Article

Comparison of methods to estimate historic species richness of mammals for tests of faunal relaxation in Canadian parks

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Abstract

AimSome recent tests of faunal change in reserves have relied on, but been limited to, estimates of species richness from random samples of historic range maps. We evaluated a different, Geographic Information Systems (GIS)-based, approach to count species directly, as the latter method might facilitate rapid estimation of historic species richness as well as composition for samples of the same size, shape and exact location as present-day reserves.LocationNational parks throughout Canada.Methods Geographic Information System.ResultsThe GIS-based method tended to count, in exact locations of modern parks, fewer species (on average, seven disturbance intolerant and five disturbance tolerant) present historically than extrapolated from randomly sampled sites, but the differences were not greater than expected by chance. However, correlations between number of species lost and park size were weaker than reported previously, suggesting a greater potential for other factors to influence a change in species richness (and composition) than inferred earlier.Main conclusionsDirect counts of historic range maps using GIS tools provided a quick means to estimate site-specific historic species richness not statistically different from estimates produced by random sampling. As well, the GIS-based method could yield data about historic species composition for the specific location and size of modern reserves, which may be more ecologically meaningful in terms of assessing what factors may have contributed to the observed species losses.

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... Terrestrial reserves have been shown to undergo faunal relaxation consistent with the principles of island biogeography. Small parks that have become isolated from their surrounding habitat matrix have been shown to have lost more of their historical complement of disturbance-sensitive mammals than large parks that are surrounded by intact habitat (Glenn & Nudds, 1989;Newmark, 1995;Gurd & Nudds, 1999;Brashares et al., 2001;Gurd et al., 2001;Wiersma & Nudds, 2001). Tests for extinctions of mammals from Canadian reserves have used two approaches to estimate historical baselines of species richness. ...
... Species richness has also been directly sampled from these maps using geographical information systems (GIS) (e.g. Wiersma & Nudds, 2001). Estimates of the numbers of species that might have been present in reserves prior to insularization were compared with current species richness (Glenn & Nudds, 1989;Gurd & Nudds, 1999;Gurd et al., 2001;Wiersma & Nudds, 2001) to estimate the extent of faunal relaxation that each park had undergone. ...
... Wiersma & Nudds, 2001). Estimates of the numbers of species that might have been present in reserves prior to insularization were compared with current species richness (Glenn & Nudds, 1989;Gurd & Nudds, 1999;Gurd et al., 2001;Wiersma & Nudds, 2001) to estimate the extent of faunal relaxation that each park had undergone. ...
Article
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Aim Tests for faunal relaxation in reserves, particularly for mammals, have relied on comparisons of current species richness with estimates of species richness derived from historical range maps. However, any range map reflects the extent of occurrence of species and not necessarily the area of occupancy. Thus, estimates of historical species richness might be prone to error introduced by ‘false positives’, that is, a species might be considered to have been present in locations where it actually was not. The effect of such ‘false positives’ could bias statistical tests of faunal relaxation to type I error, and result in estimates of the extent of faunal relaxation in reserves greater than was actually the case. We evaluated the potential for errors in historical range maps to generate inflated estimates of historical species richness of mammals at sites that are reserves today.
... Island biogeography (MacArthur and Wilson 1967) has long provided a theoretical framework to study species losses in protected areas (Diamond 1975;Soulé, Wilcox and Holtby 1979;Glenn and Nudds 1989;Burkey 1995;Newmark 1995Newmark , 1996Woodroffe and Ginsberg 1998;Gurd and Nudds 1999;Wiersma and Nudds 2001). In Canada, estimated species' distributions of terrestrial mammals prior to widespread European settlement (Banfield 1974) have been used as the reference state to which presentday species richness in parks has been compared. ...
... For example, species requiring large tracts of wilderness, such as grizzly bears (Ursus arctos) were classified as disturbance-sensitive, while those found near human settlements or in agricultural landscapes, such as coyote (Canis latrans) were classified as disturbancetolerant. The work of Glenn and Nudds (1989) and subsequent studies in Canadian protected areas has shown that some parks do not contain their historical complement of species, and the reduction in the number of disturbance-sensitive mammal species, in particular, is more severe in national and provincial parks in the southeast region of the country, which is also the region most densely populated by humans (Glenn and Nudds 1989;Gurd and Nudds 1999;Wiersma and Nudds 2001). A significant correlation between the difference in present and historical species richness of disturbance-sensitive mammals and park size (r 2 = 0.83, p = 0.0001) has been reported for these parks (Gurd and Nudds 1999). ...
... However, in a further study that measured changes between historical and present-day composition of disturbance-sensitive mammals within 24 national parks, a much weaker correlation between park size and species losses in the southeastern parks was found (r 2 = 0.15, p = 0.30; Wiersma and Nudds 2001). Thus, while island biogeography theory has provided a useful framework for thinking about species losses from protected areas, the discrepancies between the studies of mammals in Canadian parks cited here illustrate that island biogeography theory alone may not be sufficient to explain extirpations from parks (see also Rivard et al. 2000). ...
Article
It is widely accepted that large protected areas are required to effectively conserve historical species composition. However, recent analyses of mammal species loss in Canadian and African national parks contradict earlier conclusions that extent of local extinctions (i.e., extirpations) is strongly inversely related to park size, suggesting that park size alone is inadequate to predict reserve designs that may sustain biodiversity. To plan protected areas that will meet conservation goals, reserve-design models that incorporate other landscape-scale factors in addition to reserve area are needed; potential factors include the types and intensity of land use and habitat change, together with land cover types, in and around parks. Additionally, human population size around parks, and visitor density in parks may affect species loss. We quantified land use, land cover, and human population in and around 24 Canadian national parks to model effects of human disturbance and changes in natural habitats on known mammal extirpations.Multiple regression models were compared using the Akaike Information Criterion (AICc). The most parsimonious model (AICc weighting w i = 0.5391) emphasized effective habitat area in and around parks and not visitor numbers nor human population size around parks. Our model suggests that parks with as little as 3140 km2 of effective habitat area inside may be large enough to conserve historical mammal species composition if they are also surrounded by at least 18 000 km2 of effective habitat within 50 km of park boundaries.
... Such datasets are also essential to identify priority areas for biological conservation (Margules et al., 2002), to assess how effectively existing protected areas are represent- ing the present occurrence, extent or numbers of the biodi- versity features that occur in a region and to measure if the conditions of particular features within protected areas have changed through time (e.g. Gurd & Nudds, 1999;Wiersma & Nudds, 2001;Gaston et al., 2006Gaston et al., , 2008. ...
... protecting one or multiple examples of each major ecosystem or taxonomic group within a region) and buffering it from external pressures while quantifying any changes in their status through time, also require information Table 2 Relationships between range map omission and commission errors (see Methods) and the size of protected areas in the Western hemisphere. Errors were calculated using all available species lists irrespective of their completeness and for a subset of protected areas with checklists acknowledged as mostly or essentially complete & Nudds, 1999;Wiersma & Nudds, 2001). How- ever, in order successfully to detect any change in species com- position or richness in protected areas, available data must be reasonably precise. ...
Article
We use data based on species checklists and geographical range maps for 3096 amphibian, 4100 bird and 1878 terrestrial mammal species to explore possible discrepancies in estimates of species richness for protected areas. We predict substantial differences between checklist and range map richness for smaller areas, where higher errors of commission from range maps are likely to occur. We also explore how discrepancies between species inventories and range maps may affect assessments of reserve prioritization and representation.
... In the southern parts of Canada, protected areas oft en function as habitat "islands," surrounded by a sea of human development (agriculture, suburbanization, intensive resource activities, roads). Research in these parks has shown that many of them have lost species compared to similarly sized areas that are still located within a connected landscape (Gurd and Nudds 1999;Wiersma and Nudds 2001). This is consistent with theory from Island Biogeography (MacArthur and Wilson 1967), which demonstrates that oceanic islands consistently have lower species richness than an equivalent sized area of mainland. ...
... Scores were given for dispersal ability, dependence on complex (i.e., old-growth forest) habitat, and sensitivity to habitat fragmentation. Scores were assigned based on life history information given in Banfi eld (1974), and on data from Wiersma and Nudds (2001) on the number and identity of species that had gone missing from national parks across Canada, south of the sixtieth parallel, where habitat fragmentation has already taken place around many parks. The average number of mammals present in each ecoregion divided by the maximum number of mammals in all the ecoregions was used to calculate the percentage of co-occurring species. ...
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The science of conservation biology has made many contributions to improving biodiversity conservation within protected areas around the globe. Northern ecosystems are unique, and principles for protected areas design developed for temperate and tropical ecoregions may not readily be extrapolated to northern regions. Recent increases in ecological threats to the Canadian North have spurred interest in improving conservation and representation of northern ecosystems. Here, I present an overview of issues relevant to protected areas planning in the Canadian North, with a focus on the Yukon. I highlight recent Northern Research Institute- supported research on protected areas design in the Yukon, with a particular focus on the issue of representation and an examination of the potential utility of so-called "focal" species in identifying the location of representative protected areas. I show how Geographic Information Systems (GIS) may be applied to test questions of how many protected areas may be required to adequately represent mammal diversity in the ecoregions of the Yukon. I also use two different approaches to identify focal species for the Yukon to show that there is a great deal of ambiguity involved in how these species are identified.
... We took historical mammal species composition (prior to wide- spread European settlement) for each of 24 national parks from Wiersma and Nudds (2001). The accuracy of historical estimates is never fully known, and may contribute to errors in inferring species extirpations. ...
Article
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Recent analyses of mammal species loss in protected areas around the world suggest that habitat loss and human population density outside of park boundaries may be better predictors of species loss and biodiversity patterns than absolute area of parks themselves. In North American parks, there have been confl icting studies about the relative impact of habitat versus human population density on the loss of mammals. These differences may be due to scale effects, as past studies in Canadian national parks have not examined the effect of spatial scale on species loss since the time of widespread European settlement. Here, we build on previous work and look at the effects of habitat area and human population density in buffer regions that are 10, 25, 50, and 100 km (6.2, 15.5, 31.1, and 62.1 mi) outside of the boundaries of 24 national parks in Canada on the loss of disturbance-sensitive mammals. We also examine whether the relative importance of predictors is correlated with species body size. As in previous work, we fi nd that the amount of effective habitat area is a more signifi cant predictor than human population density and that scale effects are not signifi cant, at least for the scales and species examined.
... deforestation) literally up to parks' statutory boundaries ( Figure 3). Increasingly, human activities surrounding reserves impact the management of populations inside park boundaries (Wiersma and Nudds 2001;Sinclair and Byrom 2006). Given the already limited potential of reserves to effectively conserve biodiversity without the additional threats from climate change, conservation outside traditional reserve networks is essential (Ricketts 2001;Ricketts et al. 2001). ...
Article
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Canadian biodiversity is especially high in temperate southern regions, where human-dominated land uses are both intensive and widespread. As a result, endangered species are also disproportionately concentrated in these areas. Climate change presents a new threat across most of Canada, including areas of intensive human land use, which creates conditions for substantial shifts in species composition and potential losses of many rare species. Protected areas is one adaptation strategy but, in Canada, parks suffer from severe limitations in their distribution, size, and because they have static boundaries. Land use changes around several protected areas in Canada are leading increasingly to their effective isolation, a trend we demonstrate using high resolution satellite data. Little published research has yet addressed this issue in the Canadian context, although some models now forecast ecological changes in the next century. Adaptation to global change impacts will necessitate refocusing conservation strategies beyond the boundaries of protected areas to include broader landscape perspectives. Necessary responses to these challenges include validated models predicting future biotic responses to global change, expanded biodiversity monitoring across Canada, improvements to the patchwork of federal and provincial legislation protecting species, and preemptive conservation strategies that recognize impending transitions to unprecedented environmental conditions.
... Species-area curves are useful not only to predict species numbers after habitat loss, but also to estimate historical species numbers, when natural habitats were more extensive (Wiersma & Nudds, 2001). The estimated past species number (PastSN) can be compared by the potential historical species numbers (PotSN) calculated from historical data and from extrapolations of recent species ranges and trends, which can reveal the long-term trend of present species richness. ...
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