Comparison and Assessment of Aerial and Ground Estimates of Waterbird Colonies

Abstract: Aerial surveys are often used to quantify sizes of waterbird colonies; however, these surveys would benefit from a better understanding of associated biases. We compared estimates of breeding pairs of waterbirds, in colonies across southern Louisiana, USA, made from the ground, fixed-wing aircraft, and a helicopter. We used a marked-subsample method for ground-counting colonies to obtain estimates of error and visibility bias. We made comparisons over 2 sampling periods: 1) surveys conducted on the same colonies using all 3 methods during 3–11 May 2005 and 2) an expanded fixed-wing and ground-survey comparison conducted over 4 periods (May and Jun, 2004–2005). Estimates from fixed-wing aircraft were approximately 65% higher than those from ground counts for overall estimated number of breeding pairs and for both dark and white-plumaged species. The coefficient of determination between estimates based on ground and fixed-wing aircraft was ≤0.40 for most species, and based on the assumption that estimates from the ground were closer to the true count, fixed-wing aerial surveys appeared to overestimate numbers of nesting birds of some species; this bias often increased with the size of the colony. Unlike estimates from fixed-wing aircraft, numbers of nesting pairs made from ground and helicopter surveys were very similar for all species we observed. Ground counts by one observer resulted in underestimated number of breeding pairs by 20% on average. The marked-subsample method provided an estimate of the number of missed nests as well as an estimate of precision. These estimates represent a major advantage of marked-subsample ground counts over aerial methods; however, ground counts are difficult in large or remote colonies. Helicopter surveys and ground counts provide less biased, more precise estimates of breeding pairs than do surveys made from fixed-wing aircraft. We recommend managers employ ground counts using double observers for surveying waterbird colonies when feasible. Fixed-wing aerial surveys may be suitable to determine colony activity and composition of common waterbird species. The most appropriate combination of survey approaches will be based on the need for precise and unbiased estimates, balanced with financial and logistical constraints. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):697–706; 2008)     

A Comparison of Remote Sensing and Ground-Based Methods for Monitoring Wetland Restoration Success

Abstract Efficient and accurate vegetation sampling techniques are essential for the assessment of wetland restoration success. Remotely acquired data, used extensively in many locations, have not been widely used to monitor restored wetlands. We compared three different vegetation sampling techniques to determine the accuracy associated with each method when used to determine species composition and cover in restored Pacific coast wetlands dominated by Salicornia virginica (perennial pickleweed). Two ground‐based techniques, using quadrat and line intercept sampling, and a remote sensing technique, using low altitude, high resolution, color and color infrared photographs, were applied to estimate cover in three small restoration sites. The remote technique provided an accurate and efficient means of sampling vegetation cover, but individual species could not be identified, precluding estimates of species density and distribution. Aerial photography was determined to be an effective tool for vegetation monitoring of simple (i.e., single‐species) habitat types or when species identities are not important (e.g., when vegetation is developing on a new restoration site). The efficiency associated with these vegetation sampling techniques was dependent on the scale of the assessment, with aerial photography more efficient than ground‐based sampling methods for assessing large areas. However, the inability of aerial photography to identify individual species, especially mixed‐species stands common in southern California salt marshes, limits its usefulness for monitoring restoration success. A combination of aerial photography and ground‐based methods may be the most effective means of monitoring the success of large wetland restoration projects.     

Use of time‐lapse photography and digital image analysis to estimate breeding success of a cliff‐nesting seabird

Development of new methods for obtaining basic demographic data from difficult‐to‐access breeding colonies and easily disturbed species is an important challenge in studies of seabirds. We describe a method that can generate data concerning annual breeding success of cliff‐nesting seabirds or other colonial birds with open nests. Our method requires only a single visit to a colony every second or third year, and is based on the use of automated time‐lapse photography. To test our method, we used time‐lapse photos to examine the breeding success of Thick‐billed Murres (Uria lomvia) in two breeding colonies in Greenland during the years 2011, 2012, and 2014. Based on the analysis of time‐lapse photos, we found that hatching success during the 3 yr of our study ranged from 60% to 81%, fledging success from 89% to 95%, and breeding success from 53% to 74% (Table 1). Use of digital image analysis made it possible to differentiate between breeding and non‐breeding birds and determine if and when breeding attempts failed or succeeded. The key to making our method a realistic long‐term monitoring technique is the use of an automated, formal image analysis to process the thousands of photos from the time‐lapse cameras and, more specifically, to reduce a large number of photos to a manageable number. Using our method, we needed 12–22 h per study plot, depending on the number of breeding sites per plot (range = 47–127) and whether it was the first or the second time the plot was analyzed, to obtain our estimates of hatching, fledging, and breeding success. This included time for data preparation, image analyses, visual inspections, and summarizing data in a spreadsheet. We found that our estimates of breeding success were comparable to those obtained by direct observation in the field. An important aspect of using time‐lapse technology is to foresee potential reasons why time‐lapse cameras might stop taking pictures, for example, equipment failure (camera, timer, or battery) or interference by visitors (e.g., vandalism or theft). As such, thorough testing of time‐lapse systems and selecting camera locations less likely to be disturbed are most important. We believe that use of time‐lapse photography in combination with digital image analysis to estimate breeding success can be useful for determining the breeding success of other cliff‐nesting seabirds and, more generally, other birds that breed in colonies, especially those located in remote areas and where direct observation may disturb birds.     

Computer‐automated bird detection and counts in high‐resolution aerial images: a review

Bird surveys conducted using aerial images can be more accurate than those using airborne observers, but can also be more time‐consuming if images must be analyzed manually. Recent advances in digital cameras and image‐analysis software offer unprecedented potential for computer‐automated bird detection and counts in high‐resolution aerial images. We review the literature on this subject and provide an overview of the main image‐analysis techniques. Birds that contrast sharply with image backgrounds (e.g., bright birds on dark ground) are generally the most amenable to automated detection, in some cases requiring only basic image‐analysis software. However, the sophisticated analysis capabilities of modern object‐based image analysis software provide ways to detect birds in more challenging situations based on a variety of attributes including color, size, shape, texture, and spatial context. Some techniques developed to detect mammals may also be applicable to birds, although the prevalent use of aerial thermal‐infrared images for detecting large mammals is of limited applicability to birds because of the low pixel resolution of thermal cameras and the smaller size of birds. However, the increasingly high resolution of true‐color cameras and availability of small unmanned aircraft systems (drones) that can fly at very low altitude now make it feasible to detect even small shorebirds in aerial images. Continued advances in camera and drone technology, in combination with increasingly sophisticated image analysis software, now make it possible for investigators involved in monitoring bird populations to save time and resources by increasing their use of automated bird detection and counts in aerial images. We recommend close collaboration between wildlife‐monitoring practitioners and experts in the fields of remote sensing and computer science to help generate relevant, accessible, and readily applicable computer‐automated aerial photographic census techniques.     

Estimating the relative abundance of emperor penguins at inaccessible colonies using satellite imagery

Emperor penguin (Aptenodytes forsteri) populations are useful environmental indicators due to the bird’s extreme reliance on sea ice. We used remote sensing technology to estimate relative adult bird abundance at two inaccessible emperor penguin colonies in the Ross Sea, Antarctica. We performed supervised classification of 12 panchromatic satellite images of the seven known Ross Sea colonies. We used regression to predict adult bird counts at the inaccessible colonies by relating the number of pixels classified as “penguin” in the satellite images of the accessible colonies to corresponding known adult bird counts from aerial photographs or ground counts. While our analysis was hampered by excessive guano and shadows, we used satellite imagery to differentiate between relatively small (<3,000 adult birds) and larger colonies (>5,000 adult birds). Remote sensing technology is logistically less intense and less costly than aerial or ground censuses when the objective is to document penguin presence and/or large emperor penguin population changes (e.g., catastrophic changes). Improvements expected soon in the resolution of the satellite images should allow for more accurate abundance estimates.     

Bias in aerial estimates of the number of nests in White Ibis and Great Egret colonies

ABSTRACT Estimating detection error, as well as the magnitude of other potential survey biases, is essential when sampling efforts play a role in the estimation of population size and management of wildlife populations. We quantified visual biases in aerial surveys of nesting wading birds (Ciconiiformes) in colonies in the Florida Everglades using a negative binomial count regression model to compare numbers of nests in quadrats counted on the ground with numbers estimated from aerial photographs of the same quadrats. The model also allowed the determination of degree of difference between monitoring results based upon such factors as nest density, vegetative cover, and nest turnover rates. Aerial surveys of White Ibis (Eudocimus albus) colonies underestimated the true number of nests found during ground counts by 11.1%, and underestimates were significantly greater (P= 0.047) in a colony with high nest turnover. Error rates did not differ for quadrats that varied in the density of White Ibis nests did not differ, and visual bias did not increase with vegetative complexity (P= 0.73). Estimates of nest density in colonies of Great Egrets (Ardea alba) based on aerial surveys were higher than ground counts for 38% of the quadrats sampled, and mean visual bias was 23.1%. Species misidentification likely contributed to visibility bias for Great Egrets in our study, with some Snowy Egrets almost certainly mistaken for Great Egrets in aerial photos. Biases of the magnitude we observed fro Great Egrets and White Ibises can mask true population trends in long‐term monitoring and, therefore, we recommend that detection probability be explicitly evaluated when conducting aerial surveys of nesting birds.     

The impacts of introduced House Mice on the breeding success of nesting seabirds on Gough Island

Invasive species are the main threat to island biodiversity; seabirds are particularly vulnerable and are one of the most threatened groups of birds. Gough Island, a UNESCO World Heritage Site in the South Atlantic Ocean, is an Important Bird and Biodiversity Area, and one of the most important seabird colonies globally. Invasive House Mice Mus musculus depredate eggs and chicks of most seabird species on the island, but the extent of their impact has not been quantified. We used field data and bootstrapped normal distributions to estimate breeding success and the number of surviving chicks for 10 seabird species on Gough Island, and compared estimates with those of analogous species from predator‐free islands. We examined the effects of season and nest‐site location on the breeding success of populations on Gough Island, predicting that the breeding success of Gough birds would be lower than that of analogues, particularly among small burrow‐nesting species. We also predicted that winter‐breeding species would exhibit lower breeding success than summer‐breeding species, because mice have fewer alternative food sources in winter; and below‐ground nesters would have lower breeding success than surface nesters, as below‐ground species are smaller so their chicks are easier prey for mice. We did indeed find that seabirds on Gough Island had low breeding success compared with analogues, losing an estimated 1 739 000 (1 467 000–2 116 000) eggs/chicks annually. Seven of the 10 focal species on Gough Island had particularly high chick mortality and may have been subject to intense mouse predation. Below‐ground and winter breeders had lower breeding success than surface‐ and summer‐breeders. MacGillivray's Prion Pachyptila macgillivrayi, Atlantic Petrel Pterodroma incerta and Tristan Albatross Diomedea dabbenena are endemic or near‐endemic to Gough Island and are likely to be driven to extinction if invasive mice are not removed.     

Use of aerial imagery to assess habitat suitability and predict site occupancy for a declining wetland-dependent bird

Remote sensing is a valuable tool for wetland habitat quantification, monitoring and assessment. Here we show that habitat assessment via aerial image inspection is useful in predicting wetland site occupancy by black terns (Chlidonias niger), an imperiled and declining species throughout much of North America. We used Google Earth^® images and National Wetlands Inventory maps to rank 390 candidate wetlands throughout Wisconsin (USA) according to their apparent suitability as nesting habitat for black terns and quantified habitat features associated with the suitability rankings. We then conducted ground-based suitability assessments and point counts of terns at most wetlands from May to July 2010. Pre-survey assessment resulted in 123 wetlands classified as suitable, 81 as marginal, and 186 as unsuitable. Wetlands ranked as suitable were more likely to be in the hemi-marsh stage, part of a wetland complex and relatively undisturbed. Black terns were present at 47 % of the wetlands considered suitable but only 11 % of the sites considered marginal or unsuitable. Of the 42 sites where nesting was confirmed, 79 % were at wetlands classified as suitable; no nesting was recorded in any wetlands deemed unsuitable. We found strong concordance in wetland suitability rankings between the two assessment methods (remote sensing, site surveys). We propose that remote sensing is an efficient and inexpensive way to predict site occupancy by wetland birds, such as black terns, that prefer a specific kind of habitat discernible from aerial imagery. This method may be particularly useful in areas, such as the Prairie Pothole region of North America, where ground surveys of all wetlands are not feasible.     

Large-scale spatial variation in palm fruit abundance across a tropical moist forest estimated from high-resolution aerial photographs

Fruit abundance is a critical factor in ecological studies of tropical forest animals and plants, but difficult to measure at large spatial scales. We tried to estimate spatial variation in fruit abundance on a relatively large spatial scale using low altitude, high‐resolution aerial photography. We measured fruit production for all 555 individuals of the arborescent palm Astrocaryum standleyanum across 25 ha of mapped tropical moist forest on Barro Colorado Island, Panama, by visually counting fruits from the ground. Simultaneously, we used high‐resolution aerial photographs to map sun‐exposed crowns of the palm across the same area, which were then linked to ground‐mapped stems. First, we verified that the fruit crop size of individual trees was positively associated with both crown presence on aerial photos and crown area visible on aerial photos. Then, we determined how well spatial variation in Astrocaryum fruit density across the study area was predicted by spatial densities of photo‐detected crowns and crown area compared to spatial densities of ground‐mapped stems and stem diameters. We found a positive association of fruit crop size with crown visibility on aerial photographs. Although representing just one third of all individuals in the study area, photo‐detected crowns represented 57% of all fruits produced. The spatial pattern of photo‐detected crowns was strongly correlated with the spatial pattern of fruit abundance based on direct fruit counts, and correctly showed the areas with the highest and lowest fruit abundances. The spatial density of photo‐detected crowns predicted spatial variation in fruit abundance equally well as did the spatial density of ground‐mapped stems. Photo‐detected crown area did not yield a better prediction. Our study indicates that remote sensing of crowns can be a reliable and cost‐effective method for estimating spatial variation in fruit abundance across large areas for highly distinctive canopy species. Our study is also among the few to provide empirical evidence for a positive relationship between crown exposure of forest trees and fruit production.     

基于多源遥感数据的大豆叶面积指数估测精度对比

近年来遥感技术的革新促使遥感源越来越丰富.为分析多源遥感数据的叶面积指数(LAI)估测精度,本文以大豆为研究对象,利用比值植被指数(RVI)、归一化植被指数(NDVI)、土壤调整植被指数(SAVI)、差值植被指数(DVI)、三角植被指数(TVI)5种植被指数,结合地面实测LAI构建经验回归模型,比较3类遥感数据(地面高光谱数据、无人机多光谱影像以及高分一号WFV影像)对大豆LAI的估测能力,并从传感器几何位置和光谱响应特性以及像元空间分辨率三方面分析讨论了3类遥感数据的LAI反演差异.结果表明： 地面高光谱数据模型和无人机多光谱数据模型都可以准确预测大豆LAI(在α=0.01显著水平下,R²均>0.69,RMSE均<0.40);地面高光谱RVI对数模型的LAI预测能力优于无人机多光谱NDVI线性模型,但两者差异不大(E_A相差0.3%,R²相差0.04,RMSE相差0.006);高分一号WFV数据模型对研究区内大豆LAI的预测效果不理想(R²<0.30,RMSE>0.70).针对星、机、地三类遥感信息源,地面高光谱数据在反演LAI方面较传统多光谱数据有优势但不突出;16 m空间分辨率的高分一号WFV影像无法满足田块尺度作物长势监测的需求;在保证获得高精度大豆LAI预测值和高工作效率的前提条件下,基于无人机遥感的农情信息获取技术不失为一种最佳试验方案.在当今可用遥感信息源越来越多的情况下,农业无人机遥感信息可成为指导田块精细尺度作物管理的重要依据,为精准农业研究提供更科学准确的信息.     

UAVs as remote sensing platforms in plant ecology: review of applications and challenges

无人机遥感在植物生态学中的应用与挑战 无人机为获取高时空分辨率的遥感数据提供了经济灵活的工具,为植物生态学家开展从个体到区域尺度的生态学研究提供了新的机遇和手段。但作为一种新兴的技术手段,当前无人机遥感在植物生态学中的应用仍充满了挑战,植物生态学的科研需求与无人机遥感的生态应用需要更为深入的融合。本文综述了无人机遥感技术在植物生态学中的应用,展望了无人机在植物生态学研究中的应用前景。在所综述的400篇文献中,59%的文章发表于非植物生态学领域的遥感类期刊,遥感学者与生态学者的关注点存在较大差异。当前的研究集中在无人机遥感的技术层面,如数据处理和遥感反演方法等,对生态学问题本身的关注较少。综述的文献中,61%的研究案例集中在群落尺度,可见光(RGB)相机和多光谱相机是最常用的传感器类型(75%)。无人机遥感数据中蕴藏着诸多待挖掘的、有重要意义的生态参数,这些参数有助于我们识别林窗的几何特征,构建林冠的化学组合,更好地了解林冠表面的不规则性和群落的异质性。无人机遥感技术在植物生态学研究中的深入应用,需要集植物生态学家和遥感专家之合 力共同推进。     

Seevögel im Südpolarmeer

Seabirds face a number of challenges in Antarctic and Subantarctic breeding areas, such as variations in food supply and changes in the ecosystem due to climate change, aggregation and competition from the limited number of suitable breeding areas, and inaccessible breeding areas due to changes in precipitation. Adjustments to irregular food supplies, such as flexible incubation and nesting times, and torpor in chicks can help seabirds keep pace with changes occurring due to climate change. Data loggers are used to better understand the responses of adult birds to changes in the ecosystem and the resulting population trends. Various data loggers are used for a variety of applications, from one gram light loggers for long‐term tracking across large scales to high‐resolution GPS loggers with acquisition of acceleration data that can also be used to model energy expenditure. In order to fully understand the movement ecology of seabirds, several approaches need to be considered: ecological segregation in space, time and choice of food, energy landscapes and the non‐lethal effects of predation (landscapes of fear).     

Choosing the appropriate spatial resolution for monitoring coral bleaching events using remote sensing

Bleached corals provide a strong optical signal that suggests that remote sensing investigations of major bleaching events are feasible using airborne or satellite sensors. However, patchy coral cover, varying intensities of bleaching, and water depths are likely to limit the application of remote sensing techniques in monitoring and mapping coral bleaching. Today, satellite multispectral sensors routinely provide images of reefs from 4 m (Ikonos) to 30 m resolution (Landsat); however, the adequacy of these sensors for monitoring and mapping bleaching events remains unclear. To clarify these issues, scanned aerial photographs acquired during the 1998 bleaching event over the Great Barrier Reef (Australia) were analyzed at various spatial resolutions, from 10 cm to 5 m. We found that the accuracy of mapping bleaching is highly sensitive to spatial resolution. Highest accuracy was obtained at 10 cm resolution for detection of totally bleached colonies. At 1 m resolution, as much as 50% of the 10-cm resolution signal is lost, though the spatial patterns remain correctly described. Partially bleached (pale) corals are difficult to detect even in aerial surveys, leading to an underestimation of overall bleaching levels (total and partial bleaching) in aerial photos compared to in-situ surveys. If data volume and processing time are limiting factors, local variance analysis suggests that the optimal resolution necessary to capture spatial patterns of bleaching is in the range 40-80 cm.     

Coastal saltpans are a good alternative breeding habitat for Kentish plover Charadrius alexandrinus when umbrella species are present

The loss and degradation of natural habitats in coastal areas worldwide has adversely affected many waterbird species, changing their breeding distribution and reducing their productivity. Anthropogenic habitats such as saltpans can provide alternative or complementary habitats for waterbirds and mitigate the increasing human impact on natural coastal habitats. Unvegetated linear paths between salt ponds are used by ground‐nesting waterbird species to breed but their linear structure may facilitate the detection of nests by predators. This negative effect may, however, be counterbalanced by the advantages of breeding in mixed colonies. To evaluate the importance and the risks of breeding in saltpans we used the Kentish plover Charadrius alexandrinus breeding in coastal saltpans of southern Portugal as a model species. Specifically, we assessed the role of nest‐site characteristics, predation and nesting proximity to species with aggressive antipredator behaviour (black‐winged stilt Himantopus himantopus) on their breeding success. Kentish plovers selected nest‐sites on the edges of paths, with about 20% of water around the nest, and a mean visibility of more than 72%; however nest‐site characteristics were not correlated with nesting success. Predation was the main cause of nest loss in saltpans (42%); carrion crows Corvus corone were responsible for most daylight nest predation (58%) and red fox (Vulpes vulpes; 73%) for night predation. An 8‐yr monitoring plan of a Kentish plover population showed a linear increase in their breeding success as their breeding season increasingly overlapped with that of the black‐winged stilt. An experiment with artificial nests showed a significant increase in the number of exposure days (7 to 12) when these nests were within close distance of black‐winged stilt nests. Overall, our results showed that saltpans are an important alternative breeding habitat for the Kentish plover, especially for the maintenance of mixed species colonies.     

Response of breeding European Storm Petrels Hydrobates pelagicus to habitat change

Mainly through trampling and manuring, ground-nesting seabirds induced significant habitat changes both on vegetation cover and soil in one of the largest French colonies of European Storm Petrel Hydrobates pelagicus, Habitat deterioration led to a high level of erosion and the collapse of many former Rabbit Oryctolagus cuniculus burrows previously occupied by breeding Storm Petrels. The loss of burrows accelerated in recent years since Great Cormorants Phalacrocorax carbo bred on the islet with growing numbers. The main consequence of this disturbance was at first shifting of breeding Storm Petrels from burrows to rocky sites, reflecting some behavioral plasticity to buffer environmental variability. But over 18 years, a significant decrease in breeding numbers of Storm Petrels was recorded and attributed to continuous nest site destruction. Thus, other behavioral responses were also suspected, such as temporary non-breeding or emigration of birds that have to find a new nest site. Such a problem of heavy erosion and loss of nesting habitat could induce serious detrimental effects on burrowing seabirds breeding in a limited number of colonies.     

Remote sensing of upland vegetation: the potential of high spatial resolution satellite sensors

Aim Traditional methodologies of mapping vegetation, as carried out by ecologists, consist primarily of field surveying or mapping from aerial photography. Previous applications of satellite imagery for this task (e.g. Landsat TM and SPOT HRV) have been unsuccessful, as such imagery proved to have insufficient spatial resolution for mapping vegetation. This paper reports on a study to assess the capabilities of the recently launched remote sensing satellite sensor Ikonos, with improved capabilities, for mapping and monitoring upland vegetation using traditional image classification methods. Location The location is Northumberland National Park, UK. Methods Traditional remote sensing classification methodologies were applied to the Ikonos data and the outputs compared to ground data sets. This enabled an assessment of the value of the improved spatial resolution of satellite imagery for mapping upland vegetation. Post‐classification methods were applied to remove noise and misclassified pixels and to create maps that were more in keeping with the information requirements of the NNPA for current management processes. Results The approach adopted herein for quick and inexpensive land cover mapping was found to be capable of higher accuracy than achieved with previous approaches, highlighting the benefits of remote sensing for providing land cover maps. Main conclusions Ikonos imagery proved to be a useful tool for mapping upland vegetation across large areas and at fine spatial resolution, providing accuracies comparable to traditional mapping methods of ground surveys and aerial photography.     

中国亚热带4个森林动态监测样地无人机可见光遥感影像数据集

常绿阔叶林是我国亚热带地区的地带性植被类型。由于亚热带森林植物群落垂直结构复杂、林冠郁闭度高, 对常绿阔叶林冠层的研究尚缺乏高质量的监测数据。本数据集包含浙江天童山、浙江百山祖、广东车八岭、广东鼎湖山4个大于20 ha的森林动态监测样地2014年8月或2016年9月采集的无人机可见光遥感影像。本数据集是通过将无人机影像、地面控制点和地面调查数据相结合而获得的。每个样地的数据集包括4个文件: ~5 cm空间分辨率的正射影像图和数字表面模型、1 m空间分辨率的森林冠层高度数据和正射影像质量报告。本数据集可为常绿阔叶林的林冠生态学、生物多样性、生态系统功能等研究提供数据支撑。     

One House Two Families: Petrel Squatters Get a Sniff of Low‐Cost Breeding Opportunities

Burrowing is a widespread nesting behaviour, found in vertebrates and invertebrates. It is particularly common in small procellariiform seabirds such as blue petrels (Halobaena caerulea) and Antarctic prions (Pachyptila desolata), two closely related petrel species. However, digging a burrow is costly and alternative strategies may evolve. Accordingly, blue petrel males can adopt two alternative nesting strategies: digging a new burrow or squatting in an empty one. Importantly, a blue petrel squatter arriving at the colony to breed is more likely to find empty Antarctic prion burrows than empty blue petrel burrows, since the former species only start breeding a month later. However, squatting in a prion’s burrow is risky for blue petrels as the legitimate owner very often returns and claims the burrow back, thus ruining the squatter’s breeding attempt. We present here results of a survey of two sympatric colonies of blue petrels and Antarctic prions on Kerguelen Island. Our data show that blue petrel squatters preferentially occupy blue petrel empty burrows. To investigate potential underlying mechanisms behind this preference, we used a simple Y‐maze design to show that blue petrels can discriminate and prefer their specific odour over the prion odour. Our results confirm the existence of alternative burrowing strategies in blue petrels and suggest that squatters could use olfaction to avoid the less suitable Antarctic prion burrows.     

Mate fidelity and coloniality in waterbirds: a comparative analysis

Increased opportunities for information are one potential benefit of sociality. We apply this idea to the advantages of colonial breeding in bird species that are typically monogamous within a breeding season but often form new pair-bonds in subsequent seasons. Individuals may benefit from nesting in colonies at high density by identifying good-quality potential alternative mates among their neighbours. The opportunities for finding a better mating option are likely to increase with colony size and density. We tested this prediction with a comparative analysis of the association between mate fidelity and coloniality in waterbirds (wading birds and seabirds), where there is wide variation in both the degree of mate retention over consecutive breeding seasons and the degree of coloniality. We used two comparative statistical analyses, one based upon generalized least squares and the other based upon a continuous-time Markov model, to test whether the pattern of association between divorce rate and degree of coloniality was evidence for correlated evolutionary change in the two characters. We found a significant and positive association between divorce rate and the degree of coloniality in waterbirds. The probable ancestral state corresponds to a combination of a high degree of coloniality with no, or weak, mate fidelity. The reconstruction of the historical pattern of character origin and evolution indicates that the transition from a high to a low degree of coloniality occurred before the transition to higher mate fidelity. Received: 5 January 1998 / Accepted: 20 April 1998     

Seabird predation by great skuas Stercorarius skua– intra‐specific competition for food?

Competition for food is widely cited as an important cost of coloniality among birds and much of the evidence in support of this hypothesis comes from studies of colonial piscivorous seabirds. However, for generalist seabirds able to switch between different prey types, the role of food availability in relation to colony size is unclear. Here we investigate patterns of the consumption of seabird prey in relation to colony size in a generalist seabird, the great skua Stercorarius skua, in Shetland, UK. At the population level skuas feed mainly on sandeels Ammodytes marinus and fishery discards, but respond to declines in fish availability to facultatively prey on other seabirds. By comparing the consumption of seabirds among seven different sized colonies, including one colony with artificially reduced numbers of skuas (Fair Isle), we investigate whether consumption of seabird prey is influenced by skua population size, while simultaneously measuring seabird prey availability. Data from five years also enables us to investigate the influence of annual variation in environmental conditions on seabird consumption. Using measures of body condition and reproductive performance we investigate the consequences of living in different sized colonies, which may provide insight into ultimate costs of nesting at high population density. Skua diets varied among colonies and the proportion of seabird prey in the diet was inversely related to skua colony size, despite similar per capita numbers of seabirds across colonies. At the colony where their numbers were artificially suppressed, skuas consumed a greater proportion of seabirds per capita. Highly significant year effects in seabird predation were observed but the pattern among colonies remained consistent over time. Two measures of adult body condition (pectoral muscle index and mean corpuscular volume) revealed that adult great skuas were in poorer condition at the largest colony (Foula), but reproductive performance did not alter significantly among colonies. This study provides evidence that intra‐specific competition among skuas may limit opportunities for obtaining seabird prey, which may be particularly important during periods of poor availability of sandeels and fishery discards, and has implications for assessing the impact of skuas on seabird populations.