The design of artificial nestboxes for the study of secondary hole-nesting birds: a review of methodological inconsistencies and potential biases

The widespread use of artificial nestboxes has led to significant advances in our knowledge of the ecology, behaviour and physiology of cavity nesting birds, especially small passerines Nestboxes have made it easier to perform routine monitoring and experimental manipulation of eggs or nestlings, and also repeatedly to capture, identify and manipulate the parents However, when comparing results across study sites the use of nestboxes may also Introduce a potentially significant confounding variable in the form of differences in nestbox design amongst studies, such as their physical dimensions, placement height, and the way in which they are constructed and maintained However, the use of nestboxes may also introduce an unconsidered and potentially significant confounding variable clue to differences in nestbox design amongst studies, such as their physical dimensions, placement height, and the way in which they are constructed and maintained Here we review to what extent the characteristics of artificial nestboxes (e g size, shape, construction material, colour) are documented in the 'methods' sections of publications involving hole-nesting passerine birds using natural or excavated cavities or artificial nestboxes for reproduction and roosting Despite explicit previous recommendations that authors describe in detail the characteristics of the nestboxes used, we found that the description of nestbox characteristics in most recent publications remains poor and insufficient We therefore list the types of descriptive data that should be included in the methods sections of relevant manuscripts and justify this by discussing how variation in nestbox characteristics can affect or confound conclusions from nestbox studies We also propose several recommendations to improve the reliability and usefulness of research based on long-term studies of any secondary hole-nesting species using artificial nestboxes for breeding or roosting.

Predation in Ground-Nesting Birds: an Experimental Study Using Natural Egg-Color Variation

We tested the hypothesis that cryptically colored eggs would suffer less predation than conspicuous eggs in the ground-nesting red-legged partridge, Alectoris rufa. We used A. rufa as a model species because it has a wide range of natural egg colors, the eggs are widely available from breeding farms, and nests are easily mimicked because they are scrapes containing no vegetation. The study was conducted in the spring of 2001 in forest and fallow fields of central Spain in Castilla La Mancha, Ciudad Real. We used 384 clutches of natural eggs that were white, white spotted, brown, or brown spotted. Within clutches, eggs were consistent in color and size; among clutches, color differences were distributed across habitats. Clutches were checked once after 2 wk of exposure. Cryptic coloration had a survival advantage that was dependent on the local suite of predators. Rodent predation was nonselective with respect to clutch color; however, avian predation was significantly higher for conspicuous clutches. In addition, there was an interaction of landscape and egg color for avian predation. In forest landscapes, the clutches with highest survival were brown spotted, whereas in fallow landscapes, brown and brown spotted clutches had higher survival than white and white potted clutches. Thus, both the predator suite and the landscape had significant effects on the value of cryptic egg coloration. Our study is relevant for conservationists and managers in charge of restocking programs in hunting areas. The release of other partridge species or their hybrids could result in hybridization with wild partridges, potentially leading to nonoptimal clutch pigmentation and reduced survival of the native species. We therefore recommend that local authorities, managers, and conservationists be cautious with the use of alien species and hybrids and release only autochthonous species of partridges within their natural ranges.

How many of Australia's ground-nesting birds are likely to be at risk from the invasive Cane Toad (Rhinella marina)?

Cane Toads (Rhinella marina; hereafter 'toads') are large, toxic American anurans that were introduced to Australia in 1935. Research on their ecological impact has focussed on the lethal ingestion of toxic toads by native frog-eating predators. Less attention has been paid to the potential impacts of Cane Toads as predators, although these large anurans sometimes eat vertebrates, such as nestling birds and bird eggs. We review published and unpublished data on interactions between Cane Toads and Australian ground-nesting birds, and collate distributional and breeding information to identify the avian taxa potentially at risk of having eggs or chicks eaten by Cane Toads. Cane Toads are currently sympatric with 80 ground-nesting bird species in Australia, and five additional species of bird occur within the predicted future range of the toad. Although many species of bird are potentially at risk, available data suggest there is minimal impact of Cane Toads on ground-nesting species. Future research could usefully address both direct and indirect impacts of the invasion by Cane Toads, ideally with detailed field observations of these impacts on nesting success and of changes in bird breeding success as a function of invasion by toads.

Regional drivers of clutch loss reveal important trade-offs for beach-nesting birds

Coastal birds are critical ecosystem constituents on sandy shores, yet are threatened by depressed reproductive success resulting from direct and indirect anthropogenic and natural pressures. Few studies examine clutch fate across the wide range of environments experienced by birds; instead, most focus at the small site scale. We examine survival of model shorebird clutches as an index of true clutch survival at a regional scale (∼200 km), encompassing a variety of geomorphologies, predator communities, and human use regimes in southeast Queensland, Australia. Of the 132 model nests deployed and monitored with cameras, 45 (34%) survived the experimental exposure period. Thirty-five (27%) were lost to flooding, 32 (24%) were depredated, nine (7%) buried by sand, seven (5%) destroyed by people, three (2%) failed by unknown causes, and one (1%) was destroyed by a dog. Clutch fate differed substantially among regions, particularly with respect to losses from flooding and predation. ‘Topographic’ exposure was the main driver of mortality of nests placed close to the drift line near the base of dunes, which were lost to waves (particularly during storms) and to a lesser extent depredation. Predators determined the fate of clutches not lost to waves, with the depredation probability largely influenced by region. Depredation probability declined as nests were backed by higher dunes and were placed closer to vegetation. This study emphasizes the scale at which clutch fate and survival varies within a regional context, the prominence of corvids as egg predators, the significant role of flooding as a source of nest loss, and the multiple trade-offs faced by beach-nesting birds and those that manage them.

Selecting umbrella species for conservation: a test of habitat models and niche overlap for beach-nesting birds

Umbrella species are rarely selected systematically from a range of candidate species. On sandy beaches, birds that nest on the upper beach or in dunes are threatened globally and hence are prime candidates for conservation intervention and putative umbrella species status. Here we use a maximum-likelihood, multi-species distribution modeling approach to select an appropriate conservation umbrella from a group of candidate species occupying similar habitats. We identify overlap in spatial extent and niche characteristics among four beach-nesting bird species of conservation concern, American oystercatchers (Haematopus palliatus), black skimmers (Rynchops niger), least terns (Sterna antillarum) and piping plovers (Charadrius melodus), across their entire breeding range in New Jersey, USA. We quantify the benefit and efficiency of using each species as a candidate umbrella on the remaining group. Piping plover nesting habitat encompassed 86% of the least tern habitat but only 15% and 13% of the black skimmer and American oystercatcher habitat, respectively. However, plovers co-occur with all three species across 66% of their total nesting habitat extent (~ 649 ha), suggesting their value as an umbrella at the local scale. American oystercatcher nesting habitat covers 100%, 99% and 47% of piping plover, least tern and black skimmer habitat, making this species more appropriate conservation umbrellas at a regional scale. Our results demonstrate that the choice of umbrella species requires explicit consideration of spatial scale and an understanding of the habitat attributes that an umbrella species represents and to which extent it encompasses other species of conservation interest. Notwithstanding the attractiveness of the umbrella species concept, local conservation interventions especially for breeding individuals in small populations may still be needed.

Is bird incidence in Atlantic forest fragments influenced by landscape patterns at multiple scales?

The degree to which habitat fragmentation affects bird incidence is species specific and may depend on varying spatial scales. Selecting the correct scale of measurement is essential to appropriately assess the effects of habitat fragmentation on bird occurrence. Our objective was to determine which spatial scale of landscape measurement best describes the incidence of three bird species (Pyriglena leucoptera, Xiphorhynchus fuscus and Chiroxiphia caudata) in the fragmented Brazilian Atlantic forest and test if multi-scalar models perform better than single-scalar ones. Bird incidence was assessed in 80 forest fragments. The surrounding landscape structure was described with four indices measured at four spatial scales (400-, 600-, 800- and 1,000-m buffers around the sample points). The explanatory power of each scale in predicting bird incidence was assessed using logistic regression, bootstrapped with 1,000 repetitions. The best results varied between species (1,000-m radius for P. leucoptera; 800-m for X. fuscus and 600-m for C. caudata), probably due to their distinct feeding habits and foraging strategies. Multi-scale models always resulted in better predictions than single-scale models, suggesting that different aspects of the landscape structure are related to different ecological processes influencing bird incidence. In particular, our results suggest that local extinction and (re)colonisation processes might simultaneously act at different scales. Thus, single-scale models may not be good enough to properly describe complex pattern-process relationships. Selecting variables at multiple ecologically relevant scales is a reasonable procedure to optimise the accuracy of species incidence models.

Pileated Woodpecker Occupancy and the Occurrence and Recruitment of Key Habitat Attributes in a Managed Forest

Thesis (Master's)--University of Washington, 2016-08

Origin and Availability of Large Cavities for Barrow’s Goldeneye (Bucephala islandica), a Species at Risk Inhabiting the Eastern Canadian Boreal Forest

Large secondary-nesting birds such as ducks rely on appropriate cavities for breeding. The main objective of this study was to assess the availability of large cavities and the potential of a managed boreal coniferous landscape to provide nesting trees within the breeding area of the eastern population of Barrow’s Goldeneye (Bucephala islandica), a cavity-nesting species at risk in Canada. Woodpecker surveys were conducted in both conifer and mixed-wood landscapes, and cavities were sought in line transects distributed in unharvested and linear remnant stands of balsam fir (Abies balsamea) and black spruce (Picea mariana) as well as in cutblocks. No Pileated Woodpeckers (Dryocopus pileatus) were detected in the breeding area of Barrow’s Goldeneye, but the species was present in the nearby lowland area in which trembling aspen (Populus tremuloides) is abundant. Only 10 trees (0.2% of those sampled) supported cavities considered suitable for Barrow’s Goldeneye in terms of dimensions and canopy openness. Most of the suitable cavities found during this study were nonexcavated apical (chimney) cavities in relatively short snags that showed advanced states of decay. A diameter-at-breast-height threshold was determined for each tree species, after which the probability of cavity occurrence was enhanced in terms of potential cavity trees for Barrow’s Goldeneye. Remnant linear forest sites had lower potential tree densities than did their unharvested equivalents. Large cavities were thus a rare component in this boreal landscape, suggesting that they may be a limiting factor for this population at risk. Current even-aged forest management that mainly relies on clear-cut practices is likely to further reduce the potential of this landscape to provide trees with suitable cavities.

Birds and beaches, dogs and leashes : dog owners' sense of obligation to leash dogs on beaches in Victoria, Australia

Domesticated dogs threaten the conservation of beach-nesting birds in Australia through disturbance, and destruction of eggs and chicks. Leashing of dogs can improve conservation outcomes, but few dogs are leashed on beaches. We surveyed dog owners to explore their sense of obligation to leash dogs on beaches. Dog owners were more likely to feel obliged to leash their dog when they believed other people expected dogs to be leashed, and when they believed their dog was a threat to wildlife or people. Dog owners were less likely to feel obliged to leash their dog if they considered unleashed dog recreation to be important. Improved compliance may be achieved through community-based approaches to foster social norms for dog control, tailoring information products to emphasize the risk that all unleashed dogs may pose to beach-nesting birds and raising awareness of designated off-leash exercise dog recreation areas.

The influence of cover on nesting Red-capped Plovers: a trade-off between thermoregulation and predation risk?

 Some ground-nesting birds adopt a mixed strategy of nesting in the open, or under cover (e.g. vegetation). This may represent a trade-off between thermally favourable nest sites (covered) and those that enable the early detection and avoidance of predators (open). This study examined whether such a trade-off exists for Redcapped Plover Charadrius ruficapillus, whose eggs are preyed upon principally by Little Raven Corvus mellori. For real and artificial nests, nest temperatures under cover (real, 25.9 ± 0.1°C; false, 16.2 ± 0.5°C) were cooler than those in the open (real, 26.8 ± 0.1°C; false, 17.4 ± 0.9°C). Covered nests had more visual obstructions than open nests (covered, 65.5% ± 11.4%; open, 7.4% ± 2.8%) and a standardised measure of incubator escape distance, initiated by experimental human approaches, indicated incubators fled open nests at longer distances than for covered nests. Nests under cover showed a slightly (non-significant) higher probability of surviving one day (Daily Survival Rate [DSR] = 0.978) than those in the open (DSR = 0.950). For false nests containing model eggs, covered nests exhibited better survival to 10 days compared with open nests (20.4% vs. 4.7%). Thus, covered nests are associated with enhanced thermal environments and egg survival, but predators can approach the incubator more closely. Overall, the proposed trade-off between thermal and predation risks associated with nest sites appears to exist and explains the ongoing occurrence of nests in open and covered locations.

Cover, not caging, influences chronic physiological stress in a ground-nesting bird

Predator exclosures ('nest cages') around nests are increasingly used to enhance hatching success of declining ground-nesting birds. However, such exclosures are contentious and have been suggested to have detrimental effects on the species which they aim to protect. This study examines whether exclosures increase physiological stress of incubating birds, a hitherto unrecognised and untested potential drawback of exclosures. Red-capped plover Charadrius ruficapillus hatching success was radically altered and significantly higher for nests with exclosures (96.2%) compared with those without (6.8%). Chronic physiological stress in parents (as measured by the heterophil/lymphocyte [H/L] ratio in blood) did not vary between nests with and without exclosures, or between the sexes. However the absence of vegetative cover at the nest site was associated with a 62.7% elevation in H/L ratio, indicating that incubating birds which place their nests in the open are subject to increased levels of chronic stress. The results from this study demonstrate the fundamental importance of predation for the nesting success of this species and confirm that chronic stress levels are not a detrimental side effect of exclosure use.

Intense predation of non-colonial, ground-nesting bird eggs by corvid and mammalian predators

Context Loss of eggs to predators is a major cause of reproductive failure among birds. It is especially pronounced among ground-nesting birds because their eggs are accessible to a wide range of predators. Few studies document the main causes of clutch fate of ground-nesting birds. Aims The main objective of the present study was to identify the major egg predator of red-capped plovers (Charadrius ruficapillus). We also investigated the effectiveness of the following two primary strategies available to the plovers to avoid egg predation: (1) the placement of clutches under vegetative cover and (2) avoiding predators by nesting outside the peak season of predator occurrence. Methods Remote-sensing cameras were deployed on plover nests to identify egg predators and nests were monitored over four breeding seasons to document reproductive success and fate. An experiment using false clutches with model eggs investigated the influence of nest cover on the risk of egg predation throughout the year. Line-transect surveys were conducted to estimate the abundance of egg predators in and around the wetlands. Key results The little raven (Corvus mellori) was the major egg predator identified in 78.6% of red-capped plover clutches and in 92.4% of false clutches that were camera-monitored. The hatching success of plover eggs was not influenced by nest cover (P≤0.36), but model egg survival in false clutches improved significantly with the presence of nest cover (P≤0.02). The abundance of little ravens increased during the plover breeding season and was highly negatively correlated with false clutch survival (rpearson≤-0.768, P≤0.005). Conclusions Little ravens were the major predator of red-capped plover eggs and their abundance increased significantly during the plover breeding season. Any influence of nest cover on hatching success of eggs may have been masked by the extremely high rate of egg loss associated with the increased little raven abundance during the plover breeding season. Implications The high rate of egg predation is likely to have negative consequences on the local red-capped plover population, suggesting management is warranted. Little raven populations have expanded and, thus, their impact as egg predators needs to be investigated especially on threatened species. Journal compilation

(Table 1) Nesting characteristics of greater snow geese (Chen caerulescens atlanticus) on Bylot Island between 1995-2005

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3-5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.