Density,diet and productivity of Long-eared Owls Asio otus in the Italian Alps: the importance of Microtus voles

Capsule Relatively large populations, feeding predominantly upon voles, were present at higher elevations.

Aims To determine the density, productivity and diet composition of Long-eared Owls breeding at higher elevations.

Methods Population census and breeding biology were investigated, and dietary analysis performed for 32 Long-eared Owl territories that were occupied over a six-year period (2000–05), in a 155 km² study area located in the Noce Valley of the central-eastern Italian Alps.

Results Territories were mainly situated at the edge of large pine forests, near large patches of grassland and/or open-structured apple orchards, at elevations ranging between 540 and 1210 m. Density varied between 10 and 15 pairs/100 km². Mean intraspecific nest spacing averaged 1727 m and territories were either solitary or clumped in loose aggregations of one to five pairs. Mean laying date was 27 March and the mean number of fledged young was 0.95 and 2.13 per territorial and successful pair, respectively. Adults and nestlings were preyed upon by Eagle Owls Bubo bubo and Common Buzzards Buteo buteo, both of them abundant in the study area. Diet was dominated by Common Voles Microtus arvalis, complemented by Apodemus mice and thrushes. Annual variations in owl density, productivity and diet breadth varied in parallel with the occurrence of Common Voles in the diet, suggesting that the owls responded numerically to the availability of their main prey. This confirms earlier analyses on other European populations, but without the marked three- to four-year cycles observed in such areas.

Conclusion The conservation status of the species in these Alpine habitats seemed currently satisfactory. However, the unregulated use of rodenticides in apple orchards and the loss of open habitats associated with land abandonment may represent long-term threats for the species in these habitats.     

Diet specificity is not associated with increased reproductive performance of Golden Eagles Aquila chrysaetos in Western Scotland

Amongst raptor species, individuals with specialized diets are commonly observed to have higher reproductive output than those with general diets. A suggested cause is that foraging efficiency benefits accrue to diet specialists. This diet specificity hypothesis thus predicts that diet breadth and reproductive success should be inversely related within species. We highlight, however, that a prey availability hypothesis also makes the same prediction in some circumstances. Hence, when high diet specificity results from high encounter rates with an abundant, preferred prey, then prey availability may affect reproductive success, with diet specialization as an incidental correlate. Using three insular study areas in western Scotland, we examine diet specificity and reproductive success in Golden Eagles Aquila chrysaetos. Diet breadth and breeding productivity were not negatively related in any of our study areas, even though birds with specific diets did tend to have a higher incidence of preferred prey (grouse and lagomorphs) in the diet. Indeed, in two study areas there was evidence that diet generalists had higher breeding productivity. Our results therefore failed to support the diet specificity hypothesis but were consistent with the prey availability hypothesis. We highlight that although many other studies are superficially consistent with the diet specificity hypothesis, our study is not alone in failing to provide support and that the hypothesis does not provide a generic explanation for all relevant results. Diet specificity in predators can be at least partially a response to prey diversity, availability and distribution, and benefits associated with different prey types, so that being a generalist is not necessarily intrinsically disadvantageous. We suggest that the available evidence is more consistent with variation in prey abundance and availability as a more influential factor explaining spatial and temporal variation in breeding productivity of ‘generalist’ species such as the Golden Eagle. Under this argument, prey abundance and availability are the main drivers of variation in reproductive output. Diet specificity is a consequence of variation in prey availability, rather than a substantial cause of variation in reproductive success.     

Diet–demography relationships in a long‐lived predator: from territories to populations

Understanding the mechanisms that shape animal population dynamics is of fundamental interest in ecology, evolution and conservation biology. Food supply is an important limiting factor in most animal populations and may have demographic consequences. Optimal foraging theory predicts greater consumption of preferred prey and less diet diversity when food is abundant, which may benefit key fitness parameters such as productivity and survival. Nevertheless, the correspondence between individual resource use and demographic processes in populations of avian predators inhabiting large geographic areas remains largely unexplored, particularly in complex ecosystems such as those of the Mediterranean basin. Based on a long‐term monitoring program of the diet and demography of Bonelli's eagle Aquila fasciata in western Europe, here we test the hypothesis that a predator's diet is correlated to its breeding productivity and survival at both the territorial and population levels, and ultimately to its population growth rate. At the territorial level, we found that productivity increased with greater consumption of European rabbits Oryctolagus cuniculus, the Bonelli's eagle's preferred prey, and pigeons, an important alternative prey for this predator. The survival of territorial pairs was negatively affected by higher diet diversity, which probably reflected the inability to find sufficient high quality prey. Diet effects at the population level were similar but more noticeable than at the territorial level, i.e. a greater consumption of rabbits, together with lesser consumption of small‐to‐medium avian species (‘other birds’; non‐preferred prey), increased productivity, while greater diet diversity and lower consumption of rabbits was associated with reduced survival and population growth rate. Overall, our study illustrates how the diet of a predator species can be closely related to key individual vital rates, which, in turn, leave a measurable fingerprint on population dynamics within and among populations across large spatial scales.     

The role of pigeon consumption in the population dynamics and breeding performance of a peregrine falcon (<Emphasis Type="Italic">Falco peregrinus</Emphasis>) population: conservation implications

In this paper, we describe and analyze the diet of peregrine falcons during a long-term period (1982–2002). A combination of direct observations of prey brought to nests, prey remains, and regurgitated pellets were used to calculate diet diversity and dietary overlap between peregrine pairs. We also examined diet diversity in relation to breeding performance. All peregrine pairs fed mainly on birds, with pigeons the most common prey. An increase in pigeon availability has been associated with both an increase in population size and an increase in breeding performance (measured as the average productivity of pairs per year) of a small peregrine falcon population in eastern Spain. Average productivity was lower when dietary breadth was higher. We speculate that our results were the synergistic effect of declining persecution and increased pigeon availability through increased popularity of keeping racing pigeons. There is a conflict of interests between pigeon fanciers and peregrine conservation. As a consequence, this could result to an increased risk of mortality by direct persecution. In accordance with this, conservation measures aimed at preventing direct persecution are encouraged.     

Diet of age-0 tarpon (<Emphasis Type="Italic">Megalops atlanticus</Emphasis>) in anthropogenically-modified and natural nursery habitats along the Indian River Lagoon,Florida

As human development in coastal areas increases, the role of anthropogenically-created habitats in the life history of marine organisms is becoming increasingly important. We examined the diet of age-0 tarpon, Megalops atlanticus, in and around man-made mosquito control impoundments along the Indian River Lagoon in east-central Florida, with a particular focus on identifying dietary patterns associated with tarpon size and nursery habitat type (i.e., between perimeter pool habitats in established impoundments and newly-created restoration marsh habitats). Age-0 tarpon were found to consume a wide variety of prey organisms, and exhibited considerable dietary variation among study sites. Smaller juvenile tarpon consumed a limited number of small prey taxa, while larger individuals fed on a greater range of prey taxa and sizes. Overall, copepods and fishes were the dominant prey items; however, the consumption of these organisms varied considerably among size classes and sites. There was no clear difference in tarpon diet between the two types of habitat we examined. The ability of juvenile tarpon to utilize such a diverse range of prey organisms may allow populations to inhabit a variety of habitats, including man-made marshes. When natural systems have been degraded or destroyed, human-altered habitats can assume a nursery role for the species.     

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.     

Breeding performance, age effects and territory occupancy in a Bonelli's Eagle Hieraaetus fasciatus population

Bonelli's Eagle Hieraaetus fasciatus is one of the most endangered birds of prey in Europe. Despite mounting interest and research, several questions regarding the conservation implications of territory occupancy and site-dependent population regulation remain insufficiently explored for this species. Here, we report on a 12-year study of the territorial structure of a Bonelli's Eagle population in southeastern Spain. No signals of population decline were found in the breeding population, as mean annual productivity was stable and the presence of mixed-age pairs in the population decreased with the years. However, the average proportion of subadults occupying territories was larger than that observed in other Spanish populations. Contrary to the predictions of a despotic distribution model, we found no significant relationship between occupancy rates and breeding parameters. Our results showed significant variations in productivity attributable to differences in the quality of individuals (i.e. mixed versus adult pairs), but no variability among territories per se (i.e. caused by habitat heterogeneity). Moreover, coexistence with intraguild species did not have any significant effect on productivity, although the proximity of Eagle Owls Bubo bubo affected the occupation rate of territories. Finally, our population does not appear to experience site-dependent population regulation, as a positive relation between mean annual productivity and density was found. The threat posed by changes in land use in the study area leads us to suggest that strict protection of current territories is necessary to ensure population persistence, and we suggest that a significant population increase is only likely if new or deserted territories become available.     

Spatial and temporal variations in the White-tailed Eagle Haliaeetus albicilla breeding diet revealed by prey remains

ABSTRACT

Capsule: Despite its population increase, the White-tailed Eagle Haliaeetus albicilla has not become food-limited, and does not pose a serious threat to birds of conservation concern in the region east of the Baltic Sea.

Aims: We aimed to test variations in the main prey groups and dietary niche breadth of White-tailed Eagles through the years and along an environmental gradient as well as to evaluate the extent of predation on bird species of conservation concern.

Methods: Prey remains were collected during nestling ringing from successful nests located in Lithuania.

Results: The frequency of the prey groups varied along an environmental gradient, with a general pattern of increasing avian prey consumed by pairs breeding in areas with less abundant aquatic habitats. The frequency of alternative prey (i.e. terrestrial birds, mammals) and dietary niche breadth did not increase between 2005 and 2018 and during which time the White-tailed Eagle population nearly doubled. Instead, the occurrence of Northern Pike Esox lucius remains, which was the most frequent single prey species, increased throughout the study period. Bird species of conservation concern constituted only a small fraction of all the prey identified.

Conclusions: The White-tailed Eagle has not become more dependent on alternative prey despite its population increase.     

Breeding density,habitat selection and reproductive rates of the Peregrine Falcon Falco peregrinus in Álava (northern Spain)

A population of 33–35 pairs of Peregrine Falcon in álava (north Spain) was studied with the aim of assessing the habitat attributes that influence breeding density, habitat selection and breeding success. A strong relationship was found between density of the species in each UTM (Universal Transverse Mercator) square of 10 × 10 km and cliff availability. Habitat selection was analysed by comparing 15 variables in 33 occupied and 25 unoccupied cliffs located at least 2 km from the nearest Peregrine pair. Significant differences were found in five variables: cliff dominance, distance to the nearest Golden Eagle Aquila chrysaetos nesting cliff, steepness and altitude (all showing larger values in occupied cliffs), and cliff orientation, with occupied cliffs facing preferably south and east. Orientation, dominance and distance to the nearest Golden Eagle pair, and the distance to the nearest Eagle Owl Bubo bubo nesting cliff, were included in a discriminant analysis which classified 82.76% of the cliffs correctly. The productivity of the studied population was 1.44 young/territorial pair (n = 45), and no consistent relationship was found between breeding success and habitat variables.     

Using molecular and crowd-sourcing methods to assess breeding ground diet of a migratory brood parasite of conservation concern

Breeding ground food availability is critical to the survival and productivity of adult birds. The common cuckoo Cuculus canorus is a brood-parasitic Afro-Palearctic migrant bird exhibiting long-term (breeding) population declines in many European countries. Variation in population trend between regions and habitats suggests breeding ground drivers such as adult food supply. However, cuckoo diet has not been studied in detail since before the most significant population declines in Europe began in the mid-1980s. 20th century studies of cuckoo diet largely comprised field observations likely to carry bias towards larger prey taxa. Here we demonstrate the potential value of 1) using high-throughput DNA sequencing of invertebrate prey in faeces to determine cuckoo diet with minimal bias towards large prey taxa, and 2) using crowd-sourced digital photographs from across Britain to identify lepidopteran cuckoo prey taxa during recent years post-decline (2005–2016). DNA analysis found a high frequency of Lepidoptera, including moths of family Lasiocampidae, prominent within the past literature, but also grasshoppers (Orthoptera) and flies (Diptera) that may be overlooked by field observation methodologies. The range of larval lepidopteran prey identified from photographs largely agreed with those previously documented, with potential signs of reduced diversity, and identities of key adult prey taxa were supported by molecular results. Notably, many identified cuckoo prey taxa have shown severe declines due to agricultural intensification, suggesting this has driven spatial patterns of cuckoo loss. Landscape-scale, lowland rewilding interventions provide opportunities to understand the scale of reversal of previous agricultural intensification that may be necessary to restore prey populations sufficiently to permit recolonization by cuckoos.     

The effect of elevation and habitat cover on nest box occupancy and diet composition of Boreal Owls Aegolius funereus

Capsule: Diet composition of Boreal Owls Aegolius funereus was not affected by habitat cover, but it changed along the elevational gradient.

Aims: To assess the effect of elevation and habitat cover on nest box occupancy and diet composition of a central European population of Boreal Owls.

Methods: A Boreal Owl population was studied in the ?umava Mountains, Czech Republic, at elevations from 500 to 1300?m above sea level (asl), during 1984–2005.

Results: Boreal Owls occupied more frequently nest boxes above 600?m asl, but they did not clearly prefer any elevational band. Habitat cover did not affect the number of nesting attempts. There was also no relationship between habitat cover and diet composition. However, diet composition significantly changed along the elevational gradient. In particular, the proportion of alternative prey of Boreal Owls, i.e. birds and shrews Sorex sp., rose with increasing elevation. The proportion of voles Myodes and mice Apodemus in the diet decreased with increasing elevation. Among bird prey, the proportion of finches Fringillidae positively correlated with elevation.

Conclusions: Central European Boreal Owls did not show a clear preference for any habitat cover or elevational band, but the quality of the owls’ diet significantly decreased with increasing elevation.     

Main stem and off‐channel habitat use by juvenile Chinook salmon in a sub‐Arctic riverscape

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Shifts in species richness, herbivore specialization, and plant resistance along elevation gradients

Environmental gradients have been postulated to generate patterns of diversity and diet specialization, in which more stable environments, such as tropical regions, should promote higher diversity and specialization. Using field sampling and phylogenetic analyses of butterfly fauna over an entire alpine region, we show that butterfly specialization (measured as the mean phylogenetic distance between utilized host plants) decreases at higher elevations, alongside a decreasing gradient of plant diversity. Consistent with current hypotheses on the relationship between biodiversity and the strength of species interactions, we experimentally show that a higher level of generalization at high elevations is associated with lower levels of plant resistance: across 16 pairs of plant species, low-elevation plants were more resistant vis-à-vis their congeneric alpine relatives. Thus, the links between diversity, herbivore diet specialization, and plant resistance along an elevation gradient suggest a causal relationship analogous to that hypothesized along latitudinal gradients.     

PREY OF GIANT EAGLE OWLS IN THE DE HOOP NATURE RESERVE,CAPE PROVINCE,AND SOME OBSERVATIONS ON HUNTING STRATEGY

Avery, G., Robertson, A. S., Palmer, N.G. & Prins, A. J. 1985. Prey of Giant Eagle Owls in the De Hoop Nature Reserve, Cape Province, and some observations on hunting strategy. Ostrich 56: 117–122.

Prey remains of Giant Eagle Owls Bubo lacteus in the De Hoop Nature Reserve were analysed in terms of numerical composition and mass. Thirteen species of bird contributed 57,8 % of the mass, six species of mammal 41,4 %, four species of invertebrate less than 0,4 % and one species of amphibian 0;4% Black-headed Heron Ardea melanocephala was the most common bird representing 14,8 % of the mass. Accipitridae (buzzards and harriers) and Tytonidae (owls represented 10,1 % of the mass. Leporidae (hares) were the most common mammals represented 10,7% of the mass. A single bone of an adult Grysbok Raphicerus melanoris represented 20,1% of the mass and was almost certanly taken as carrion. Similarities with diet data from other areas are apparent. The Giant Eagle Owl is omnivorous, its diet being determined bayrhabitat and local abundance. Comparison with the diet of Cape Eagle Owl Bubo capensis confirms that separation of habitats, with associated differences in the accessibility and availability of prey, explains the differences in prey taken by these species.     

Spatial structure in the diet of imperial eagles Aquila heliaca in Kazakhstan

We evaluated the relationship between spatial variability in prey and food habits of eastern imperial eagles Aquila heliaca at a 90,000 ha national nature reserve in north-central Kazakhstan. Eagle diet varied greatly within the population and the spatial structure of eagle diet within the population varied according to the scale of measurement. Patterns in dietary response were inconsistent with expectations if either ontogenetic imprinting or competition determined diet choice, but they met expectations if functional response determined diet. Eagles nesting near a high-density prey resource used that resource almost exclusively. In contrast, in locations with no single high-density prey species, eagles' diet was more diverse. Our results demonstrate that spatial structuring of diet of vertebrate predators can provide important insight into the mechanisms that drive dietary decisions.     

Habitat selection, diet, arthropod availability and growth of a moorland wader: the ecology of European Golden Plover Pluvialis apricaria chicks

Habitat use, diet and food supply of European Golden Plover Pluvialis apricaria chicks were studied on blanket bog in the South Pennines, UK. The home ranges occupied by chicks until fledging averaged 40 ha; they contained relatively more cotton grass and bare peat than was available generally, but less heather and grassland. Use of Bilberry Vaccinium myrtillus and Crowberry Empetrum nigrum increased with age, whereas that of cotton grass Eriophorum spp. declined. Dietary composition, as assessed by faecal analysis, was similar to that obtained from the crops of dead birds. The diet of younger chicks, assessed by dry weight of prey, consisted of about 30% each of adult and larval tipulids, whereas for chicks older than 16 days, about 70% was tipulid larvae. Beetles, spiders and caterpillars each comprised 5–20% of the diet, depending on age. Older chicks took larger prey. The exploitation of larval tipulids was correlated positively with the chicks' use of cotton grass and bare peat areas, whereas caterpillars, beetles and spiders were more often taken from dwarf shrubs, reflecting variation in prey abundance. Bare peat was an important foraging habitat, whose exploitation was associated positively with the growth rate of young chicks, and with mean prey size. Both weight gain and survival of young chicks were positively correlated with the abundance of tipulid adults, confirming the hypothesis that the flush of tipulid hatching is an important determinant of breeding success. Golden Plover chicks are capable of considerable movements to track variation in prey availability. This behaviour is likely to be an important survival strategy facilitating the utilization of a mosaic of cotton grass and dwarf shrub habitats. Appropriate rotational strip-burning or grazing management could be used to create and maintain such habitats.     

Population trend and breeding biology of Montagu's Harrier Circus pygargus in a natural vegetation site in northeast Spain

Capsule Reproductive output in a natural habitat was higher than in birds breeding in cereal crops, highlighting the importance of natural habitats for the species.

Aims To evaluate breeding in a natural habitat in inland Castellon province, Spain, and compare breeding parameters with other European populations.

Methods Breeding population size in inland Castellon was recorded between 1981 and 2003. A logistic growth model was used to describe population increase. We also calculated clutch size, brood size, productivity, percentage of successful nests and fledging success for the period 1989–2003.

Results Population size increased from three pairs in 1981 to 98 pairs in 2003. During 1989–2003, productivity (2.74 ± 1.49 fledglings/pair) and percentage of successful nests (84.%) were higher than observed in other European populations.

Conclusions Our results show the importance of our study area for the conservation of this species in the Iberian Peninsula, as it might act as a source of colonists for other areas. Food and protection against predators might explain the high breeding output, which may in turn explain the rapid population growth in the area. The study also suggests that natural habitats might be important for the species elsewhere.     

Multi‐season occupancy models identify biotic and abiotic factors influencing a recovering Arctic Peregrine Falcon Falco peregrinus tundrius population

Critical information for evaluating the effectiveness of management strategies for species of concern include distinguishing seldom occupied (or low‐quality) habitat from habitat that is frequently occupied and thus contributes substantially to population trends. Using multi‐season models that account for imperfect detection and a long‐term (1981–2002) dataset on migratory Arctic Peregrine Falcons Falco peregrinus tundrius nesting along the Colville River, Alaska, we quantified the effects of previous year's productivity (i.e. site quality), amount of prey habitat, topography, climate, competition and year on occupancy dynamics across two spatial scales (nest‐sites, cliffs) during recovery of the population. Initial occupancy probability was positively correlated with area of surrounding prey habitat and height of nest‐sites above the Colville River. Colonization probability was positively correlated with nest height and negatively correlated with date of snowmelt. Local extinction probability was negatively correlated with productivity, area of prey habitat and nest height. Colonization and local extinction probabilities were also positively and negatively correlated, respectively, with year. Our results suggest that nest‐sites (or cliffs) along the Colville River do not need equal protection measures. Nest‐sites and cliffs with historically higher productivity were occupied most frequently and had lower probability of local extinction. These sites were on cliffs high above the river drainage, surrounded by adequate prey habitat and with southerly aspects associated with early snowmelt and warmer microclimates in spring. Protecting these sites is likely to encourage continued occupancy by Arctic Peregrine Falcons along the Colville River and other similar areas. Our findings also illustrate the importance of evaluating fitness parameters along with climate and habitat features when analysing occupancy dynamics, particularly with a long‐term dataset spanning a range of annual climate variation.     

Population density of the greater mouse-eared bat (Myotis myotis), local diet composition and availability of foraging habitats

We studied the regional variation in population density of Myotis myotis (Borkhausen 1797) in south-eastern Bavaria, Germany, and its relations to diet composition and the availability of potential foraging habitats. We monitored colony size and juvenile mortality from 1991 to 2003, conducted faecal analyses in 1993 and determined land-use patterns around colonies. The numbers of individuals counted in the nursery colonies showed only small fluctuations over the years. However, data on colony size demonstrated a pronounced regional variation. Epigeic arthropods, mainly Carabidae, were the most important prey. The diet included prey taxa of forest as well as grassland habitats. The percentage of those prey taxa in the diet that originated in grassland managed with different intensity varied according to the availability of these potential foraging habitats around the nursery roosts. The calculated population density of the bats was positively correlated to forest area and especially to the area of mixed forest around the nursery roosts. Our results indicate that the availability of foraging habitats is a limiting factor for local population densities in the greater mouse-eared bat.     

Invertebrate prey availability limits reproductive success but not breeding population size in suburban House Sparrows Passer domesticus

Factors affecting avian demography and abundance in urban landscapes are poorly understood and this hinders attempts to manage urban bird communities. Several recent studies indicate that lack of invertebrate prey in urban landscapes may constrain avian productivity and fitness relative to that in other habitats. House Sparrow Passer domesticus populations have undergone large declines in many European urban centres and inadequate reproductive success linked to invertebrate availability has been postulated as a potential cause of these declines. We conducted a replicated supplementary feeding experiment to test whether the availability of invertebrate prey limits the breeding success and adult abundance (colony size) of House Sparrows in suburban London, where House Sparrow populations declined by 60% during the decade preceding our study. Daily mealworm provision over two successive breeding seasons, sufficient to provide 82% of chick energy requirements of House Sparrow pairs nesting within 50 m of feeders, had a large positive impact on the abundance of recently fledged birds (+62%), but only a small positive impact, limited mainly to small colonies, on the overall abundance of territorial males. Colony growth was only weakly related to fledgling abundance in the previous year and did not appear to be constrained by nest‐site availability. Conservation interventions that enhance invertebrate availability for suburban House Sparrows may increase reproductive success but are unlikely, on their own, to lead to population growth or recovery.