Environmental heterogeneity, spatial segregation of prey, and the utilization of southwest Atlantic mudflats by migratory shorebirds

We studied the relationships between the habitat use of migratory shorebirds and the spatial distributions of the Southwestern Atlantic Fiddler Crab Uca uruguayensis , polychaetes, sediment characteristics and tidal levels in the Río de La Plata estuary, Argentina, where U. uruguayensis is one of the most important intertidal species. Crabs have a well-defined patchy distribution that is segregated spatially from that of polychaetes. Crab density on the surface varied across the tidal cycle, reaching maximum values during low tide. Polychaete density decreased with depth but showed no change through the tidal cycle; however, given that sediment penetrability did change during the same period, their availability to probing shorebirds is expected to change. Habitat use by shorebirds followed the spatial distribution of prey; shorebirds that foraged on polychaetes (White-rumped Sandpiper Calidris fuscicollis , Two-banded Plover Charadrius falklandicus and Hudsonian Godwit Limosa haemastica ) focused their attention on the areas with the highest densities of polychaetes, whereas species that preyed mostly on crabs (Ruddy Turnstone Arenaria interpres , Whimbrel Numenius phaeopus and Grey Plover Pluvialis squatarola ) predominantly used areas with crabs. This segregation occurred particularly during low tide, a period in which polychaetes became fully available. Results show that the spatial and temporal heterogeneity of mudflats in relation to the types and availability of prey has a strong effect on shorebird habitat use.     

Distribution of shorebirds at the Berg River estuary, South Africa, in relation to foraging mode, food supply and environmental features

Foraging densities of Curlew Sandpipers Calidris ferruginea and Grey Plovers Pluvialis squa-tarola at the Berg River estuary, South Africa, are examined in relation to prey and habitat attributes.
There is a substantial overlap in the prey species eaten by Curlew Sandpipers and Grey Plovers, but the distribution patterns of the two species within the estuary could not be explained by the same prey attributes.
The distribution of tactilely foraging Curlew Sandpipers was determined largely by the numerical abundance of nereid worms. They foraged preferentialiy on muddy rather than sandy substrata, but their distribution could not be explained on the basis of substratum penetrability. The density of visually foraging Grey Plovers was correlated with the biomass, rather than abundance, of the larger of the two common nereids on the estuary. The favoured foraging area of Grey Plovers had a high vegetation cover, but their foraging dispersion was not influenced by substratum type.
The energy intake rates of Curlew Sandpipers were broadly inversely related to their foraging density, although the rate of prey capture was maximal at high prey density. These birds may use neighbour success rates in selecting foraging sites. Energy intake rates of Grey Plovers were very consistent across the estuary, suggesting that individuals occupy or defend areas which provide access to comparable food resources.     

Position of horseshoe crabs in estuarine food webs: N and C stable isotopic study of foraging ranges and diet composition

To discern the position of horseshoe crabs as a potentially important predator in estuarine food webs, we determined where they foraged and what they ate. We used N and C stable isotopes to link adult horseshoe crabs to their oraging locations and potential food sources in Pleasant Bay, Cape Cod. The δ¹⁵N in tissues of horseshoe crabs and their potential foods suggest crabs were loyal to local foraging sites and did not forage substantially in subestuaries receiving >110 kg N ha⁻¹ year⁻¹. Among locations where crabs foraged, δ¹³C values in potential foods showed that food webs in subestuaries subject to higher N loads were supported by algal producers, while food webs in subestuaries with lower N loads were also supported by Spartina. δ¹³C values in horseshoe crab tissue did not change with load, suggesting they ate a mixed diet, regardless of N load. N and C isotopes in horseshoe crab feces were similar to signatures of estimated diet, suggesting low assimilation efficiency, perhaps due to ingestion of low quality organic matter. Although horseshoe crabs were relatively opportunistic in foraging habits, conservation or culture of horseshoe crabs may require habitats with higher water quality, ample particulate organic matter, and supporting a variety of prey.     

Growth-related changes in diet and foraging behavior of the yellow wrasseThalassoma lutescens at Kuchierabu Island

Ontogenetic changes in diet and foraging behavior ofThalassoma lutescens were examined in shallow reef habitats around Kuchierabu Island, southern Japan. This species mainly took small benthic invertebrates, including gammarids, polychaetes, sipunculids, chitons, crabs, gastropods, pelecypods and urchins from algal mats. Larger fish consumed correspondingly larger prey, although most of the latter were armored with hard exoskeletons, shells or body plates (e.g., crabs, gastropods, pelecypods and urchins). Such hard parts were crushed with the molar-like, pharyngeal teeth which develop with fish growth, allowing exploitation of such larger, hard-bodied prey. Because the densities of larger prey species were relatively low in the initial habitats foraged, larger fish shifted their foraging attention to rock and coral crevices, where the prey species dwelt in greater numbers, as well as foraging over larger areas. Such behavioral changes maintained high foraging efficiency in larger fish.     

Insect prey foraging strategies in Callicebus oenanthe in northern Peru

Titi monkeys (genus Callicebus) are small-bodied platyrrhines that supplement their predominantly frugivorous diet with variable amounts of leaves, seeds, and/or arthropod prey. Notable interspecific variation in the amount of insect prey in the diet has been observed in Callicebus, ranging from 0% to 20%. In this study, I investigate the degree and type of prey foraging in a little-known species, Callicebus oenanthe inhabiting a fragmented, secondary forest on the foothills of the Andes in northern Peru. I present data on prey type, prey search and capture techniques, substrate/vegetation use, foraging height, prey capture efficiency, and seasonal variation of insect prey foraging in one group of C. oenanthe observed from January to August 2005. Insect prey accounted for 22% of the diet, the highest amount reported for any Callicebus species to date, and insects from at least six different orders were included. C. oenanthe was mainly an investigative forager of hidden prey, manipulating easy-to-open substrates such as rolled up leaves, and hunted ant swarms and larger insects opportunistically. Insect foraging was predominant during the dry season (26%) and decreased during the wet season (13%). The study group foraged mostly in the understory (2-6 m) within vine-laden shrubs and trees, which may conform to an anti-predator strategy of crypticity. Overall the group had an 83% insect capture success rate. These data suggest that insect prey is an important part of the diet of C. oenanthe and may be especially notable during periods of resource scarcity. This study adds to the knowledge concerning insect prey foraging in Callicebus, which can have an important role in defining ecological strategies in the selection of secondary protein food resources within a given ecosystem.     

Day and night feeding territoriality in Willets Catoptrophorus semipalmatus and Whimbrel Numenius phaeopus during the non-breeding season in the tropics

This study was carried out to determine whether or not, and under what conditions, Willets Catoptrophorus semipalmatus and Whimbrel Numenius phaeopus continue to defend and use their daytime feeding territories at night in a tropical environment. The study was conducted in coastal Venezuela by registering, by night and by day, the behaviour and the position of colour-marked and radio-tagged birds. Night observations were made with the use of a light intensifier.
In Willets, both territorial and non-territorial birds were observed. The use and defence of territories was observed only on sandy mud areas where Fiddler Crabs Uca cumulanta were plentiful; the soft mud sites were used only by non-territorial birds. All Whimbrel were territorial. Individuals of both species defending a given space during daytime continued to occupy and defend the same area during the night. The proportion of birds foraging at night was higher on moonlit than on moonless nights. Territorial defence involved alert postures, parallel walks, calls and the chasing of intruders. Agonistic encounters between Willets and Whimbrel and other species were rare. Willets were territorial at all states of the tide except when high tides flooded the territories. Tide had no effect on the time of feeding in Whimbrel. Moonlight was clearly the factor conditioning the occurrence of nocturnal foraging on territories by both species. Foraging strategies and the type of prey and substrata explain why the incidence of night foraging varied with moonlight in territorial Willets and Whimbrels and not in non-territorial individuals.     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Green crab (Carcinus maenas) foraging efficiency reduced by fast flows

Predators can strongly influence prey populations and the structure and function of ecosystems, but these effects can be modified by environmental stress. For example, fluid velocity and turbulence can alter the impact of predators by limiting their environmental range and altering their foraging ability. We investigated how hydrodynamics affected the foraging behavior of the green crab (Carcinus maenas), which is invading marine habitats throughout the world. High flow velocities are known to reduce green crab predation rates and our study sought to identify the mechanisms by which flow affects green crabs. We performed a series of experiments with green crabs to determine: 1) if their ability to find prey was altered by flow in the field, 2) how flow velocity influenced their foraging efficiency, and 3) how flow velocity affected their handling time of prey. In a field study, we caught significantly fewer crabs in baited traps at sites with fast versus slow flows even though crabs were more abundant in high flow areas. This finding suggests that higher velocity flows impair the ability of green crabs to locate prey. In laboratory flume assays, green crabs foraged less efficiently when flow velocity was increased. Moreover, green crabs required significantly more time to consume prey in high velocity flows. Our data indicate that flow can impose significant chemosensory and physical constraints on green crabs. Hence, hydrodynamics may strongly influence the role that green crabs and other predators play in rocky intertidal communities.     

Integrating disparate datasets to model the functional response of a marine predator: A case study of harbour porpoises in the southern North Sea

1. Quantifying consumption and prey choice for marine predator species is key to understanding their interaction with prey species, fisheries, and the ecosystem as a whole. However, parameterizing a functional response for large predators can be challenging because of the difficulty in obtaining the required data on predator diet and on the availability of multiple prey species.
2. This study modeled a multi‐species functional response (MSFR) to describe the relationship between consumption by harbour porpoises (Phocoena phocoena) and the availability of multiple prey species in the southern North Sea. Bayesian methodology was employed to estimate MSFR parameters and to incorporate uncertainties in diet and prey availability estimates. Prey consumption was estimated from stomach content data from stranded harbour porpoises. Prey availability to harbour porpoises was estimated based on the spatial overlap between prey distributions, estimated from fish survey data, and porpoise foraging range in the days prior to stranding predicted from telemetry data.
3. Results indicated a preference for sandeels in the study area. Prey switching behavior (change in preference dependent on prey abundance) was confirmed by the favored type III functional response model. Variation in the size of the foraging range (estimated area where harbour porpoises could have foraged prior to stranding) did not alter the overall pattern of the results or conclusions.
4. Integrating datasets on prey consumption from strandings, predator foraging distribution using telemetry, and prey availability from fish surveys into the modeling approach provides a methodological framework that may be appropriate for fitting MSFRs for other predators.

     

Nereidid polychaetes as the major diet of migratory shorebirds on the estuarine tidal flats at Fujimae-higata in Japan

The dietary items of five migratory shorebirds, Dunlin (Calidris alpina), Red-necked Stint (C. ruficollis), Grey Plover (Pluvialis squatarola), Whimbrel (Numenius phaeopus) and Black-headed Gull (Larus ridibundus), were examined by analyses of fecal droppings during the birds' migration or wintering and by surveys of macrobenthic fauna around their foraging sites on the tidal flats of Fujimae-higata, Nagoya, central Japan. Body parts of nereidid, capitellid, and spionid polychaetes and crustaceans were found in fecal droppings from all of these shorebirds. Two nereidid species (Hediste diadroma and Neanthes succinea) with relatively large body sizes seemed to be the majority dietary items. At one site, H. diadroma was dominant in terms of biomass (40-370 g/m(2)) throughout year except, for less than 1 g/m(2) in March and May (within or just after reproduction of this species). Monthly changes in the occurrence of food items in fecal droppings of C. alpina were examined in 1999 and 2000. Most (85-100%) of the fecal droppings contained nereidid body parts, including Hediste-specific simple chaetae from November to April, whereas only 23% of the droppings contained them in May. Chaetae of capitellid or spionid polychaetes were frequently found from January to April (38-86% of droppings). Crustacean body parts, including amphipod appendages, were frequently found from March to May (86-100% of droppings). The relationship between foraging habits of the shorebirds and the life history of their major prey nereidid species is discussed.     

The effect of sward height and drainage on Common Starlings Sturnus vulgaris and Northern Lapwings Vanellus vanellus foraging in grassland habitats

Agricultural change is often cited as a causal factor in the decline of the UK's farmland birds because bird declines have mirrored changes in agricultural practices. Although much is known about the mechanisms driving population declines on arable systems, mechanisms in grassland systems are relatively poorly studied, despite receiving a similar degree of intensification. Agricultural intensification may affect bird declines by reducing food abundance or accessibility, forager mobility or predation risk. Here we examine experimentally the effects of sward height on the foraging behaviour of adult Common Starlings Sturnus vulgaris , and the effects of sward height and drainage on the behaviour of Northern Lapwing Vanellus vanellus chicks. Both species are declining across the UK and both forage in farmed grassland habitats, but they differ in their foraging methods because Starlings probe for soil invertebrates whereas Lapwing chicks glean prey from surfaces. Overall, after controlling for prey abundance, short swards were found to be more productive for both species. Prey capture rate within foraging bouts did not differ with sward height for Starlings, but Starlings spent more time foraging on short swards and captured 33.2% more prey. Starlings walked more steps on short swards. Lapwing chick foraging rates declined as sward height increased. Soil moisture was not found to be a predictor of Lapwing chick foraging rates within the observed range. Our results suggest that short swards are a more profitable foraging habitat for soil and surface invertebrate feeders. Short swards may facilitate surface prey detection, improve forager mobility and increase foraging time by altering vigilance patterns. Provision of short swards in areas where these are lacking could be simple method of improving foraging habitats for grassland birds.     

Spatial and trophic niche differentiation in two sympatric lizards (Tropidurus torquatus and Cnemidophorus ocellifer) with different foraging tactics

Abstract In a ‘restinga’ habitat of southeastern Brazil, we studied the food habits and the microhabitat use of two lizards with distinct foraging modes: the tropidurid Tropidurus torquatus, a sit-and-wait predator, and the teiid Cnemidophorus ocellifer, a wide forager. The diet of the two species differed strongly, indicating a low level of similarity in their trophic niche. The sit-and-wait predator fed mainly on mobile prey, whereas the wide forager fed mainly on sedentary prey (larvae). The spatial niche breadth of T. torquatus was larger than that of C. ocellifer. Despite interspecific differences, the two species overlapped greatly in micro-habitat use. The data indicate that at Linhares the two lizard species differed more in food resources than in microhabitat, and that most of the food differences reflect the foraging patterns of the species.     

The foraging ecology of sympatric marine fish in the genus Embiotoca (Embiotocidae): Importance of foraging behavior in prey size selection

Summary Two locally sympatric temperate marine reef fishes, Embiotoca jacksoni and E. lateralis (Embiotocidae), have high taxonomic similarity in diets. Subdivision of gammarid amphipods, their principal prey, was found. E. jacksoni took more tubicolous gammarid amphipods whereas E. lateralis consumed mostly free-living individuals. The species differed considerably with respect to between-individual variability in taxonomic compositions of their diets. Each E. jacksoni closely resembled other conspecifics in this regard while individual E. lateralis displayed very high between-fish variation. The principal interspecific difference in fish diets concerned the sizes of prey items taken. E. jacksoni ate small but very common items and the mean prey weight in their guts did not differ from random collections of available prey. E. lateralis concentrated on large, rarer sizes such that the average prey weight in their guts was much heavier than available or in the diet of E. jacksoni of the same length. Disparate foraging behaviors was a much better indicator of the relative differences in diets of these two fishes than was external fish morphology. E. jacksoni, which can winnow prey items from unwanted debris, was a relatively indiscriminant forager. E. lateralis did not winnow but actively searched for prey. This species was a much more discriminating forager, but displayed much variability in foraging behavior.     

Post-migratory care of young by Crab Plovers Dromas ardeola

The Crab Plover Dromas ardeola is the only waterbird species known to provision offspring well after the post-reproductive migration and through overwintering. A few previous quantitative studies have reported inconclusively that juveniles begged rarely, and also indiscriminately at other juveniles. Here, we describe the feeding behaviour of adult and juvenile Crab Plovers during the first part of the wintering period. Juveniles begged frequently for food, always toward adults, and obtained 0.1 prey items/min from them. Begging birds obtained the largest prey items of those captured by the adults. When foraging alone, juveniles captured prey at the same rate as adults, but captured smaller crabs. The feeding success of adults was not altered by the presence of the begging juvenile. Juveniles depended partly on adults during the first part of the overwintering period, but were almost independent towards the end. Crab Plovers may adopt such unusually prolonged care because they need to abandon their breeding areas quickly, when environmental conditions are extreme. Large numbers of Crab Plovers overwinter in a few areas, now subject to human alteration. Isolated adults may forage on other areas such as narrow and disturbed shores, but juveniles may require wide beaches, suitable for group foraging, that should be considered as key areas for the recruitment of juveniles into the population.     

Foraging tactics of a terrestrial salamander: Sustained yield in territories

Territorial red-backed salamanders (Plethodon cinereus) were given a high density of two prey types differing in size and caloric profitability. They chose a diet that approximated optimal foraging when they foraged in areas previously marked with advertisement pheromones. However, when foraging in previously unmarked or conspecific-marked areas they did not forage optimally, in a short time frame. In the latter two situations they evinced a lower rate of net energy due to their failure to specialize on the more profitable prey type and their longer intercapture intervals while time was devoted either to marking behaviour or submissive posturing. The salamanders appeared to opt for a strategy of sacrificing initial caloric yield until they had established marked territories and then switching to a higher sustained caloric yield. The data show that experimental handling of a predator per se can alter its foraging behaviour. Also a predator just initiating a territory may forage quite differently from one that has already established ownership, making the time frame during which observations are made important to understanding the predator's foraging strategy.     

Foraging time allocation in relation to sex by the gulf coast fiddler crab (Uca panacea)

Summary Schoener (1971) proposed that the reproductive demands of animals should be important in shaping their foraging behavior because fitness is affected. He defined two forager types: energy maximizers (reproductive success depends on energetic intake) and time minimizers (reproductive success depends on time spent in activities other than foraging), and suggested that females most often illustrate the former and males the latter. We tested whether mating activities influence the foraging behavior of Uca panacea, and the predictions that females would be energy maximizers because of their reproductive strategy and that males would also be energy maximizers because of their courtship activity. Time allocated to foraging by 800 male and female fiddler crabs (at two sites) was quantified; no significant difference in foraging time was found between the sexes. Both male and female crabs allotted a large portion of their time to foraging because both sexes depend on stored energy during their reproductive bouts. Our results show that the particular forager type can be predicted based on reproductive demands, but a forager type can not always be assigned to a particular sex without consideration of all important ecological and physiological factors determining reproductive success.     

Predatory fish sounds can alter crab foraging behaviour and influence bivalve abundance

The risk of predation can have large effects on ecological communities via changes in prey behaviour, morphology and reproduction. Although prey can use a variety of sensory signals to detect predation risk, relatively little is known regarding the effects of predator acoustic cues on prey foraging behaviour. Here we show that an ecologically important marine crab species can detect sound across a range of frequencies, probably in response to particle acceleration. Further, crabs suppress their resource consumption in the presence of experimental acoustic stimuli from multiple predatory fish species, and the sign and strength of this response is similar to that elicited by water-borne chemical cues. When acoustic and chemical cues were combined, consumption differed from expectations based on independent cue effects, suggesting redundancies among cue types. These results highlight that predator acoustic cues may influence prey behaviour across a range of vertebrate and invertebrate taxa, with the potential for cascading effects on resource abundance.     

Intraspecific differences in the diet of Antarctic fur seals at Nyrøysa, Bouvetøya

Intraspecific differences in the diets of many species of pinnipeds are to be expected in view of the great differences in morphology, life history and foraging behaviour between the sexes of many species. We examined the diet of the Antarctic fur seal population at Bouvetøya, Southern Ocean, to assess intersexual differences. This was made possible by the analysis of prey remains extracted from scats and regurgitations collected in areas used primarily by one or the other sex. The results indicate that both males and females feed primarily on Antarctic krill Euphausia superba with several species of fish and squid being taken, likely opportunistically given their prevalence. Significant differences were identified in the frequency of occurrence of otoliths in scats and the percentage numerical abundance of the major fish prey species in the diet. Adult males ate a smaller quantity of fish overall, but ate significantly more of the larger fish species. The greater diving capabilities of males and the fact that they are not limited in the extent of their foraging area by having to return regularly to feed dependant offspring may play a role in the differences found between the diets of males and females. Additionally, females might be more selective, favouring myctophids because they are richer in energy than krill. The absence of major differences in the diet between the sexes at this location is likely due to the high overall abundance of prey at Bouvetøya.     

Quantifying the effects of habitat structure on prey detectability and accessibility to farmland birds

For species that rely on visual cues to detect prey items, increasing the structural complexity of a patch can greatly influence forager behaviour through consequent reductions in prey detectability and accessibility. These effects are likely to manifest themselves in terms of foraging site selection and there is plentiful evidence for preferential site selection for a suite of taxa. However, the underlying effects of habitat structure on foraging behaviour, which are likely to drive these observed site selections, are much less well understood. We present the results of two studies designed to quantify the effects of vegetation structure on prey detectability and accessibility to avian invertebrate feeders and granivores on farmland. There was a significant negative relationship between potential prey detectability and both distance and vegetation height in cereal crops and stubbles for Northern Lapwings Vanellus vanellus . The interscan distance travelled by Lapwings differed significantly between habitats, with longer distances travelled in cereal crops and harrowed compared with ploughed soil and grasses. The peck rate, head-up rate and mean search period of foraging Chaffinches Fringilla coelebs were not affected by increasing vegetation structure but forager mobility was significantly reduced. We hope that by quantifying the effects of vegetation structure on prey detectability and accessibility we can highlight the importance of considering these factors, as well as prey abundance, when developing management strategies for farmland birds.     

Red-gartered Coot Fulica armillata feeding on the grapsid crab Cyrtograpsus angulatus: advantages and disadvantages of an unusual food resource

The behaviour of Red-gartered Coots feeding on an unusual food source was examined at Mar Chiquita Coastal Lagoon, Argentina. The grapsid crab Cyrtograpsus angulatus made up all observed prey items, and 61% were small. Both handling and foraging duration increased with the size of captured crabs, but foraging efficiency decreased. Crab availability affected both the dive duration of the Coots and their foraging decisions with regard to prey-size selection. Two species of gull were observed kleptoparasitizing Coots, especially when the Coot was handling medium or large crabs. Feeding by Coots on Cyrtograpsus angulatus has not been previously documented and may be a feeding innovation. Our estimations suggest that Coots were foraging optimally, since smaller crabs were more energetically profitable.