Distance software: design and analysis of distance sampling surveys for estimating population size

1. Distance sampling is a widely used technique for estimating the size or density of biological populations. Many distance sampling designs and most analyses use the software Distance. 2. We briefly review distance sampling and its assumptions, outline the history, structure and capabilities of Distance, and provide hints on its use. 3. Good survey design is a crucial prerequisite for obtaining reliable results. Distance has a survey design engine, with a built-in geographic information system, that allows properties of different proposed designs to be examined via simulation, and survey plans to be generated. 4. A first step in analysis of distance sampling data is modeling the probability of detection. Distance contains three increasingly sophisticated analysis engines for this: conventional distance sampling, which models detection probability as a function of distance from the transect and assumes all objects at zero distance are detected; multiple-covariate distance sampling, which allows covariates in addition to distance; and mark–recapture distance sampling, which relaxes the assumption of certain detection at zero distance. 5. All three engines allow estimation of density or abundance, stratified if required, with associated measures of precision calculated either analytically or via the bootstrap. 6. Advanced analysis topics covered include the use of multipliers to allow analysis of indirect surveys (such as dung or nest surveys), the density surface modeling analysis engine for spatial and habitat-modeling, and information about accessing the analysis engines directly from other software. 7. Synthesis and applications. Distance sampling is a key method for producing abundance and density estimates in challenging field conditions. The theory underlying the methods continues to expand to cope with realistic estimation situations. In step with theoretical developments, state-of- the-art software that implements these methods is described that makes the methods accessible to practicing ecologists.

A Model-Based Approach for Making Ecological Inference from Distance Sampling Data

We consider a fully model-based approach for the analysis of distance sampling data. Distance sampling has been widely used to estimate abundance (or density) of animals or plants in a spatially explicit study area. There is, however, no readily available method of making statistical inference on the relationships between abundance and environmental covariates. Spatial Poisson process likelihoods can be used to simultaneously estimate detection and intensity parameters by modeling distance sampling data as a thinned spatial point process. A model-based spatial approach to distance sampling data has three main benefits: it allows complex and opportunistic transect designs to be employed, it allows estimation of abundance in small subregions, and it provides a framework to assess the effects of habitat or experimental manipulation on density. We demonstrate the model-based methodology with a small simulation study and analysis of the Dubbo weed data set. In addition, a simple ad hoc method for handling overdispersion is also proposed. The simulation study showed that the model-based approach compared favorably to conventional distance sampling methods for abundance estimation. In addition, the overdispersion correction performed adequately when the number of transects was high. Analysis of the Dubbo data set indicated a transect effect on abundance via Akaike’s information criterion model selection. Further goodness-of-fit analysis, however, indicated some potential confounding of intensity with the detection function.

Evaluation of line-transect sampling for estimating koala abundance in the Pine Rivers Shire, south-east Queensland

Distance sampling using line transects has not been previously used or tested for estimating koala abundance. In July 2001, a pilot survey was conducted to compare the use of line transects with strip transects for estimating koala abundance. Both methods provided a similar estimate of density. On the basis of the results of the pilot survey, the distribution and abundance of koalas in the Pine Rivers Shire, south-east Queensland, was determined using line-transect sampling. In total, 134 lines (length 64 km) were used to sample bushland areas. Eighty-two independent koalas were sighted. Analysis of the frequency distribution of sighting distances using the software program DISTANCE enabled a global detection function to be estimated for survey sites in bushland areas across the Shire. Abundance in urban parts of the Shire was estimated from densities obtained from total counts at eight urban sites that ranged from 26 to 51 ha in size. Koala abundance in the Pine Rivers Shire was estimated at 4584 (95% confidence interval, 4040-5247). Line-transect sampling is a useful method for estimating koala abundance provided experienced koala observers are used when conducting surveys.

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups.

Densidade, tamanho populacional e conservação de primatas em fragmento de Mata Atlântica no sul do Estado de Minas Gerais, Brasil

Este trabalho teve como objetivo estimar a densidade e o tamanho populacional de quatro espécies de primatas [Alouatta clamitans Cabrera, 1940; Callicebus nigrifrons (Spix, 1823); Callithrix aurita (É. Geoffroy, 1812); Cebus nigritus (Goldfuss, 1809)] que ocorrem em um fragmento de Mata Atlântica de aproximadamente 350 ha, localizado no município de Pouso Alegre, estado de Minas Gerais e reunir subsídios para a conservação dessas espécies na região. O levantamento populacional foi realizado através do método de amostragem de distâncias em transecções lineares (Distance Sampling). Os dados foram coletados entre os meses de abril e agosto de 2008 a partir de quatro transecções implantadas na área de estudo. Os cálculos de densidade e tamanho populacional foram realizados empregando-se o programa Distance 5.0. As densidades foram estimadas em 23,83 ± 9,78 ind./km² para Callicebus nigrifrons, 14,76 ± 5,92 ind./km² para Callithrix aurita e 7,71 ± 2,13 ind./km² para Cebus nigritus. O tamanho populacional foi estimado em 83,0 ± 34,0 indivíduos para C. nigrifrons, 52,0 ± 20,8 indivíduos para Callithrix aurita e 27,0 ± 7,4 indivíduos para Cebus nigritus. Com relação ao bugio (A. guariba clamitans), constatou-se que apenas um grupo com seis indivíduos sobrevive na área. Conclui-se que, no caso de continuarem isoladas, essas populações têm poucas chances de sobrevivência no futuro frente aos riscos de eventos estocásticos. A criação de corredores ecológicos conectando a área de estudo aos outros fragmentos em seu entorno e a translocação de indivíduos de outras áreas da Mata Atlântica para esta região poderão constituir alternativas para garantir a viabilidade dessas populações em longo prazo. Para tanto, é necessário que se consolide uma política pública no município de Pouso Alegre voltada à criação, ampliação e gestão de Unidades de Conservação, e ao incentivo para a adoção de práticas produtivas sob critérios de sustentabilidade no entorno dessas áreas de interesse ecológico.

Study of baleen whales’ ecology and interaction with maritime traffic activities to support management of a complex socio-ecological system

Thesis written in co-mentorship with Robert Michaud.

Effects of Gunnison Sage-Grouse habitat treatment efforts on associated avifauna and vegetation structure

Conservation efforts over the last 20 years for the Gunnison Sage-Grouse (Centrocercus minimus) have involved extensive habitat manipulations done predominantly to improve brood rearing habitat for the grouse. However, the effects of Gunnison Sage-Grouse habitat treatments on sympatric avifauna and responses of vegetation to manipulations are rarely measured, and if they are, it is immediately following treatment implementation. This study examined the concept of umbrella species management by retrospectively comparing density and occupancy of eight sagebrush associated songbird species and six measures of vegetation in treated and control sites. Our results suggested that songbird densities and occupancy changed for birds at the extreme ends of their association with sagebrush and varied with fine-scale habitat structure. We found Brewer’s Sparrows (Spizella breweri) decreased in density on treated sites and Vesper Sparrows (Pooecetes gramineus) increased. Occupancy estimation revealed that Brewer’s Sparrows and Green-tailed Towhees (Pipilo chlorurus) occupied significantly fewer treated points whereas Vesper Sparrows occupied significantly more. Vegetation comparisons between treated and control areas found shrub cover to be 26% lower in treated sites. Lower shrub cover in treated areas may explain the differences in occupancy and densities of the species sampled based on known habitat needs. The fine-scale analysis showed a negative relationship to forb height and cover for the Sage Sparrow (Amphispiza belli) indicating, from vegetation measures showing grass and forb cover during a good precipitation year covered significantly more area in the treatment than the control sites, that Sage Sparrows may also not respond favorably to Gunnison Sage-Grouse habitat treatments. While the concept of an umbrella species is appealing, evidence from this study suggests that conservation efforts aimed at the Gunnison Sage-Grouse may not be particularly effective for conserving other sagebrush obligate species of concern. This is probably due to Gunnison Sage-Grouse habitat management being focused on the improvement of brood rearing habitat which reduces sagebrush cover and promotes development of understory forbs and grasses.

Population estimates and habitat use by the critically endangered Taita Apalis Apalis fuscigularis in south-eastern Kenya

The Taita Apalis Apalis fuscigularis (IUCN category: Critically Endangered) is a species endemic to south-eastern Kenya. We assessed population size and habitat use in the three forest sites in which it is known to occur (Ngangao, Chawia and Vuria, totalling 257 ha). The estimate of total population size, derived from distance sampling at 412 sample points, ranged from 310 to 654 individuals, with the northern section of Ngangao fragment having 10-fold higher densities than Chawia (2.47-4.93 versus 0.22-0.41 birds ha(-1)). Ngangao north alone hosted 50% of the global population of the species. The highly degraded Vuria fragment also had moderately high densities (1.63-3.72 birds ha(-1)) suggesting that the species tolerates some human disturbance. Taita Apalis prefers vegetation with abundant climbers, but the predictive power of habitat use models was low, suggesting that habitat structure is not a primary cause for the low density of the species in Chawia. Protecting the subpopulation in the northern section of Ngangao is a priority, as is identifying factors responsible of the low abundance in Chawia, because ameliorating conditions in this large fragment could substantially increase the population of Taita Apalis.

Aerial line transect survey to estimate abundance of marsh deer (Blastocerus dichotomus) (Illiger, 1815)

A população de cervo-do-pantanal (Blastocerus dichotomus) está drasticamente reduzida no Brasil. O nosso objetivo foi o de estimar a abundância do cervo-do-pantanal na bacia do Rio Paraná e discutir a metodologia aplicada. Os resultados darão suporte para uma análise do impacto do enchimento da represa de Porto Primavera sobre essa população. Sessenta e nove animais foram registrados através de sobrevôo utilizando-se a metodologia de transecção linear com amostragem das distâncias. Os dados não corrigidos resultaram em uma densidade estimada de 0,0035ind/ha e uma população de 636 indivíduos. A correção de g para os animais que não foram vistos apresentou uma densidade de 0,0049 ind/ha e uma abundância de 896 (CV=0,27) indivíduos. A metodologia foi aplicada com sucesso na estimativa de cervo-do-pantanal. Esse resultado é importante para avaliarmos a população do cervo-do-pantanal na área e para futuramente analisarmos o impacto do enchimento da represa.

How well will Brazil's system of atlantic forest reserves maintain viable bird populations?

It is crucial for biodiversity conservation that protected areas are large and effective enough to support viable populations of their original species. We used a point count distance sampling method to estimate population sizes of a range of bird species in three Atlantic forest protected areas of size 5600, 22,500, and 46,050 ha. Population sizes were generally related to reserve area, although in the mid-sized reserve, there were many rare species reflecting a high degree of habitat heterogeneity. The proportions of forest species having estimated populations > 500 ranged from 55% of 210 species in the largest reserve to just 25% of 140 species in the smallest reserve. All forest species in the largest reserves had expected populations > 100, but in the small reserve, 28% (38 species) had populations < 100 individuals. Atlantic forest endemics were no more or less likely to have small populations than widespread species. There are 79 reserves (> 1000 ha) in the Atlantic forest lowlands. However, all but three reserves in the north of the region (Espirito Santo and states north) are smaller than 10,000 ha, and we predict serious levels of local extinction from these reserves. Habitat heterogeneity within reserves may promote species richness within them, but it may also be important in determining species loss over time by suppressing populations of individual species. We suggest that most reserves in the region are so small that homogeneity in the habitat/altitude within them is beneficial for maintenance of their (comparatively small) original species compliment. A lack of protection in the north, continued detrimental human activity inside reserves, and our poor knowledge of how well the reserve system protects individual taxa, are crucial considerations in biodiversity management in the region.

Density and Spatial Distribution of Buffy-tufted-ear Marmosets (Callithrix aurita) in a Continuous Atlantic Forest

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Density and population size of mammals introduced on a land-bridge island in southeastern Brazil

The introduction of alien species is one of the main threats to the conservation of native species, especially in island ecosystems. Here, we report on the population growth of 15 species of mammals introduced in 1983 on the island of Anchieta, an 828 ha land-bridge island in southeastern Brazil. We estimated the density of mammals through 296 km of line transect census. Five species introduced became extinct (coypu, brocket deer, six-banded armadillo, nine-banded armadillo, maned three-toed sloth); six became over-abundant (marmoset, coati, agouti, seven-banded armadillo, and capybara); one has a stable population (capuchin monkey). Anchieta Island has the highest density of mammals in the entire Atlantic forest (486.77 ind/km(2)), especially nest predators (232.83 ind/km(2)) and herbivores (253.58 ind/km(2)). Agoutis (Dasyprocta spp.) and marmosets (Callithrix penicillata) were, by far, the species with the highest population growth. The high density of mammals in this island may have strong consequences for plant recruitment and bird diversity.

Severe population decline of marsh deer, Blastocerus dichotomus (Cetartiodactyla: Cervidae), a threatened species, caused by flooding related to a hydroelectric power plant

Balancing power production and environmental conservation can be problematic. The objective of this study was to investigate the abundance of marsh deer in the Paraná River Basin, above the Sergio Motta (Porto Primavera) Dam, before and after the impact of the dam closure. A fixed-wing, flat window aircraft was used to survey study transects. Observations were recorded based on the distance sampling line transect method, assuming that the detection probability decreases with increased distance. The abundance of marsh deer in the survey region prior to flooding was estimated to be 974 individuals (CV = 0.23). The overall abundance dropped from 974 to 444 (CV = 0.26) individuals after flooding, an overall reduction of 54%. This reduction can be attributed to the direct impact of the flooding process, but it was likely exacerbated by indirect effects, such as increased disease, hunting, and reduction in food availability. Prior to flooding, the marsh deer was distributed widely throughout the dam's catchment area; however, the marsh deer habitat was almost completely destroyed by the flooding process. This situation highlights the need to implement management strategies that ensure the survival of the remaining fragmented population.

Double-Observer Line Transect Methods: Levels of Independence

Double-observer line transect methods are becoming increasingly widespread, especially for the estimation of marine mammal abundance from aerial and shipboard surveys when detection of animals on the line is uncertain. The resulting data supplement conventional distance sampling data with two-sample mark–recapture data. Like conventional mark–recapture data, these have inherent problems for estimating abundance in the presence of heterogeneity. Unlike conventional mark–recapture methods, line transect methods use knowledge of the distribution of a covariate, which affects detection probability (namely, distance from the transect line) in inference. This knowledge can be used to diagnose unmodeled heterogeneity in the mark–recapture component of the data. By modeling the covariance in detection probabilities with distance, we show how the estimation problem can be formulated in terms of different levels of independence. At one extreme, full independence is assumed, as in the Petersen estimator (which does not use distance data); at the other extreme, independence only occurs in the limit as detection probability tends to one. Between the two extremes, there is a range of models, including those currently in common use, which have intermediate levels of independence. We show how this framework can be used to provide more reliable analysis of double-observer line transect data. We test the methods by simulation, and by analysis of a dataset for which true abundance is known. We illustrate the approach through analysis of minke whale sightings data from the North Sea and adjacent waters.