Improving the accuracy of population size estimates for burrow‐nesting seabirds

Seabird numbers can change rapidly as a result of environmental processes, both natural and anthropogenic. Informed management and conservation of seabirds requires accurate and precise monitoring of population size. However, for burrow‐nesting species this is rarely achieved due to spatial and temporal heterogeneity in burrow occupancy. Here, we describe a novel method for deriving more accurate population size estimates that employs mark‐recapture methods to correct for unknown variation in nest occupancy throughout a breeding season. We apply it to estimate breeding numbers of a colonial, burrowing seabird, the Little Penguin Eudyptula minor, on the Summerland Peninsula, Phillip Island, Australia. Estimates of active burrow numbers during the September 2008 to February 2009 breeding season were adjusted to numbers of breeding birds based on burrow occupation and modelled population demographics at six, fortnightly monitored reference sites. The population was estimated to be 26 100 (95% CI: 21 100–31 100) and 28 400 (23 800–33 000) breeding Penguins in two temporally separated surveys within one breeding season. We demonstrate using simulation that the method is robust to variation in burrow occupancy throughout the breeding season, providing consistent and more accurate estimates of population size. The advantage of using the corrected method is that confidence intervals will include the true population size. Confidence limits widened as burrow occupancy declined, reflecting the increased uncertainty as larger adjustments for low burrow occupancy were required. In contrast, the uncorrected method that uses burrow occupancy alone as a measure of breeding numbers was inconsistent and significantly underestimated population size across much of the breeding season. Although requiring considerably more survey effort, the corrected approach provides a more accurate means for monitoring population changes in colonially breeding animals while collecting demographic data that can help diagnose the drivers of population change.     

Corvids congregate to breeding colonies of a burrow‐nesting seabird

Egg predation is a major cause of reproductive failure among birds, and can compromise the viability of affected populations. Some egg predators aggregate near colonially breeding birds to exploit the seasonal increase of prey resources. We investigated spatial and temporal variations in the abundance of an egg predator (little raven Corvus mellori; Corvidae) to identify whether ravens aggregate spatially or temporally to coincide with any of three potential prey species: burrow‐nesting little penguin (Eudyptula minor; Spheniscidae), short‐tailed shearwater (Ardenna tenuirostris; Procellariidae), and surface‐nesting silver gull (Chroicocephalus novaehollandiae; Laridae). We derived spatially explicit density estimates of little ravens using distance sampling along line transects throughout a calendar year, which encompassed little penguin, short‐tailed shearwater and silver gull breeding and non‐breeding seasons. High raven abundance coincided temporally with penguin and gull egg laying periods but not with that of shearwaters. The spatial distribution of raven density corresponded with the little penguin colony but not with shearwater or gull colonies. Thus, the presence of little penguin eggs in burrows correlated strongly with little raven activity, and this implies that little ravens may have learnt to exploit the plentiful subsurface food resource of little penguin eggs. Corvid management may be required to maintain the viability of this socially and economically important penguin colony.     

Preparation for flight: pre‐fledging exercise time is correlated with growth and fledging age in burrow‐nesting seabirds

Chicks of many burrow‐nesting seabirds are known to repeatedly emerge from their nests (these trips being termed ‘excursions’) and exercise their wings prior to fledging, but this behavior is poorly documented in the literature, and thus the relationship between growth and exercise remains unclear. Here, we used infrared video cameras placed in front of streaked shearwater Calonectris leucomelas nests during the chick‐emergence period to examine correlations between chick excursions and parameters known to be important for juvenile survival after fledging. In addition, we also attached acceleration‐temperature loggers to several chicks in order to evaluate the relationship between excursion time and time spent exercising the wing muscles (i.e. flapping). Chicks that undertook longer excursions exhibited more rapid increases in wing length and larger body masses at fledging, and also fledged earlier. Correlations between fitness‐related parameters and excursion time indicate that excursions during the emergence period might offer insights into the various relationships between growth and behavior and/or the mechanisms underlying offspring survival following fledging.     

On the statistical mechanics of species abundance distributions

A central issue in ecology is that of the factors determining the relative abundance of species within a natural community. The proper application of the principles of statistical physics to species abundance distributions (SADs) shows that simple ecological properties could account for the near universal features observed. These properties are (i) a limit on the number of individuals in an ecological guild and (ii) per capita birth and death rates. They underpin the neutral theory of Hubbell (2001), the master equation approach of  [Volkov et?al., 2003] and [Volkov et?al., 2005] and the idiosyncratic (extreme niche) theory of Pueyo et al. (2007); they result in an underlying log series SAD, regardless of neutral or niche dynamics. The success of statistical mechanics in this application implies that communities are in dynamic equilibrium and hence that niches must be flexible and that temporal fluctuations on all sorts of scales are likely to be important in community structure.     

Testing the use of infra‐red video cameras to census a nocturnal burrow‐nesting seabird,the European Storm Petrel Hydrobates pelagicus

Accurate population estimates are important for monitoring the conservation status of species, but nocturnal burrow‐nesting seabirds are notoriously difficult to count. Storm petrel species (Hydrobatidae) are particularly challenging, as burrow entrances are indistinct and response rates to playback low and variable, leading to imprecise population estimates and poor ability to detect trends. Playback is also labour‐intensive, requiring several days of calibration trials to determine site‐ and year‐specific response rates. To test the viability of using infra‐red digital video technology to census storm petrels, we conducted a trial on European Storm Petrels Hydrobates pelagicus at their largest UK colony, Mousa (Shetland). Hourly activity rates recorded from infra‐red filming accurately predicted the number of Apparently Occupied Sites (AOS), but this relationship was stronger in natural habitats (boulder‐beach, rocks and scree) than in walls, where between‐night variation was high due to poor and variable image quality, as well as the flight behaviour of Storm Petrels around vertical structures. Few filming attempts failed completely, but image quality was poor on dark nights with rain or fog, and deteriorated during the season as night‐length increased. Playback accurately predicted AOS density in natural habitats, but overestimated total AOS in walls by 38%. Simulations suggest that for the same fieldwork sampling effort, infra‐red filming will generate slightly more precise AOS estimates compared with playback. Better illumination using infra‐red floodlights would increase detection rates on dark nights, probably resulting in greater precision, but the large amount of expensive equipment and reviewing time required currently make infra‐red filming costly and inefficient relative to playback. However, we recommend its use at sites that cannot otherwise be surveyed safely, or where disturbance is a concern.     

Estimating the abundance of Long‐billed Curlews in Nebraska

ABSTRACT Long‐billed Curlews (Numenius americanus) are shorebirds of high conservation concern in North America. Populations have declined in the last 150 yr primarily due to habitat loss and conversion. We conducted a 2‐yr study to estimate the density and statewide abundance of breeding Long‐billed Curlews in Nebraska during 2008 and 2009. Surveys were conducted during the prenesting period in April when Long‐billed Curlews were likely to be detected. We used a simple random sample of roadside survey routes (N= 39), each consisting of 40 5‐min point‐counts at 800‐m intervals. We modeled detection probability and found that wind speed negatively affected detectability, but found no evidence of either year effects or observer bias. We estimated there were 0.0038 Long‐billed Curlews per hectare (0.38 Long‐billed Curlews per km²) along survey routes and, by extrapolation, estimated there were 23,909 (SE = 1685; 95% CI: 20,810—27,471) Long‐billed Curlews in Nebraska. Our population estimate suggests that ～15 to 22% of the United States population of Long‐billed Curlews is found in Nebraska. Curlews were not evenly distributed within our survey area, with the highest densities in the central Sandhills, an area dominated by grass‐stabilized sand dunes and shallow wetlands, and the lowest densities in predominantly agricultural areas. Because Long‐billed Curlews in Nebraska face many potential threats resulting from land‐use changes, monitoring of the breeding population should be continued. Our survey method was efficient and yielded interpretable results; other states or regions should consider adopting this approach for estimating curlew abundance.     

Low incubation investment in the burrow‐nesting Crab Plover Dromas ardeola permits extended foraging on a tidal food resource

We used GPS data‐loggers, video‐recordings and dummy eggs to assess whether foraging needs may force the low incubation attentiveness (< 55%) of the Crab Plover Dromas ardeola, a crab‐eating wader of the Indian Ocean that nests colonially in burrows. The tidal cycle was the major determinant of the time budget and some foraging trips were more distant from the colony than previously known (up to 26 km away and lasting up to 45 h). The longest trips were mostly made by off‐duty parents, but on‐duty parents also frequently left the nest unattended while foraging for 1–7 h. However, the time spent at the colony area (47%) and the time spent roosting on the foraging grounds (16%) would have allowed almost continuous incubation, as in other species with shared incubation. Therefore, the low incubation attentiveness is not explained by the need for long foraging trips but is largely dependent on a high intermittent rhythm of incubation with many short recesses (5.8 ± 2.6 recesses/h) that were not spent foraging but just outside the burrow or thermoregulating at the seashore. As a result, the eggs were warmed on average only 1.7 °C above burrow temperature, slightly more during high tide periods and when burrow temperature was lower between 20:00 and 10:00 h, only partly counteracting the temperature fluctuations of the incubation chamber. These results suggest that low incubation attentiveness is due to the favourable thermal conditions provided by safe nesting burrows and by the hot tropical breeding season, a combination that allows simultaneous foraging by parents and the exploitation of distant foraging grounds. Why Crab Plovers engage in many short recesses from incubation still remains to be clarified but the need to thermoregulate at the seashore and to watch for predators may play a role.     

Estimating the number of species in a stochastic abundance model

Consider a stochastic abundance model in which the species arrive in the sample according to independent Poisson processes, where the abundance parameters of the processes follow a gamma distribution. We propose a new estimator of the number of species for this model. The estimator takes the form of the number of duplicated species (i.e., species represented by two or more individuals) divided by an estimated duplication fraction. The duplication fraction is estimated from all frequencies including singleton information. The new estimator is closely related to the sample coverage estimator presented by Chao and Lee (1992, Journal of the American Statistical Association 87, 210-217). We illustrate the procedure using the Malayan butterfly data discussed by Fisher, Corbet, and Williams (1943, Journal of Animal Ecology 12, 42-58) and a 1989 Christmas Bird Count dataset collected in Florida, U.S.A. Simulation studies show that this estimator compares well with maximum likelihood estimators (i.e., empirical Bayes estimators from the Bayesian viewpoint) for which an iterative numerical procedure is needed and may be infeasible.     

Estimating nests not present at the time of breeding surveys: an important consideration in assessing nesting populations

ABSTRACT.   Counts of nest starts are often used as indicators of the size of avian nesting populations, or of avian productivity. However, the accuracy of single or repeated counts of unmarked nests over time for estimating seasonal numbers of nests may be strongly affected by nest events that fall in between survey dates, or that occurred prior to or after the survey period. Accuracy may also be affected by uncertainty in the interpretation of counts due to overlap between starting and ending dates of asynchronous nests during the intervisit interval. To measure the combined magnitude of these effects on survey accuracy, we overlaid a monthly "survey" regime on known initiation and ending dates of 2055 nests of ciconiiform birds. Assuming all nests present on the date of simulated survey were counted, monthly surveys underestimated the true number of nest starts by 24–64%, depending on species and year. Using a simple model, we also demonstrate that accuracy does not increase much as survey frequency increases, and that significant estimation error can occur over a wide range of nest success values and degrees of asynchrony. We suggest that (1) these biases can be significant for surveys of many kinds of nesting birds including some territorial passerines, (2) this bias cannot be addressed by increasing survey frequency, and (3) the degree of renesting may be of critical interest for inferring breeding population size from nest count data. We suggest three possible approaches for modeling this error.     

Comparisons of abundance of coral-reef fish: Catch and effort surveys vs visual census

Abstract Catch per unit effort (CPUE) in fisheries science and visual counts in marine ecology are widely used to provide estimates of relative abundance. Concurrent use of these techniques therefore offers an opportunity for cross-validation. This study compares CPUE to underwater visual census (UVC) estimates of relative species abundance in a multispecies fishery: coral-reef fish in the Solomon Islands. Multivariate analyses showed large differences between CPUE and UVC estimates of abundance. The families Acanthuridae and Scandae tended to be the primary cause of differences between the techniques when the full assemblage offish was analysed. However, the relationship between CPUE and UVC did not improve when these families were excluded from the data set and the analyses repeated on families (Serranidae, Lutjamdae, Lethrinidae) caught by the predominant gear type, handlining. This result highlights the point that the choice and use of particular methods require careful consideration in conjunction with the nature of factors being investigated. Many problems of sampling are specific to particular methods and some investigations may benefit from a more pluralistic approach.     

Detection probability and abundance estimation of epiphytic lichens based on height‐limited surveys

Questions: What are the detection probabilities of epiphytic crustose lichens on oak (Quercus robur) when only the lowest 2 m of the trunk are surveyed? How does the abundance of lichen species change with height above the ground, and is the change related to tree age? How well can total abundance (0‐6 m) be predicted based on data from the lowest 2 m? Which tree characteristics explain the vertical distribution of the study species? Location: Southeast Sweden. Methods: The occurrence and abundance of eight crustose lichen species were recorded on 35 oaks, 0‐6 m from the ground. Results: The detection probability was high (>96%) for seven out of the eight species. The abundance of six species declined significantly with increasing height. For five species, >69% of the total abundance (0‐6 m) was recorded on the lowest 2 m. The proportion of the total abundance present above 2 m increased significantly with age for three species. Models predicting total abundance based on data from the lowest 2 m and diameter explained >80% of the deviance for all except one species. The vertical distribution of the study species was explained by bark fissure depth, area and cover of macrolichens. Conclusions: For crustose lichens associated with old oaks, surveying only the lowest 2 m of the trunk yields reliable occurrence data and fairly good estimates of total abundance. However, before interpreting data from the lowest 2 m, knowledge of species vertical distribution, and how the distribution changes with tree age is essential.     

Plasticity in moult speed and timing in an arctic‐nesting goose species

Environmental constraints are strong in migratory species that breed in the Arctic. In addition to breeding, Anatidae have to renew all their flight feathers during the short arctic summer. We examine how temporal constraints and climate affect the phenology of flight feather moult in the greater snow goose Chen caerulescens atlantica, a High Arctic nesting species. We used a database of 1412 moulting adult females measured over 15 yr on Bylot Island, Nunavut. Ninth (9th) primary length was used to determine the moult stage and speed of feather growth. We found a positive relationship between median annual hatching and moult initiation dates and the slope did not differ from 1. The interval between hatching and moult initiation was thus rather fixed and geese did not initiate moult earlier when reproductive phenology was delayed. Nonetheless, there was no relationship between median hatching date and the date at which birds regained flight capacity, suggesting that date of end of moult is independent of the reproductive phenology. There was a trend for an increase in the speed of flight feather growth in years with delayed hatching date. This is the most likely mechanism that could explain moult phenology adjustment in this species. Finally, we found a positive relationship between 9th primary length (corrected for inter‐annual variations) and body condition, suggesting a delay in moulting for individuals in poor condition. These results suggest that moult plasticity is primarily governed by variations in feather growth speed. This phenotypic plasticity could be necessary to complete flight feather renewal before the end of the arctic summer, independently of reproductive phenology and spring environmental conditions. Our novel results suggest possible phenological adjustments through moult speed, which was considered constant in geese until now.     

Restoring Sage‐grouse nesting habitat through removal of early successional conifer

Conifer woodlands have expanded into sagebrush (Artemisia spp.) ecosystems and degrade habitat for sagebrush obligate species such as the Greater Sage‐grouse (Centrocercus urophasianus). Conifer management is increasing despite a lack of empirical evidence assessing outcomes to grouse and their habitat. Although assessments of vegetation recovery after conifer removal are common, comparisons of successional trends with habitat guidelines or actual data on habitat used by sage‐grouse is lacking. We assessed impacts of conifer encroachment on vegetation characteristics known to be important for sage‐grouse nesting. Using a controlled repeated measures design, we then evaluated vegetation changes for 3 years after conifer removal. We compared these results to data from 356 local sage‐grouse nests, rangewide nesting habitat estimates, and published habitat guidelines. We measured negative effects of conifer cover on many characteristics important for sage‐grouse nesting habitat including percent cover of forbs, grasses, and shrubs, and species richness of forbs and shrubs. In untreated habitat, herbaceous vegetation cover was slightly below the cover at local nest sites, while shrub cover and sagebrush cover were well below cover at the nest sites. Following conifer removal, we measured increases in herbaceous vegetation, primarily grasses, and sagebrush height. Our results indicate that conifer abundance can decrease habitat suitability for nesting sage‐grouse. Additionally, conifer removal can improve habitat suitability for nesting sage‐grouse within 3 years, and trajectories indicate that the habitat may continue to improve in the near future.     

Decaying trees improve nesting opportunities for cavity‐nesting birds in temperate and boreal forests: A meta‐analysis and implications for retention forestry

Many studies have dealt with the habitat requirements of cavity‐nesting birds, but there is no meta‐analysis on the subject and individual study results remain vague or contradictory. We conducted a meta‐analysis to increase the available evidence for nest‐site selection of cavity‐nesting birds. Literature was searched in Web of Science and Google Scholar and included studies that provide data on the habitat requirements of cavity‐nesting birds in temperate and boreal forests of varying naturalness. To compare nest and non‐nest‐tree characteristics, the following data were collected from the literature: diameter at breast height (DBH) and its standard deviation (SD), sample size of trees with and without active nest, amount of nest and available trees described as dead or with a broken crown, and amount of nest and available trees that were lacking these characteristics. Further collected data included bird species nesting in the cavities and nest‐building type (nonexcavator/excavator), forest type (coniferous/deciduous/mixed), biome (temperate/boreal), and naturalness (managed/natural). From these data, three effect sizes were calculated that describe potential nest trees in terms of DBH, vital status (dead/alive), and crown status (broken/intact). These tree characteristics can be easily recognized by foresters. The results show that on average large‐diameter trees, dead trees, and trees with broken crowns were selected for nesting. The magnitude of this effect varied depending primarily on bird species and the explanatory variables forest type and naturalness. Biome had lowest influence (indicated by ΔAIC). We conclude that diameter at breast height, vitality, and crown status can be used as tree characteristics for the selection of trees that should be retained in selectively harvested forests.     

Seasonal abundance of livestock‐associated Culicoides species in northeastern Brazil

Culicoides biting midges (Diptera: Ceratopogonidae) are of great medical and veterinary importance because the haematophagous females of some species can transmit diseases to humans and animals. In order to determine the presence and seasonal abundance of the bluetongue virus (BTV) vector Culicoides insignis Lutz at domestic animal sheds in northeastern Brazil, insects were collected once a month between January and December 2010. Light traps were set from 18.00 to 06.00 hours at a pigsty, chicken coop and bovine corral. Culicoides insignis accounted for 81% of the 22 316 specimens collected. Other well‐represented species were: Culicoides paucienfuscatus Barbosa (3246 individuals), Culicoides diabolicus Hoffman (308), Culicoides leopoldoi Ortiz (224) and Culicoides duartei Tavares and Luna Dias (221). The remainder accounted for 4% of the total sample. Culicoides insignis occurred mostly at the cattle corral, 98.2% in the rainy season. This study confirms the presence and close association of C. insignis with cattle in Maranhão state, northeastern Brazil and emphasizes the risk of bluetongue infections spreading in the area.     

A meta‐analysis of species–abundance distributions

The species–abundance distribution (SAD) describes the abundances of all species within a community. Many different models have been proposed to describe observed SADs. Best known are the logseries, the lognormal, and a variety of niche division models. They are most often visualized using either species richness – log abundance class (Preston) plots or abundance – species rank order (Whittaker) plots. Because many of the models predict very similar shapes, model distinction and testing become problematic. However, the variety of models can be classified into three basic types: one that predicts a double S‐shape in Whittaker plots and a unimodal distribution in Preston plots (the lognormal type), a second that lacks the mode in Preston plots (the logseries type), and a third that predicts power functions in both plotting types (the power law type). Despite the interest of ecologists in SADs no formal meta‐analysis of models and plotting types has been undertaken so far. Here we use a compilation of 558 species–abundance distributions from 306 published papers to infer the frequency of the three SAD shapes in dependence of environmental variables and type of plotting. Our results highlight the importance of distinguishing between fully censused and incompletely sampled communities in the study of SADs. We show that completely censused terrestrial or freshwater animal communities tend to follow lognormal type SADs more often than logseries or power law types irrespective of species richness, spatial scale, and geographic position. However, marine communities tend to follow the logseries type, while plant communities tend to follow the power law. In incomplete sets the power law fitted best in Whittaker plots, and the logseries in Preston plots. Finally our study favors the use of Whittaker over Preston plots.