Stephen T. Buckland’s research while affiliated with University of St Andrews and other places

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Publications (137)


Figure 1. The relationship between the log number (+1) of Parrot Crossbills against log number (+1) of Common Crossbills at Falsterbo, southern Sweden, from 1973 to 2018 (r = 0.898, n = 46, P < 0.001).
Figure 2. Auto-correlation coefficients at different time lags for national indices for Norway Spruce and Scots Pine cone (seed) production in Sweden from 1910 to 2020, and seed indices for Finland from 1960 to 2000 for Norway Spruce and 1960 to 2004 for Scots Pine. The dashed lines show the critical values where P = 0.05.
Figure 4. Fitted smooth relationships between crossbill numbers at Falsterbo and Norway Spruce and Scots Pine seed indices in the current (top) and previous (bottom) year, as estimated from the HGAM (Table 4). In each subfigure, the two left-hand plots show the fitted smooth relationships between Common (dark lines) and Parrot Crossbills (light lines) and Norway Spruce and Scots Pine indices, with 95% confidence regions in the corresponding dashed lines. The right-hand two interaction plots show the estimated marginal relationships between crossbill numbers and the Norway Spruce seed index, for the cases where the corresponding Scots Pine seed index was low, medium or high.
The association between irruption years by Common and Parrot Crossbills from 1800 to 2008. Numbers in paren- theses are the expected number of years for each irruption type. Yates' corrected χ 2 = 29.9, P < 0.001.
Spearman correlation coefficients for cone indices comparing different conifers and pairs of five regions of Sweden.
Irruptions of crossbills Loxia spp. in northern Europe – patterns and correlations with seed production by key and non‐key conifers
  • Article
  • Full-text available

April 2024

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77 Reads

Ibis

Ron W. Summers

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Jonas Fridman

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Stephen T. Buckland

Irruptions by boreal seed‐eating and frugivorous birds are assumed to be driven by the production of seeds and fruits, crops of which are highly variable between years. Using data from Sweden, we tested whether irruptions of Common Crossbills Loxia curvirostra were correlated with low Norway Spruce Picea abies seed production in the same year as the irruption and/or high seed production in the year prior to an irruption. Similar tests were made for Parrot Crossbill Loxia pytyopsittacus irruptions in relation to Scots Pine Pinus sylvestris seed production. In northern Europe, these conifers represent the key food species of the two crossbill species, respectively. Despite differing times that seeds take to mature and asynchronous seed production between the two conifer species, including a 3‐year cycle for Norway Spruce, the two crossbill species often irrupted in the same year as one another. Analyses showed that irruptions into Britain and other parts of western Europe by both crossbill species were correlated with low seed production by Norway Spruce in Sweden. Low seed production by Scots Pine had a marginally non‐significant additive effect on both crossbill species. In a second set of analyses, the best‐fitting model was one in which low seed production by both conifers in a given year and high seed production in the previous year were each correlated with large numbers of irrupting Common and Parrot Crossbills. The models indicate that the incidental co‐occurrence of low seed production of Norway Spruce and Scots Pine in a given year, after a year of high seed production, may result in an irruption. The seed production of Norway Spruce and Scots Pine in Sweden was correlated with production by the same species in Finland, indicating widespread synchrony of cropping across northern Europe.

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Estimating effective survey duration in camera trap distance sampling surveys

October 2023

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485 Reads

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6 Citations

Among other approaches, camera trap distance sampling (CTDS) is used to estimate animal abundance from unmarked populations. It was formulated for videos and observation distances are measured at predetermined ‘snapshot moments’. Surveys recording still images with passive infrared motion sensors suffer from frequent periods where animals are not photographed, either because of technical delays before the camera can be triggered again (i.e. ‘camera recovery time’) or because they remain stationary and do not immediately retrigger the camera following camera recovery time (i.e. ‘retrigger delays’). These effects need to be considered when calculating temporal survey effort to avoid downwardly biased abundance estimates. Here, we extend the CTDS model for passive infrared motion sensor recording of single images or short photo series. We propose estimating ‘mean time intervals between triggers’ as combined mean camera recovery time and mean retrigger delays from the time interval distribution of pairs of consecutive pictures, using a Gamma and Exponential function, respectively. We apply the approach to survey data on red deer, roe deer and wild boar. Mean time intervals between triggers were very similar when estimated empirically and when derived from the model‐based approach. Depending on truncation times (i.e. the time interval between consecutive pictures beyond which data are discarded) and species, we estimated mean time intervals between retriggers between 8.28 and 15.05 s. Using a predefined snapshot interval, not accounting for these intervals, would lead to underestimated density by up to 96% due to overestimated temporal survey effort. The proposed approach is applicable to any taxa surveyed with camera traps. As programming of cameras to record still images is often preferred over video recording due to reduced consumption of energy and memory, we expect this approach to find broad application, also for other camera trap methods than CTDS.


Accounting for spatial habitat and management boundaries when estimating forest bird population distribution and density: inferences from a soap film smoother

June 2023

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12 Reads

Birds are often obligate to specific habitats which can result in study areas with complex boundaries due to sudden changes in vegetation or other features. This can result in study areas with concave arcs or that include holes of unsuitable habitat such as lakes or agricultural fields. Spatial models used to produce species’ distribution and density estimates need to respect such boundaries to make informed decisions for species conservation and management. The soap film smoother is one model for complex study regions which controls the boundary behaviour, ensuring realistic values at the edges of the region. We apply the soap film smoother to account for boundary effects and compare it with thin plate regression spline (TPRS) smooth and design-based conventional distance sampling methods to produce abundance estimates from point-transect distance sampling collected data on Hawai‘i ‘Ākepa Loxops coccineus in the Hakalau Forest Unit of the Big Island National Wildlife Refuge Complex, Hawai‘i Island, USA. The soap film smoother predicted zero or near zero densities in the northern part of the domain and two hotspots (in the southern and central parts of the domain). Along the boundary the soap film model predicted relatively high densities where ‘Ākepa occur in the adjacent forest and near zero elsewhere. The design-based and soap film abundance estimates were nearly identical. The width of the soap film confidence interval was 16.5% and 0.8% wider than the width of the TPRS smooth and design-based confidence intervals, respectively. The peaks in predicted densities along the boundary indicates leakage by the TPRS smooth. We provide a discussion of the statistical methods, biological findings and management implications of applying soap film smoothers to estimate forest bird population status.


Review of potential line-transect methodologies for estimating abundance of dolphin stocks in the eastern tropical Pacific

January 2023

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244 Reads

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11 Citations

IWC Journal of Cetacean Research and Management

A twelve-year hiatus in fishery-independent marine mammal surveys in the eastern tropical Pacific Ocean (ETP), combined with a mandate to monitor dolphin stock status under international agreements and the need for reliable stock status information to set dolphin bycatch limits in the tuna purse-seine fishery, has renewed debate about how best to assess and monitor ETP dolphin stock status. The high cost of replicating previous ship-based surveys has intensified this debate. In this review, transect methods for estimating animal abundance from dedicated research surveys are considered, with a focus on both contemporary and potential methods suitable for surveying large areas for dolphin species that can form large, multi-species aggregations. Covered in this review are potential improvements to the previous ship-based survey methodology, other ship-based methods, alternative approaches based on high-resolution imagery and passive acoustics, and combinations of ship-based and alternative approaches. It is concluded that for immediate management needs, ship-based surveys, with some suggested modifications to improve precision, are the only reliable option despite their high cost. However, it is recommended that a top research priority should be development of composite methods. Pilot studies on the use of high-resolution imagery and passive acoustics for development of indices of relative abundance to be used in composite methods should be part of any future ship-based survey efforts.


Multi-species population indices for sets of species including rare, disappearing or newly occurring species

July 2022

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221 Reads

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6 Citations

Ecological Indicators

Multi-species indices (MSI) are widely used as ecological indicators and as instruments to inform environmental policies. Many of these indices combine species-specific estimates of relative population sizes using the geometric mean. Because the geometric mean is not defined when values of zero occur, usually only commoner species are included in MSIs and zero values are replaced by a small non-zero value. The latter can exhibit an arbitrary influence on the geometric mean MSI. Here, we show how the compound Poisson and the negative binomial model can be used in such cases to obtain an MSI that has similar features to the geometric mean, including weighting halving and doubling of a species’ population equally. In contrast to the geometric mean, these two statistical models can handle zero values in population sizes and thus accommodate newly occurring and temporarily or permanently disappearing species in the MSI. We compare the MSIs obtained by the two statistical models with the geometric mean MSI and measure sensitivity to changes in evenness and to population trends in rare and abundant species. Additionally, we outline sources of uncertainty and discuss how to measure them. We found that, in contrast to the geometric mean and the negative binomial MSI, the compound Poisson MSI is less sensitive to changes in evenness when total abundance is constant. Further, we found that the compound Poisson model can be influenced more than the other two methods by trends of species showing a low interannual variance. The negative binomial MSI is less sensitive to trends in rare species compared with the other two methods, and similarly sensitive to trends in abundant species as the geometric mean. While the two new MSIs have the advantage that they are not arbitrarily influenced by rare, newly appearing and disappearing species, both do not weight all species equally. We recommend replacing the geometric mean MSI with either compound Poisson or negative binomial when there are species with a population size of zero in some years having a strong influence on the geometric mean MSI. Further, we recommend providing additional information alongside the MSIs. For example, it is particularly important to give an evenness index in addition to the compound Poisson MSI and to indicate the number of disappearing and newly occurring species alongside the negative binomial MSI.




FIG. 1 Study area in the trans-Himalayan region and subalpine region of the Upper Bhagirathi basin, with the locations of camera traps used for estimating densities of the bharal Pseudois nayaur, and Himalayan musk deer Moschus leucogaster, respectively. The inset map shows the location of the Bhagirathi basin in Uttarakhand State, Western Himalaya, India.
FIG. 2 Kernel density estimates of daily activity pattern of the bharal and the musk deer in summer and winter in the Upper Bhagirathi basin.
Using distance sampling with camera traps to estimate the density of group-living and solitary mountain ungulates

April 2021

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686 Reads

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17 Citations

Oryx

Throughout the Himalaya, mountain ungulates are threatened by hunting for meat and body parts, habitat loss, and competition with livestock. Accurate population estimates are important for conservation management but most of the available methods to estimate ungulate densities are difficult to implement in mountainous terrain. Here, we tested the efficacy of the recent extension of the point transect method, using camera traps for estimating density of two mountain ungulates: the group-living Himalayan blue sheep or bharal Pseudois nayaur and the solitary Himalayan musk deer Moschus leucogaster. We deployed camera traps in 2017–2018 for the bharal (summer: 21 locations; winter: 25) in the trans-Himalayan region (3,000–5,000 m) and in 2018–2019 for the musk deer (summer: 30 locations; winter: 28) in subalpine habitats (2,500–3,500 m) in the Upper Bhagirathi basin, Uttarakhand, India. Using distance sampling with camera traps, we estimated the bharal population to be 0.51 ± SE 0.1 individuals/km2 (CV = 0.31) in summer and 0.64 ± SE 0.2 individuals/km2 (CV = 0.37) in winter. For musk deer, the estimated density was 0.4 ± SE 0.1 individuals/km2 (CV = 0.34) in summer and 0.1 ± SE 0.05 individuals/km2 (CV = 0.48) in winter. The high variability in these estimates is probably a result of the topography of the landscape and the biology of the species. We discuss the potential application of distance sampling with camera traps to estimate the density of mountain ungulates in remote and rugged terrain, and the limitations of this method.


Fig. 2. Global portrait of linked changes in fire and biodiversity. Examples of documented and predicted fire-driven changes in biodiversity are shown. Details of the anthropogenic drivers associated with each of these changes are provided in the main text or table S1, following the numbered key. Examples are overlaid on a map of the number of times a fire was recorded from 2000 to 2019 in a given 500 m by 500 m MODIS pixel averaged across the 10 km by 10 km pixels displayed on the map.
Fig. 3. Some tropical ecosystems are experiencing too much fire and others not enough. (A) Frequent fires are a key aspect of African savanna ecosystems that support a large portion of the world's remaining wild large mammals. However, fire activity in the Serengeti-Mara of Tanzania has been reduced, and some areas no longer experience fire. This could increase shrub encroachment (top left; photo by S. Archibald) and the displacement of wild herbivores that prefer open areas (top right; photo by D. D'Auria) (18, 42). (B) The Amazon basin is home to ~10 to 15% of the world's terrestrial biodiversity. In southeast Amazonia (bottom left; photo by P. M. Brando), human drivers increase deforestation fires and uncontrolled fires. This is driving shifts from humid forest to drier forests or derived savannas. Cascading effects on fauna include the decline of forest butterfly species such as the leaf wing butterfly (Zaretis itys) [bottom right; photo by Morales/agefotostock (40)].
Fig. 4. Modeling ecosystems in transition in the Mediterranean Basin. Integrating data on land use, climate change, and fire dynamics [(A); photo by L. Brotons] with empirical bird occurrence data [(B); photo by F. Veronesi, Francesco Veronesi from Italy/CC BY-SA (https://creativecommons.org/licenses/by-sa/2.0)] is helping to predict the impact of social and ecological changes on species distributions. (C) Comparison of management actions showed that the Dartford warbler, an open-country species, will benefit from managed wildfire that creates new open spaces (47). Box plots show the median change in Dartford warbler habitat and the interquartile range from 10 simulations. A2 climate scenarios were associated with a lower number of large wildfires than B2 climate scenarios.
Fig. 5. Pyrodiversity with purpose in temperate forests of the western United States. (A) The Klamath-Siskiyou region is home to Indigenous peoples with different languages and histories. After more than a century of policies that promoted fire suppression, newly developed collaborations led by Indigenous communities and including scientists and local stakeholders are being formed to reinstate Indigenous fire practices. This cultural burning
Fire and biodiversity in the Anthropocene

November 2020

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4,131 Reads

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423 Citations

Science

Fire's growing impacts on ecosystems Fire has played a prominent role in the evolution of biodiversity and is a natural factor shaping many ecological communities. However, the incidence of fire has been exacerbated by human activity, and this is now affecting ecosystems and habitats that have never been fire prone or fire adapted. Kelly et al. review how such changes are already threatening species with extinction and transforming terrestrial ecosystems and discuss the trends causing changes in fire regimes. They also consider actions that could be taken by conservationists and policy-makers to help sustain biodiversity in a time of changing fire activity. Science , this issue p. eabb0355


Study area showing survey points (open circles) separated into north and south strata (polygons). The location (black dot) of Hakalau Forest National Wildlife Refuge is shown on Hawai‘i Island, Hawaiian Islands. Base map from World Geodetic System 1984 (WGS84) zone 5; coastline from U.S. Geological Survey's National Elevation Dataset (USGS, U.S. Geological Survey 2014).
Predicted trends derived from the spatio–temporal GAM for the ‘ākepa population across Hakalau (top panel), north (bottom left panel) and south (bottom right panel) strata of Hakalau. GAM generated density (birds ha⁻¹) estimates (black line) with detection probability variance propagated uncertainty (95% CI gray ribbon). Density estimate with 95% CI from the design‐based method (diamond and whisker bar). Surveys were not conducted in 2009 (vertical bar).
Predicted spatio–temporal surfaces of ‘ākepa densities in Hakalau between 1987 and 2017. Densities range from 0 (dark blue) to 6.5 birds ha⁻¹ (yellow) within the study area.
Using density surface models to estimate spatio‐temporal changes in population densities and trend

April 2020

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254 Reads

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17 Citations

Precise measures of population abundance and trend are needed for species conservation; these are most difficult to obtain for rare and rapidly changing populations. We compare uncertainty in densities estimated from spatio–temporal models with that from standard design‐based methods. Spatio–temporal models allow us to target priority areas where, and at times when, a population may most benefit. Generalised additive models were fitted to a 31‐year time series of point‐transect surveys of an endangered Hawaiian forest bird, the Hawai‘i ‘ākepa Loxops coccineus. This allowed us to estimate bird densities over space and time. We used two methods to quantify uncertainty in density estimates from the spatio–temporal model: the delta method (which assumes independence between detection and distribution parameters) and a variance propagation method. With the delta method we observed a 52% decrease in the width of the design‐based 95% confidence interval (CI), while we observed a 37% decrease in CI width when propagating the variance. We mapped bird densities as they changed across space and time, allowing managers to evaluate management actions. Integrating detection function modelling with spatio–temporal modelling exploits survey data more efficiently by producing finer‐grained abundance estimates than are possible with design‐based methods as well as producing more precise abundance estimates. Model‐based approaches require switching from making assumptions about the survey design to assumptions about bird distribution. Such a switch warrants consideration. In this case the model‐based approach benefits conservation planning through improved management efficiency and reduced costs by taking into account both spatial shifts and temporal changes in population abundance and distribution.


Citations (70)


... If a photo contained multiple species, it was divided into separate records for each species. Only independent valid photos were counted, defined as photos (or groups) of different individuals or groups of the same species taken at a monitoring point within 30 min (Kühl et al. 2023). ...

Reference:

Utilization Patterns and Optimization Suggestions for Wildlife Passages in Xinjiang Nature Reserves
Estimating effective survey duration in camera trap distance sampling surveys

... One may again refer to Scott et al. (2018) for a detailed explanation of the study. A few other attempts at Wyrtki (1966Wyrtki ( , 1967 and (Fiedler and Talley (2006)) incorporating different sampling techniques to capture animal movements are by Lennert-Cody et al. (2018), where the authors explain a line-transect sampling technique to estimate the ETP-Dolphin school sizes, Glennie et al. (2021), where the authors use a suitable distance sampling technique to estimate wild animal movements, or Gerrodette et al. (2019), where the authors develop a hierarchical Bayesian framework to estimate dolphin school size. Now, in order to verify if these dolphin clusters are indeed length-biased, we adopt a simple exploratory approach. ...

Review of potential line-transect methodologies for estimating abundance of dolphin stocks in the eastern tropical Pacific

IWC Journal of Cetacean Research and Management

... Franzén & Johannesson, 2007). We calculated so called Multi-Species Change Indices (MSCIs) applying statistical models that allow to include species' extinctions and colonisations between compared periods (Korner-Nievergelt et al., 2022). Species Change Indices (SCIs) and ...

Multi-species population indices for sets of species including rare, disappearing or newly occurring species

Ecological Indicators

... Animals in this area are recorded at a set of predetermined snapshot moments, separated by the time interval t. This is done to ensure that the distribution of detections is not biased by animal movement (Pal et al. 2021). Time interval t is chosen based on the movement speed of the species, and is here set at one second. ...

Using distance sampling with camera traps to estimate the density of group-living and solitary mountain ungulates

Oryx

... The majority of policy frameworks [10] evidence of their practical application, focus predominantly on either fire prevention, suppression [89,106] and hazard reduction by professionals or on biodiversity conservation through reducing fire frequency, inroads [6,90] or area, usually in unison with reducing other drivers of unsuitable disturbance [37,92,107]. Consequently, while these policies and legal frameworks, including their subsequent legislation [93,105,108] influence and may support the use of many different non-industrial land management practices to achieve various goals of biodiversity conservation by promoting, for example [94,109] traditional management, recycling, and grazing, wildfire protection practiced at local or land management levels is not specifically designed to support the creation of strong co-benefits, outcomes, and indicators that meet both sets of goals [95,105,110]. However, such joint-focused outcomes and indicators linking wildfire protection to biodiversity conservation have been developed at different spatial levels, in other policy frameworks, and in formal conservation agreements [96,111,112]. ...

Fire and biodiversity in the Anthropocene

Science

... studies have also found that model-based adaptive sampling designs can have important benefits for model performance (e.g. Camp et al., 2020;Flint et al., 2023;Shanahan et al., 2021, but see Bird et al., 2022). Interestingly, we found that the type of modelbased method had little effect on model performance: provided some form of model-based sampling was used, the benefits to SDM performance were similar. ...

Using density surface models to estimate spatio‐temporal changes in population densities and trend

... Diseases can also spread from provisioned wildlife to provisioning humans (Lawson et al., 2014). Elevated wildlife mortality rates around provisioned food may occur (Pavisse et al., 2019;Swallow et al., 2019), with predation by domestic pets as well as window collisions exacerbated by encouraging wildlife into the vicinity of human settlements (Kummer and Bayne, 2015;Pavisse et al., 2019). The reproductive success of recipients might also be negatively affected if they become dependent upon provisioned food resources of poor nutritive value (Chamberlain et al., 2009;Plummer et al., 2013), which can affect blood chemistry (Ishigame et al., 2006;Støstad et al., 2019) with unknown health impacts. ...

Assessing factors associated with changes in the numbers of birds visiting gardens in winter: Are predators partly to blame?

... The covariates included in the Distance analyses included species, cluster size, time of day, terrain ruggedness index (TRI) and vegetation density index (VDI) (mean site TRI and VDI shown in Table S1). The key function and series expansion model best fitting the data was chosen by the engine for analysis (Marques et al. 2007). Due to the distance accuracy limitation of the laser rangefinder, models were truncated to exclude sightings with a perpendicular distance beyond 600 m before running analyses. ...

Improving Estimates of Bird Density Using Multiple- Covariate Distance Sampling

Ornithology

... Some studies recommend camera traps programmed with timed or video settings when data are analyzed with distance sampling 25,26 . Since motion detection sensors are more sensitive at closer distances, camera traps may collect images of animals nearer to the cameras, causing biases in distance measures 25,26 . ...

Model selection with overdispersed distance sampling data
  • Citing Article
  • August 2018

... In fact, the behaviours that we described may have been amplified by the polar bears' shift to a land-based lifestyle along this coastline in Svalbard. Although polar bear distribution on sea ice is extremely patchy 86,104,105 , opportunities to avoid others might be particularly reduced when residing on land strips in a landscape free of sea ice. ...

The number and distribution of polar bears in the western Barents Sea

Polar Research