Development and evaluation of predictive habitat models to assist the conservation planning of a threatened lucanid beetle, Hoplogonus simsoni, in north-east Tasmania

The use of predictive habitat distribution models by land managers in the conservation management of threatened species is increasing. Few models, however, are subsequently field-checked and evaluated. This study evaluates the statistical strength and usefulness for conservation purposes of three predictive habitat models developed for a threatened stag beetle, Hoplogonus simsoni, found in the wet eucalypt forests and mixed/rainforests of north-east Tasmania. The relationship between various environmental variables for which spatial (GIS) information was available and the density, frequency of occurrence and presence/absence of the species was investigated using generalised linear modelling. Models developed were coupled with the GIS data to develop maps of predicted occurrence within the species’ range, grouped into categories of habitat quality. The models found that altitude, aspect, slope, distance to nearest stream and overstorey tree height were significantly associated with the occurrence of the species. Evaluation of the statistical strength of the models with independent data of species’ occurrence collected at 95 sites found that the density model performed poorly with little correlation between predicted and observed densities of the species. The frequency of occurrence model, however, showed a moderate ability to predict both species’ abundance and presence/absence. The presence/absence model had a similar discriminatory ability in predicting presence or absence of H. simsoni, but also showed some potential as an indirect predictor of species’ abundance. Assuming a correlation between relative abundance and habitat quality, the frequency of occurrence predictive model appeared to be the better and more direct discriminator of high quality habitat relative to the other models. The value of species’ habitat models and the need to evaluate their utility in the development of conservation strategies are discussed.     

Modelling habitat suitability of the swamp antechinus (Antechinus minimus maritimus) in the coastal heathlands of southern Victoria, Australia

In recent years, predictive habitat distribution models, derived by combining multivariate statistical analyses with Geographic Information System (GIS) technology, have been recognised for their utility in conservation planning. The size and spatial arrangement of suitable habitat can influence the long-term persistence of some faunal species. In southwestern Victoria, Australia, populations of the rare swamp antechinus (Antechinus minimus maritimus) are threatened by further fragmentation of suitable habitat. In the current study, a spatially explicit habitat suitability model was developed for A. minimus that incorporated a measure of vegetation structure. Models were generated using logistic regression with species presence or absence as the dependent variable and landscape variables, extracted from both GIS data layers and multi-spectral digital imagery, as the predictors. The most parsimonious model, based on the Akaike Information Criterion, was spatially extrapolated in the GIS. Probability of species presence was used as an index of habitat suitability. A negative association between A. minimus presence and both elevation and habitat complexity was evidenced, suggesting a preference for relatively low altitudes and a vegetation structure of low vertical complexity. The predictive performance of the selected model was shown to be high (91%), indicating a good fit of the model to the data. The proportion of the study area predicted as suitable habitat for A. minimus (Probability of occurrence ?0.5) was 11.7%. Habitat suitability maps not only provide baseline information about the spatial arrangement of potentially suitable habitat for a species, but they also help to refine the search for other populations, making them an important conservation tool.     

Inferring species distributions in the absence of occurrence records: An example considering wolverine (Gulo gulo) and Canada lynx (Lynx canadensis) in New Mexico

Information about geographic distributions is required for species conservation and management. Ultimately, this information is derived from records of occurrence. However, the reliability and availability of occurrence records are variable. A conceptual framework for evaluating the reliability of occurrence records is provided. Only records associated with physical evidence, especially a museum voucher specimen, are considered verified. However, errors in species identification or location are possible even for verified records. In addition, biases exist in occurrence records because they generally are collected haphazardly. Other sources of bias include sampling error associated with small areas or range limits and aspects of the species’ biology that make it unlikely to be documented. A practical method is provided for interpreting a species’ distribution in a particular area given a paucity of reliable occurrence records. Factors that must be considered for including such areas of interest within the range of a species include: (1) plausible reason for the paucity of records; (2) continuous suitable habitat between the area of interest and localities of reliable occurrence; and (3) absence of biogeographic breaks in the distribution of other organisms with similar evolutionary histories. The possible distribution of wolverine (Gulo gulo) and Canada lynx (Lynx canadensis) in New Mexico provides a case study of this approach. It is concluded that the mountains of north-central New Mexico should be considered within the natural range of wolverine and Canada lynx.     

Incorporating uncertainty into analyses of red-crested pochard habitat selection

Studies describing habitat use in animal species need to take into account detectability of individuals in order to reach more robust conclusions. However, the importance of detectability in habitat selection analyses of rare and cryptic species has received little investigation, although robust methods for estimating detectability have been made recently available. Understanding habitat requirements should be an important management tool for the conservation of the red-crested pochard (Netta rufina), a rare duck species in France and Europe. Three different lake samples (82 lakes in total) were conjointly used in 2000 and 2001 to assess the species habitat requirements, using its presence, density and hatching dates as response variables. Risk of false absence was estimated using the double-observer approach at less than 0.001. A robust-design approach produced estimates of individual brood detectability (0.545, SE = 0.053). Observed red-crested pochard densities were adjusted to time dependent detectabilities, and modelled as a function of habitat variables. Habitat fragmentation and low permanent water levels negatively affected brood densities. Interestingly, these variables were not retained when modelling the unadjusted densities. This analysis showed that investigating temporal variation in brood detectability was a crucial prerequisite in the study of this rare species’ habitat requirements. More generally, it strongly suggests that integrating detection probability and its variations in habitat use analyses of cryptic species of conservation concern may be an essential methodological step to reach more valid conclusions on habitat management.     

Using ecological niche modelling to infer past, present and future environmental suitability for Leiopelma hochstetteri, an endangered New Zealand native frog

Leiopelma hochstetteri is an endangered New Zealand frog now confined to isolated populations scattered across the North Island. A better understanding of its past, current and predicted future environmental suitability will contribute to its conservation which is in jeopardy due to human activities, feral predators, disease and climate change. Here we use ecological niche modelling with all known occurrence data (N = 1708) and six determinant environmental variables to elucidate current, pre-human and future environmental suitability of this species. Comparison among independent runs, subfossil records and a clamping method allow validation of models. Many areas identified as currently suitable do not host any known populations. This apparent discrepancy could be explained by several non exclusive hypotheses: the areas have not been adequately surveyed and undiscovered populations still remain, the model is over simplistic; the species’ sensitivity to fragmentation and small population size; biotic interactions; historical events. An additional outcome is that apparently suitable, but frog-less areas could be targeted for future translocations. Surprisingly, pre-human conditions do not differ markedly highlighting the possibility that the range of the species was broadly fragmented before human arrival. Nevertheless, some populations, particularly on the west of the North Island may have disappeared as a result of human mediated habitat modification. Future conditions are marked with higher temperatures, which are predicted to be favourable to the species. However, such virtual gain in suitable range will probably not benefit the species given the highly fragmented nature of existing habitat and the low dispersal ability of this species.     

Unexpected high levels of genetic variability and the population structure of an island endemic rodent (Oryzomys couesi cozumelae)

Oryzomys couesi cozumelae is an endemic, threatened rodent from Cozumel Island, Mexico. We estimated its genetic diversity and structure by analyzing microsatellite loci in 228 samples from 12 sampling sites widely distributed throughout the island. Unexpected high levels of genetic and allelic diversity were found: a total of 54 alleles, an average of 10.8 alleles per locus, and high heterozygosity values (mean H_O = 0.624, H_E = 0.690 and H_Nei = 0.689). These values are higher than those reported for small sized insular mammals, higher than that found in 37 individuals of the mainland O. couesi from southern Mexico (H_O = 0.578) that we analyzed for comparative purposes, and similar to those of other mainland small mammal populations. Despite factors that affect Cozumel’s biota, such as exotic predators and competitors, hurricanes, seasonal population fluctuations and anthropogenic activities, no evidence of genetic bottlenecks was found. A significant population structure was observed and a model of isolation-by-distance was supported. Our findings render O. c. cozumelae a high conservation value, not only for its high genetic diversity and structure, but because available data suggests that its population has declined significantly in recent years. Further habitat fragmentation and population isolation could result in a higher genetic structure and loss of genetic diversity. The protection of habitat, the maintenance of habitat connectivity and the removal of introduced competitors and predators are a conservation priority. Acknowledging that the genetic structure of populations has crucial conservation implications, the present genetic information should be taken into account in management plans for the conservation of O. c. cozumelae.     

Endangered crayfish in northern Japan: Distribution, abundance and microhabitat specificity in relation to stream and riparian environment

Japan’s only native crayfish species Cambaroides japonicus has been declining dramatically in the past few decades. For the purpose of conservation planning, twenty-two coastal streams were surveyed to investigate summer distributions of crayfish in relation to stream and riparian environment. Classification and regression trees were used to predict the occurrence and abundance of crayfish. The classification tree model with stream variables as predictors showed that crayfish would occur in swift or high gradient streams (correct classification rate = 91%). Within those streams, however, crayfish only inhabited depositional microhabitats, in which the areas are limited in availability. Crayfish were not found in gentle, low gradient streams containing abundant depositional microhabitats. This paradoxical distribution pattern was attributed to availability of boulder substrates in swift or high gradient streams. The regression tree model indicated that crayfish abundance was determined primarily by the percentage of boulder substrates and the presence of fish (observed vs. predicted r = 0.64).The classification tree model using only riparian variables indicated that the total woody plant (mainly broadleaf species) density followed by the percentage of early successional species such as alder and willow determined the splits of the tree model (correct classification rate = 95%). A leaf processing experiment on 10 riparian plant species suggested that crayfish preferred high nitrogen (or low C/N) leaves.These results suggest that swift or high gradient fishless streams associated with abundant cover in dense broadleaf forest serve conservation areas for this endangered crayfish, and that consideration of riparian composition may facilitate conservation efforts.     

A habitat model for brown bear conservation and land use planning in the central Apennines

We investigated the brown bear habitat suitability in an 8000 km² study area encompassing Abruzzo, Latium, and Molise regions in central-southern Italy. Based on long-term field surveys and published records, we classified bear habitat as occupied or unoccupied in 92 out of 320 sample squares (5 × 5 km). For each sample square 36 habitat variables were measured from topographic maps and Corine land-cover III level digital maps. The influence of habitat features on bear presence was investigated by multivariate and one-way analyses of variance and by logistic regression analysis. The logistic model correctly classified 95.5% of sample squares of bear presence and 93.8% of those where bears were absent. Average altitude, deciduous woodlands and ecotone length, showed a positive relationship with bear presence, whereas vineyard-olive groves and shrublands were negatively correlated with bear presence. No specific land management guidelines or strategies exist for bear conservation in central Italy, based on knowledge of habitat-population relationships. The landscape scale habitat model we developed could be useful to predict bear occurrence, to identify critical areas for a brown bear conservation strategy, and to enhance the arrangement of the protected areas network for the conservation of this species.     

Impacts of land use, anthropogenic disturbance, and harvesting on an African medicinal liana

African medicinal plant species are increasingly threatened by overexploitation and habitat loss, but little is known about the conservation status and ecology of many medicinal species. Mondia whitei (Apocynaceae, formerly Asclepiadaceae), a medicinal liana found in Sub-Saharan Africa, has been subject to intensive harvesting and habitat loss. We surveyed M. whitei in Kakamega Forest, the largest of three remnant Kenyan forests known to contain the species. In 174 100 m² plots, we quantified the status of M. whitei and investigated its relationships with land use, disturbance and harvesting. With average adult densities of 101 plants/ha, M. whitei is not locally rare in Kakamega. However, the absence of flowers and fruits, together with a spatial disconnect between adults and juveniles, suggests that sexual regeneration is patchy or infrequent. Comparing among habitat types, we found that plants were most abundant in regenerating indigenous forest managed by the Forest Department, which permits some extractive uses. Conversely, plants were largest in indigenous forest managed by the Kenya Wildlife Service, which prohibits extractive uses. Most anthropogenic disturbances were not associated with M. whitei, but plant occurrence and density were higher along paths used by livestock than along other types of paths. Larger individuals appeared to be preferentially harvested, but adult plants were more likely to occur in harvested plots than un-harvested plots. This work emphasizes that restrictions on disturbance and extractive use do not automatically promote medicinal plant conservation. Moreover, harvesting may have important genetic and demographic consequences that are overlooked by studies focused on numerical losses.     

Statewide modeling of bobcat, Lynx rufus, habitat in Illinois, USA

We used sighting location and remotely sensed habitat data, multivariate statistical techniques, and a geographic information system to model bobcat (Lynx rufus) habitat in Illinois, thereby providing state wildlife managers with information to review the listing of bobcats as a state-threatened species and contribute to the development of a statewide management plan. We used canonical discriminant function analysis to model presence/absence and relative abundance of bobcats statewide. These models suggested that bobcats occurred in moderate to high abundance in 23 of 98 counties statewide (23%). We used stepwise logistic regression (SLR) analysis to model statewide habitat suitability. Spatial modeling of the SLR equation predicted that 29% of Illinois contained suitable habitat classified as P>0.50. Models were accurate when validated with an independent data set and indicated the importance of woods-related habitat variables to bobcats. In conclusion, these models provided tools to rapidly assess status that contributed to delisting bobcats as a threatened species in Illinois and provided further information to guide conservation efforts.     

Habitat correlates of distribution and local population decline of the endemic Sardinian newt Euproctus platycephalus

Habitat-based statistical models relating patterns of presence and absence of species to habitat variables could be useful to resolve conservation-related problems and highlight the causes of population declines. In this paper, we apply such a modelling approach to an endemic amphibian, the Sardinian mountain newt Euproctus platycephalus, considered by IUCN a critically endangered species. Sardinian newts inhabit freshwater habitat in streams, small lakes and pools on the island of Sardinia (Italy). Reported declines of newt populations are not yet supported by quantitative data, however, they are perceived or suspected across the species' historical range. This study represents a first attempt trying to statistically relate habitat characteristics to Sardinian newt occurrence and persistence. Linear regression analysis revealed that newts are more likely to be found in sites with colder water temperature, less riparian vegetation and, marginally, absence of fish. The implications of the results for the conservation of the species are discussed, and suggestions for the short-term management of newt inhabited sites suggested.     

Assessing critical habitat: Evaluating the relative contribution of habitats to population persistence

A principal challenge of species conservation is to identify the specific habitats that are essential for long-term persistence or recovery of imperiled species. However, many commonly used approaches to identify important habitats do not provide direct insight into the contribution of those habitats to population persistence. To assess how habitats contribute to overall population viability and characterize their relative importance, a spatially-explicit population viability model was used to integrate a species occurrence model with habitat quality and demographic information to simulate the population dynamics of the Ord’s kangaroo rat (Dipodomys ordii) in Alberta, Canada. Long-term productivity (births-deaths) in each patch was simulated and iterative patch removal experiments were conducted to generate estimates of the relative contribution of habitat types to overall population viability. Our results indicated that natural dune habitats are crucial for population viability, while disturbed/human-created habitats make a minor contribution to population persistence. The results also suggest that the habitats currently available to Ord’s kangaroo rats in Alberta are unlikely to support long-term persistence. Our approach was useful for identifying habitats that did not contribute to population viability. A large proportion of habitat (39%) represented sinks and their removal increased estimated population viability. The integration of population dynamics with habitat quality and occurrence data can be invaluable when assessing critical habitat, particularly in regions with variable habitat quality. Approaches that do not incorporate population dynamics may undermine conservation efforts by under- or over-estimating the value of habitats, erroneously protecting sink habitats, or failing to prioritize key source habitats.     

The effects of land use and climate on red kite distribution in the Iberian peninsula

The main breeding populations of the red kite (Milvus milvus L.), have been declining in the Iberian peninsula during the last decade. However, there is a lack of regional assessments of habitat suitability that identifies limiting ecological factors for the species and areas with conservation problems. In this work we present a regional model for the distribution and abundance of breeding red kites in the Iberian peninsula. The occurrence and estimated abundance in 100 km² UTM squares resulting from road censuses were modelled with broad-scale explanatory variables obtained from satellite imagery, thematic digital cartography, climatic data and spatial coordinates. The occurrence model incorporated mainly climatic variables and had a good discrimination ability, while the abundance model incorporated mainly land-use variables and had a lower explanatory power (r²=0.14). The predictions somewhat overestimated the results of the censuses, and this agrees with the decline of population size and range observed for this species in the Iberian peninsula. These models are relevant in the conservation of the species: first, they suggest the limiting factors for red kite in the Iberian peninsula, and, second, they generate predictive maps pointing out both areas in which conservation problems may be acute (suitable locations that are unoccupied), and areas where no data is available but the red kite is likely to be present (thus guiding further survey and research).     

Habitat relations of Rhea americana in an agroecosystem of Buenos Aires Province, Argentina

The maintenance of wild populations of Greater Rhea (Rhea americana) through effective management requires an understanding of their habitat requirements in terms of vegetation composition and field configuration. We studied the relative influence of some anthropogenic variables (presence of route, house and fences) and resource variables (presence of water source, composition and coverage of plant species, vegetation height and bare soil), on the habitat use by a population of rheas in a cattle ranch of Buenos Aires Province, Argentina. Habitat use was determined indirectly by documenting the number of faeces in summer, autumn-winter and spring 1999. The presence or absence of faeces was related to the measured variables through discriminant analysis that allowed the elaboration of predictive models of habitat use by this species. Contrary to what was expected, those variables related to human activity showed a low predictive value on the habitat use by rheas when compared with resource variables. Rheas preferentially selected the stream area in all seasons and sites with great percent cover of Bupleurum sp., Phyla canescens, Sida leprosa, Plantago lanceolata, Trifolium repens, Lolium multiflorum, Stipa spp., and Stenotaphrum secundatum. Low vegetation height was another important component of rhea's habitat in summer and autumn-winter. The high accuracy level obtained by validation tests of this model supports its utility for the management of rhea populations in other cattle ranches of the region, and to analyze the suitability of other ranches for reintroduction programs.     

Modelling habitat and spatial distribution of an endangered longhorn beetle - A case study for saproxylic insect conservation

Saproxylic insects are characterised by their exceptional diversity and high proportion of threatened species. No recent studies have demonstrated the validity of habitat suitability analysis for scientifically based habitat management for these species. We studied the habitat requirements of the endangered longhorn beetle Cerambyx cerdo, a species with a supposed keystone function for the saproxylic insect community living on oaks. We used species distribution modelling based on datasets from Central Europe to understand the species-habitat relationships and to find the environmental variables responsible for habitat selection of C. cerdo. Our results show that the most important parameters, insolation, presence of oak sap, bark depth and the distance from the next colonised tree, are able to predict the presence of C. cerdo very well. A spatial validation procedure revealed very similar predictive power, indicating the general validity of our model. Tree-level parameters were shown to have a stronger effect on the occurrence probability than landscape-level predictors. To improve the tree-level conditions (e.g. insolation on the trunk) habitat management in the form of semi-open pasture landscapes is recommended from which many other taxa will also draw considerable benefit. The provision of such conditions over decades is the essential key in the conservation of this longhorn beetle species. The success of the European network of conservation areas “Natura 2000” heavily depends on broad biological knowledge of the designated protected species. The present paper shows that species distribution models can give valuable contributions for conservation in saproxylic insects.     

Building large-scale spatially explicit models to predict the distribution of suitable habitat patches for the Greater rhea (Rhea americana), a near-threatened species

We developed large-scale spatially explicit models to predict the distribution of suitable habitat patches for the Greater rhea (Rhea americana), a near-threatened species, in two areas of central Argentina with different land use: a grassland area (ca. 4943 km²) mainly devoted to cattle grazing and an agro-ecosystem area (ca. 4006 km²) mostly used for crop production. The models were developed using logistic regression and were based on current records of Greater rhea occurrence coupled with remote sensing data, including land cover and human presence variables. The habitat suitability maps generated were used to predict the suitable habitat patch structure for wild rhea populations in each area. Fifty-one percent of the total grassland area was suitable for the species, being represented by a single large patch that included 62% of the individual locations. In the agro-ecosystem, only 28% of the total area was suitable, which was distributed among four patches. Seventy percent of rhea observations were in suitable habitat, with all rheas grouped in the largest patch. Conservation efforts for preserving wild rhea populations should be focused on maintaining habitats similar to grasslands, which are less profitable for landowners at present. Consequently, the protection of the pampas grasslands, a key habitat for this species as well as for others with similar habitat requirements, will demand strong conservation actions through the reconciliation of interests between producers and conservationists, since the proportion of croplands is increasing.     

Predicting loss and fragmentation of habitat of the vulnerable subtropical rainforest tree Macadamia integrifolia with models developed from compiled ecological data

Habitat models are widely used in ecology, however there are relatively few studies of rare species, primarily because of a paucity of survey records and lack of robust means of assessing accuracy of modelled spatial predictions. We investigated the potential of compiled ecological data in developing habitat models for Macadamia integrifolia, a vulnerable mid-stratum tree endemic to lowland subtropical rainforests of southeast Queensland, Australia. We compared performance of two binomial models—Classification and Regression Trees (CART) and Generalised Additive Models (GAM)—with Maximum Entropy (MAXENT) models developed from (i) presence records and available absence data and (ii) developed using presence records and background data. The GAM model was the best performer across the range of evaluation measures employed, however all models were assessed as potentially useful for informing in situ conservation of M. integrifolia, A significant loss in the amount of M. integrifolia habitat has occurred (p < 0.05), with only 37% of former habitat (pre-clearing) remaining in 2003. Remnant patches are significantly smaller, have larger edge-to-area ratios and are more isolated from each other compared to pre-clearing configurations (p < 0.05). Whilst the network of suitable habitat patches is still largely intact, there are numerous smaller patches that are more isolated in the contemporary landscape compared with their connectedness before clearing. These results suggest that in situ conservation of M. integrifolia may be best achieved through a landscape approach that considers the relative contribution of small remnant habitat fragments to the species as a whole, as facilitating connectivity among the entire network of habitat patches.     

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.     

Biogeography and conservation of taxa from remote regions: An application of ecological-niche based models and GIS to North-African canids

In North Africa, and especially in the Sahara Desert, biodiversity is poorly known. Of the five widespread canid species present, one is Data Deficient, three are considered widespread although habitat selection could limit their area of occupancy, and distribution maps available are coarse for conservation planning. This study identifies biogeographic patterns in North-African canids through the combination of high resolution presence data with 16 environmental factors. Predictive models trained in north-west Africa are projected to all North Africa. canids exhibited distinct biogeographical affinities. GIS tools and Maximum Entropy models identify a mixture of climatic and habitat factors as main predictors of species occurrence. Suitable habitats for North-African canids are mostly fragmented: probable occurrence was identified for Canis aureus in Saharan peripheral regions and mountains, for Vulpes pallida in a narrow band along the Sahel and in southern Saharan mountains, for Vulpes rueppellii throughout the Sahara, for Vulpes vulpes in northern Africa until the Sahara northern limit, and for Vulpes zerda in almost all Sahara. Areas of potential sympatry between species with similar niches and parapatric ranges are identified along relatively narrow bands. The small pixel size of projections allows the identification of suitable refuges for species otherwise absent in the driest Saharan habitats, providing framework data for the definition of the global conservation status of V. pallida, and conservation strategies for the guild. The biological value of Saharan mountains is emphasised as they constitute isolated suitable areas. Ecological-niche based models should be developed for other endangered Saharan vertebrates.