The habitat requirements of the rufous treecreeper (Climacteris rufa). 2. Validating predictive habitat models

Examining the predictive capability of statistical models with data independent from that used to derive the model is a vital step in the iterative procedure of assessing model performance. I derived logistic regression models of the habitat use of the rufous treecreeper (Climacteris rufa) at two spatial scales: woodland (territory selection model) and territory (nest-site selection model). The performance of these models was assessed in relation to the original data collected and validated with new, independent data. When applied to the original data, the territory model had a high predictive capability correctly classifying 90% of sites (n=100) that were either occupied or unoccupied by treecreepers. Correct classification rate was reduced to 70% (n=50) when the model was applied to the validation data. Model performance was generally robust when probability of occurrence values for the species were varied. In contrast, the nest-site model had lower predictive capabilities correctly classifying between 66 and 68% of sites, and performed relatively poorly when probability values were varied. The performance of the models differed slightly between the original and validation data, and substantially between the spatial scales examined. Territory use by rufous treecreepers could be predicted with some confidence indicating that the territory model may be a useful tool for habitat management. Nest-site use could not be predicted with confidence probably as a result of the high abundance of suitable, but unused, nest sites in the study area.     

Differences in the reproductive success and survival of the rufous treecreeper (Climacteris rufa) between a fragmented and unfragmented landscape

The rufous treecreeper (Climacteris rufa) has declined in abundance in the agricultural regions of southwestern Australia. The patterns of decline are well documented, but the processes that threaten population persistence are poorly understood. I compared the reproductive success and survival of the treecreeper between three sites in an unfragmented landscape and four remnant categories (large, small, grazed and ungrazed) in a fragmented, agricultural landscape. Nest success and annual productivity were significantly higher in the unfragmented landscape, but varied between sites and remnant categories within landscapes. Nest success was lowest in grazed remnants and annual productivity was positively associated with territory size in the fragmented landscape. Fledgling survival rates did not differ between landscapes, but there was a trend for juvenile survival rates to be higher in the unfragmented landscape. I used artificial nests to compare relative predation rates between landscapes, and provisioning rates and prey biomass brought to nestlings to assess differences in food availability. There were no landscape differences in predation rates, but provisioning rates to nestlings and total prey biomass were significantly lower in the fragmented landscape. Mean habitat quality was also lower in the fragmented landscape, although it differed between remnant categories. Reduced reproductive success, juvenile survival, food availability and habitat quality may threaten the viability of the rufous treecreeper population living in the fragmented landscape. Limiting the modification of remaining habitat (e.g. removing stock grazing) and improving habitat quality are required to assist in the conservation of this species.     

Independent effects of woodland loss and fragmentation on Brown Treecreeper distribution

We examined the influence of local habitat and the surrounding landscape on the distribution of Brown Treecreepers in a matrix of woodlands and pastures. Our goals were to: (1) determine the importance and scale of the independent effects of woodland cover and fragmentation on treecreeper distribution, and (2) employ landscape variables to improve models of treecreeper distribution based on local habitat features. Woodland fragmentation was important at a large scale while both woodland cover and fragmentation were important at a smaller scale. Excluding unoccupied sites in highly fragmented landscapes improved the ability of local habitat features to explain Brown Treecreeper distribution, which appeared to be constrained by cavity density. Brown Treecreepers' response to fragmentation at the larger scale may occur because fragmentation disrupts dispersal. Alternatively, their response may be an example of a general phenomenon of fragmentation effects only arising when < 20% of woodland cover remains at a given scale. As fragmentation increases, so does the need to incorporate landscape patterns into wildlife-habitat models.     

Thresholds in selection of breeding habitat by the Eurasian treecreeper (Certhia familiaris)

Assessment of habitat thresholds is a topical issue in ecology, both from theoretical and applied perspectives. We examined how forest structure influences selection of breeding habitat in the Eurasian treecreeper (Certhia familiaris). It is an old-growth forest passerine, which can be considered an umbrella species. Habitat selection data, covering five breeding seasons, were collected from a study area across three spatial scales: (a) territory core scale (a radius of 30 m), (b) territory scale (a radius of 200 m) and (c) large scale (a radius of 500 m). Logistic regression analyses revealed that the radii of 30 and 200 m from the nest were the most important spatial scales for the forest patch occupancy. A loss of forest habitat and a low circumference of stems within the territory decreased the probability of occupancy. There was a distinct threshold in the amount of forest cover on forest patch occupancy when the volume of timber was taken into account. At low timber volumes (0-151 m³/ha) the amount of forest cover was negatively related to the occupancy rate of forest patches. This negative relationship changed abruptly when the volume of timber exceeded 152 m³/ha, after which forest cover had a positive effect on the occupancy rate of forest patches. This is a new way to examine habitat thresholds in relation to forest cover. Furthermore, as debated in recent literature, treecreepers also tend to respond to habitat loss rather than to changes in habitat configuration, which stresses the need for habitat restoration and conservation. Moreover, our results emphasize the importance of within-territory structure over the characteristics of the habitat matrix in selection of breeding habitat.     

Regional conservation priorities for a large predator: golden eagles (Aquila chrysaetos) in the Alpine range

We compared the density of golden eagles (Aquila chrysaetos) and their nest dispersion, productivity and diet in Alpine and pre-Alpine areas. The comparison was made at two spatial scales: (1) at the territory level within two contiguous populations, and (2) at the population level, using published data for 22 populations scattered across four countries (France, Switzerland, Germany and Italy). Availability of golden eagle's main prey species was higher in Alpine than in pre-Alpine regions. The potential foraging ranges of pairs of eagles in the pre-Alps had larger amounts of woodland and lower amounts of suitable foraging habitat than those of Alpine pairs. At both spatial scales, density and productivity were lower in the pre-Alps. Also, pre-Alpine populations were characterised by a lower percentage of main prey species in the diet, which was significantly correlated with productivity, and by a higher breadth of diet, which was negatively correlated with density. Overall, higher prey availability and habitat suitability in the Alps resulted in higher density and breeding success, in a manner compatible with theoretical metapopulation models. We suggest that priority be given to conservation of eagle populations in high quality Alpine habitat, and that conservation action be carried out by a mixture of site-protection measures and conservation of the wider environment.     

Habitat selection by the pale-headed brush-finch (Atlapetes pallidiceps) in southern Ecuador: implications for conservation

The pale-headed brush-finch (Atlapetes pallidiceps) is threatened with extinction due to habitat loss, but very little is known about its ecological requirements. We used multiple logistic regression to study habitat selection of this species at landscape, territory, and nest site scales in order to make recommendations about effective management. Habitat selection by the sympatric stripe-headed brush-finch (Buarremon torquatus) was examined with the same methods in order to analyse interspecific resource partitioning and potential competition. The pale-headed brush-finch selected semi-open habitat types with intermediate scrub heights, and avoided forests. Nest sites depended on the presence of vines or bamboo. By contrast, the stripe-headed brush-finch chose dense habitat with low ground cover under tall vegetation and avoided semi-open habitat. The two species had overlapping territories but differed significantly in microhabitat use and the use of vegetation strata. We found no convincing evidence that the stripe-headed brush-finch displaces the pale-headed brush-finch from optimal habitat. The preservation of semi-open scrubland maintained by low-intensity grazing is suggested for future conservation of the pale-headed brush-finch.     

Thresholds in landscape parameters: occurrence of the white-browed treecreeper Climacteris affinis in Victoria, Australia

The concept of critical thresholds of habitat loss has recently received considerable attention in conservation biology and landscape ecology, yet empirical examples of thresholds are scarce. Threatened species management could benefit from recognition of thresholds because conditions under which populations are at risk can be specified. In this study, 56 woodland patches in north-west Victoria were surveyed for the white-browed treecreeper Climacteris affinis, a threatened insectivorous bird of the semi-arid zone of southern Australia. Comparisons with historic records indicate the species’ range is contracting in Victoria. Using logistic regression and hierarchical partitioning, two models of patch occupancy were developed. Tree species composition was an important factor in both models, confirming the treecreepers’ affinity for belah Casuarina pauper and slender cypress-pine Callitris gracilis-buloke Allocasuarina luehmannii woodlands in north-west Victoria. The first model emphasized the importance of demographic isolation: probability of patch occupancy decreased with distance to the nearest occupied patch. A threshold response in demographic isolation was apparent. In agricultural landscapes, most suitable woodland patches within 3 km of an occupied patch were occupied, whereas patches beyond the threshold were vacant. The threshold distance increased to a minimum of 8 km in a matrix of native vegetation, suggesting landscape context affects the response of white-browed treecreepers to habitat fragmentation. Demographic isolation is a quasi-dependent variable and therefore a second model was developed using surrogate variables for demographic isolation. A positive relationship with the proportion of woodland cover in the landscape (100 km²) emerged as the pre-eminent explanatory factor. Depending on woodland quality, a threshold of patch occupancy was apparent at levels of woodland cover between 15 and 25%. However, belah and slender cypress-pine-buloke woodlands now cover only 10% of their original extent in the region. These results highlight the inter-dependence of patch isolation with the amount and quality of habitat in the landscape and the implications this has for maintaining functional connectivity. The retention (or restoration) of suitable habitat is the critical issue for conservation of the white-browed treecreeper, but in landscapes below the threshold of habitat cover, viability of local populations may be influenced by the configuration and quality of remaining habitat.     

Habitat management and patterns of predation of Northern Lapwings on wet grasslands: The influence of linear habitat structures at different spatial scales

In managed landscapes, habitat structure is frequently manipulated through the creation of features such as tracks, hedges, and waterways. If predator and prey activity are concentrated around these features, levels of predation may be elevated in these landscapes. This issue is of particular importance when habitat structures are used to attract species of conservation concern. For example, the installation of linear waterways in wet grasslands is a common form of habitat management to benefit breeding waders and wader nests and foraging chicks tend to be aggregated around wet features. If predator activity is also focused around these features, and if their linearity increases the probability of prey being located, then the conservation benefits of this management technique may be eliminated. We explore predator movement in relation to the structure and complexity of linear wet features within a lowland wet grassland landscape. We examine patterns of nest and chick predation in lapwing (Vanellus vanellus) at the whole-site, between-field and within-field scales. Mammalian predators were responsible for the majority of nest predation. However, we found no evidence that mammalian predators used linear wet features disproportionately within the landscape, or that wet feature distribution influenced the probability of nest or chick predation. At the whole-site scale, nest predation rates were significantly higher in areas with greater predator presence and lowest where the number of breeding neighbours was high. Thus, predation levels were influenced by large-scale patterns of predator presence and lapwing density but not by the use of linear wet features as a habitat management tool. Managing predator impacts is therefore likely to require empirical assessments of local predator distribution and abundance in order to target measures effectively.     

Conspecific attraction in loggerhead shrikes: implications for habitat conservation and reintroduction

The loggerhead shrike, Lanius ludovicianus, is a declining songbird that forms breeding aggregations. Despite such reports from several populations, only one statistical analysis of loggerhead shrike territory distribution has been published to date. I use a spatio-temporal simulation technique to test for deviations from randomness in the spatial distribution of point data that takes into account date of nest establishment. I apply this model to data on the distribution of shrike nests in Comanche County, Oklahoma, USA, collected over the 1998 and 1999 breeding seasons. When the data are considered without regard to date of nest-establishment, the results are equivocal; nests are significantly aggregated in 1999, but not 1998. However, when order of nest establishment is taken into account, later nests are significantly more likely to be closer than expected to conspecifics in both years. This is true even when the distribution of simple resources, such as suitable nest-trees, is included in the model. These results are consistent with the hypothesis that loggerhead shrikes ‘prospect’ for suitable habitat using the distribution of breeding conspecifics. This aspect of loggerhead shrike breeding ecology should be considered for both habitat conservation and captive breeding and reintroduction programs for this species.     

Hunting habitat selection by hen harriers on moorland: Implications for conservation management

We examine habitat use by hunting hen harriers Circus cyaneus at three study sites in Scotland to evaluate whether foraging patterns differ between sexes, sites, and stages of the breeding period. We modelled time spent hunting in focal plots as a function of habitat and nest proximity. Male hunting intensity (time spent hunting per hour of observation and km²) varied between sites and breeding periods, being lower during the nestling than the incubation period. Habitat use patterns were mostly consistent among study sites, which is important for developing species management recommendations applicable over the species’ range. Males avoided improved grassland, and selected areas of mixed heather and rough grass (with an optimum at ca. 50% heather cover). The effect of nest proximity was small. In contrast, females hunted mainly within 300-500 m of the nest, with a small additive effect of vegetation cover, areas of fragmented heather being preferred. Habitat management to benefit foraging harriers will involve creating (or maintaining) mosaics of heather/grassland around nest areas. Additionally, it might be possible to manipulate habitat to reduce conflict in areas where harrier predation on red grouse is important by segregating areas holding highest grouse densities (with high heather cover) from those favoured for harrier foraging (heather-grass mosaics). However, it would be necessary to test whether these manipulations might also influence harrier nest distribution, an effect which could negate any benefits from this strategy.     

Landscape-level thresholds of habitat cover for woodland-dependent birds

Theory suggests that a disproportionate loss of species occurs when total habitat cover decreases to 10-30% of the landscape. To date, little empirical evidence has been collected to test for such thresholds in habitat cover, especially at the landscape scale. Here, we present empirical data on the species richness of woodland-dependent birds collected systematically from 24 landscapes (each 100 km²) that sample a gradient in habitat cover from <2% to 60%. To compare the relative effects of habitat cover and habitat configuration, landscapes with similar amounts of habitat but contrasting configuration (i.e., aggregated versus dispersed) were surveyed and the richness of woodland-dependent birds collated for each landscape. The relationship between species richness, habitat cover and habitat configuration was examined using analysis of co-variance (ANCOVA), multiple linear regression and univariate non-linear modelling. There was a significant effect of habitat cover (co-variate) in the ANCOVA, but the main treatment effect of configuration was not significant. However, comparison of non-linear models indicated that the shape of the response curve of species loss with decreasing habitat cover differed between aggregated and dispersed landscapes. Species richness was significantly related to habitat cover in all analyses, explaining between 55% and 60% of the variance in regression models. Mean patch shape complexity and the extent of habitat aggregation were also significant explanatory variables, but explained less than 10% of the variance in richness of woodland birds. Biogeographic variables (range in elevation and geographic location) explained up to 14% of the variance in species richness. There was strong evidence for a threshold response in species richness: non-linear models (broken-stick, exponential, inverse) exhibiting a sharp decline in species richness in landscapes with less than 10% habitat cover provided a better fit to the observed data than linear models. To our knowledge, this is the first empirical demonstration of landscape-level thresholds in species richness. We emphasise that thresholds in species richness denote multiple species’ extinction events, the end point of the process of species decline. For viable populations, habitat cover must be maintained well above the threshold level. Finally, thresholds of assemblage measures, such as species richness, potentially mask compositional changes in the avifauna community and may also conceal the loss of species with greater sensitivity to landscape change.     

Multi-scale patterns of habitat use by re-introduced mammals: A case study using medium-sized marsupials

Knowledge of the habitat requirements of threatened species at both local and landscape scales is crucial for maintaining viable populations and for making conservation and management decisions. Here, we use live trapping and radio-tracking to investigate habitat use by four species of threatened marsupials - burrowing bettongs (Bettongia lesueur), brush-tailed bettongs (B. penicillata), greater bilbies (Macrotis lagotis), and bridled nailtail wallabies (Onychogalea fraenata). The study populations had been re-introduced to Scotia Sanctuary in western New South Wales, Australia, within a predator-proof area. All showed preferences for particular macrohabitats while resting by day, with M. lagotis and B. penicillata selecting Eucalyptus woodland with Triodia understorey and B. lesueur and O. fraenata selecting Eucalyptus woodland with shrubs. However, they showed no such partiality at night. Bettongia penicillata used areas with Triodia and litter but few herbs for shelter, while burrows of M. lagotis avoided shrubs. Habitat components that influenced trap captures were: crust cover and herb layer cover (negative) for B. penicillata, trees <5 m in height and number of shrubs (both negative) for B. lesueur, crust cover for M. lagotis, and crust cover and trees <5 m high for O. fraenata (both negative). There was also a negative association at this scale between B. penicillata and both B. lesueur and M. lagotis, suggesting the possibility of competition. Our results support the idea that studies at multiple spatial scales are crucial to understand the habitat use and requirements of threatened fauna, and should therefore be incorporated into future re-introduction programs.     

Factors and scales potentially important for saproxylic beetles in temperate mixed oak forest

The influence of environmental factors on species richness and species composition may be manifested at different spatial levels. Exploring these relationships is important to understand at which spatial scales certain species and organism groups become sensitive to fragmentation and changes in habitat quality. At different spatial scales we evaluated the potential influence of 45 factors (multiple regression, PCA) on saproxylic oak beetles in 21 smaller broadleaved Swedish forests of conservation importance (woodland key habitats, WKH). Local amount of dead wood in forests is often assumed to be important, but two landscape variables, area of oak dominated woodland key habitats within 1 km of sites and regional amount of dead oak wood, were the main (and strong) predictors of variation in local species richness of oak beetles. The result was similar for red-listed beetles associated with oak. Species composition of the beetles was also best predicted by area of oak woodland key habitat within 1 km, with canopy closure as the second predictor. Despite suitable local quality of the woodland key habitats, the density of such habitat patches may in many areas be too low for long-term protection of saproxylic beetles associated with broadleaved temperate forests. Landscapes with many clustered woodland key habitats rich in oak should have high priority for conservation of saproxylic oak beetles.     

Landscape composition affects habitat use and foraging flight distances in breeding European starlings

The recent decline of the European starling in northern Europe has been associated with changes in agricultural land-use. We investigated how agricultural land-use affected starling habitat use and home-range size in a heterogeneous landscape in southern Sweden. Breeding starlings primarily foraged in pastures, a habitat with a high availability of potential food for nestlings. When the availability of pasture close to the nest was low, starlings increased their use of other permanently grass-covered habitats such as field margins and ditches. They also spent a larger proportion of their daytime flying because they visited more distant foraging areas. The further from the nest parents foraged, the higher was the probability that they foraged on a pasture. Feeding frequency of nestlings was negatively related to the distance from the nest parents foraged, but no consequences on nestling survival or growth were found. A lower breeding density at low local availability of pasture may compensate for lack of good foraging habitat. It is suggested that recent changes in the agricultural landscape may have increased the mismatch between the availability of good foraging areas and nest-sites. This might have contributed to the recent population decline of the starling in northern Europe by affecting both foraging habitat quality and flight costs paid by parents during breeding.     

Multi-scale effect of landscape processes and habitat quality on newt abundance: Implications for conservation

Recent studies in population dynamics suggest that landscape processes and habitat quality act at different scales on population abundances, but very few have modelled their simultaneous effects. However, at a time of large declines in natural populations, it is essential to understand such multivariate components. We tested the hypothesis that natural populations of palmate newts (Triturus helveticus) are affected on three scales: breeding patch (pond), habitat complementation (terrestrial cover), and metapopulation structure (density of ponds, surrounding populations). We conducted our survey in 130 ponds from southern France (Larzac) and analysed data with generalized additive models (GAM). Two main novel results emerge from these models: (1) the three landscape scales have significant effects on newt abundance, with more newts in deep, vegetated ponds, devoid of fish and surrounded by wooded areas and inhabited ponds; (2) the quality of the surrounding breeding patches is of primary importance in determining the abundance at core sites in a complex way: high abundances are associated positively with high densities of inhabited ponds, but negatively with the number of surrounding ponds. Deforestation, invasive species and abandonment of ponds all have negative impacts on the persistence of palmate newt populations. Future studies should encompass landscapes at different scales and incorporate the habitat quality in surrounding sites to better understand population dynamics and provide adequate conservation measures.     

Eucalypt plantations as habitat for birds on previously cleared farmland in south-eastern Australia

In parts of Australia, extensive areas of cleared land are now being planted with commercial plantations of native eucalypts. Questions arise about the extent to which such plantations can rectify previous loss of habitat and contribute to biodiversity conservation. This study assessed abundance of bird species (as one aspect of biodiversity) on 105 sites (25 cleared land, 58 plantations and 22 native forest) in two regions of rural Victoria, south-eastern Australia. Generalised linear modelling was used to assess some of the landscape and habitat variables that contributed to the value of plantation sites for particular groups of bird species. These models demonstrated the importance of on-site habitat variables in explaining the abundance of groups of bird species, with landscape context making small additional contributions.Mean abundance of forest and woodland birds was higher in eucalypt plantations than cleared farmland, and marginally lower than in native forest. Patterns differed between bird guilds. For example, insectivores that forage in the canopy and tall shrub layers were at least as common in plantations as in native forest, with birds in the latter group using young eucalypts as if they were tall shrubs. Birds that forage from open ground among trees were more common in plantations than native forest, and may benefit substantially from the new habitat fortuitously provided for them. This group includes several species that have declined in natural woodland habitats. Nectarivores, carnivores and birds that forage among low shrubs were less common in plantations than in native forest. Insectivores that forage from eucalypt bark made little use of plantations. Different approaches to plantation design and management would be needed to cater for groups such as these. Specific measures include planting of rough-barked eucalypts in addition to smooth-barked species, and provision of artificial hollows. Retention of existing remnants of native forest (e.g. old trees and forest patches) is a priority, to supply habitat elements that would otherwise be missing for long periods.     

At which spatial and temporal scales does landscape context affect local density of Red Data Book and Indicator species?

The landscape context is crucial for forest conservation in regions where the natural forest is fragmented. The focus of practical conservation is currently shifting from local stands to a landscape perspective, but few studies have tested the relative effect of different spatial and temporal scales for occurrence and persistence of species of conservation concern. We studied Red Data Book and Indicator species (the latter proposed to indicate presence of Red Data Book species) of vascular plants, lichens, bryophytes and wood-inhabiting fungi in 22 old temperate broadleaved forests in southern Sweden. We analysed at which scales these species respond to habitat proportion in surrounding landscape. The proportion of suitable habitat was measured at two temporal scales (present-day and historic) and at two spatial scales (about 0-1 km and 1-5 km of study sites). Local density of Red Data Book species increased with increasing proportion of suitable habitat in the current landscape (within 1-5 km of study sites) while Indicator species were unaffected. The response to landscape differed between organism groups. Vascular plants (near significantly) and wood-inhabiting fungi showed a time delay of 120 years in their response, indicating a possible regional extinction debt. An appropriate minimum landscape scale for conservation of Red Data Book species in temperate broadleaved forests in Sweden seems to be about 5 km (radius), but smaller landscapes may be important for vascular plants and wood-inhabiting fungi of conservation concern. In addition, restoration is urgent to counteract the effect of time delays in species responses to recent habitat loss.     

Testing the importance of spatial configuration of winter habitat for woodland caribou: An application of graph theory

Conservation of remaining woodland caribou (Rangifer tarandus caribou) populations requires land management strategies that not only maintain caribou habitat, but also favour habitat connectivity. This study presents and field tests graph theory-based measures of landscape connectivity, and demonstrate an association between the distribution of woodland caribou and well-connected winter habitat. Cost values for the intervening land cover types were determined based on the probability of selection relative to high-quality winter habitat. Habitat connectivity was then represented by linking high-quality habitat patches along least-cost paths through this parameterized cost surface. A randomization procedure was used to assess the animal’s association with habitat connected at increasing distance thresholds to identify appropriate scales of response. A strong relationship was obtained between large clusters of high-quality winter habitat patches and winter GPS telemetry location points (November 1-March 15) from two woodland caribou herds in Manitoba, Canada. This relationship was stronger when only late winter location points (January 1-March 15) were used. Our results highlight the importance of accounting for the spatial configuration of habitat on the landscape and the intervening land cover types when assessing range quality for woodland caribou. They also provide support for the use of graph theory to assist in identifying core activity areas for woodland caribou and key linkages between these areas and other parts of the landscape.     

The occurrence of primary burrowing crayfish in terrestrial habitat

A researcher’s perception of a target species’ landscape strongly influences the design of habitat studies conducted at broad spatial scales. Consequently, researcher-dependent perceptions may misguide conservation efforts. Although the life histories of some crayfish (i.e., primary burrowers) are centered on a fossorial existence independent of surface water, all North American crayfish are viewed in an aquatic context. This paradigm restricts the range of habitats that are typically sampled and managed for crayfish conservation. This study used presence/absence of the primary burrower Distocambarus crockeri at 137 locations within the Long Cane Ranger District of the Sumter National Forest, South Carolina, USA, to model the habitat association of the species across a GIS-based landscape. Logistic regression indicated that D. crockeri presence was most strongly associated with a terrestrial habitat defined by a set of morphologically similar soils located along ridge tops. Furthermore, the species was negatively associated with aquatic habitats such as streams and floodplains. The results indicate that D. crockeri is a terrestrial habitat specialist and should be modeled and managed at the landscape as a terrestrial organism. When viewed as a subset of the total United States cambarid fauna, primary burrowers are disproportionately imperiled. Primary burrowers comprise only 15% of the total crayfish fauna, while they account for 32% of those crayfish ranked critically imperiled. Habitat loss and an aquatic bias that restricted sampling to aquatic and semi-aquatic habitats might explain the group’s disproportionate imperilment.