Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

Molecular studies on the ouabain binding site of the Na+, K+-ATPase in milkweed butterflies

Summary. The Na⁺, K⁺-ATPase of the Monarch butterfly (Danaus plexippus) is insensitive to the inhibition by cardiac glycosides due to an amino acid replacement: histidine instead of asparagine at position 122 of the α-subunit representing the ouabain binding site. By PCR amplification of the DNA sequence of this site, a PCR product of 270 bp was obtained from DNA extracted from Danainae species (Danaus plexippus, D. chrysippus, D. gillipus, D. philene, D. genutia, Tirumala hamata, Euploea spp., Parantica weiskei, P. melusine), Sphingidae (Daphnis nerii) and mimics of milkweed butterflies (Hypolimnas missipus, Limenitis archippus and L. arthemis, Nymphalidae). Analysis of the nucleotide sequences revealed that the single point mutation in the ouabain binding domain (AAC-Asn for CAC-His) was present only in Danaus plexippus, but not in the other species investigated. Since these milkweed butterflies also store cardenolides, other structural modifications of the Na⁺, K⁺-ATPase may have occurred or other strategies of cardenolide tolerance have been developed. Received 15 May 2000; accepted 29 June 2000     

Chemical defence in chewing and sucking insect herbivores: Plant-derived cardenolides in the monarch butterfly and oleander aphid

Summary Cardenolide sequestration by a hemimetabolous aphid and a holometabolous butterfly from the neotropical milkweed,Asclepias curassavica L., is compared. The oleander aphid,Aphis nerii B. de F., sequestered a similarly narrow range of cardenolide concentrations to the monarch butterfly,Danaus plexippus (L.), from the wide range of concentrations available in leaves of A.curassavica. However, A.nerii sequestered significantly less cardenolide (269 µg/0.1 g) thanD. plexippus (528 µg/0.1 g). The honeydew excreted by A.nerii was comprised of 46% cardenolide. The complete polarity range of 25 cardenolides detected by thin layer chromatography in A.curassavica was represented in the 17 whole aphid cardenolides and the 20 aphid honeydew cardenolides detected. D.plexippus sequestered a narrower polarity range of 11 cardenolides, having eliminated low polarity cardenolide genins and glycosides. It is suggested that these chemical differences may be related to interactions among the broad feeding tactics of sucking or chewing milkweed leaves, life history constraints of holometabolyversus hemimetaboly, the distribution of milkweed food resources in space and time, and the dynamics of natural enemies.     

Grassland Bird Responses to Land Management in the Largest Remaining Tallgrass Prairie

Abstract: Extensive habitat loss and changing agricultural practices have caused widespread declines in grassland birds throughout North America. The Flint Hills of Kansas and Oklahoma—the largest remaining tallgrass prairie—is important for grassland bird conservation despite supporting a major cattle industry. In 2004 and 2005, we assessed the community, population, and demographic responses of grassland birds to the predominant management practices (grazing, burning, and haying) of the region, including grasslands restored under the Conservation Reserve Program (CRP). We targeted 3 species at the core of this avian community: the Dickcissel (Spiza americana), Grasshopper Sparrow (Ammodramus savannarum), and Eastern Meadowlark (Sturnella magna). Bird diversity was higher in native prairie hayfields and grazed pastures than CRP fields, which were dominated by Dickcissels. Although Dickcissel density was highest in CRP, their nest success was highest and nest parasitism by Brown‐headed Cowbirds (Moluthrus ater) lowest in unburned hayfields (in 2004). Conversely, Grasshopper Sparrow density was highest in grazed pastures, but their nest success was lowest in these pastures and highest in burned hayfields, where cowbird parasitism was also lowest (in 2004). Management did not influence density and nest survival of Eastern Meadowlarks, which were uniformly low across the region. Nest success was extremely low (5–12%) for all 3 species in 2005, perhaps because of a record spring drought. Although the CRP has benefited grassland birds in agricultural landscapes, these areas may have lower habitat value in the context of native prairie. Hayfields may provide beneficial habitat for some grassland birds in the Flint Hills because they are mowed later in the breeding season than elsewhere in the Midwest. Widespread grazing and annual burning have homogenized habitat—and thus grassland‐bird responses—across the Flint Hills. Diversification of management practices could increase habitat heterogeneity and enhance the conservation potential of the Flint Hills for grassland birds.     

First-instar monarch larval growth and survival on milkweeds in southern California: effects of latex, leaf hairs and cardenolides

Growth rate and survival of first-instar larvae of Danaus plexippus, a milkweed specialist, depended on milkweed species, and was related to the amount of latex produced from wounds, leaf cardenolide concentrations and the presence of leaf hairs. Larval growth was more rapid and survival was higher on leaves of Asclepias californica with experimentally reduced latex, and this species has characteristically high latex, low- to mid-range cardenolide concentrations, and very hirsute leaves. Similarly, growth was higher on reduced latex leaves of both A. eriocarpa (a high latex/high cardenolide, hirsute species) and A. erosa (glabrous fleshy leaves, high latex/high cardenolides). There were no differences in either survival or growth rate between larvae on reduced latex or control leaves of the low latex/low cardenolide A. fascicularis with soft glabrous leaves and both survival and growth rate were higher on this species than the other species tested. Larval growth rates on leaves with reduced latex were similar among ten milkweed species tested to date but differed from growth rates on intact leaves suggesting that latex and possibly included cardenolides are both important in first-instar monarch larval growth, development and survival. We show for a range of ecologically important milkweeds that experiments on cut plant material (no latex outflow) lead to higher growth rates compared to intact plants. Such laboratory assays based on detached leaves will be misleading if the objective is to determine the impact of treatments such as Bt-maize pollen on monarchs on field plants.     

Sex-biased costs in nest defence behaviours by lesser snow geese (Chen caerulescens): consequences of parental roles?

Nest defence behaviours, such as attacking predators and defending against predator attacks, expose birds to risk of injury and death. However, direct costs of such behaviours are poorly documented. To evaluate potential costs of nest defence behaviours in lesser snow geese (Chen caerulescens), we (1) estimated the proportion of interactions between arctic foxes (Alopex lagopus) and geese that resulted in physical contact and (2) examined how nest defence behaviours varied between male and female geese. We separated interactions into attacks initiated by foxes (attacks by foxes) and attacks initiated by geese (attacks by geese). Risks associated with attacks by geese were considerably lower than the risks associated with attacks by foxes; only 1 of 1,179 attacks by geese resulted in physical contact between foxes and geese, whereas 26 of 89 attacks by foxes involved such contact (two female geese were killed during these attacks). Attacks by geese were made almost exclusively by male geese (>97%), whereas female geese were involved in 75% of all attacks by foxes that resulted in physical contact with geese. There was, thus, a considerable difference in risks associated with male and female nest defence behaviours. We suggest that parental roles during nesting (i.e., females incubate and males guard) expose female geese to greater risk of injury and death. Male geese may, however, reduce the risk of injury or death to their mates with pre-emptive attacks on foraging foxes.     

Differential parasitism of least flycatchers and yellow warblers by the brown-headed cowbird

Summary Brown-headed cowbirds (Molothrus ater) parasitized yellow warblers (Dendroica petechia) six times more frequently than least flycatchers (Empidonax minimus) nesting in the same riparian habitat in southern Manitoba. This difference was surprising because least flycatchers were higher quality hosts; flycatchers always accepted cowbird eggs while warblers did so on only about half the occasions. Both hosts were equally good foster parents, at least until fledging; thus, once an egg was accepted there was probably no further advantage in parasitizing one species over the other. The probability that a nest was parasitized decreased with increasing nest height, perhaps as a consequence of the cowbird's habit of searching for nests close to the ground. Since least flycatchers nested higher in the canopy than yellow warblers, a greater proportion of flycatcher nests probably were not detected by cowbirds. Nevertheless, nest height alone could not account fully for the lower incidence of parasitism on flycatchers. Clutch initiation in flycatchers peaked 6 days after warbler clutch initiations and 5 days after that of cowbirds. Despite the later peak in initiations, flycatcher nests were always available as potential hosts over the entire cowbird laying season and it was not until new clutch initiations by warblers declined in late summer that flycatchers were actually used as hosts. Because least flycatchers responded more aggressively than yellow warblers to a model female cowbird at the nest, we concluded that greater nest defense by flycatchers may have also reduced the rate of brood parasitism in this host. Together, our results suggest the large difference in the frequency of parasitism between these two hosts was primarily a product of nest location but that differences in host nest-defense and breeding season asynchrony may have contributed to preferential host selection.Offprint requests to: J.V. Briskie     

The organization of work in Polybia occidentalis: costs and benefits of specialization in a social wasp

Summary Nest construction, a complex social activity requiring the coordination of 3 tasks (Fig.2), was compared in large (<350 adults) and small (<50 adults) colonies of Polybia occidentalis. The 3 tasks—water foraging, pulp foraging, and building—are performed by 3 separate groups of workers (Fig.4). Of the 8 acts comprising the 3 tasks, 5 regularly involve the transfer of water or pulp from one worker to another on the nest.Small colonies required nearly twice as long (35.4 worker-min) as large colonies (20.1 workermin) to complete a unit amount of construction work. Behavioral acts involving material transfer among workers were responsible for most of the increase in small colonies. In other words, the waiting times experienced by material donors and recipients were greater in small colonies. In small colonies workers switched among the three tasks more frequently than in large colonies (Fig. 4). This was the result of more frequent switching by generalists (workers that performed 2 or 3 of the tasks), rather than by a decrease in the proportion of specialists (workers performing only 1 task type) (Fig. 3).The series-parallel system by which Polybia occidentalis organizes nest construction has a major advantage over the series operation of solitary wasps. Pulp foragers collect and carry loads that are 6.1 times as large as builders can work with at the nest, and water foragers bring in loads that appear to be limited only by crop capacity and that provide all the moisture necessary for the complete processing of 0.74 of a foraged pulp load. As a result P. occidentalis can collect and process a given amount of nest material using 2.6 times fewer foraging trips than would be required by the series system. This in turn means that P. occidentalis not only achieves an energy saving that probably more than offsets the increased costs of material handling at the nest, but it reduces the exposure of its foragers to predators in the field.     

Forager specialization and the control of nest repair in Polybia occidentalis Olivier (Hymenoptera : Vespidae)

We measured patterns of individual forager specialization and colony-wide rates of material input during periods of response to experimental nest damage and during control periods in three colonies of the tropical social wasp Polybia occidentalis.

Effects of an Alien Ant Invasion on Abundance,Behavior, and Reproductive Success of Endemic Island Birds

Abstract: Biological invaders can reconfigure ecological networks in communities, which changes community structure, composition, and ecosystem function. We investigated whether impacts caused by the introduced yellow crazy ant (Anoplolepis gracilipes), a pantropical invader rapidly expanding its range, extend to higher‐order consumers by comparing counts, behaviors, and nesting success of endemic forest birds in ant‐invaded and uninvaded rainforest on Christmas Island (Indian Ocean). Point counts and direct behavioral observations showed that ant invasion altered abundances and behaviors of the bird species we examined: the Island Thrush (Turdus poliocephalus erythropleurus), Emerald Dove (Chalcophaps indica natalis), and Christmas Island White‐eye (Zosterops natalis). The thrush, which frequents the forest floor, altered its foraging and reproductive behaviors in ant‐invaded forest, where nest‐site location changed, and nest success and juvenile counts were lower. Counts of the dove, which forages exclusively on the forest floor, were 9–14 times lower in ant‐invaded forest. In contrast, counts and foraging success of the white‐eye, a generalist feeder in the understory and canopy, were higher in ant‐invaded forest, where mutualism between the ant and honeydew‐secreting scale insects increased the abundance of scale‐insect prey. These complex outcomes involved the interplay of direct interference by ants and altered resource availability and habitat structure caused indirectly by ant invasion. Ecological meltdown, rapidly unleashed by ant invasion, extended to these endemic forest birds and may affect key ecosystem processes, including seed dispersal.     

Behaviorally mediated spatial and temporal refuges from a cleptoparasite,Argochrysis armilla (Hymenoptera: Chrysididae), attacking a ground-nesting wasp,Ammophila dysmica (Hymenoptera: Sphecidae)

Summary It has been suggested that parasite pressure favors the evolution of sociality within the Hymenoptera. I analyzed the impact of a chrysidid nest parasite, Argochrysis armilla, on its host, the solitary ground-nesting wasp, Ammophila dysmica, to assess the role of parasitism in favoring two steps towards sociality: aggregated nesting and delayed nest provisioning. The foraging strategy of Argochrysis armilla involves discovering host nests during excavation, learning the locations of discovered nest, and returning to oviposit in nests during nest provisioning; I therefore assessed the influence of host behavior on (1) parasite discovery of nests and (2) parasite oviposition in nests. Significantly fewer parasites discovered host nests that were excavated during the early morning and late afternoon, due to partial asynchrony of host and parasite activity. Nests excavated in areas of low nest density were also less likely to be discovered; use of low density nest sites increased during periods of high parasite activity. Due to a rapidly decelerating rate of parasite recruitment to nests under excavation, the duration of nest digging had only a limited influence on nest discovery by parasites. The probability of parasite oviposition in a host nest was determined by the number of parasites discovering the nest during excavation and by the time between nest excavation and provisioning; delaying nest provisioning reduced the risk of parasite oviposition. Delayed provisionings primarily appeared to be a result of the stochastic process of hunting and prey encounter. The number of provisions placed in a nest (one vs two) had no effect on the probability of nest parasitism. Spatial patterns of parasitism were directly density dependent in 1984 and density independent in 1986. In this system parasite pressure acts against the formation of nesting aggregations and in favor of delayed nest provisioning. The dependence of these results on species-specific aspects of the parasite's foraging strategy and the host's defensive strategy suggests, however, that different parasite species may generate qualitatively different selection pressures, potentially contributing to the diversity of nesting behavior in the Hymenoptera.     

Love thy neighbour? Social nesting pattern,host mass and nest size affect ectoparasite intensity in Darwin’s tree finches

Social nesting behaviour is commonly associated with high prevalence and intensity of parasites in intraspecific comparisons. Little is known about the effects of interspecific host breeding density for parasite intensity in generalist host–parasite systems. Darwin’s small tree finch (Camarhynchus parvulus) on Santa Cruz Island, Galápagos Islands, nests in both heterospecific aggregations and at solitary sites. All Darwin finch species on Santa Cruz Island are infested with larvae of the invasive blood-sucking fly Philornis downsi. In this study, we test the prediction that total P. downsi intensity (the number of parasites per nest) is higher for nests in heterospecific aggregations than at solitary nests. We also examine variation in P. downsi intensity in relation to three predictor variables: (1) nest size, (2) nest bottom thickness and (3) host adult body mass, both within and across finch species. The results show that (1) total P. downsi intensity was significantly higher for small tree finch nests with many close neighbours; (2) finches with increased adult body mass built larger nests (inter- and intraspecific comparison); (3) parasite intensity increased significantly with nest size across species and in the small tree finch alone; and (4) nest bottom thickness did not vary with nest size or parasite intensity. These results provide evidence for an interaction between social nesting behaviour, nest characteristics and host mass that influences the distribution and potential impact of mobile ectoparasites in birds.     

Aerial defense of the nest by workers of the stingless bee Trigona (Tetragonisca) angustula (Latreille) (Hymenoptera: Apidae)

Summary The stingless bee Trigona (Tetragonisca) angustula has a sophisticated defense strategy against flying insect predators at the entrance of its nest. Groups of worker bees hover on both sides in front of the nest entrance tube, facing a flight corridor leading to the nest. Intruders which enter this corridor are attacked by these bees from the side and from behind and are forced to the ground by biting bees clinging to their wings. T. angustula is subject to predation by Lestrimelitta limao, a cleptobiotic stingless bee which performs organized raids on other nests to rob food supplies, larval provisions and nest constructing material. The presence of citral, released by L. limao during the raids, leads to a rapid increase in the number of hovering guard bees in front of T. angustula nests. This recruitment in response to citral suggests that the defense behavior in T. angustula has evolved under the pressure of L. limao raids and that citral functions in T. angustula as an alarm kairomone.     

Food and the deceptive acquisition of mates by polygynous male harriers

Summary Female northern harriers Circus cyaneus are polygynous, marsh-nesting raptors, whose mate choices are enigmatic. I determined the mate choice cues employed by females by correlating the order in which males were chosen with characters that 1) significantly influenced reproductive success; 2) were assessable prior to settlement; and 3) varied between breeding situations. Only nest sites and male provisioning performance met all these conditions: wet nest sites significantly (P<0.05) increased nest survival and high provisioning rates significantly (P<0.01) enhanced brood survival. The order in which females settled was strongly correlated with provisioning performance in both years (r _s -0.65, and r _s -0.84), but not with nest site quality. Females thus apperared to choose males principally on provisioning performance. Despite using the same cue, however, females choosing mated males reared only 0.28 young for every nestling raised by concurrently settling monogamous females (the lowest ratio recorded for any avian species). Extrapolation of male courtship provisioning patterns from clutch sizes and laying date indicated that females received similar proportions prior to and during egg-laying, but that males later preferentially fed females. Secondary females therefore chose mated males on the basis of a temporally changing and unreliable cue. The polyterritoriality, cooperative harems, skewed sex ratio, and sexy-son hypotheses were all inadequate in explaining polygyny in harriers. Female choice of mated males among harriers is best explained by the deceitful provisioning of food by males.     

Nest site limitation and facultative polygyny in the ant Leptothorax longispinosus

Summary The ant L. longispinosus displays geographic variation in its pattern of facultative polygyny (Fig. 2). In nature, nest density and frequency of multiple queening are positively associated over three sites. A putative causal relation between availability of vacant nest sites and polygyny was examined in New York, where a plot was seeded with additional nest sites and monitored for 24 months. Both queen number and worker number per nest on the experimental plot were reduced relative to controls (Fig. 4, Fig. 5), indicating that scarcity of available nest sites influences the pattern of polygyny in this species. The observed demographic changes resulted from fractionation of existing colonies; adding nest sites induced polydomy. Although numbers of adult ants changed with addition of nest sites, the numbers of immatures were no different after 2 years (Table 1), suggesting that the population was undergoing growth to expand into the additional sites. These results are the first direct experimental evidence linking polygyny to an ecological parameter for any ant species.     

Pheromones and colonization: reassessment of the milkweed bug migration model (Heteroptera: Lygaeidae: Lygaeinae)

Summary. Research on insect migration has justifiably emphasized females – the so-called “oogenesis-flight syndrome”– since it is the females that place the eggs into new habitats. The large and small milkweed bugs, Oncopeltus fasciatus and Lygaeus kalmii, respectively, have featured prominently in studies of insect migration and sequestration of host plant toxins for chemical defense. Here we report that males of these species, and males of another well-studied lygaeine (Neacoryphus bicrucis), produce pheromones in glands usually considered to serve only a defensive role in Heteroptera (the metathoracic scent glands), and that these pheromones are exploited by a tachinid parasitoid as a host-finding kairomone. The pheromones are mixtures of C₆ and C₈ saturated and unsaturated esters reminiscent of lepidopteran pheromones, and the key compound of the O. fasciatus pheromone has now been correctly identified as (E)-2,7-octadienyl acetate. It is proposed that the concept of the oogenesis-flight syndrome for these kinds of insects should accommodate the role of males in the migration process. The hypothesis is presented that male-produced pheromones play a significant role in guiding colonization of new habitats in many heteropteran species. In addition, data are presented suggesting that there is a trade-off between the amount of pheromone produced by colonizing males and the host breadth of the species. Received 21 December 1998; accepted 15 February 1999.     

Brood parasitism disproportionately increases nest provisioning and helper recruitment in a cooperatively breeding bird

Obligate avian brood parasites lay their eggs in nests of other species (hosts), which raise parasitic young. Parasitic nestlings are likely to influence host’s parental behaviours as they typically beg for food more vigorously than young host for a given hunger level. However, few studies have tested this idea, with conflicting results. These prior studies were largely limited to biparental hosts, but little is known about the effect of brood parasitism on parental behaviours in hosts that breed cooperatively. We followed a multimodel approach to examine the effect of brood parasitism on nest provisioning and helper recruitment in the baywing (Agelaioides badius), a cooperative breeder parasitised by screaming (Molothrus rufoaxillaris) and shiny (Molothrus bonariensis) cowbirds. Multimodel inference results indicated that feeding visits increased with nestling age, cooperative group size and number of cowbird nestlings in the brood. Brood size had little influence on feeding visits, which further suggests that baywings adjusted their provisioning effort in response to cowbird parasitism. In addition, nests parasitised artificially with shiny cowbird eggs or hatchlings recruited more helpers than unmanipulated nests having only host or screaming cowbird young. Our results provide novel evidence that brood parasitism and cooperative breeding interact in determining the levels of nest provisioning.     

Red-winged blackbird parental investment following brood parasitism by brown-headed cowbirds: is parentage important?

Parental investment by red-winged blackbirds (Agelaius phoeniceus) in response to natural and experi‐mental parasitism by brown-headed cowbirds (Molothrus ater), and in response to freeze-dried, female cowbird mounts presented near redwing nests during the egg-laying period was measured. Two measures of redwing parental investment were used: nest defense effort toward a model predator, and rate of feeding nestlings. There were no significant differences in levels of parental investment among unparasitized nests, naturally parasitized nests, or experimentally parasitized nests. Similarly, parental investment did not differ between redwings that were exposed to the cowbird mount and those that were not exposed to the mount, or among redwings exposed to the cowbird mount at different distances from the nest. This suggests that red-winged blackbirds do not recognize when they have been parasitized, and hence do not associate parasitism with a decrease in their parentage, or that parentage is not an important predictor of parental investment in this species. Received: 24 January 1997 / Accepted after revision: 7 June 1997     

American robin nestlings compete by jockeying for position

Summary We investigated whether nestling American robins (Turdus migratorius) were capable of influencing food distribution in their nests by perceiving that certain sectors of the nest received a relatively high proportion of feedings and positioning themselves accordingly. Feeding observations were obtained from videotape recordings taken at different stages of the nestling period. Parents generally arrived at a predictable location on the nest rim and allocated proportionally more food to nestlings in the central position. The degree of nestling movement was significantly positively correlated with variation in the predictability of parental arrival locations on the nest rim. Furthermore, nestlings moved more in broods suffering brood reduction. This suggests that when competition for food is intense and the location of parental arrival is predictable, nestlings respond by jockeying for access to the most favorable (i.e., central) position in the nest. We conclude that jockeying for position by nestlings can influence the pattern of food allocation by parents, and that hungry nestlings can improve their competitive standing against nestmates by moving to positions where parents are more likely to feed them. Correspondence to: S.B. McRae     

Influence of subtle substrate differences on feeding by shorebirds on intertidal mudflats

Shallow-feeding shorebirds, dowitchers (Limnodromus griseus and L. scolopaceus), western sandpipers (Calidris mauri), dunlin (C. alpina) and American avocets (Recurvirostra americana), reduced the density of their prey in mudflats with little sand but not in mudflats with a moderate admixture of sand. An experiment in Upper Newport Bay, Southern California, during October and November 1979 to explain the difference in density is described. The effect of sand on shorebird feeding was compared in the field by measuring the times spent feeding in plots where sand had or had not been added, respectively. Increasing the sand content to 14% from 2% in the top centimeter decreased the time spent in treated plots by all species compared to adjacent untreated plots. The prey species were small polychaete and oligochaete worms (0.25 to 1.25 mm wide) similar in diameter to sand grains (0.5 to 1.0 mm). In plots where sand had been added, avocets fed by pecking at the surface in addition to scything, the more common method of feeding on muddy substrates. The results suggest that sand interferes with the detection and or capture of prey that are similar in diameter to small sand grains and explains the differences in the effects of predation by these birds seen on mudflats with a moderate admixture of sand compared to the effects on mudflats with little sand. Differential success in prey capture between one microhabitat and the nest (rather than a reduction in competition, as suggested by some authors) might explain the different use of such habitats.