Felis catus ) on public lands: estimating density, activity, and diet in the Florida Keys

Feral and free-ranging domestic cats ( Felis catus ) can have strong negative effects on small mammals and birds, particularly in island ecosystems. We deployed camera traps to study free-ranging cats in national wildlife refuges and state parks on Big Pine Key and Key Largo in the Florida Keys, USA, and used spatial capture–recapture models to estimate cat abundance, movement, and activities. We also used stable isotope analyses to examine the diet of cats captured on public lands. Top population models separated cats based on differences in movement and detection with three and two latent groups on Big Pine Key and Key Largo, respectively. We hypothesize that these latent groups represent feral, semi-feral, and indoor/outdoor house cats based on the estimated movement parameters of each group. Estimated cat densities and activity varied between the two islands, with relatively high densities (~4 cats/km 2 ) exhibiting crepuscular diel patterns on Big Pine Key and lower densities (~1 cat/km 2 ) exhibiting nocturnal diel patterns on Key Largo. These differences are most likely related to the higher proportion of house cats on Big Pine relative to Key Largo. Carbon and nitrogen isotope ratios from hair samples of free-ranging cats (n = 43) provided estimates of the proportion of wild and anthropogenic foods in cat diets. At the population level, cats on both islands consumed mostly anthropogenic foods (>80% of the diet), but eight individuals were effective predators of wildlife (>50% of the diet). We provide evidence that cat groups within a population move different distances, exhibit different activity patterns, and that individuals consume wildlife at different rates, which all have implications for managing this invasive predator.

Florida↗

Feeding habits of an endemic fish, Oxygymnocypris stewartii , in the Yarlung Zangbo River in Tibet, China

Feeding habits of Oxygymnocypris stewartii were investigated based on monthly sampling in the Yarlung Zangbo River from August 2008 to August 2009. The gut contents of 194 individuals were analysed and quantified with numerical and gravimetric methods. This species can be considered a generalized and opportunistic predator feeding both on teleosts and aquatic insects. A multivariate analysis revealed noticeable variation in O. stewartii diet composition with fish size and season. The smaller specimens fed primarily on Cobitidae and Hydropsychidae. As they grew, Cyprinidae and Chironomidae larvae became important prey. The preferred food items were teleosts in summer and autumn. For winter and spring, the predominant prey switched to chironomidae larvae. Diet composition did not significantly vary between the sexes. Finally, a significant and positive correlation between predator and prey length was found. These findings provide the fundamental information better understanding the role of this important endemic species in the Yarlung Zangbo River food web.

Tibet↗

Bacterioplankton communities of Crater Lake, OR: Dynamic changes with euphotic zone food web structure and stable deep water populations

The distribution of bacterial and archaeal species in Crater Lake plankton varies dramatically over depth and with time, as assessed by hybridization of group-specific oligonucleotides to RNA extracted from lakewater. Nonmetric, multidimensional scaling (MDS) analysis of relative bacterial phylotype densities revealed complex relationships among assemblages sampled from depth profiles in July, August and September of 1997 through 1999. CL500-11 green nonsulfur bacteria (Phylum Chloroflexi) and marine Group I crenarchaeota are consistently dominant groups in the oxygenated deep waters at 300 and 500 m. Other phylotypes found in the deep waters are similar to surface and mid-depth populations and vary with time. Euphotic zone assemblages are dominated either by ??-proteobacteria or CL120-10 verrucomicrobia, and ACK4 actinomycetes. MDS analyses of euphotic zone populations in relation to environmental variables and phytoplankton and zooplankton population structures reveal apparent links between Daphnia pulicaria zooplankton population densities and microbial community structure. These patterns may reflect food web interactions that link kokanee salmon population densities to community structure of the bacterioplankton, via fish predation on Daphnia with cascading consequences to Daphnia bacterivory and predation on bacterivorous protists. These results demonstrate a stable bottom-water microbial community. They also extend previous observations of food web-driven changes in euphotic zone bacterioplankton community structure to an oligotrophic setting. ?? 2007 Springer Science+Business Media B.V.

Hydrobiologia↗

Lake trout ( Salvelinus namaycush ) suppression for bull trout ( Salvelinus confluentus ) recovery in Flathead Lake, Montana, North America

Non-native lake trout Salvelinus namaycush displaced native bull trout Salvelinus confluentus in Flathead Lake, Montana, USA, after 1984, when Mysis diluviana became abundant following its introduction in upstream lakes in 1968–1976. We developed a simulation model to determine the fishing mortality rate on lake trout that would enable bull trout recovery. Model simulations indicated that suppression of adult lake trout by 75% from current abundance would reduce predation on bull trout by 90%. Current removals of lake trout through incentivized fishing contests has not been sufficient to suppress lake trout abundance estimated by mark-recapture or indexed by stratified-random gill netting. In contrast, size structure, body condition, mortality, and maturity are changing consistent with a density-dependent reduction in lake trout abundance. Population modeling indicated total fishing effort would need to increase 3-fold to reduce adult lake trout population density by 75%. We conclude that increased fishing effort would suppress lake trout population density and predation on juvenile bull trout, and thereby enable higher abundance of adult bull trout in Flathead Lake and its tributaries.

Montana↗

Theory and application of semiochemicals in nuisance fish control

Controlling unwanted, or nuisance, fishes is becoming an increasingly urgent issue with few obvious solutions. Because fish rely heavily on semiochemicals, or chemical compounds that convey information between and within species, to mediate aspects of their life histories, these compounds are increasingly being considered as an option to help control wild fish. Possible uses of semiochemicals include measuring their presence in water to estimate population size, adding them to traps to count or remove specific species of fish, adding them to waterways to manipulate large-scale movement patterns, and saturating the environment with synthesized semiochemicals to disrupt responses to the natural cue. These applications may be especially appropriate for pheromones, chemical signals that pass between members of same species and which also have extreme specificity and potency. Alarm cues, compounds released by injured fish, and cues released by potential predators also could function as repellents and be especially useful if paired with pheromonal attractants in “push-pull” configurations. Approximately half a dozen attractive pheromones now have been partially identified in fish, and those for the sea lamprey and the common carp have been tested in the field with modest success. Alarm and predator cues for sea lamprey also have been tested in the laboratory and field with some success. Success has been hampered by our incomplete understanding of chemical identity, a lack of synthesized compounds, the fact that laboratory bioassays do not always reflect natural environments, and the relative difficulty of conducting trials on wild fishes because of short field seasons and regulatory requirements. Nevertheless, workers continue efforts to identify pheromones because of the great potential elucidated by insect control and the fact that few tools are available to control nuisance fish. Approaches developed for nuisance fish also could be applied to valued fishes, which suffer from a lack of powerful management tools.

Journal of Chemical Ecology↗

Modeling control of Common Carp (Cyprinus carpio) in a shallow lake–wetland system

The introduction of Common Carp (Cyprinus carpio) into North American waterways has led to widespread alteration of aquatic ecosystems. Control of this invader has proven extremely difficult due to its capacity for rapid population growth. To help understand how Common Carp can potentially be controlled we developed a population dynamics model (CarpMOD) to explore the efficacy of active and passive control measures that impose mortality on multiple life stages (embryos, juveniles and adults). We applied CarpMOD to Common Carp in Malheur Lake, a large shallow lake in Southeast Oregon, USA. Simulated control measures included commercial harvest of adults, trapping of juveniles, embryo electroshocking, and passive removal imposed via avian predation. Results from CarpMOD suggest that no single active removal method would decrease Common Carp biomass below the targeted 50 kg/ha threshold. Combinations of two or all three active removal methods could, however, reduce biomass below the desired threshold due to cumulative mortality on multiple life stages. CarpMOD simulations suggest that the level of carp removal necessary to reach the desired biomass threshold is approximately 40% at each life-stage, which may be unrealistic to maintain over longer time scales. Passive removal via avian predation may also contribute to suppression of Common Carp, but was not sufficient in isolation to reduce biomass below the desired threshold. Collectively, our results indicate control of Common Carp as a sole means of ecosystem restoration is unlikely to be effective in the system we modeled. This suggests additional means of restoration may be warranted, perhaps in combination with control of Common Carp, or development of more effective control measures.

Oregon↗

Seasonal assembly of arthropod communities on milkweeds experiencing simulated herbivory

The seasonal assembly of arthropod communities is shaped by biotic and abiotic aspects of the habitat that limit the appearance or activity phenology of potential community members. In addition, previous interactions within the community, such as herbivore-induced plant defensive responses, aggregation, and predator avoidance likely affect the assembly of arthropod communities on individual plants. We observed the phenology of arthropod communities and defensive plant traits on 100 milkweed ( Asclepias eriocarpa ) individuals at monthly intervals over a growing season. We experimentally wounded a subset of plants each month (April–August) to observe the effect of simulated added herbivore damage on the seasonal assembly of these arthropod communities. All plant traits and measures of arthropod communities changed over the season. The observed response to experimental leaf damage suggested a trend of induced susceptibility in early months, but not late months. Plants receiving early-season simulated herbivory experienced more subsequent leaf damage than unmanipulated plants. We observed several lagged correlations in our study indicating that blue milkweed beetle ( Chrysochus cobaltinus ) abundance was lower in months following high natural leaf damage, and that the abundance of a secondary omnivore ( Lygaeus kalmii ) and total predator abundance tended to follow months with high C. cobaltinus abundance. Ahistorical habitat factors determined much of the observed seasonality of arthropod communities, but induced responses to simulated herbivory also contributed historical effects that influenced arthropod community assembly.

Arthropod-Plant Interactions↗

Quantifying site-specific physical heterogeneity within an estuarine seascape

Quantifying physical heterogeneity is essential for meaningful ecological research and effective resource management. Spatial patterns of multiple, co-occurring physical features are rarely quantified across a seascape because of methodological challenges. Here, we identified approaches that measured total site-specific heterogeneity, an often overlooked aspect of estuarine ecosystems. Specifically, we examined 23 metrics that quantified four types of common physical features: (1) river and creek confluences, (2) bathymetric variation including underwater drop-offs, (3) land features such as islands/sandbars, and (4) major underwater channel networks. Our research at 40 sites throughout Plum Island Estuary (PIE) provided solutions to two problems. The first problem was that individual metrics that measured heterogeneity of a single physical feature showed different regional patterns. We solved this first problem by combining multiple metrics for a single feature using a within-physical feature cluster analysis. With this approach, we identified sites with four different types of confluences and three different types of underwater drop-offs. The second problem was that when multiple physical features co-occurred, new patterns of total site-specific heterogeneity were created across the seascape. This pattern of total heterogeneity has potential ecological relevance to structure-oriented predators. To address this second problem, we identified sites with similar types of total physical heterogeneity using an across-physical feature cluster analysis. Then, we calculated an additive heterogeneity index, which integrated all physical features at a site. Finally, we tested if site-specific additive heterogeneity index values differed for across-physical feature clusters. In PIE, the sites with the highest additive heterogeneity index values were clustered together and corresponded to sites where a fish predator, adult striped bass ( Morone saxatilis ), aggregated in a related acoustic tracking study. In summary, we have shown general approaches to quantifying site-specific heterogeneity.

Massachusetts↗

Top-down and bottom-up interactions influence fledging success at North America’s largest colony of Caspian terns (Hydroprogne caspia)

Our study investigated the influence of bottom-up and top-down drivers on the declining fledging success at a once thriving breeding colony of Caspian terns ( Hydroprogne caspia ). Situated at the mouth of the Columbia River, OR, East Sand Island (ESI) is home to the largest Caspian tern breeding colony in North America. Since 2001, the decline in fledging success of Caspian terns at ESI has been associated with a significant increase in average river discharge during May and June. During the years 2001–2011, the abundance of forage fish available to terns in the estuary was inversely related to river discharge. This relationship also apparently affected the reliance of nest predators on the tern colony as a food source, resulting in increased disturbance and decreased fledging success at the tern colony in years of higher river discharge. There was a significant longitudinal increase in disturbance rates by bald eagles ( Haliaeetus leucocephalus ) during June for terns nesting at the ESI colony, and eagle disturbance rates were positively associated with May river discharge. We also found a significant increase in kleptoparasitism rates of terns by hybrid glaucous-winged/western gulls ( Larus glaucescens x Larus occidentalis ) since 2001, and Caspian tern fledging success at ESI decreased with increasing average annual rates of gull kleptoparasitism. Our results support the hypothesis that the decline in Caspian tern fledging success at this large estuarine colony was primarily driven by the interaction of bottom-up and top-down factors, influencing tern fledging success through the food supply and triggering potential predators to identify the tern breeding colony as an alternative source of prey.

East Sand Island↗

Stable isotope analysis enhances our understanding of diamondback terrapin Malaclemys terrapin foraging ecology

Dietary studies on generalist predators may provide valuable information on spatial or temporal changes in the structure of ecological communities. We initiated this study to provide baseline data and determine the utility of stable isotope analysis (SIA) to evaluate the foraging strategies of an opportunistic reptilian predator, the diamondback terrapin ( Malaclemys terrapin ), which specializes in salt marshes and mangrove estuaries along the Atlantic and Gulf coasts. We evaluated stable carbon (δ 13 C) and nitrogen (δ 15 N) isotope values of multiple tissues from terrapins inhabiting mainland and island mangrove habitats in south Florida and potential food sources to examine spatial and temporal variations in terrapin resource use. We fit linear regression models to determine the best predictors of isotopic values for both terrapins and their prey, and Stable Isotope Bayesian Ellipses in R (SIBER) analysis to examine terrapin isotopic niche space and overlap between groups. We identified differences in terrapin isotopic δ 13 C and δ 15 N values among all sites. Blood and scute tissues revealed different isotopic compositions and niche overlap between sites, suggesting diets or foraging locations may change over time, and amount of variation is site specific. Niche overlap between size classes was larger for blood (short term) versus scute (long term), suggesting greater variability in food habits or resource isotopes over the long term versus short term. These results demonstrate the usefulness of SIA in examining the spatial and temporal variability in diamondback terrapin resource use within estuary systems and further define their niche within these dynamic food webs.

Estuaries and Coasts↗

Salt marsh habitats and diamondback terrapins in a rapidly changing climate: A review

The impacts associated with global climate change (e.g., sea-level rise, tropical storms, and warming temperatures) are expected to alter predator–prey interactions, foundation species, and plant community structure in coastal ecosystems. While the complex dynamics of these habitats have been examined under future climate predictions, few ecosystem models incorporate influences from fauna, such as the diamondback terrapin, the only estuarine turtle native to North America. This review examines the impacts of climate change on diamondback terrapins ( Malaclemys terrapin ) and the role that terrapins play as higher trophic level predators and keystone species in driving the dynamics of these ecosystems. We also review the potential implications of changes to terrapin populations on coastal ecosystems as a conservation challenge and suggest solutions to advance our understanding of those complex systems. Because of their role as a keystone and area-sensitive species that helps maintain healthy coastal habitats by foraging on herbivorous periwinkle snails, alterations to terrapin life history from climate change are expected, which could have significant impacts to the conservation of coastal habitats. Life history alterations could occur due to individual stressors, such as warming temperatures altering terrapin sex ratios. However, because of the complexity of these coastal systems, these stressors could also act additively or synergistically. Inclusion of faunal taxa such as the diamondback terrapin in modeling efforts examining climate change impacts to coastal ecosystems would better represent the complexity of these habitats thereby providing a more comprehensive evaluation of the entire ecosystem, resulting in more effective conservation strategies.

Estuaries and Coasts↗

Assessing environmental drivers and protist community dynamics that shaped the historic August 2022 Heterosigma akashiwo bloom in San Francisco Bay, California

San Francisco Bay, California, typically has chlorophyll a (chl- a ) concentrations below 10 µg L −1 , despite nutrient loadings exceeding those in many estuaries with recurring harmful algal blooms (HABs). However, in August 2022 there was a Heterosigma akashiwo (raphidophyte) bloom with chl- a concentrations exceeding 450 µg L −1 , resulting in widespread hypoxia and fish die-off. We used protist community (18S ribosomal ribonucleic acid gene metabarcoding) and environmental data collected 2–3 times per month from 2015–2023 to assess differences between 2022 and other years, and capture dynamics pre-, during, and post-bloom. The 2022 protist community clustered separately from other years in non-metric multidimensional scaling and hierarchical cluster analyses. A diatom bloom in June generated the highest median June chl- a concentration (15 µg L −1 ) recorded between 1977–2023 and drew down nutrients (nitrogen, silicate), leading to the establishment of a heterotrophic community in July with low dinoflagellate abundance (0.9% of protists versus 5.9% average in other years). The absence of dinoflagellates, potentially including H. akashiwo predators , plus suitable habitat (13–14 daylight hours, temperatures 17–21˚C, nitrate concentrations 17–24 µmol L −1 , neap tides) likely provided opportunity for H. akashiwo to proliferate from 0.03% to 48% of protists between July 20 and August 25. As the bloom collapsed, the genus Gyrodinium (dinoflagellate predator of H. akashiwo ) increased from 3.0% to 52% of protists between August 25 and August 31, likely exerting grazing pressure. Results show that antecedent protist community dynamics play important roles in shaping HABs, and interactions between biotic and abiotic factors govern the development of blooms.

California↗

Group foraging by a stream minnow: shoals or aggregations?

The importance of social attraction in the formation of foraging groups was examined for a stream-dwelling cyprinid, the rosyside dace, Clinostomus funduloides. Dace arrivals and departures at natural foraging sites were monitored and tested for (1) tendency of dace to travel in groups, and (2) dependency of arrival and departure rates on group size. Dace usually entered and departed foraging sites independently of each other. Group size usually affected neither arrival rate nor departure probability. Thus, attraction among dace appeared weak; foraging groups most often resulted from dace aggregating in preferred foraging sites. The strongest evidence of social attraction was during autumn, when dace departure probability often decreased with increasing group size, possibly in response to increased threat of predation by a seasonally occurring predator. Dace also rarely avoided conspecifics, except when an aggressive individual defended a foraging site. Otherwise, there was little evidence of exploitative competition among dace for drifting prey or of foraging benefits in groups, because group size usually did not affect individual feeding rates. These results suggest that the benefits of group foraging demonstrated under laboratory conditions in other studies may not always apply to field conditions.

Animal Behaviour↗