Consumer-specific responses to riverine subsidy pulses in a riparian arthropod assemblage

1. Aquatic resource fluxes from streams can provide significant subsidies for riparian consumers. Because aquatic resource fluxes can be highly variable in space and time, the subsidy efficiency (i.e. transfer to the recipient food web) is controlled by the short‐term aggregative response of riparian consumers. 2. Field manipulations of stream‐derived invertebrate prey subsidies were used to examine specific aggregative responses of ground‐dwelling arthropods to riverine subsidy pulses in a braided‐river (Tagliamento River, NE Italy). Subsidy manipulation comprised short‐term reductions of natural stream‐derived subsidies and increased subsidies of stream‐derived invertebrate prey during four seasons. 3. We hypothesised that specific aggregative responses of riparian arthropods depend on their specialisation on aquatic insects which was inferred from stable isotope analysis. Natural riverine subsidy sources including aquatic insect emergence and surface‐drifting organisms were quantified. 4. Arthropods responded significantly with a reduction in abundance by 51%, at reduced subsidies and an increase by 110% at increased subsidies, when averaged over all seasons. Different arthropod taxa responded differently to subsidy manipulations in relation to their specialisation on aquatic subsidies: ground beetles with a diet consisting predominantly of aquatic insects responded only to subsidy reductions, indicating that their local abundance was not limited by natural stream‐derived subsidies; lycosid spiders with a partly aquatic diet showed no significant response; and ants, although relying on a terrestrial diet, responded positively to added stream‐derived invertebrate prey, indicating that stranding of surface‐drifting terrestrial invertebrates represented an important subsidy pathway. 5. Ground beetles and lycosid spiders were seasonally separated in their use of aquatic subsidies. Results indicate that the life‐history characteristics of riparian consumers can control the subsidy efficiency for the recipient community. By the effective uptake of pulsed riverine‐derived subsidies, riparian arthropods can enhance the transfer of riverine food sources to the riparian food web.     

Multi-Scale Effects of Nestling Diet on Breeding Performance in a Terrestrial Top Predator Inferred from Stable Isotope Analysis

Inter-individual diet variation within populations is likely to have important ecological and evolutionary implications. The diet-fitness relationships at the individual level and the emerging population processes are, however, poorly understood for most avian predators inhabiting complex terrestrial ecosystems. In this study, we use an isotopic approach to assess the trophic ecology of nestlings in a long-lived raptor, the Bonelli’s eagle Aquila fasciata, and investigate whether nestling dietary breath and main prey consumption can affect the species’ reproductive performance at two spatial scales: territories within populations and populations over a large geographic area. At the territory level, those breeding pairs whose nestlings consumed similar diets to the overall population (i.e. moderate consumption of preferred prey, but complemented by alternative prey categories) or those disproportionally consuming preferred prey were more likely to fledge two chicks. An increase in the diet diversity, however, related negatively with productivity. The age and replacements of breeding pair members had also an influence on productivity, with more fledglings associated to adult pairs with few replacements, as expected in long-lived species. At the population level, mean productivity was higher in those population-years with lower dietary breadth and higher diet similarity among territories, which was related to an overall higher consumption of preferred prey. Thus, we revealed a correspondence in diet-fitness relationships at two spatial scales: territories and populations. We suggest that stable isotope analyses may be a powerful tool to monitor the diet of terrestrial avian predators on large spatio-temporal scales, which could serve to detect potential changes in the availability of those prey on which predators depend for breeding. We encourage ecologists and evolutionary and conservation biologists concerned with the multi-scale fitness consequences of inter-individual variation in resource use to employ similar stable isotope-based approaches, which can be successfully applied to complex ecosystems such as the Mediterranean.     

More than a barrier: The complex effects of ecotone vegetation type on terrestrial consumer consumption of an aquatic prey resource

Community structure and dynamics can be influenced by resource transfers between ecosystems, yet little is known about how boundary structure determines both the magnitude of exchanges and their effects on recipient and donor communities. Aquatic and terrestrial ecosystems are often linked by resource fluxes and riparian vegetation is commonly affected by anthropogenic alterations to land use or river hydrological regime. I investigated whether shrubs at the freshwater–terrestrial interface alter the supply, distribution and importance of aquatic prey resources to terrestrial consumers. Shrubs were predicted to alter the larval community composition of aquatic insects and the emergence of winged adults, thus affecting aquatic prey subsidies to terrestrial consumers. In addition, shrubs were hypothesized to alter the microclimatic suitability of the riparian zone for adult aquatic insects, act as a physical barrier to their dispersal and affect terrestrial community composition, particularly the abundance and type of predators that could benefit from the aquatic prey resource. Stable isotope dietary analyses and a survey of shrub‐dominated and open grassland riparian habitats revealed that larval densities of aquatic insects (EPTM: Ephemeroptera, Plecoptera, Trichoptera and Megaloptera) were higher in shrub than grassland habitats; however, reduced emergence and lateral dispersal in shrub areas led to lower densities of adults. The temperature and relative humidity of the riparian zone did not differ between the habitats. Ground‐active terrestrial invertebrate communities had a higher proportion of cursorial spiders in grassland, coinciding with greater abundances of aquatic prey. Aquatic prey contribution to cursorial spider diet matched adult aquatic insect abundances. Overall, riparian shrubs reduced the magnitude, or at least altered the timing, of cross‐ecosystem subsidy supply, distribution and use by consumers through mechanisms operating in both the aquatic and terrestrial ecosystems. Thus, the structure of ecosystem boundaries has complex effects on the strength of biological interactions between adjacent systems.     

Do seasonally fluctuating aquatic subsidies influence the distribution pattern of birds between riparian and upland forests?

Seasonal fluctuation of allochthonous subsidies influences food web structure and dynamics in recipient communities. This study investigated whether aquatic subsidies influence the dynamics of insectivorous birds in entire catchment. We estimated the prey biomass and bird density in riparian and upland habitats in three catchments in temperate deciduous forests in Hokkaido, Japan. Aquatic prey was found only in riparian forests and the biomass peaked in early spring, while terrestrial prey was equally distributed between habitats and increased in biomass in late spring. Bird density was higher in riparian than in upland forests before bud break, when the biomass of aquatic insects peaked, but was similar in both forests during the rest of the seasons. These results suggest that aquatic prey subsidies are used not only by birds inhabiting riparian forests, but also by birds associated with upland forests. Aquatic prey subsidies may be particularly important in the spring as a critical food resource for survival and the breeding activities of birds, thereby, influencing the population dynamics of bird communities.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

Riparian contributions to the diet of terapontid grunters (Pisces: Terapontidae) in wet–dry tropical rivers

The diets of terapontid assemblages in 22 catchments across Australia's wet–dry tropics were investigated in relation to the direct use of terrestrial‐riparian inputs, as well as the role of ontogeny and morphology in mediating consumption of allocthonous material. The diet of several species was restricted almost entirely to instream trophic resources throughout their life history. In contrast, ontogenetic diet shifts towards increasing consumption of terrestrial prey types were a prominent feature of the dietary ecology of some terapontids, with collective allocthonous dietary items making a significant contribution (up to 42%) to diet in larger size classes of several species. For those species consuming terrestrial‐riparian material in their diet, terrestrial invertebrates were the most common prey item; however, terrestrial vegetation, principally riparian fruits, and terrestrial vertebrates were also important dietary inclusions in the larger size classes of particular species. A large mouth gape was the morphological feature most strongly associated with consumption of terrestrial food resources within the Terapontidae. Results indicate that the direct consumption of terrestrially derived food sources in northern Australian aquatic systems may be more important than previously asserted, and that additional research is required to better clarify the role of terrestrial subsidies to these ecosystems.     

Cross‐ecosystem disturbance influences on the life history and population size structure of a riparian predator

Cross‐ecosystem transfers of resources could alter the life history traits of consumers in adjacent systems by changing the nature and availability of prey. However, large‐scale influences, such as natural disturbances, that control the magnitude of prey subsidies are likely to modify these effects. To investigate impacts of cross‐ecosystem subsidies on the life history traits of a riparian predator we measured the size, sex and condition of riparian fishing spiders (Dolomedes aquaticus) across a gradient of flooding frequency and intensity. These spiders rely on adult aquatic insects for a large proportion of their diet and previous research demonstrated that increased flooding decreased the abundance of aquatic insect prey. In this study, laboratory experiments indicated that increased prey availability hastened the first moult of the spiders after winter and decreased the propensity for cannibalistic interactions of individuals of the same size. However, despite the likely positive influences of increased food supply, in the field the highest abundance and proportion of large, potentially reproductive females occurred at the most flood‐prone rivers, where aquatic prey availability was the lowest. It is likely that other factors modified by the disturbance regime, such as habitat availability, flood‐related mortality and intra‐specific interaction rates, altered the influence of cross‐ecosystem subsidies on the life history traits of these spiders. Thus, our results indicate that disturbance‐related effects can flow across ecosystem boundaries and alter the life history traits of predators relying on allochthonous resources.     

Dietary changes in predators and scavengers in a nocturnally illuminated riparian ecosystem

Aquatic and terrestrial ecosystems are linked by fluxes of carbon and nutrients in riparian areas. Processes that alter these fluxes may therefore change the diet and composition of consumer communities. We used stable carbon isotope (δ¹³C) analyses to test whether the increased abundance of aquatic prey observed in a previous study led to a dietary shift in riparian consumers in areas illuminated by artificial light at night (ALAN). We measured the contribution of aquatic‐derived carbon to diets in riparian arthropods in experimentally lit and unlit sites along an agricultural drainage ditch in northern Germany. The δ¹³C signature of the spider Pachygnatha clercki (Tetragnathidae) was 0.7‰ lower in the ALAN‐illuminated site in summer, indicating a greater assimilation of aquatic prey. Bayesian mixing models also supported higher intake of aquatic prey under ALAN in spring (34% versus 21%). In contrast, isotopic signatures for P. clercki (0.3‰) and Pardosa prativaga (0.7‰) indicated a preference for terrestrial prey in the illuminated site in spring. Terrestrial prey intake increased in spring for P. clercki under ALAN (from 70% to 74%) and in spring and autumn for P. prativaga (from 68% to 77% and from 67% to 72%) and Opiliones (from 68% to 72%; 68% to 75%). This was despite most of the available prey (up to 80%) being aquatic in origin. We conclude that ALAN changed the diet of riparian secondary consumers by increasing the density of both aquatic and terrestrial prey. Dietary changes were species‐ and season‐specific, indicating that the effects of ALAN may interact with phenology and feeding strategy. Because streetlights can occur in high density near freshwaters, ALAN may have widespread effects on aquatic–terrestrial ecosystem linkages.     

Growth and development rates in a riparian spider are altered by asynchrony between the timing and amount of a resource subsidy

Rapid growth in response to increased prey abundance may be induced by environmental variability associated with resource subsidies. Spiders living in riparian areas are subject to frequent, episodic bursts of aquatic prey (subsidies). These periods of high resource abundance may occur at different points in recipient consumers’ development through variation in emergence patterns of prey between years or across a landscape. We examine how variable timing of subsidy abundance intersects with life history scheduling to produce different growth and development outcomes for individuals within a population. Through a series of controlled feeding experiments, we tested the hypotheses that the spider Tetragnatha versicolor: (1) exhibits compensatory growth in response to subsidy variability, (2) that rapid increases in mass may result in a greater risk of mortality, and (3) that the timing of subsidy resources relative to the development schedule of this spider may produce different outcomes for individual growth patterns and adult condition. Spiders fed at very high rates grew fastest but also showed evidence of increased mortality risk during moulting. T. versicolor is capable of exhibiting strong growth compensation—individuals suffering initial growth restriction were able to catch up completely with animals on a constant diet utilising the same amount of food. Spiders that received an early pulse of resources (simulating an early arrival of an aquatic insect subsidy to riparian forests) did worse on all measures of development and fitness than spiders that received either a constant supply of food or a late pulse of resources. Importantly, receiving large amounts of food early in life appears to actually confer relative disadvantages in terms of later performance compared with receiving subsidies later in development. Subsidies may provide greater benefits to individuals or age cohorts encountering this resource abundance closer to the onset of reproductive efforts than subsidies arriving early in development.     

Reduced breeding success of Pied Flycatchers Ficedula hypoleuca along regulated rivers

Most large rivers in northern Sweden are regulated to produce hydropower, with subsequent effects on flow dynamics and aquatic insect communities. Several studies have shown that aquatic and terrestrial systems are intimately connected via the export of emergent aquatic insects, but few have assessed how human modifications of aquatic habitats may influence this connection. We compared breeding success of the insectivorous Pied Flycatcher Ficedula hypoleuca in near‐riparian upland forests along two regulated and two free‐flowing large rivers in northern Sweden over 3 years. The regulated rivers showed lower aquatic insect export to the surroundings, as a consequence of regulation‐induced loss of suitable aquatic insect habitats. Survival of Pied Flycatcher nestlings was 10–15% greater along the free‐flowing rivers. Females breeding near the free‐flowing rivers also started egg‐laying earlier and with greater synchrony than those at the regulated rivers, and showed a smaller decrease in weight during breeding than did females along the regulated rivers. However, there were no differences in occupation rate, clutch size or number of successfully hatched juveniles between regulated and free‐flowing rivers. As regulated rivers showed lower abundance of flying aquatic insects, which may also reduce the abundance of terrestrial invertebrate prey, regulation‐induced changes in the export of emergent aquatic insects may explain both directly and indirectly the observed reduction in Pied Flycatcher breeding success along regulated rivers. Large‐scale river regulation may therefore impair the breeding success of insectivorous birds through impacts on prey availability.     

Trophic overlap between fish and riparian spiders: potential impacts of an invasive fish on terrestrial consumers

Studies on resource sharing and partitioning generally consider species that occur in the same habitat. However, subsidies between linked habitats, such as streams and riparian zones, create potential for competition between populations which never directly interact. Evidence suggests that the abundance of riparian consumers declines after fish invasion and a subsequent increase in resource sharing of emerging insects. However, diet overlap has not been investigated. Here, we examine the trophic niche of native fish, invasive fish, and native spiders in South Africa using stable isotope analysis. We compared spider abundance and diet at upstream fishless and downstream fish sites and quantified niche overlap with invasive and native fish. Spider abundance was consistently higher at upstream fishless sites compared with paired downstream fish sites, suggesting that the fish reduced aquatic resource availability to riparian consumers. Spiders incorporated more aquatic than terrestrial insects in their diet, with aquatic insects accounting for 45–90% of spider mass. In three of four invaded trout rivers, we found that the average proportion of aquatic resources in web‐building spider diet was higher at fishless sites compared to fish sites. The probability of web‐building and ground spiders overlapping into the trophic niche of invasive brown and rainbow trout was as high as 26 and 51%, respectively. In contrast, the probability of spiders overlapping into the trophic niche of native fish was always less than 5%. Our results suggest that spiders share resources with invasive fish. In contrast, spiders had a low probability of trophic overlap with native fish indicating that the traits of invaders may be important in determining their influence on ecosystem subsidies. We have added to the growing body of evidence that invaders can have cross‐ecosystem impacts and demonstrated that this can be due to niche overlap.     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

Differential reliance on aquatic prey subsidies influences mercury exposure in riparian arachnids and songbirds

Cross‐ecosystem subsidies move substantial amounts of nutrients between ecosystems. Emergent aquatic insects are a particularly important prey source for riparian songbirds but may also move aquatic contaminants, such as mercury (Hg), to riparian food webs. While many studies focus on species that eat primarily emergent aquatic insects, we instead study riparian songbirds with flexible foraging strategies, exploiting both aquatic and terrestrial prey sources. The goal in this study is to trace reliance on aquatic prey sources and correlate it to Hg concentrations in common riparian arachnids (Families Tetragnathidae, Opiliones, and Salticidae) and songbirds (Common Yellowthroat Geothlypis trichas, Spotted Towhee Pipilo maculatus, Swainson''s Thrush Catharus ustulatus, Song Sparrow Melospiza melodia, and Yellow Warbler Setophaga petechia). We used stable isotopes of δ¹³C and δ¹⁵N and Bayesian mixing models in MixSIAR to determine the reliance of riparian predators on aquatic prey sources. Using mixed effects models, we found that arachnid families varied in their reliance on aquatic prey sources. While songbird species varied in their reliance on aquatic prey sources, songbirds sampled earlier in the season consistently relied more on aquatic prey sources than those sampled later in the season. For both arachnids and songbirds, we found a positive correlation between the amount of the aquatic prey source in their diet and their Hg concentrations. While the seasonal pulse of aquatic prey to terrestrial ecosystems is an important source of nutrients to riparian species, our results show that aquatic prey sources are linked with higher Hg exposure. For songbirds, reliance on aquatic prey sources early in the breeding season (and subsequent higher Hg exposure) coincides with timing of egg laying and development, both of which may be impacted by Hg exposure.     

Species-specific use of allochthonous resources by ground beetles (Carabidae) at a river–land interface

A variety of organisms mediate river–terrestrial linkages through spatial subsidies. However, most empirical studies have classified organisms rather broadly (e.g., by functional group or taxonomic family) and have dismissed species-level linkages at the interface of ecosystems. Here, we show how allochthonous resource use varies among taxonomically similar species of ground beetles (family Carabidae) across seasons (June–September). We investigated seasonal shifts in the distribution of five beetle species and their dietary responses to spatial subsidies (emerging aquatic insects) in a Japanese braided river. Despite their taxonomic closeness, the ground beetles showed species-specific responses to spatial subsidies, and beetle distribution patterns tended to coincide with their diets. Overall, 1–56% of ground beetle diets were derived from aquatic prey. One genus (Bembidion spp.) mainly consumed aquatic prey, while three species fed primarily on terrestrial prey across all seasons. However, one species (Lithochlaenius noguchii) showed shifts in its diet from aquatic to terrestrial prey according to subsidy availability. The observed variation in allochthonous resource use was likely related to species-specific foraging modes, physiological tolerance to dry conditions, and interspecific competition. Our findings suggest that considering species-specific interactions is necessary to fully understand cross-system interactions and recipient food-web dynamics.     

Temporal changes in prey composition and biomass delivery to African Crowned Eagle nestlings in urban areas of KwaZulu-Natal,South Africa

Globally urban areas are expanding rapidly and this usually has negative effects on biodiversity. Despite this, some species manage to persist in urban areas, as is the case with African Crowned Eagles Stephanoaetus coronatus in KwaZulu-Natal, South Africa. As relatively little is known about African Crowned Eagle nestling diet, especially about how it changes with nestling age, we investigated this with nest camera-traps. We analysed temporal changes in prey composition and biomass delivery during the nestling stage. We also recorded which adults provisioned and attended the nest. The main prey fed to nestlings were Rock Hyrax Procavia capensis and Hadeda Ibis Bostrychia hagedash. Adult males did most of the food provisioning, especially at the start of the nestling period. We found a decrease in total prey number and biomass with nestling age. This may be caused by changing requirements of nestlings. Furthermore, delivering fewer prey at later nestling stages may be a facilitating mechanism to enhance fledging of the nestling. Although the total number of prey brought to the nest decreased, we found an increase in numbers of Vervet Monkey Chlorocebus pygerythrus in the diet with nestling age. This indicated an increase in larger prey being delivered to the nests as the nestling aged. We suggest that this could be caused by increased participation in hunting by the larger female as her nest attendance time decreased as the nestling aged. We conclude with emphasising the importance of protecting the Durban Metropolitan Open Space System (D’MOSS) zones for the persistence of this Near Threatened raptor species, and populations of its prey in urban areas for its breeding success.     

Responses to River Inundation Pressures Control Prey Selection of Riparian Beetles

Background

Riparian habitats are subjected to frequent inundation (flooding) and are characterised by food webs that exhibit variability in aquatic/terrestrial subsidies across the ecotone. The strength of this subsidy in active riparian floodplains is thought to underpin local biodiversity. Terrestrial invertebrates dominate the fauna, exhibiting traits that allow exploitation of variable aquatic subsidies while reducing inundation pressures, leading to inter-species micro-spatial positioning. The effect these strategies have on prey selection is not known. This study hypothesised that plasticity in prey choice from either aquatic or terrestrial sources is an important trait linked to inundation tolerance and avoidance.

Method/Principal Findings

We used hydrological, isotopic and habitat analyses to investigate the diet of riparian Coleoptera in relation to inundation risk and relative spatial positioning in the floodplain. The study examined patch scale and longitudinal changes in utilisation of the aquatic subsidy according to species traits. Prey sourced from terrestrial or emerging/stranded aquatic invertebrates varied in relation to traits for inundation avoidance or tolerance strategies. Traits that favoured rapid dispersal corresponded with highest proportions of aquatic prey, with behavioural traits further predicting uptake. Less able dispersers showed minimal use of aquatic subsidy and switched to a terrestrial diet under moderate inundation pressures. All trait groups showed a seasonal shift in diet towards terrestrial prey in the early spring. Prey selection became exaggerated towards aquatic prey in downstream samples.

Conclusions/Significance

Our results suggest that partitioning of resources and habitat creates overlapping niches that increase the processing of external subsidies in riparian habitats. By demonstrating functional complexity, this work advances understanding of floodplain ecosystem processes and highlights the importance of hydrological variability. With an increasing interest in reconnecting rivers to their floodplains, these invertebrates represent a key functional element in ensuring that such reconnections have demonstrable ecological value.     

Age-Related Changes in Nestling Diet of the Cooperatively Breeding Green Woodhoopoe

In many socially monogamous bird species, parents of altricial young respond to the increasing demands of growing nestlings by increasing their feeding rate and the size of prey items delivered and by altering the types of prey provided. In some cooperatively breeding species, similar changes in feeding rate and prey size have been documented. However, potential changes in the types of prey delivered, both as nestlings age and by different group members, remain largely unexplored. Moreover, studies rarely compare the diet fed to nestlings with that eaten by the provisioning adults themselves. Here, I show that green woodhoopoe ( Phoeniculus purpureus ) nestlings receive a smaller proportion of spiders and larger proportions of caterpillars and centipedes as they grow older. Both male and female adults delivered a higher proportion of spiders to young nestlings than they ate while self-feeding, probably in response to particular nutritional requirements of the chicks. However, only males altered the proportions of caterpillars and centipedes delivered, providing smaller proportions to young nestlings than eaten themselves. These prey items may be too large for young nestlings to handle, and males may make a greater adjustment in provisioning diet than females because they collect more caterpillars and centipedes than do females. Although there were sex differences in provisioning diet, there were no differences between same-sex breeders and helpers in terms of the overall proportions of prey delivered or the changes with nestling age. Hence, individuals of different reproductive status may be following the same provisioning rules, at least in terms of prey type.     

The significance of seed food in chick development re‐evaluated by tracking day‐to‐day dietary variation in the nestlings of a granivorous passerine

The dietary adjustment of nestlings of granivorous birds to a seed diet and the different morphological characteristics of ingested food have rarely been examined in natural conditions. It has been suggested that the provision of cereal grains to nestlings of some seed‐eating bird species in modern agroecosytems is the result of poor food conditions after agricultural intensification. We analysed the abundance of invertebrate prey in the main foraging habitat of parent birds, daily changes (from hatching to fledging) in the efficiency of cereal seed digestion, and the dietary characteristics, diet composition and prey type delivered to nestlings of the Yellowhammer Emberiza citrinella. Analysis of faecal sacs from nests located in breeding habitat with an abundant invertebrate fauna revealed no relationship between the proportion of cereal seeds in the diet of nestlings and the food supply in the main foraging sites of the parents. Neonate nestlings (1 day old) exclusively received weakly chitinized invertebrate prey (arachnids and flies), but from the second day of life the nestlings were fed a variety of highly chitinized invertebrate prey, the percentage biomass of which did not change for the remainder of the nesting period. Cereal grains started to be delivered to 3‐day‐old nestlings and were already efficiently digested, and the percentage biomass of this food type increased progressively with nestling age. We suggest that the provisioning of cereal grains to nestlings is not forced by external factors, such as modern agricultural intensification; rather, it is an intentional behaviour of parent birds aimed at achieving physiological adjustment to seed food in the early stages of ontogeny.     

Interpopulation resource partitioning of Lesser Frigatebirds and the influence of environmental context

Conspecific individuals inhabiting nearby breeding colonies are expected to compete strongly for food resources owing to the constraints imposed by shared morphology, physiology, and behavior on foraging strategy. Consequently, colony‐specific foraging patterns that effectively partition the available resources may be displayed. This study aimed to determine whether intraspecific resource partitioning occurs in two nearby colonies of Lesser Frigatebirds (Fregata ariel). A combination of stable isotope analysis and GPS tracking was used to assess dietary and spatial partitioning of foraging resources during the 2013 and 2014 breeding seasons. These results were compared to vessel‐derived estimates of prey availability, local primary productivity, and estimates of reproductive output to suggest potential drivers and implications of any observed partitioning. Isotopic data indicated a more neritic source of provisioned resources for near‐fledged chicks at an inshore colony, whereas their offshore counterparts were provisioned with resources with a more pelagic signal. Deep pelagic waters (>200 m) had higher availability of a preferred prey type despite a trend for lower primary productivity. Differences in foraging ecology between the two populations may have contributed to markedly different reproductive outputs. These findings suggest environmental context influences dietary and spatial aspects of foraging ecology. Furthermore, the effect of colony‐specific foraging patterns on population demography warrants further research.     

Importance of niche quality for Yellowhammer Emberiza citrinella nestling survival, development and body condition in its native and exotic ranges: the role of diet

The niche hypothesis predicts that some introduced species establish and spread successfully because their new environment provides expanded niche opportunities compared with their native environments. By investigating nestling survival, growth and body condition in relation to diet composition and prey abundance, we tested the prediction that the success of Yellowhammers Emberiza citrinella in New Zealand could be explained by the availability of better quality food resources in its introduced range compared with its native range. We found that Yellowhammer nestlings in New Zealand were larger and heavier than those in Britain, but that nestling survival and body condition did not differ between countries. The preferred prey items, which had some influence on nestling development and survival, were more prevalent in the diet of nestlings in New Zealand, suggesting that nestlings in the introduced range had a superior quality diet. However, there was little evidence to support the prediction that the preferred prey were more abundant in the introduced range, implying that some other factor such as prey accessibility or adult fitness may account for the differences in diet between countries.