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Perspectives and challenges for the use of radar in biological conservation

Radar is at the forefront for the study of broad‐scale aerial movements of birds, bats and insects and related issues in biological conservation. Radar techniques are especially useful for investigating species which fly at high altitudes, in darkness, or which are too small for applying electronic tags. Here, we present an overview of radar applications in biological conservation and highlight its future possibilities. Depending on the type of radar, information can be gathered on local‐ to continental‐scale movements of airborne organisms and their behaviour. Such data can quantify flyway usage, biomass and nutrient transport (bioflow), population sizes, dynamics and distributions, times and dimensions of movements, areas and times of mass emergence and swarming, habitat use and activity ranges. Radar also captures behavioural responses to anthropogenic disturbances, artificial light and man‐made structures. Weather surveillance and other long‐range radar networks allow spatially broad overviews of important stopover areas, songbird mass roosts and emergences from bat caves. Mobile radars, including repurposed marine radars and commercially dedicated ‘bird radars’, offer the ability to track and monitor the local movements of individuals or groups of flying animals. Harmonic radar techniques have been used for tracking short‐range movements of insects and other small animals of conservation interest. However, a major challenge in aeroecology is determining the taxonomic identity of the targets, which often requires ancillary data obtained from other methods. Radar data have become a global source of information on ecosystem structure, composition, services and function and will play an increasing role in the monitoring and conservation of flying animals and threatened habitats worldwide.

Ecography

Life-history variation of a neotropical thrush challenges food limitation theory

Since David Lack first proposed that birds rear as many young as they can nourish, food limitation has been accepted as the primary explanation for variation in clutch size and other life-history traits in birds. The importance of food limitation in life-history variation, however, was recently questioned on theoretical grounds. Here, we show that clutch size differences between two populations of a neotropical thrush were contrary to expectations under Lack's food limitation hypothesis. Larger clutch sizes were found in a population with higher nestling starvation rate (i.e. greater food limitation). We experimentally equalized clutches between populations to verify this difference in food limitation. Our experiment confirmed greater food limitation in the population with larger mean clutch size. In addition, incubation bout length and nestling growth rate were also contrary to predictions of food limitation theory. Our results demonstrate the inability of food limitation to explain differences in several life-history traits: clutch size, incubation behaviour, parental feeding rate and nestling growth rate. These life-history traits were better explained by inter-population differences in nest predation rates. Food limitation may be less important to life history evolution in birds than suggested by traditional theory. ?? 2005 The Royal Society.

Proceedings of the Royal Society B: Biological Sci

Variation in selective regimes drives intraspecific variation in life-history traits and migratory behaviour along an elevational gradient

Comparative studies, across and within taxa, have made important contributions to our understanding of the evolutionary processes that promote phenotypic diversity. Trait variation along geographic gradients provides a convenient heuristic for understanding what drives and maintains diversity. Intraspecific trait variation along latitudinal gradients is well-known, but elevational variation in the same traits is rarely documented. Trait variation along continuous elevational gradients, however, provides compelling evidence that individuals within a breeding population may experience different selective pressures. Our objectives were to quantify variation in a suite of traits along a continuous elevational gradient, evaluate whether individuals in the population experience different selective pressures along that gradient and quantify variation in migratory tendency along that gradient. We examined variation in a suite of 14 life-history, morphological and behavioural traits, including migratory tendency, of yellow-eyed juncos along a continuous 1000-m elevational gradient in the Santa Catalina Mountains of Arizona. Many traits we examined varied along the elevational gradient. Nest survival and nestling growth rates increased, while breeding season length, renesting propensity and adult survival declined, with increasing elevation. We documented the migratory phenotype of juncos (partial altitudinal migrants) and show that individual migratory tendency is higher among females than males and increases with breeding elevation. Our data support the paradigm that variation in breeding season length is a major selective pressure driving life-history variation along elevational gradients and that individuals breeding at high elevation pursue strategies that favour offspring quality over offspring quantity. Furthermore, a negative association between adult survival and breeding elevation and a positive association between nest survival and breeding elevation help explain both the downslope and reciprocal upslope seasonal migratory movements that characterize altitudinal migration in many birds. Our results demonstrate how detailed studies of intraspecific variation in suites of traits along environmental gradients can lend new insights into the evolutionary processes that promote diversification and speciation, the causes of migratory behaviour, and how animal populations will likely respond to climate change.

Journal of Animal Ecology

Are migrating raptors guided by a geomagnetic compass?

We tested whether routes of raptors migrating over areas with homogeneous topography follow constant geomagnetic courses more or less closely than constant geographical courses. We analysed the routes taken over land of 45 individual raptors tracked by satellite-based radiotelemetry: 25 peregrine falcons, Falco peregrinus, on autumn migration between North and South America, and seven honey buzzards, Pernis apivorus, and 13 ospreys, Pandion haliaetus, on autumn migration between Europe and Africa. Overall, migration directions showed a better agreement with constant geographical than constant geomagnetic courses. Tracks deviated significantly from constant geomagnetic courses, but were not significantly different from geographical courses. After we removed movements directed far from the mean direction, which may not be migratory movements, migration directions still showed a better agreement with constant geographical than constant geomagnetic courses, but the directions of honey buzzards and ospreys were not significantly different from constant geomagnetic courses either. That migration routes of raptors followed by satellite telemetry are in closer accordance with constant geographical compass courses than with constant geomagnetic compass courses may indicate that geographical (e.g. based on celestial cues) rather than magnetic compass mechanisms are of dominating importance for the birds' long-distance orientation.

Animal Behaviour

Effects of geolocation archival tags on reproduction and adult body mass of sooty shearwaters (Puffinus griseus)

We attached 11 g (1.4% body-mass equivalent) global location sensing (GLS) archival tag packages to tarsi of 25 breeding sooty shearwaters (Puffinus griseus, titi) on Whenua Hou (Codfish Island), New Zealand during the chick-rearing period in 2005. Compared with chicks reared by non-handled adults that did not carry tags, deployment of tags on one or both adult parents ultimately resulted in 35% reduction in chick body mass and significantly reduced chick skeletal size preceding fledging (19 April). However, body mass between chick groups was not significantly different after controlling for skeletal size. Effects on chicks were more pronounced in six pairs where both parents carried tags. Chick mass was negatively related to the duration that adults carried tags. In this study, none of the chicks reared by pairs where both parents were tagged, 54% of chicks reared by pairs where one parent was tagged, and 83% of chicks reared by non-handled and non-tagged parents achieved a previously determined pre-fledging mass threshold (564 g; Sagar & Horning 1998). Body mass of adults carrying tags and returning from transequatorial migration the following year were 4% lighter on average than non-tagged birds, but this difference was not statistically significant. Reduced mass among chicks reared by adults carrying tags during the chick-provisioning period indicated that adults altered "normal" provisioning behaviours to maintain their own body condition at the expense of their chicks. Population-level information derived from telemetry studies can reveal important habitat-linked behaviours, unique aspects of seabird foraging behaviours, and migration ecology. Information for some species (e.g., overlap with fisheries) can aid conservation and marine ecosystem management. We advise caution, however, when interpreting certain data related to adult provisioning behaviours (e.g., time spent foraging, provisioning rates, etc.). If effects on individuals are of concern, we suggest shorter-term deployments, smaller and lighter tags, and alternative attachment techniques, especially when investigating threatened or endangered species. ?? The Royal Society of New Zealand 2009.

New Zealand Journal of Ecology

Phylogeny and species traits predict bird detectability

Avian acoustic communication has resulted from evolutionary pressures and ecological constraints. We therefore expect that auditory detectability in birds might be predictable by species traits and phylogenetic relatedness. We evaluated the relationship between phylogeny, species traits, and field‐based estimates of the two processes that determine species detectability (singing rate and detection distance) for 141 bird species breeding in boreal North America. We used phylogenetic mixed models and cross‐validation to compare the relative merits of using trait data only, phylogeny only, or the combination of both to predict detectability. We found a strong phylogenetic signal in both singing rates and detection distances; however the strength of phylogenetic effects was less than expected under Brownian motion evolution. The evolution of behavioural traits that determine singing rates was found to be more labile, leaving more room for species to evolve independently, whereas detection distance was mostly determined by anatomy (i.e. body size) and thus the laws of physics. Our findings can help in disentangling how complex ecological and evolutionary mechanisms have shaped different aspects of detectability in boreal birds. Such information can greatly inform single‐ and multi‐species models but more work is required to better understand how to best correct possible biases in phylogenetic diversity and other community metrics.

Ecography

Mate desertion in the snail kite

Mate desertion during the breeding cycle was documented at 28 of 36 (78%) snail kite, Rostrhamus sociabilis nests in Florida between 1979 and 1983. Offspring mortality occurred at only one deserted nest, however. Parents that were deserted by their mates continued to care for their young until independence (3?5 additional weeks) and provided snails at a rate similar to that of both parents combined before desertion. Males and females deserted with nearly equal frequency, except in 1982 when more females deserted. No desertion occurred during drought years, whereas desertion occurred at nearly every nest during favourable conditions. The occurrence of mate desertion was generally related to indirect measures of snail abundance: foraging range, snail delivery rates to the young and growth rates. Small broods were deserted more frequently by females than by males and tended to be deserted earlier than large ones. After desertion, deserters had the opportunity to re-mate and nest again since breeding seasons were commonly lengthy, but whether they did so was impossible to determine conclusively in most cases. The deserted bird sometimes incurred increased energetic costs and lost breeding opportunities during periods of monoparental care.

Animal Behaviour

Do parents synchronise nest visits as an antipredator adaptation in birds of New Zealand and Tasmania?

Birds with altricial offspring need to feed them regularly, but each feeding visit risks drawing attention to the nest and revealing its location to potential predators. Synchronisation of visits by both parents has been suggested as a behavioural adaptation to reduce the risk of nest predation. Under this hypothesis, higher risk of nest predation favours greater synchrony of parental feeding visits. We investigated this prediction over three timescales using nestling provisioning data from 25 passerine species in Tasmania and New Zealand. We estimated the extent to which parents actively synchronised their visits to the nest by comparing observed patterns of synchrony with those expected to occur at random. We found that in general, species did not synchronise visits more often than expected by chance. Species varied in the tendency to synchronise visits, but this variation was not explained by likely predation pressure in the distant evolutionary past: New Zealand endemic species, which evolved in the absence of mammalian nest predators, synchronised their visits as often as species which evolved with more diverse predatory guilds. Nest predation risk has increased over time in New Zealand due to introduced predators, but synchrony in visits also was not explained by manipulated predation risk: visit synchrony was equivalent between a predator-removal site and a site where predators remained. However, within one New Zealand species, visit synchrony was higher for mainland populations, which have been exposed to predatory mammals for c .800 years, than for a population on an offshore island to which predatory mammals were never introduced. We conclude that breeding birds may have some capacity to adapt the synchrony with which they provision over short evolutionary timescales. However, the lack of synchrony in most species suggests that either asynchrony provides benefits that outweigh the greater risk of predation, or synchrony incurs costs not compensated by reduced predation.

Tasmania

Nest predation research: Recent findings and future perspectives

Nest predation is a key source of selection for birds that has attracted increasing attention from ornithologists. The inclusion of new concepts applicable to nest predation that stem from social information, eavesdropping or physiology has expanded our knowledge considerably. Recent methodological advancements now allow focus on all three players within nest predation interactions: adults, offspring and predators. Indeed, the study of nest predation now forms a vital part of avian research in several fields, including animal behaviour, population ecology, evolution and conservation biology. However, within nest predation research there are important aspects that require further development, such as the comparison between ecological and evolutionary antipredator responses, and the role of anthropogenic change. We hope this review of recent findings and the presentation of new research avenues will encourage researchers to study this important and interesting selective pressure, and ultimately will help us to better understand the biology of birds.

Journal of Ornithology

Central-West Siberian-breeding Bar-tailed Godwits (Limosa lapponica) segregate in two morphologically distinct flyway populations

Long-distance migratory species often include multiple breeding populations, with distinct migration routes, wintering areas and annual-cycle timing. Detailed knowledge on population structure and migratory connectivity provides the basis for studies on the evolution of migration strategies and for species conservation. Currently, five subspecies of Bar-tailed Godwits Limosa lapponica have been described. However, with two apparently separate breeding and wintering areas, the taxonomic status of the subspecies L. l . taymyrensis remains unclear. Here we compare taymyrensis Bar-tailed Godwits wintering in the Middle East and West Africa, respectively, with respect to migration behaviour, breeding area, morphology and population genetic differentation in mitochondrial DNA. By tracking 52 individuals from wintering and staging areas over multiple years, we show that Bar-tailed Godwits wintering in the Middle East bred on the northern West-Siberian Plain ( n = 19), whilst birds from West Africa bred further east, mostly on the Taimyr Peninsula ( n = 12). The two groups differed significantly in body size and shape, and also in the timing of both northward and southward migrations. However, they were not genetically differentiated, indicating that the phenotypic (i.e. geographic, morphological and phenological) differences arose either very recently or without current reproductive isolation. We conclude that the taymyrensis taxon consists of two distinct populations with mostly non-overlapping flyways, which warrant treatment as separate taxonomic units. We thus propose to distinguish a more narrowly defined taymyrensis subspecies (i.e. the Bar-tailed Godwits wintering in West Africa and breeding on Taimyr), from a new yamalensis subspecies (i.e. the birds wintering in the Middle East and breeding on the northern West-Siberian Plain).

Ibis

Potential threats facing a globally important population of the magnificent frigatebird Fregata magnificens

Tracking of seabirds has been used to identify foraging hotspots, migratory routes and to assess at-sea threats facing populations. One such threat is the potential negative interaction between seabirds and fisheries through incidental by-catch. In 2012, 60 magnificent frigatebirds Fregata magnificens were found dead, entangled in fishing line, at the globally important breeding site in the British Virgin Islands (BVI). To assess the potential relationship between foraging behaviour and fishing activity, data loggers were deployed on breeding magnificent frigatebirds to record foraging movements. In addition, a survey of local fishers was conducted to assess the scale of incidental by-catch. We recorded 28 complete foraging trips from GPS and GPS-GSM loggers, and 1758 PTT locations. Birds travelled up to 3.3–1067 km from their breeding colony and entered the waters of 10 neighbouring territories. A high percentage of fishers (93% n = 28) reported catching at least one seabird annually, of which the most common were magnificent frigatebirds and brown boobies Sula leucogaster . There are estimated to be at least 1112 vessels in the recreational and artisanal fishing fleets of BVI and its neighbouring islands. Thus, this substantial fishery may have potentially profound effects on seabird populations in the region.

Tropical Zoology

Global positioning system and associated technologies in animal behaviour and ecological research

Biologists can equip animals with global positioning system (GPS) technology to obtain accurate (less than or equal to 30 m) locations that can be combined with sensor data to study animal behaviour and ecology. We provide the background of GPS techniques that have been used to gather data for wildlife studies. We review how GPS has been integrated into functional systems with data storage, data transfer, power supplies, packaging and sensor technologies to collect temperature, activity, proximity and mortality data from terrestrial species and birds. GPS 'rapid fixing' technologies combined with sensors provide location, dive frequency and duration profiles, and underwater acoustic information for the study of marine species. We examine how these rapid fixing technologies may be applied to terrestrial and avian applications. We discuss positional data quality and the capability for high-frequency sampling associated with GPS locations. We present alternatives for storing and retrieving data by using dataloggers (biologging), radio-frequency download systems (e.g. very high frequency, spread spectrum), integration of GPS with other satellite systems (e.g. Argos, Globalstar) and potential new data recovery technologies (e.g. network nodes). GPS is one component among many rapidly evolving technologies. Therefore, we recommend that users and suppliers interact to ensure the availability of appropriate equipment to meet animal research objectives.

Philosophical Transactions of the Royal Society B:

Within-group relatedness and patterns of reproductive sharing and cooperation in the tropical chestnut-crested yuhina

In cooperatively breeding animals, genetic relatedness among group members often determines the extent of reproductive sharing, cooperation and competition within a group. Studies of species for which cooperative behaviour is not entirely based on kinship are key for understanding the benefits favouring the evolution and maintenance of cooperative breeding among nonrelatives. In the cooperatively breeding chestnut-crested yuhina, Yuhina everetti , a songbird endemic to Borneo, we tested whether unrelated helpers are more likely to gain parentage than are related helpers consistent with the hypothesis that inbreeding risk constrains reproduction by related helpers. We also examined whether related or unrelated helpers provision broods more because of differences in their potential indirect or direct fitness benefits of helping. Kin structure of breeding groups (breeding pair and up to eight helpers of both sexes, median = 2 helpers, 96% of 57 pairs had helpers) based on genetic analysis was mixed; 48% of 76 breeder/helper dyads were first-order (26%) or second-order (22%) relatives of one or both members of the breeding pair, and 52% were nonrelatives. Only unrelated male and female helpers gained parentage, and helpers did not differ in their provisioning rate according to their relatedness to the broods. We documented quasi-parasitism or co-breeding by female helpers in 14% of 29 broods and extrapair paternity by male helpers in 21% of 47 broods. This rate of extrapair paternity is relatively high among the few tropical species examined but fit with predictions for mixed-kin groups where inbreeding is avoided. These findings support the emerging pattern for cooperative breeding in birds with mixed-kin groups, wherein unrelated helpers are more likely to gain parentage than are related helpers and helping effort is not necessarily predicted by kinship.

Animal Behaviour

Comparative life history of the south temperate Cape Penduline Tit (Anthoscopus minutus) and north temperate Remizidae species

We studied the breeding biology of the south temperate Cape Penduline Tit ( Anthoscopus minutus ) in order to compare its life history traits with those of related north temperate members of the family Remizidae, namely the Eurasian Penduline Tit ( Remiz pendulinus ) and the Verdin ( Auriparus flaviceps ). We used this comparison to test key predictions of three hypotheses thought to explain latitudinal variation in life histories among bird species—the seasonality and food limitation hypothesis, nest predation hypothesis and adult mortality hypothesis. Contrary to the general pattern of smaller clutch size and lower adult mortality among south-temperate birds living in less seasonal environments, the Cape Penduline Tit has a clutch size larger than that of the Verdin and similar to that of the Eurasian Penduline Tit, and higher adult mortality than both of the other two species. The most notable difference between the Cape Penduline Tit and the two other species is in parental behavioural strategy, with the former exhibiting bi-parental care at all stages of nesting together with facultative cooperative breeding, whereas the Eurasian Penduline Tit has uni-parental care and the Verdin has a combination of female-only incubation but bi-parental nestling care. Consequently, in comparison to the other two species, the Cape Penduline Tit exhibits greater nest attentiveness during incubation, a similar per-nestling feeding rate and greater post-fledging survival. Its relatively large clutch size, high parental investment and associated high adult mortality in a less seasonal environment are consistent with key predictions of the adult mortality hypothesis but not with key predictions of the seasonality and food limitation hypothesis in explaining life history variation among Remizidae species. These results add to a growing body of evidence of the importance of age-specific mortality in shaping life history evolution.

Journal of Ornithology

Energetic solutions of Rock Sandpipers to harsh winter conditions rely on prey quality

Rock Sandpipers Calidris ptilocnemis have the most northerly non-breeding distribution of any shorebird in the Pacific Basin (upper Cook Inlet, Alaska; 61°N, 151°W). In terms of freezing temperatures, persistent winds and pervasive ice, this site is the harshest used by shorebirds during winter. We integrated physiological, metabolic, behavioural and environmental aspects of the non-breeding ecology of Rock Sandpipers at the northern extent of their range to determine the relative importance of these factors in facilitating their unique non-breeding ecology. Not surprisingly, estimated daily energetic demands were greatest during January, the coldest period of winter. These estimates were greatest for foraging birds, and exceeded basal metabolic rates by a factor of 6.5, a scope of increase that approaches the maximum sustained rate of energetic output by shorebirds during periods of migration, but far exceeds these periods in duration. We assessed the quality of their primary prey, the bivalve Macoma balthica , to determine the daily foraging duration required by Rock Sandpipers to satisfy such energetic demands. Based on size-specific estimates of M. balthica quality, Rock Sandpipers require over 13 h/day of foraging time in upper Cook Inlet in January, even when feeding on the highest quality prey. This range approaches the average daily duration of mudflat availability in this region ( c . 18 h), a maximum value that annually decreases due to the accumulation of shore-fast ice. Rock Sandpipers are likely to maximize access to foraging sites by following the exposure of ice-free mudflats across the upper Cook Inlet region and by selecting smaller, higher quality M. balthica to minimize foraging times. Ultimately, this unusual non-breeding ecology relies on the high quality of their prey resources. Compared with other sites across their range, M. balthica from upper Cook Inlet have relatively light shells, potentially the result of the region's depauperate invertebrate predator community. Given the delicate balance between environmental and prey conditions that currently make Cook Inlet a viable wintering area for Rock Sandpipers, small variations in these variables may affect the suitability of the site in the future.

Alaska

Fifty-fifth Christmas Bird Count. 159. Ocean City, Md

The distribution of dynamic pressure behind a Harris' hawk's wing was sampled using a wake rake consisting of 15 pitot tubes and one static tube. The hawk was holding on to a perch, but at an air speed and gliding angle at which it was capable of gliding. The perch was instrumented, so that the lift developed by the wing was known and the lift coefficient could be calculated. The mean of 92 estimates of profile drag coefficient was 0.0207, with standard deviation 0.0079. Lift coefficients ranged from 0.51 to 1.08. Reynolds numbers were nearly all in the range 143000-194000. The estimates of profile drag coefficient were reconcilable with previous estimates of the wing profile drag of the same bird, obtained by the subtractive method, and also with values predicted by the `Airfoil-ii? program for designing aerofoils, based on a digitized wing profile from the ulnar region of the wing. The thickness of the wake suggested that the boundary layer was mostly or fully turbulent in most observations and separated in some, possibly as an active means of creating drag for control purposes. It appears that the bird could momentarily either increase or decrease the profile drag of specific parts of the wing, by active changes of shape, and it appeared to use the carpo-metacarpal region especially for such control movements. Further investigation in a low turbulence wind tunnel would help to resolve doubts about the possible influence of airstream turbulence on the behaviour of the boundary layer.

Journal of Experimental Biology

Fifty-fifth Christmas Bird Count. 159. Ocean City, Md

The distribution of dynamic pressure behind a Harris' hawk's wing was sampled using a wake rake consisting of 15 pitot tubes and one static tube. The hawk was holding on to a perch, but at an air speed and gliding angle at which it was capable of gliding. The perch was instrumented, so that the lift developed by the wing was known and the lift coefficient could be calculated. The mean of 92 estimates of profile drag coefficient was 0.0207, with standard deviation 0.0079. Lift coefficients ranged from 0.51 to 1.08. Reynolds numbers were nearly all in the range 143000-194000. The estimates of profile drag coefficient were reconcilable with previous estimates of the wing profile drag of the same bird, obtained by the subtractive method, and also with values predicted by the ?Airfoil-ii? program for designing aerofoils, based on a digitized wing profile from the ulnar region of the wing. The thickness of the wake suggested that the boundary layer was mostly or fully turbulent in most observations and separated in some, possibly as an active means of creating drag for control purposes. It appears that the bird could momentarily either increase or decrease the profile drag of specific parts of the wing, by active changes of shape, and it appeared to use the carpo-metacarpal region especially for such control movements. Further investigation in a low turbulence wind tunnel would help to resolve doubts about the possible influence of airstream turbulence on the behaviour of the boundary layer.

Audubon Field Notes

Pacific Barrow’s Goldeneye refine migratory phenology in response to overwintering temperatures and annual snow melt

Timing of seasonal bird migrations is broadly determined by internal biological clocks, which are synchronized by photoperiod, but individuals often refine their migratory timing decisions in response to external factors. Using 11 years of satellite telemetry data, we show that Pacific Barrow’s Goldeneye ( Bucephala islandica ), at higher latitudes, initiated spring and molt migrations later and fall migration earlier than individuals at lower latitudes. We further show that individuals refined migratory timing in response to interannual variation in environmental conditions. Individual Barrow’s Goldeneye initiated spring migration earlier in years with warmer springs at their overwintering locations and concluded spring migration earlier in years with earlier annual snow melt on their breeding grounds. Because individuals respond to conditions both where they initiate and where they conclude spring migration, our results suggest that Barrow’s Goldeneye update their migratory decisions en route. For all three migrations in their annual cycle, birds delayed initiating migration if they had been captured and tagged prior to that migration. Birds that initiated migration late for their latitude were less likely to include a stopover and completed that migration faster, partially compensating for delayed departures. Our results are consistent with the hypothesis that Barrow’s Goldeneye use a combination of endogenous cues and environmental cues in migratory decision making. Sensitivity to environmental cues suggests that Barrow’s Goldeneye may have behavioural plasticity that is adaptive when faced with ongoing climate change.

Alaska, Alberta, British Columbia