Avian Diversity and Functional Insectivory on North-Central Florida Farmlands

Abstract:  We studied the potential for native birds to control insect pests on farms. We assessed habitat factors correlated with diversity, distribution, and insect-foraging activity of native birds on farms in north-central Florida and then characterized common bird species that consumed insect biomass in crops as "functional insectivores" (birds most likely to contribute to pest control). Analyses of point-count survey data and foraging observations collected over 2 years on paired organic and conventional farm sites indicated that (1) farms supported most (82–96%) land birds known to breed in the region; (2) bird species richness and abundance varied significantly with matrix habitat and field border type (but not with year or farm management type); (3) the highest bird abundances were associated with mixed crop plantings, field borders, and adjacent matrix composed of forest and hedge; and (4) abundances of 10 species identified as functional insectivores were primarily influenced by crop type (mixed crops attracted significantly more insect foragers into fields than monocrops). We documented birds eating pest insects in crops and did not observe substantive crop damage by birds during growing-season observations. We advocate use of the term functional insectivore to emphasize the potential positive role of avian insectivory on farms during the growing season.     

Birds as predators in tropical agroforestry systems

Insectivorous birds reduce arthropod abundances and their damage to plants in some, but not all, studies where predation by birds has been assessed. The variation in bird effects may be due to characteristics such as plant productivity or quality, habitat complexity, and/or species diversity of predator and prey assemblages. Since agroforestry systems vary in such characteristics, these systems provide a good starting point for understanding when and where we can expect predation by birds to be important. We analyze data from bird exclosure studies in forests and agroforestry systems to ask whether birds consistently reduce their arthropod prey base and whether bird predation differs between forests and agroforestry systems. Further, we focus on agroforestry systems to ask whether the magnitude of bird predation (1) differs between canopy trees and understory plants, (2) differs when migratory birds are present or absent, and (3) correlates with bird abundance and diversity. We found that, across all studies, birds reduce all arthropods, herbivores, carnivores, and plant damage. We observed no difference in the magnitude of bird effects between agroforestry systems and forests despite simplified habitat structure and plant diversity in agroforests. Within agroforestry systems, bird reduction of arthropods was greater in the canopy than the crop layer. Top-down effects of bird predation were especially strong during censuses when migratory birds were present in agroforestry systems. Importantly, the diversity of the predator assemblage correlated with the magnitude of predator effects; where the diversity of birds, especially migratory birds, was greater, birds reduced arthropod densities to a greater extent. We outline potential mechanisms for relationships between bird predator, insect prey, and habitat characteristics, and we suggest future studies using tropical agroforests as a model system to further test these areas of ecological theory.     

Hierarchical patch choice by an insectivorous bat through prey availability components

Food availability does not only refer to the abundance of edible items; accessibility and detectability of food are also essential components of the availability concept. Constraints imposed by a habitat’s physical structure on the accessibility and detectability of food have been seldom treated simultaneously to the abundance of prey at the foraging patch level in observational studies. We designed a research that allowed decoupling the effects of microhabitat structure and prey abundance on foraging patch selection of the trawling insectivorous long-fingered bat (Myotis capaccinii). The use of different patches of river was surveyed by radiotelemetry during three periods of the bat’s annual cycle, and prey abundance was accordingly measured in and out of the hunting grounds of the tracked bats by insect traps emulating the species’ foraging. Bats preferentially used river stretches characterised by an open course and smooth water surfaces, i.e. they used the most suitable patches in terms of prey accessibility and detectability, respectively. In addition, prey abundance in the selected river stretches was higher than in others where bat activity was not recorded, although the latter also offered good access and prey detection possibilities. Bats also shifted foraging stretches seasonally, likely following the spatiotemporal dynamics of prey production over the watershed. We suggest that the decisions of bats during the patch choice process fitted a hierarchical sequence driven first by the species’ morphological specialisations and ability to hunt in unobstructed spaces, then by the detectability of prey on water surfaces and, finally, by the relative abundance of prey.     

Long-term declines of European insectivorous bird populations and potential causes

Evidence of declines in insect populations has recently received considerable scientific and societal attention. However, the lack of long-term insect monitoring makes it difficult to assess whether declines are geographically widespread. By contrast, bird populations are well monitored and often used as indicators of environmental change. We compared the population trends of European insectivorous birds with those of other birds to assess whether patterns in bird population trends were consistent with declines of insects. We further examined whether declines were evident for insectivores with different habitats, foraging strata, and other ecological preferences. Bird population trends were estimated for Europe (1990–2015) and Denmark (1990–2016). On average, insectivores declined over the study period (13% across Europe and 28% in Denmark), whereas omnivores had stable populations. Seedeaters also declined (28% across Europe; 34% in Denmark), but this assessment was based on fewer species than for other groups. The effects of insectivory were stronger for farmland species (especially grassland species), for ground feeders, and for cold-adapted species. Insectivory was associated with long-distance migration, which was also linked to population declines. However, many insectivores had stable populations, especially habitat generalists. Our findings suggest that the decline of insectivores is primarily associated with agricultural intensification and loss of grassland habitat. The loss of both seed and insect specialists indicates an overall trend toward bird communities dominated by diet generalists.     

Ecological and Economic Services Provided by Birds on Jamaican Blue Mountain Coffee Farms

Abstract: Coffee farms can support significant biodiversity, yet intensification of farming practices is degrading agricultural habitats and compromising ecosystem services such as biological pest control. The coffee berry borer (Hypothenemus hampei) is the world's primary coffee pest. Researchers have demonstrated that birds reduce insect abundance on coffee farms but have not documented avian control of the berry borer or quantified avian benefits to crop yield or farm income. We conducted a bird‐exclosure experiment on coffee farms in the Blue Mountains, Jamaica, to measure avian pest control of berry borers, identify potential predator species, associate predator abundance and borer reductions with vegetation complexity, and quantify resulting increases in coffee yield. Coffee plants excluded from foraging birds had significantly higher borer infestation, more borer broods, and greater berry damage than control plants. We identified 17 potential predator species (73% were wintering Neotropical migrants), and 3 primary species composed 67% of migrant detections. Average relative bird abundance and diversity and relative resident predator abundance increased with greater shade‐tree cover. Although migrant predators overall did not respond to vegetation complexity variables, the 3 primary species increased with proximity to noncoffee habitat patches. Lower infestation on control plants was correlated with higher total bird abundance, but not with predator abundance or vegetation complexity. Infestation of fruit was 1–14% lower on control plants, resulting in a greater quantity of saleable fruits that had a market value of US$44–$105/ha in 2005/2006. Landscape heterogeneity in this region may allow mobile predators to provide pest control broadly, despite localized farming intensities. These results provide the first evidence that birds control coffee berry borers and thus increase coffee yield and farm income, a potentially important conservation incentive for producers.     

Birds and bats reduce insect biomass and leaf damage in tropical forest restoration sites

Both birds and bats are important insect predators in tropical systems. However, the relative influence of birds and bats on insect populations and their indirect effects on leaf damage have not previously been investigated in tropical forest restoration sites. Leaf damage by herbivorous insects can negatively affect the growth and survival of tropical plants and thus can influence the success of tropical forest restoration efforts. We used an exclosure experiment to examine the top-down effects of birds and bats on insects and leaf damage in a large-scale forest restoration experiment. Given the potential influence of tree planting design on bird and bat abundances, we also investigated planting design effects on bird and bat insectivory and leaf damage. The experiment included two planting treatment plots: islands, where trees were planted in patches, and plantations, where trees were planted in rows to create continuous cover. In both types of plots, insect biomass was highest on tree branches where both birds and bats were excluded from foraging and lowest on branches without exclosures where both birds and bats were present. In the island plots, birds and bats had approximately equal impacts on insect populations, while in plantations bats appeared to have a slightly stronger effect on insects than did birds. In plantations, the levels of leaf damage were higher on branches where birds and bats were excluded than on branches where both had access. In island plots, no significant differences in leaf damage were found between exclosure treatments although potential patterns were in the same direction as in the plantations. Our results suggest that both birds and bats play important roles as top predators in restoration systems by reducing herbivorous insects and their damage to planted trees. Tropical restoration projects should include efforts to attract and provide suitable habitat for birds and bats, given their demonstrated ecological importance.     

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.     

Persistent bimodal activity patterns in wild and captive black-tailed godwit <Emphasis Type="Italic">Limosa limosa</Emphasis> under different environmental conditions

There is a large literature dealing with daily foraging routines of wild birds during the non-breeding season. While different laboratory studies have showed that some bird activity patterns are a persistent property of the circadian system, most of field studies preclude the potential role of an endogenous circadian rhythm in controlling bird’s foraging routines. In this study we compared the patterns of diurnal foraging activity and intake rates of migrating black-tailed godwits, Limosa limosa (radio-tagged and non-tagged individuals) at two stopover sites (habitats) with different environmental characteristics, aiming at identifying proximate factors of bird activity routines. To gain insights into the role of food availability in control of such foraging routines, we also estimated foraging activity patterns in captive godwits subjected to constant food availability. Captive and wild black-tailed godwits showed a persistent bimodal activity pattern through daylight period. Food availability had a significant effect on the intake rates, but had a subtler effect on foraging and intake rate rhythms. Temperature and wind speed (combined in a weather index) showed non-significant effects on both rhythms. Although we could not discard a role for natural diurnal changes in light intensity, an important timing cue, our findings support the idea that an endogenous circadian rhythm could be an important proximate factor regulating foraging activity and food items taken per unit time of wild black-tailed godwits during migration.     

Optimal movement strategies for social foragers in unpredictable environments

Spatial movement models often base movement decision rules on traditional optimal foraging theories, including ideal free distribution (IFD) theory, more recently generalized as density-dependent habitat selection (DDHS) theory, and the marginal value theorem (MVT). Thus optimal patch departure times are predicted on the basis of the density-dependent resource level in the patch. Recently, alternatives to density as a habitat selection criterion, such as individual knowledge of the resource distribution, conspecific attraction, and site fidelity, have been recognized as important influences on movement behavior in environments with an uncertain resource distribution. For foraging processes incorporating these influences, it is not clear whether simple optimal foraging theories provide a reasonable approximation to animal behavior or whether they may be misleading. This study compares patch departure strategies predicted by DDHS theory and the MVT with evolutionarily optimal patch departure strategies for a wide range of foraging scenarios. The level of accuracy with which individuals can navigate toward local food sources is varied, and individual tendency for conspecific attraction or repulsion is optimized over a continuous spectrum. We find that DDHS theory and the MVT accurately predict the evolutionarily optimal patch departure strategy for foragers with high navigational accuracy for a wide range of resource distributions. As navigational accuracy is reduced, the patch departure strategy cannot be accurately predicted by these theories for environments with a heterogeneous resource distribution. In these situations, social forces improve foraging success and have a strong influence on optimal patch departure strategies, causing individuals to stay longer in patches than the optimal foraging theories predict.     

Central place food caching: a field experiment with red-backed shrikes (Lanius collurio L.)

Summary Red-backed shrikes (Lanius collurio L.) stored experimentally presented mice (Mus musculus L) by reimpaling them on thorns of sloe bushes (Prunus spinosa L.) in the vicinity of the nest. Large mice (20 g) were stored further away from the nest than small mice (4 g), while the smallest mice (1 g) were transported directly to the nest and cut up there. Large prey required more round trips to deliver than smaller prey (4 g). Time to immobilize, load and deliver the prey and the proportion of undigestable tissue increased with prey size. Birds used stored prey as an alternative to hunting in other patches as expected from patch use models. The birds maintained a high rate of food delivery to the young by using stored mice during periods when their foraging success of natural prey was low. Several aspects of the shrike's food storing behaviour are in qualitative accord with suggestions derived from models assuming maximization of energy delivery rate.     

Foraging impact on zooplankton by age-0 walleye pollock (<Emphasis Type="Italic">Theragra chalcogramma</Emphasis>) around a front in the southeast Bering Sea

The waters around the Pribilof Islands in the southeast Bering Sea are a center of abundance for age-0 walleye pollock (Theragra chalcogramma). Each spring and summer a tidal front is formed around the islands separating a well-mixed inshore habitat from a stratified offshore habitat. The objective of this study was to assess the foraging impact on zooplankton by age-0 pollock in the vicinity of this frontal structure. A bioenergetic model was used to estimate age-0 pollock food consumption from field estimates of water temperature, age-0 pollock density, diet and growth. Sampling of field variables took place over three hydrographic habitats along an inshore–offshore transect located north of the islands. The bioenergetics analysis was applied for a 2-week period during the late summer of four consecutive years, 1994–1997. Model results of age-0 pollock food consumption indicated variable levels of food depletion, changing with prey type, year and habitat. The foraging impact of age-0 pollock on copepods and euphausiids (most common prey) ranged from about 3% to 77% of the biomass available at the start of the simulation. Copepod depletion was typically greater than euphausiid depletion. Consequently, juvenile pollock <60 mm in standard length were more likely to experience food limitation due to the greater proportion of copepods in their diet. We present evidence of severe foraging impact during 1996, when one of the primary prey items of juvenile pollock (i.e. large copepods) was scarcely represented both in their diet and in the water column. In all years, most instances of prey depletion were found at the inshore and front habitats; age-0 pollock densities were too low relative to their prey to severely impact the offshore zooplankton populations. We discuss these results with respect to modeling assumptions and in the context of previously acquired knowledge of fish behavior around frontal regions.Communicated by T. Ikeda, Hakodate     

A patch use model to separate effects of foraging costs on giving-up densities: an experiment with white-tailed deer (<Emphasis Type="Italic">Odocoileus virginianus</Emphasis>)

The giving-up density of food (GUD), the amount of food remaining in a patch when a forager ceases foraging there, can be used to compare the costs of foraging in different food patches. But, to draw inferences from GUDs, specific effects of foraging costs (predation risk, metabolic and missed opportunities costs) on GUDs have to be identified. As high predation risk, high metabolic costs and abundant food all should produce high GUDs, this does not allow us to infer directly the quality of a habitat. In order to separate the effect of each foraging cost, we developed an optimal foraging model based on food supplementation. We illustrate the use of our model in a study where we assessed the impact of a power line right-of-way in a white-tailed deer (Odocoileus virginianus) winter yard by determining whether the negative effects of cover loss outweigh the positive effects of browse regeneration.     

Living in a ghetto within a local population: an empirical example of an ideal despotic distribution

Merging patterns and processes about the way individuals should be distributed in a habitat is a key issue in the framework of spatial ecology. Here the despotic distribution of individuals in two distinct and neighboring patches within a local population of a long-lived colonial bird, the Yellow-legged Gull (Larus michahellis), was assessed. There was no density dependence for suitable habitat at the study population, but behavioral data suggested that birds from the good patch precluded birds from the bad patch from breeding in their patch. Younger breeders were almost exclusively found in the bad patch, where individuals were probably attracted by conspecific attraction from the good patch. Most breeding parameters were lower in the bad patch, resulting mainly from a higher vulnerability to environmental perturbations and a higher rate of intraspecific nest predation. Attempts at breeding dispersal between the two patches were only observed from the bad to the good patch. Strikingly, adult survival and large-scale dispersal, two life history parameters that are very conservative in long-lived organisms, were also more affected at the bad patch when catastrophic predation occurred. The study was consistent with an ideal despotic distribution at small spatial scale, and suggests that individual behavior can influence local population dynamics.     

Risk-aversion,relative abundance of resources and foraging preference

Summary Foraging theory depicts dietary choice as a function of prey quality and absolute abundance. Ecological processes, however, can depend on the way foragers respond to the relative abundances of available prey types; several models for frequency-dependent foraging adequately describe these responses. Our laboratory experiments with white-throated sparrows investigated preferential choice of two food rewards as we manipulated both reward quality and relative abundance. In any single experiment the two rewards provided the same mean food quantity, but the variances differed. Average energy budgets predicted risk-aversion, so that foraging preference should decrease as reward variance increases. We presented each two-reward pairing at availability ratios of 1:2, 1:1, and 2:1 for three consecutive days. By the third day risk-aversion exceeded preference for reward variance significantly. When relative abundances of the low and high variance rewards were not equal, the birds tended to prefer the rare over the common reward. This response began before the birds had thoroughly sampled the reward distributions. Preference for rarity apparently constrained the birds' economic response to reward variance levels.     

Birds as Suppliers of Seed Dispersal in Temperate Ecosystems: Conservation Guidelines from Real‐World Landscapes

Abstract: Seed dispersal by animals is considered a pivotal ecosystem function that drives plant‐community dynamics in natural habitats and vegetation recovery in human‐altered landscapes. Nevertheless, there is a lack of suitable ecological knowledge to develop basic conservation and management guidelines for this ecosystem service. Essential questions, such as how well the abundance of frugivorous animals predicts seeding function in different ecosystems and how anthropogenic landscape heterogeneity conditions the role of dispersers, remain poorly answered. In three temperate ecosystems, we studied seed dispersal by frugivorous birds in landscape mosaics shaped by human disturbance. By applying a standardized design across systems, we related the frequency of occurrence of bird‐dispersed seeds throughout the landscape to the abundance of birds, the habitat features, and the abundance of fleshy fruits. Abundance of frugivorous birds in itself predicted the occurrence of dispersed seeds throughout the landscape in all ecosystems studied. Even those landscape patches impoverished due to anthropogenic disturbance received some dispersed seeds when visited intensively by birds. Nonetheless, human‐caused landscape degradation largely affected seed‐deposition patterns by decreasing cover of woody vegetation or availability of fruit resources that attracted birds and promoted seed dispersal. The relative role of woody cover and fruit availability in seed dispersal by birds differed among ecosystems. Our results suggest that to manage seed dispersal for temperate ecosystem preservation or restoration one should consider abundance of frugivorous birds as a surrogate of landscape‐scale seed dispersal and an indicator of patch quality for the dispersal function; woody cover and fruit resource availability as key landscape features that drive seedfall patterns; and birds as mobile links that connect landscape patches of different degrees of degradation and habitat quality via seed deposition.     

Temporal patterns of foraging activity in some wood warblers in relation to the availability of insect prey

Summary I monitored the temporal pattern of diurnal feeding activity in several wood warbler (Parulidae) species and concomitantly recorded the numbers of active (flying) insects in 2 willow habitats in the western United States. At one site the temporal relationship between the density of active and inactive (nonflying) insects was investigated. The diurnal patterns of insect and bird activity were inversely related and each pattern was significantly nonuniform throughout the day; the wood warblers were largely inactive during the middle of the day when insects were most active.As foliage-gleaning birds, wood warblers depend primarily on the availability of inactive (nonflying) insects that they pick from the foliage, and they appear to be limited in their foraging activity by the unavailability of such insects during midday. Interestingly, the duration of midday inactivity for a given bird species varied inversely with the proportion of time that species spent flycatching. Thus, food availability may play an important role in determining the temporal patterns of feeding activity in these insectivorous bird species.     

A Mechanistic Approach to Evaluation of Umbrella Species as Conservation Surrogates

Abstract:  Although species with large area requirements are sometimes used as umbrella species, their general utility as conservation tools is uncertain. We surveyed the species diversity of birds, butterflies, carabids, and forest-floor plants in forest sites across an area (1600 km²) in which we delineated large breeding home ranges of Northern Goshawk ( Accipiter gentilis ). We tested whether protection of the home ranges could serve as an effective umbrella to protect sympatric species of the four taxa. We also used an empirical habitat model of occupancy of home range to examine mechanisms by which the Northern Goshawk acts as an umbrella species. Among species richness, abundance, and species composition of the four taxa, only abundance and species composition of birds differed between sites located inside and outside home ranges, which was due to greater abundance of bird species that were prey of Northern Goshawks inside the home ranges. Thus, although home range indicated areas with high abundance of certain bird prey species, it was not effective as an indicator of the species diversity of all four taxa. We also did not find any difference in species richness, abundance, and species composition between sites predicted as occupied and unoccupied using the habitat model. In contrast, when we selected sites on the basis of each habitat variable in the model, habitat variables that selected sites either in agricultural or forested landscapes encompassed sites with high species richness or particular species composition. This result suggests that the low performance of the Northern Goshawk as an umbrella species is due to this species' preference for habitat in both agricultural and forested landscapes. Species that can adjust to changes in habitat conditions may not act as effective umbrella species despite having large home ranges.     

Functional Equivalency between Rice Fields and Seminatural Wetland Habitats

Abstract: Evaluating the potential for anthropogenic habitats to act as surrogates for the natural habitats they replace is a key issue in conservation biology. In California, flooded rice fields are used by numerous aquatic birds during winter. If this habitat functions similarly to more natural wetlands, increased flooding may help replace the extensive wetlands that occurred in the region prior to agricultural development. I tested whether food abundance, perceived predation threat, foraging performance, and the way in which birds allocate their time to different behaviors differed between flooded rice fields and seminatural wetlands for several species of aquatic bird. When appropriate, I also compared flooded and unflooded fields. Invertebrate densities did not differ among habitats. Seminatural wetlands had less rice grain but more seeds from other plants than the two rice habitats. The frequency with which predators passed over a feeding area was lower in flooded fields than in unflooded fields or seminatural wetlands. Most differences in feeding performance and time allocation among habitats were small and statistically insignificant. For some species, feeding efficiency was greater in seminatural wetlands than in flooded fields. Increasing attack rates and the amount of time spent feeding when in flooded fields, however, may allow birds to compensate for reduced efficiency. Multivariate analyses showed that group size, predation threat, time of day, date, and water depth often were associated with behaviors, but that these variables rarely accounted for habitat differences. Flooded fields apparently provide equivalent foraging habitat to seminatural wetlands and, because of reduced predation threat, may be a safer habitat for waterbirds. Thus, if managed appropriately, one of the world's dominant forms of agriculture can provide valuable waterbird habitat.     

Assessment of predation risk via illumination level: facultative central place foraging in the cricetid rodent Phyllotis darwini

It is well known that the risk of predation affects prey decision making. However, few studies have been concerned with the cues used by prey to assess this risk. Prey animals may use indirect environmental cues to assess predation hazard since direct evaluation may be dangerous. I studied the assessment of predation risk, manipulated via environmental illumination level, and the trade-off between foraging and predation hazard avoidance in the nocturnal rodentPhyllotis darwini (Rodentia: Cricetidae). In experimental arenas I simulated dark and full moon nights (which in nature correlate with low and high predation risk, respectively) and measured the immediate responses of animals to flyovers of a raptor model. Second, varying illumination only, I evaluated patch use, food consumption, central place foraging, and nocturnal variation of body weight. During flyover experiments, animals showed significantly more evasive reactions under full moon illumination than in moonless conditions. In the patch use experiments, rodents significantly increased their giving-up density and decreased their total food consumption under moonlight. On dark nights, rodents normally fed in the food patch, but when illumination was high they became central place foragers in large proportion. Moreover, the body weight of individuals decreased proportionately more during bright nights. These results strongly suggest thatP. darwini uses the level of environmental illumination as a cue to the risk of being preyed upon and may sacrifice part of its energy return to avoid risky situations.     

MORPH—An individual-based model to predict the effect of environmental change on foraging animal populations

This paper describes an individual-based model, MORPH, that has been designed to predict the effect of environmental change on foraging animal populations. The key assumptions of MORPH are that individuals within populations behave in order to maximise their perceived fitness, but that perceived fitness may not always be positively related to the actual chances of survival and reproduction. MORPH has been parameterised for coastal birds on several European sites and predicted the effect of environmental change, caused by factors such as habitat loss, disturbance from humans and sea-level rise, on the survival and body condition of these species. However, MORPH contains a basic framework to describe animal physiology and foraging behaviour, and the distribution and abundance of the resources required by these animals. Therefore, MORPH is not restricted to coastal birds, and is potentially applicable to a wider range of systems. To be applied to a forager system, MORPH requires parameters describing (i) the distribution of the food supply and how food quality and abundance changes through time; (ii) the rate at which foragers consume food given the abundance of food and competitors; (iii) the amount of food the forager must consume each day to survive; (iv) the distribution and seasonal changes in other factors which influence the foraging behaviour and survival of foragers. The purpose of this paper is to (i) describe MORPH, (ii) give examples of its application, (iii) describe the types of systems to which MORPH can be applied, and (iv) publish its source code and a user guide.