Are pheasants attracted or repelled by roads? A test of a crucial assumption for transect censuses

Censuses are widely used to monitor populations. One of the most interesting modern techniques is distance sampling, which depends on some crucial assumptions, including the random distribution of animals with respect to transect lines, which are often asphalt and gravel roads. We tested the assumption of a random distribution of animals with respect to roads for females of ring-necked pheasants (Phasianus colchicus). Roads can be used to census pheasants using distance sampling, without significant bias because pheasants are randomly distributed with respect to roads. Our method is easy to apply to other field conditions where radio tracking data are available and can be used to test the assumption in other studies and/or with different species.     

The Effect of Animal Movement on Line Transect Estimates of Abundance

Line transect sampling is a distance sampling method for estimating the abundance of wild animal populations. One key assumption of this method is that all animals are detected at their initial location. Animal movement independent of the transect and observer can thus cause substantial bias. We present an analytic expression for this bias when detection within the transect is certain (strip transect sampling) and use simulation to quantify bias when detection falls off with distance from the line (line transect sampling). We also explore the non-linear relationship between bias, detection, and animal movement by varying detectability and movement type. We consider animals that move in randomly orientated straight lines, which provides an upper bound on bias, and animals that are constrained to a home range of random radius. We find that bias is reduced when animal movement is constrained, and bias is considerably smaller in line transect sampling than strip transect sampling provided that mean animal speed is less than observer speed. By contrast, when mean animal speed exceeds observer speed the bias in line transect sampling becomes comparable with, and may exceed, that of strip transect sampling. Bias from independent animal movement is reduced by the observer searching further perpendicular to the transect, searching a shorter distance ahead and by ignoring animals that may overtake the observer from behind. However, when animals move in response to the observer, the standard practice of searching further ahead should continue as the bias from responsive movement is often greater than that from independent movement.     

Measuring effects of linear obstacles on wildlife movements: accounting for the relationship between step length and crossing probability

Animal movements in the landscape are influenced by linear features such as rivers, roads and power lines. Prior studies have investigated how linear features, particularly roads, affect movement rates by comparing animal's movement rate measured as step lengths (i.e., the distance between consecutive observations such as GPS locations) before, during and after crossing of a linear feature. The null hypothesis has been that the length of crossing steps should not differ from other steps, and a deviation from this, mainly that steps are longer during crossing, has been taken as support for a disturbance effect of the linear feature. However, based on the simple relationship between the length of a step and its probability to cross a linear feature, we claim that this assumption is inappropriate to test for behavioural responses to linear features. The probability is related to the proportion of the total length of the trajectory (i.e., the path of movement) a step constitutes. Consequently, care should be taken when formulating hypotheses about how animal moves in relation to linear features in the landscape. Statistical tests should be set up with respect to the expected length based on the distribution of step lengths in the trajectory. We propose two methods that accounts for the bias in crossing frequency that is caused by step lengths, and illustrates their applications by using simulated animal trajectories as well as empirical data on reindeer in an area with a power line.     

Linear declines in exotic and native plant species richness along an increasing altitudinal gradient in the Snowy Mountains,Australia

Abstract Declines in plant species richness with increasing altitude are common, but the form of the relationship can vary, with both monotonic decreasing relationships and humped relationship recorded. However, these different richness to altitude relationships may be due to methods that used different plot sizes/areas and survey efforts. To explore native and exotic plant richness along an altitudinal gradient in the Snowy Mountains of Australia, we consistently surveyed plots that were 120 m² in area at 39 sites ranging from 540 to 2020 m. To relate exotic plant richness to disturbance, we surveyed plots at 16 sites along main roads and 23 sites along minor roads and also compared these 39 roadside plots to 120‐m² plots located in undisturbed vegetation adjacent to the roadside (native plant richness was only surveyed in 25 of these 39 adjacent plots). We found a negative linear relationship between total, exotic and native species richness and altitude for plots on the side of main roads (16 sites) and minor roads (23 sites). For adjacent plots negative linear relationships were significant for all measures of species richness except for native species adjacent to major roads. As the pattern occurred for exotics it is less likely to be due to historical constraints on the species pools. The pattern could be influenced by difference in levels of disturbance along the gradient, although any such gradient in disturbance would have to apply to roadside and adjacent plots on major and minor roads. Therefore, it may be due to other factors such as changes in climate along the altitudinal gradient, although additional sampling including direct measures of climatic conditions, soil and disturbance factors would be needed to determine if this was the case.     

Estimating Abundance From One-Dimensional Passive Acoustic Surveys

ABSTRACT Conventional distance sampling, the most-used method of estimating animal density and abundance, requires ranges to detected individuals, which are not easily measured for vocalizations. However, in some circumstances the sequential pattern of detection of vocalizations along a transect line gives information about the range of detection. Thus, from a one-dimensional acoustic point-transect survey (i.e., records of vocalizations detected or not detected at regularly spaced listening stations) it is possible to obtain a useful estimate of density or abundance. I developed equations for estimation of density for one-dimensional surveys. Using simulations I found that for the method to have little bias when both range of detection and rate of vocalization need to be estimated, stations needed to be spaced at 30–80% of the range of detection and the rate of vocalization should be >0.7. If either the range of detection or rate of vocalization is known, conditions are relaxed, and when both parameters are known the method works well almost universally. In favorable conditions for one-dimensional methods, estimated abundances have overall errors not much larger than those from conventional line-transect distance sampling. The methods appeared useful when applied to real acoustic data from whale surveys. The techniques may also be useful in surveys with nonacoustic detection of animals.     

Line Transect Sampling in Small and Large Regions

Melville and Welsh (2001, Biometrics 57, 1130-1137) consider an approach to line transect sampling using a separate calibration study to estimate the detection function g. They present a simulation study contrasting their results with poor results from a traditional estimator, labeled the "Buckland" estimator and referenced to Buckland et al. (1993, Distance Sampling: Estimating Abundance of Biological populations). The poor results from the "Buckland" estimator can be explained by the following observations: (i) the estimator is designated for untruncated distance data, but was applied by Melville and Welsh to truncated distance data; (ii) distance data were not pooled across transects, contrary to standard practice; and (iii) bias of the estimator was evaluated with respect to a fixed rather than a randomized grid of transect lines. We elaborate on the points above and show that the traditional methods perform to expectation when applied correctly. We also emphasize that the estimator labeled the "Buckland" estimator by Melville and Welsh is not an estimator recommended by Buckland et al. for practical survey applications.     

Design and Analysis of Line Transect Surveys for Primates

Line transect surveys are widely used for estimating abundance of primate populations. The method relies on a small number of key assumptions, and if these are not met, substantial bias may occur. For a variety of reasons, primate surveys often do not follow what is generally considered to be best practice, either in survey design or in analysis. The design often comprises too few lines (sometimes just 1), subjectively placed or placed along trails, so lacks both randomization and adequate replication. Analysis often involves flawed or inefficient models, and often uses biased estimates of the locations of primate groups relative to the line. We outline the standard method, emphasizing the assumptions underlying the approach. We then consider options for when it is difficult or impossible to meet key assumptions. We explore the performance of these options by simulation, focusing particularly on the analysis of primate group sizes, where many of the variations in survey methods have been developed. We also discuss design issues, field methods, analysis, and potential alternative methodologies for when standard line transect sampling cannot deliver reliable abundance estimates.     

Effects of survey methods on burrowing owl behaviors

Monitoring wildlife populations often involves intensive survey efforts to attain reliable estimates of population size. Such efforts can increase disturbance to animals, alter detection, and bias population estimates. Burrowing owls (Athene cunicularia) are declining across western North America, and information on the relative effects of potential survey methods on owl behaviors is needed. We designed a field experiment to compare burrowing owl flight distances, times displaced, and probabilities of being displaced between 4 potential population survey methods (single walking surveyor, single vehicle stop, single vehicle stop with 2 surveyors, and double vehicle stop with 2 surveyors), and an experimental control in the agricultural matrix of Imperial Valley, California. Between 25 April and 1 May 2008, we randomly applied survey methods to 395 adult male owls during daylight hours (0700 hours through 1900 hours). All survey methods increased odds of displacing owls from perches. Survey methods with observers outside the vehicle were 3 times more likely to displace an owl than a single vehicle stop where observers remained inside the vehicle. Owls were displaced farther distances by all survey methods compared to control trials, but distances and time displaced did not differ among survey methods. We recommend that surveys for counting owls during the breeding season in agroecystems like the Imperial Valley where high densities of owls nest primarily along the borders of fields be conducted using single vehicle stops with or without 2 surveyors, depending on conditions for locating owls from roads. © 2011 The Wildlife Society.     

Spoor counts as indices of large carnivore populations: the relationship between spoor frequency, sampling effort and true density

1. Studies of large carnivore populations and, particularly, reliable estimates of population density are necessary for effective conservation management. However, these animals are difficult to study, and direct methods of assessing population size and density are often expensive and time-consuming.
2. Indirect sampling, by counting spoor, could provide repeatable and inexpensive measures of some population parameters. The relationship between true population density and indirect sampling results has seldom been described in large carnivore studies.
3. In northern Namibia the population densities of leopards, lions and wild dogs were measured through recognition of individuals and groups. Spoor counts were then conducted independently, to assess the relationship between true density and the distribution of spoor.
4. Sampling effort, both in terms of the number of roads and total road distance in a sample zone, and the intensity of sampling, had a marked effect on the accuracy and precision of spoor frequency calculations.
5. In a homogeneous habitat, leopard spoor were evenly spread along different roads and spoor frequency was independent of road length. Taking into account very low sample sizes, the spoor density of leopards, lions and wild dogs showed a strong linear correlation with true density. The slope of the regression for leopards was different to that of lions and wild dogs.     

Geographical sampling bias and its implications for conservation priorities in Africa

Aim To design and apply statistical tests for measuring sampling bias in the raw data used to the determine priority areas for conservation, and to discuss their impact on conservation analyses for the region. Location Sub‐Saharan Africa. Methods An extensive data set comprising 78,083 vouchered locality records for 1068 passerine birds in sub‐Saharan Africa has been assembled. Using geographical information systems, we designed and applied two tests to determine if sampling of these taxa was biased. First, we detected possible biases because of accessibility by measuring the proximity of each record to cities, rivers and roads. Second, we quantified the intensity of sampling of each species inside and surrounding proposed conservation priority areas and compared it with sampling intensity in non‐priority areas. We applied statistical tests to determine if the distribution of these sampling records deviated significantly from random distributions. Results The analyses show that the location and intensity of collecting have historically been heavily influenced by accessibility. Sampling localities show dense, significant aggregation around city limits, and along rivers and roads. When examining the collecting sites of each individual species, the pattern of sampling has been significantly concentrated within and immediately surrounding areas now designated as conservation priorities. Main conclusions Assessment of patterns of species richness and endemicity at the scale useful for establishing conservation priorities, below the continental level, undoubtedly reflects biases in taxonomic sampling. This is especially problematic for priorities established using the criterion of complementarity because the estimated spatial costs of this approach are highly sensitive to sampling artefacts. Hence such conservation priorities should be interpreted with caution proportional to the bias found. We argue that conservation priority setting analyses require (1) statistical tests to detect these biases, and (2) data treatment to reflect species distribution rather than patterns of collecting effort.     

Little evidence of a road‐effect zone for nocturnal,flying insects

Roads and traffic may be contributing to global declines of insect populations. The ecological effects of roads often extend far into the surrounding habitat, over a distance known as the road‐effect zone. The quality of habitat in the road‐effect zone is generally degraded (e.g., due to edge effects, noise, light, and chemical pollution) and can be reflected in species presence, abundance, or demographic parameters. Road‐effect zones have been quantified for some vertebrate species but are yet to be quantified for insects. Investigating the road‐effect zone for insects will provide a better understanding of how roads impact ecosystems, which is particularly important given the role insects play as pollinators, predators, and prey for other species. We quantified the road‐effect zone for nocturnal flying insects along three major freeways in agricultural landscapes in southeast Australia. We collected insects using light traps at six points along 2‐km transects perpendicular to each highway (n = 17). We sorted the samples into order, and dried and weighed each order to obtain a measure of dry biomass. Using regression models within a Bayesian framework of inference, we estimated the change in biomass of each order with distance from the road, while accounting for environmental variables such as temperature, moon phase, and vegetation structure. The biomass of nine of the ten orders sampled did not change with distance from the freeway. Orthoptera (i.e., grasshoppers and crickets) was the only order whose biomass increased with distance from the freeway. From our findings, we suggest that the impacts of roads on insects are unlikely extending into the surrounding landscape over a distance of 2 km. Therefore, if there are impacts of roads on insects, these are more likely to be concentrated at the road itself, or on finer taxonomic scales such as family or genus level.     

Geographic sampling bias in the South African Frog Atlas Project: implications for conservation planning

Quality conservation planning requires quality input data. However, the broad scale sampling strategies typically employed to obtain primary species distribution data are prone to geographic bias in the form of errors of omission. This study provides a quantitative measure of sampling bias to inform accuracy assessment of conservation plans based on the South African Frog Atlas Project. Significantly higher sampling intensity near to cities and roads is likely to result in overstated conservation priority and heightened conservation conflicts in urban areas. Particularly well sampled protected areas will also erroneously appear to contribute highly to amphibian biodiversity targets. Conversely, targeted sampling in the arid northwest and along mountain ranges is needed to ensure that these under-sampled regions are not excluded from conservation plans. The South African Frog Atlas Project offers a reasonably accurate picture of the broad scale west-to-east increase in amphibian richness and abundance, but geographic bias may limit its applicability for fine scale conservation planning. The Global Amphibian Assessment species distribution data offered a less biased alternative, but only at the cost of inflated commission error.     

Variance of Stratified Survey Estimators With Probability of Detection Adjustments

Abstract: Estimates of wildlife population sizes are frequently constructed by combining counts of observed animals from a stratified survey of aerial sampling units with an estimated probability of detecting animals. Unlike traditional stratified survey designs, stratum-specific estimates of population size will be correlated if a common detection model is used to adjust counts for undetected animals in all strata. We illustrate this concept in the context of aerial surveys, considering 2 cases: 1) a single-detection parameter is estimated under the assumption of constant detection probabilities, and 2) a logistic-regression model is used to estimate heterogeneous detection probabilities. Naïve estimates of variance formed by summing stratum-specific estimates of variance may result in significant bias, particularly if there are a large number of strata, if detection probabilities are small, or if estimates of detection probabilities are imprecise. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):837–844; 2008)     

Large Roads Reduce Bat Activity across Multiple Species

Although the negative impacts of roads on many terrestrial vertebrate and bird populations are well documented, there have been few studies of the road ecology of bats. To examine the effects of large roads on bat populations, we used acoustic recorders to survey bat activity along ten 300 m transects bordering three large highways in northern California, applying a newly developed statistical classifier to identify recorded calls to the species level. Nightly counts of bat passes were analyzed with generalized linear mixed models to determine the relationship between bat activity and distance from a road. Total bat activity recorded at points adjacent to roads was found to be approximately one-half the level observed at 300 m. Statistically significant road effects were also found for the Brazilian free-tailed bat (Tadarida brasiliensis), big brown bat (Eptesicus fuscus), hoary bat (Lasiurus cinereus), and silver-haired bat (Lasionycteris noctivagans). The road effect was found to be temperature dependent, with hot days both increasing total activity at night and reducing the difference between activity levels near and far from roads. These results suggest that the environmental impacts of road construction may include degradation of bat habitat and that mitigation activities for this habitat loss may be necessary to protect bat populations.     

Improving inference for aerial surveys of bears: The importance of assumptions and the cost of unnecessary complexity

Obtaining useful estimates of wildlife abundance or density requires thoughtful attention to potential sources of bias and precision, and it is widely understood that addressing incomplete detection is critical to appropriate inference. When the underlying assumptions of sampling approaches are violated, both increased bias and reduced precision of the population estimator may result. Bear (Ursus spp.) populations can be difficult to sample and are often monitored using mark‐recapture distance sampling (MRDS) methods, although obtaining adequate sample sizes can be cost prohibitive. With the goal of improving inference, we examined the underlying methodological assumptions and estimator efficiency of three datasets collected under an MRDS protocol designed specifically for bears. We analyzed these data using MRDS, conventional distance sampling (CDS), and open‐distance sampling approaches to evaluate the apparent bias‐precision tradeoff relative to the assumptions inherent under each approach. We also evaluated the incorporation of informative priors on detection parameters within a Bayesian context. We found that the CDS estimator had low apparent bias and was more efficient than the more complex MRDS estimator. When combined with informative priors on the detection process, precision was increased by >50% compared to the MRDS approach with little apparent bias. In addition, open‐distance sampling models revealed a serious violation of the assumption that all bears were available to be sampled. Inference is directly related to the underlying assumptions of the survey design and the analytical tools employed. We show that for aerial surveys of bears, avoidance of unnecessary model complexity, use of prior information, and the application of open population models can be used to greatly improve estimator performance and simplify field protocols. Although we focused on distance sampling‐based aerial surveys for bears, the general concepts we addressed apply to a variety of wildlife survey contexts.     

A test of the hypothesis of compensatory upstream dispersal using a stream-dwelling waterstrider,Gerris remigis Say

Summary This paper reports the results of 2 experiments designed to determine if adult Gerris remigis (Hemiptera, heteroptera), a stream-dwelling waterstrider, tend to disperse preferentially upstream, as predicted by the colonization cycle hypothesis summarized by Muller (1982). Markrecapture observations and experimental removals were used to assess the distance and direction of movement of adult G. remigis along a small mountain stream, over a full year. These experiments indicated that adult G. remigis show a significant upstream bias in movement distance, but not in numbers of animals moving. This upstream bias is characteristic of pre-reproductive, sexually immature adults of both sexes, and occurs primarily in association with movements to and from diapause sites. Although the existence of a significant upstream bias in movement distances tends to support the colonization cycle hypothesis, the data from the removal experiments clearly show that upstream dispersal is not sufficient to compensate entirely for downstream drift.     

Nocturnal line transect sampling of wild boar (Sus scrofa) in a Mediterranean forest: long-term comparison with capture–mark–resight population estimates

Although accurate estimates of wild boar (Sus scrofa) populations are crucial for any effective resource management or pest control programme, this species is well-known to be difficult to monitor. We conducted a 10-year study in a fenced Mediterranean forest (Rome, Italy) to evaluate nocturnal line transect sampling performances. We focused on its accuracy in monitoring changes in density, which was independently estimated by capture–mark–resight (CMR) performed on counts at feeding sites. We carried out night surveys in the autumn of 2001–2010, using portable infrared cameras to detect animals. We sampled on foot to cover the whole study area and the different habitat types evenly. However, to ensure safe working conditions during night and to limit disturbance, we placed transects along paths and forest roads. Therefore, we investigated the potential impact of our convenience sampling on the detection process, using radiolocations of wild boars to assess their distribution with respect to selected transects. We found that our survey design should not have biased our estimates and that densities and coefficients of variations from line transect sampling were consistent with CMR results. Although labour-intensive, we believe that our approach can improve wild boar monitoring effectively, even in concealing habitats, providing decision makers with accurate estimates (and quantified confidence limits) which can help to develop the most appropriate management programme. Moreover, the current low price of new-generation infrared cameras can also increase strongly the cost-effectiveness of this method.     

Comparison of survey methods for an invasive plant at the subwatershed level

Invasive plant survey methods that are practical and economical are needed to describe established colonies and detect nascent invaders. We compared results from random and roadside surveys of Alliaria petiolata (Bieb.) Cavara & Grande across a 5730-ha subwatershed. The random survey included 150 1-ha plots; the roadside survey examined 0.1-mile increments (10-m deep) along paved roads (totaling 1104 0.16-ha plots). In the random survey, agriculture was the dominant land use (49% of sampled area), and most A. petiolata patches were in wooded, shaded riparian, and waste areas (34%, 34%, and 29% of patches, respectively). In the roadside survey, right-of-way land use was dominant (38% of sampled area), and most A. petiolata patches were in right-of-way, wooded, and shaded riparian areas (53%, 22%, and 19% of patches, respectively). According to generalized linear model analysis, survey methods did not differ in the overall probability of finding A. petiolata (P=0.17 and 0.11 for random and roadside surveys, respectively). Shaded riparian, wooded, and mixed-species right-of-way land uses were the dominant habitat for A. petiolata in both surveys, but only the random survey indicated waste areas as significant habitat. Alliaria petiolata occurred mostly as small patches in roadsides, but as large patches in random plots, suggesting faster spread in the roadside. Results indicated that disturbed lands along roadsides were important for invasion and spread of A. petiolata; therefore, the roadside survey was a useful, practical method for detecting nascent invasions and management planning. The random sampling lacked a land use bias, and provided data that could be generalized across the subwatershed; however, this method required at least four times more person hours to complete than the roadside survey for a similar amount of area. Although roadside sampling did not provide a completely reliable assessment of target plant populations within the landscape, it may provide an adequate approximation, depending on the specific goals of the survey. Concurrent surveys would provide the most complete information.     

Comparing iDNA from mosquitoes and flies to survey mammals in a semi-controlled Neotropical area

Ingested-derived DNA (iDNA) from insects represents a powerful tool for assessing vertebrate diversity because insects are easy to sample, have a diverse diet and are widely distributed. Because of these advantages, the use of iDNA for detecting mammals has gained increasing attention. Here we aimed to compare the effectiveness of mosquitoes and flies to detect mammals with a small sampling effort in a semi-controlled area, a zoo that houses native and non-native species. We compared mosquitoes and flies regarding the number of mammal species detected, the amount of mammal sequence reads recovered, and the flight distance range for detecting mammals. We also verified if the combination of two mini-barcodes (12SrRNA and 16SrRNA) would perform better than either mini-barcode alone to inform local mammal biodiversity from iDNA. To capture mosquitoes and flies, we distributed insect traps in eight sampling points during 5 days. We identified 43 Operational Taxonomic Units from 10 orders, from the iDNA of 17 mosquitoes and 46 flies. There was no difference in the number of species recovered per individual insect between mosquitoes and flies, but the number of flies captured was higher, resulting in more mammal species recovered by flies. Eight species were recorded exclusively by mosquitoes and 20 by flies, suggesting that using both samplers would allow a more comprehensive screening of the biodiversity. The maximum distance recorded was 337 m for flies and 289 m for mosquitoes, but the average range distance did not differ between insect groups. Our assay proved to be efficient for mammal detection, considering the high number of species detected with a reduced sampling effort.     

How do roads affect the habitat use of an assemblage of scavenging raptors?

Scavengers may benefit from the availability of dead animals along roads that result from collisions with vehicles. However, roads are also considered risky places for many species. Animal habitat selection patterns usually balance energy intake with mortality risk. In this work we analyzed the foraging space use of an assemblage of diurnal scavenging raptors in relation to distance from roads in northwest Patagonia. We selected patches at different distances from roads, and placed a sheep carcass in each patch during the night (n = 18 carcasses in total). In general, carcasses near roads were detected by diurnal scavenging raptors much faster than those far from roads. Smaller raptors such as southern caracaras (Caracara plancus), chimango caracaras (Milvago chimango), and black vultures (Coragyps atratus), were commonly associated with roads both in terms of overall detections and scavenging activities. Southern and chimango caracaras proved to be very good at detecting carcasses, were faster to land in order to feed from them, and were found in greater numbers near roads than far from them. Even though Andean condors (Vultur gryphus) and black-chested buzzard-eagles (Geranoaetus melanoleucus) flew all over the area, they chose to feed far from roads. Our work emphasizes that some scavengers have taken advantage of the novel food resources provided by roads whereas others are reluctant to feed near them. Within a scenario of an increasing number of roads, some species can extend their distributions favoring competition and biotic homogenization processes within original communities. We highlight the importance of taking into account large flying scavengers in land-use planning. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.