The use of spatial ecological modelling as a tool for improving the assessment of geographic range size of threatened species

We analysed endemic threatened tree and reptile species of Socotra Island (Yemen), characterised by different ecological requirements and spatial distribution, in order to evaluate the usefulness of spatial ecological modelling in the estimation of species extent of occurrence (EOO) and area of occupancy (AOO). Point occurrences for the entire species range were used to model their spatial distribution by Random Forest (RF) and Generalised Linear Model (GLM). For each species the suitability area (SA) was obtained by applying the 0% omission error criterion on the probability map, and compared or integrated with EOO and AOO area obtained by topological methods such as the minimum convex polygon (MCP), α-hull and 2 km × 2 km grid.RF showed a lower prediction error than GLM. Higher accuracy was achieved for species with higher number of occurrences and narrower ecological niche. SA was always greater than AOO measured with the 2 km × 2 km grid method. SA was greater than EOO, measured by both MCP and α-hull methods, for species with localised distribution, while it was smaller for widely distributed species. EOO-α-hull area was equal or smaller than that calculated by MCP depending on the spatial distribution of species. AOO measured considering the SA within the EOO-MCP was greater than that measured using the standard 2 km × 2 km grid. Conversely, AOO calculated considering the suitable area within the EOO-α-hull showed variable results, being smaller or greater than the 2 km × 2 km grid AOO depending on the ecological niche and spatial distribution of species. According to our results, SEM does not provide an effective alternative to topological methods for the estimate of EOO and AOO. However, it may be considered a useful tool to estimate AOO within the boundaries of EOO measured by the α-hull method, because it reduces some potential sources of inconsistency and bias.     

Scale dependency and area of occupancy: Tortula freibergii in north-west England

Abstract

Area of occupancy (AOO) is used commonly as a measure of species range size and the IUCN Red List Criteria provide thresholds of AOO for determining the extinction risk of species. Tortula freibergii is a rare moss globally and is considered to be a priority for conservation in the UK. This study provides the results of a comprehensive survey of the distribution of the species in north-west England, at a linear resolution of 0.5 km, which shows it to be considerably more widespread than understood previously. The data are used to create a 'species-area curve' that spans the scales of 0.1 to 10 km and examine how measures of AOO change according to the scale of measurement. AOO values declined sharply as the scale of measurement reduced, as a result of the linear and frequently fragmented distribution of the species. Implications for the application of the IUCN Red List Criteria and the monitoring of species are discussed.     

A new method to estimate areas of occupancy using herbarium data

The World Conservation Union (IUCN) Red List of Threatened Species is an important instrument to evaluate the conservation status of living organisms. However, Red List assessors have been limited by the lack of reliable methods to calculate the area of occupancy (AOO) of species, which is an important parameter for red list assessments. Here we present a new practical method to estimate AOO based on herbarium specimen data: the Cartographic method by Conglomerates (CMC). This method, which combines elements from the Areographic and Cartographic methods previously used to calculate AOO, was tested with ten cactus species from the Chihuahuan Desert Region. The results derived from this novel procedure produced in average AOO calculations 3.5 and 5.5 smaller than the Areographic and Cartographic methods, respectively. The CMC takes into account the existence of disjunctions in the distribution range of the species, producing comparatively more accurate AOO estimations. Another advantage of the CMC is that it generates results more harmonic with the current Red List criteria. In contrast, the overestimated results of the Areographic and Cartographic methods tend to artificially categorize the species, even extremely narrow endemics, in lower endangerment status.     

Threats and biodiversity in the mediterranean biome

Aim Global conservation assessments recognize the mediterranean biome as a priority for the conservation of the world's biodiversity. To better direct future conservation efforts in the biome, an improved understanding of the location, magnitude and trend of key threats and their relationship with species of conservation importance is needed. Location Mediterranean‐climate regions in California‐Baja California, Chile, South Africa, Australia and the Mediterranean Basin. Methods We undertook a systematic, pan‐regional assessment of threats in the mediterranean biome including human population density, urban area and agriculture. To realize the full implications of these threats on mediterranean biodiversity, we examined their relationship with species of conservation concern: threatened mammals at the global scale and threatened plants at the subecoregional scale in California, USA. Results Across the biome, population density and urban area increased by 13% and agriculture by 1% between 1990 and 2000. Both population density and urban area were greatest in California‐Baja California and least in Australia while, in contrast, agriculture was greatest in Australia and least in California‐Baja California. In all regions lowlands were most affected by the threats analysed, with the exception of population density in the Chilean matorral. Threatened species richness had a significant positive correlation with population density at global and subecoregional scales, while threatened species were found to increase with the amount of urban area and decrease as the amount of natural area and unfragmented core area increased. Main conclusions Threats to mediterranean biodiversity have increased from 1990 to 2000, although patterns vary both across and within the five regions. The need for future conservation efforts is further underlined by the positive correlation between species of conservation concern and the increase in population density over the last decade. Challenges to reducing threats extend beyond those analysed to include human–environmental interactions and their synergistic effects, such as urbanization and invasive species and wildfires.     

Improving the estimation of area of occupancy for IUCN Red List assessments by using a circular buffer approach

The area of occupancy (AOO) is one of the main measures used by IUCN to quantify range size for species. AOO represents the area of suitable habitat currently occupied by the taxon and is usually quantified by counting the number of occupied cells in a uniform grid that covers the entire range of a taxon. However, this methodology adds uncertainty by the location of the origin of the grid frame. In this communication paper, we tested the influence of the origin of the grid frame used to quantify AOO and found for Swiss bryophytes that 14 species (out of 1089) fall into a different Red List category when the origin of the grid frame was shifted. With this and theoretical examples we show that AOO quantified by circles around the occurrences (a circular buffer approach) would reduce uncertainty significantly because they are independent of the origin of a grid frame. A circular buffer approach to quantify AOO contribute thus to more robust and accurate Red Lists and its usage is in accordance with the IUCN criteria.     

Using species distribution and occupancy modeling to guide survey efforts and assess species status

Habitat loss and fragmentation continue to be major issues affecting the persistence and conservation of species, but identification of critical habitat remains a challenge. Species distribution modeling and occupancy modeling are both approaches that have been used to predict species distributions and can identify critical habitat characteristics associated with species occurrence. Additionally, occupancy sampling can provide measures of detectability, increasing the confidence that a species is truly absent when not detected. While increasingly popular, these methods are infrequently used in synergy, and rarely at fine spatial scales. We provide a case study of using distribution and occupancy modeling in unison to direct survey efforts, provide estimates of species presence/absence, and to identify local and landscape features important for species occurrence. The focal species for our study was Ambystoma jeffersonianum, a threatened salamander in the state of Illinois, U.S.A. We found that fine-scale distribution models accurately discriminated occupied from unoccupied breeding ponds (78–91% accuracy), and surveys could be effectively guided using a well-fit model. We achieved a high detection rate (0.774) through occupancy sampling, and determined that A. jeffersonianum never used ponds inhabited by fish, and the probability of a pond being used for breeding increased as canopy cover increased. When faced with limited resources, combining fine-scale distribution modeling with a robust occupancy sampling design can expedite survey efforts, confidently designate species occupancy status, prioritise habitat for future surveys and/or restoration, and identify critical habitat features. This approach is broadly applicable to other taxa that have specific habitat requirements.     

The desolation of Smaug: The human-driven decline of the Sungazer lizard (Smaug giganteus)

The Sungazer (Smaug giganteus) is a threatened lizard species endemic to the Highveld grasslands of South Africa. The species faces risks from habitat loss and fragmentation, and illegal harvesting for traditional medicine and the pet trade. Despite these threats, the current conservation status of the species was poorly validated. We visited 79 Sungazer populations recorded in 1978 to assess population change since the initial surveys, and surveyed an additional 164 sites to better define the distribution and estimate the current population size. We interrogated all known historical trade data of the species. One-third of Sungazer populations have been extirpated over the past 37 years. The distribution includes two allopatric populations, with the smaller Mpumalanga population experiencing a significantly higher decline. The species has an extent of occurrence (EOO) of 34 500 km², and an area of occupancy (AOO) of 1149 km². The interpreted distribution is 17 978 km², and just under 60% remains untransformed grassland. We estimate a population size of 677 000 mature individuals, down 48% from the estimated historical population, prior to commercial agricultural development. A total of 1194 live Sungazers were exported under permit from South Africa between 1985 and 2014, with a significant increase in numbers exported over the last decade. Without any evidence of captive breeding, we believe that these animals are all wild-caught. Based on the AOO, level of decline, fragmentation within the distribution and suspected level of exploitation, we recommend classification of the species as Vulnerable under IUCN Red List Criteria A2acd and B2ab(ii–v). The establishment of a protected area network, genetic research and further investigations into the pet and traditional medicine trades are urgently needed.     

Understanding bias in geographic range size estimates

Aim Estimates of geographic range size derived from natural history museum specimens are probably biased for many species. We aim to determine how bias in these estimates relates to range size. Location We conducted computer simulations based on herbarium specimen records from localities ranging from the southern United States to northern Argentina. Methods We used theory on the sampling distribution of the mean and variance to develop working hypotheses about how range size, defined as area of occupancy (AOO), was related to the inter‐specific distribution of: (1) mean collection effort per area across the range of a species (MC); (2) variance in collection effort per area across the range of a species (VC); and (3) proportional bias in AOO estimates (PBias: the difference between the expected value of the estimate of AOO and true AOO, divided by true AOO). We tested predictions from these hypotheses using computer simulations based on a dataset of more than 29,000 herbarium specimen records documenting occurrences of 377 plant species in the tribe Bignonieae (Bignoniaceae). Results The working hypotheses predicted that the mean of the inter‐specific distribution of MC, VC and PBias were independent of AOO, but that the respective variance and skewness decreased with increasing AOO. Computer simulations supported all but one prediction: the variance of the inter‐specific distribution of VC did not decrease with increasing AOO. Main conclusions Our results suggest that, despite an invariant mean, the dispersion and symmetry of the inter‐specific distribution of PBias decreases as AOO increases. As AOO increased, range size was less severely underestimated for a large proportion of simulated species. However, as AOO increased, range size estimates having extremely low bias were less common.     

South African mouse shrews (<Emphasis Type="Italic">Myosorex</Emphasis>) feel the heat: using species distribution models (SDMs) and IUCN Red List criteria to flag extinction risks due to climate change

Five species of mouse or forest shrews (Myosorex) are endemic to South Africa, Lesotho and Swaziland, four of which (Myosorex varius, Myosorex cafer, Myosorex longicaudatus and Myosorex cf. tenuis) are associated with montane or temperate grassland, fynbos and/or forest habitats while a fifth (Myosorex sclateri) is associated with lowland subtropical forests. Due to their small size, specialised habitat, low dispersal capacity, high metabolism and sensitivity to temperature extremes, we predicted that, particularly for montane species, future climate change should have a negative impact on area of occupancy (AOO) and ultimately extinction risks. Species distribution models (SDMs) indicated general declines in AOO of three species by 2050 under the A1b and A2 climate change scenarios (M. cafer, M. varius, M. longicaudatus) while two species (M. sclateri and M. cf. tenuis) remained unchanged (assuming no dispersal) or increased their AOO (assuming dispersal). While temperate species such as M. varius appear to be limited by temperature maxima (preferring cooler temperatures), the subtropical species M. sclateri appears to be limited by temperature minima (preferring warmer temperatures). Evidence for declines in AOO informed the uplisting (to a higher category of threat) of the Red List status of four Myosorex species to either vulnerable or endangered as part of a separate regional International Union for Conservation of Nature (IUCN) Red List assessment.     

Predicting spatial occurrence of beetles and pseudoscorpions in hollow oaks in southeastern Sweden

We modelled presence/absence per tree of beetles and pseudoscorpions living in tree hollows in relation to trunk circumference, habitat openness, and connectivity (= density of hollow oaks in the surrounding area), using data from 281 oaks. The presence/absence models were then used to predict species’ occurrences in a county (11,600 km²) in southeastern Sweden. For eight of the nine species, the most parsimonious occupancy model included a positive relationship with connectivity and at least one tree characteristic. Occupancy underestimates from occurrence records—the ratio of the area of occupancy based on our predictive model to the area of occupancy based on occurrence records—varied between 3 and 83 among species when using occurrence records up to 1993, with significantly larger underestimates for smaller beetle species. Today (after extensive surveys), underestimation has decreased to 1.3–25, confirming that calculations solely based on species occurrence records greatly underestimate the area of occupancy. We suggest this should be taken into account to a greater extent and in a clearer way than today when constructing red lists. The radius of the connectivity measure that generated the best fit varied between 135 and 2,857 m among species, with longer distances for more threatened species. Consequently, preservation of the most threatened species (Elater ferrugineus and Tenebrio opacus) requires conservation efforts at larger spatial scales than required to protect Osmoderma eremita, which frequently has been used as an indicator and umbrella species.     

The trait and host plant ecology of aphids and their distribution and abundance in the United Kingdom

Aim We characterized the annual populations of 170 aphid species by their log abundance, site occupancy and site continuity (i.e. the persistence of species in time) and used this information to make predictions about groups of species that displayed characteristic patterns. By doing so, we aimed to identify commonalities in functional traits (host‐alternation; mode of reproduction; life‐cycle plasticity; median body size) and host plant geographic range sizes that may indicate why some species are common and others not. Location The population dynamics of winged aphids at 27 locations in the United Kingdom were studied. Methods The annual numbers of aphids were studied using a dataset comprising over 11 million individuals across 509 site‐years. Traits and host plants were analysed using linear mixed effects models and nonlinear regression models. Results Linear mixed effects models showed that the fixed effects of host alternation and winter host plant area of occupancy were important in predicting log abundance, site occupancy and site continuity. Life‐cycle plasticity was also a significant effect, although not for log abundance. Relationships between site continuity, site occupancy and log abundance were strongly nonlinear. Site continuity always lagged site occupancy, indicating that species were less likely to retain previously occupied sites when abundances were low. Main conclusions Aphid traits are a better paradigm than taxonomic relatedness in explaining macroecological patterns. Host alternation induces an annual flux of migrants that engenders higher annual log abundances, consistent with the theory that species with high local densities tend to confer a much wider distribution than those with low densities. The abundance of aphids is monotonically related to the geographic range size of their winter host, suggestive of a strong bottom‐up effect (i.e. resource controlled). The areas of occupancy (AOO) of winter host plants constrain aphids to low abundances because a greater proportion of hosts are trees and shrubs which tend to have smaller AOO than herbs and grasses, the common summer host types.     

Spatial mismatches between plant biodiversity facets and evolutionary legacy in the vicinity of a major Mediterranean city

The analyses of congruencies among biodiversity components address the issue of conservation priorities, but previously they have been done at coarse scales with limited relevance for conservation actions. Moreover, these former studies consider only the species level components of biodiversity and not the intra-specific evolutionary legacy that influences future biodiversity. This study represents the first assessment of congruencies between various components of plant biodiversity and the evolutionary legacy of a narrow endemic taxon (Arenaria provincialis, Caryophyllaceae). Assessment is conducted in the vicinity of a Mediterranean big city (Marseille, S.E. France) where habitats and flora are threatened by mass tourism and urban sprawl. Our analyses reveal that the different plant biodiversity facets assessed are spatially mismatched and unequally protected. Moreover, by using only species-level components of biodiversity as conservation targets we ignore crucial areas for the evolutionary legacy of this narrow endemic plant. Our results highlight the crucial role of phylogeography as a criterion to target the genetic precursors of future biodiversity in conservation planning.     

Discrepancies in occupancy and abundance approaches to identifying and protecting habitat for an at‐risk species

Predicting how environmental factors affect the distribution of species is a fundamental goal of conservation biology. Conservation biologists rely on species distribution and abundance models to identify key habitat characteristics for species. Occupancy modeling is frequently promoted as a practical alternative to use of abundance in identifying habitat quality. While occupancy and abundance are potentially governed by different limiting factors operating at different scales, few studies have directly compared predictive models for these approaches in the same system. We evaluated how much occupancy and abundance are driven by the same environmental factors for a species of conservation concern, the greater short‐horned lizard (Phrynosoma hernandesi). Occupancy was most strongly dictated by precipitation, temperature, and density of ant mounds. While these factors were also in the best‐supported predictive models for lizard abundance, the magnitude of the effects varied, with the sign of the effect changing for temperature and precipitation. These discrepancies show that while occupancy modeling can be an efficient approach for conservation planning, predictors of occupancy probability should not automatically be equated with predictors of population abundance. Understanding the differences in factors that control occupancy versus abundance can help us to identify habitat requirements and mitigate the loss of threatened species.     

The importance of defining the geographic distribution of species for conservation: The case of the Bearded Wood-Partridge

Delimitation of the distribution areas of species has fundamental implications for the understanding of biodiversity and for decision-making in conservation. This is illustrated by the case of the Bearded Wood-Partridge (Dendrortyx barbatus), which is endemic to Mexico and was classified as threatened by the IUCN. Recently the discovery of this species in new locations caused an increase in the known distribution area whereupon it was reclassified in a lower risk category. In our study, delimitation and comparison of the Bearded Wood-Partridge distribution area is carried out utilising five different methods: minimum convex polygon; areographic; cartographic; ecological niche modeling; and, “free hand”. A number of locality records are also used to demonstrate the chronological order of appearance. The results show that the size and shape of the distribution area of this species vary depending on the number of records and on their spatial and environmental location, as well as on the particular delimitation method used. However, ecological niche modeling provides the best results in terms of spatial and numerical sensitivity as well as lower values of omission and a moderate extent of predicted areas. We suggest that decisions related to species conservation (categories of risk, areas of endemism, etc.), particularly those species of high geographical restriction, should be contingent on the formalised delimitation of distribution areas based on ecological niche modeling methods.     

The larval ecology of the butterfly Euphydryas desfontainii (Lepidoptera: Nymphalidae) in SW-Portugal: food plant quantity and quality as main predictors of habitat quality

Corresponding to theory, the persistence of metapopulations in fragmented landscapes depends on the area of suitable habitat patches and their degree of isolation, mediating the individual exchange between habitats. More recently, habitat quality has been highlighted as being equally important. We therefore assess the role of habitat area, isolation and quality for the occupancy of larval stages of the regionally threatened butterfly Euphydryas desfontainii occurring in grassland habitats comprising the host plant Dipsascus comosus. We put a special focus on habitat quality which was determined on two spatial scales: the landscape (among patches) and the within-patch level. On the landscape level, occupancy of caterpillars was determined by a presence-absence analysis at 28 host plant patches. On the within-patch level, oviposition site selection was studied by comparing 159 host plants with egg clutches to a random sample of 253 unoccupied host plants within six habitat patches. The occupancy of caterpillars and presence of egg clutches on host plants was then related to several predictors such as patch size and isolation on the landscape level and host plant characteristics and immediate surroundings on the within patch level. On the landscape level, only host plant abundance was related to the presence of caterpillars, while size and isolation did not differ between occupied and unoccupied patches. However, the weak discrimination of larval stages among patches changed on the within-patch level: here, several microclimatic predictors such as sunshine hours and topography, host plant morphology and phenology as well as further potential host plants in the immediate surroundings of the plant chosen for oviposition strongly determined the presence of egg clutches. We strongly suggest promoting the presence of the host plant in topographically and structurally rich habitat patches to offer potential for microclimatic compensation for a species considered threatened by climate change.     

Climax epiphytic communities in Mediterranean Spain

The Antitrichetalia curtipendulae epiphytic climax communities of Šmarda & Hadàč from the Spanish Mediterranean area have been classified by numerical and traditional floristic methods. Ninety-eight phytosociological relevés have been recorded from a number of Iberian forests. Our results indicate two clusters that can be easily related to two different alliances: Lobarion pulmonariae (Nephrometum resupinati) and Pterogonio gracile-Antitrichion californicae (Antitrichielum californicae). A phytogeographical and syntaxonomical discussion is followed by a suggested Red List of threatened lichens and bryophytes occurring in these communities in the Spanish Mediterranean area with the aim of advancing conservation of endangered and vulnerable species.     

Are there contrasted impacts of urbanization and land uses on population persistence? The case of Teucrium pseudochamaepitys, an endangered species in Southern France

Among the endangered flora of the Mediterranean basin, Teucrium pseudochamaepitys, endemic Lamiaceae, is threatened by human activities. The threats are even more important that its distribution in Mediterranean France is very limited. This study was based on the comparison of nine sites urbanized or impacted by other human activities. Particularly, the associated plant community, the density and reproductive parameters of the study species, and pollinator activities were compared to asset the effect of urbanization and land use changes on the local population persistence. Interestingly, in urban sites, the result shows more ruderal species and a higher density of T. pseudochamaepitys due to clonal growth. Surprisingly, reproductive success is low in each site despite an effective insect pollination. However, diversity of pollinators is lower in urban sites. Effects are thus contrasted considering urbanization and land use changes. Changes in land uses lead to closing landscape which threaten short term persistence of populations due to competition while the loss of pollinator diversity may impact urban populations over the long term only.     

Exploring biodiversity in a metropolitan area in the Mediterranean region: The urban and suburban flora of Rome (Italy)

This work provides an overview of plant diversity in the municipality of Rome (Italy) through an assessment of the flora in urban and suburban sectors of the city. It is aimed at providing the knowledge required to support proactive action for plant conservation. On the basis of a literature-derived catalogue and of an extensive survey campaign, the flora was investigated in terms of conservation interest; habitat types and locations that require protection measures were identified according to the occurrence of valuable native plants. Valuable species exclusive of the urban sector are threatened by the compaction trend affecting the urban fabric. Such species occur mainly in ruderal environments and fallows of archaeological sites and urban parks, near river courses and in remnants of natural forests. Valuable species exclusive of the suburban sector are threatened by urban sprawl. They occur prevalently in wet environments of the subcoastal strip and in coastal sands, Mediterranean maquis, tuffaceous gorges, sulphur springs and archaeological sites. The results highlight the need to preserve the complexity of the land mosaic, especially within the urban matrix, and to strengthen the existing environmental protection tools in the suburban area against foreseeable land cover changes.     

When Local Extinction and Colonization of River Fishes Can Be Predicted by Regional Occupancy: the Role of Spatial Scales

Background

Predicting which species are likely to go extinct is perhaps one of the most fundamental yet challenging tasks for conservation biologists. This is particularly relevant for freshwater ecosystems which tend to have the highest proportion of species threatened with extinction. According to metapopulation theories, local extinction and colonization rates of freshwater subpopulations can depend on the degree of regional occupancy, notably due to rescue effects. However, relationships between extinction, colonization, regional occupancy and the spatial scales at which they operate are currently poorly known.

Methods

And Findings: We used a large dataset of freshwater fish annual censuses in 325 stream reaches to analyse how annual extinction/colonization rates of subpopulations depend on the regional occupancy of species. For this purpose, we modelled the regional occupancy of 34 fish species over the whole French river network and we tested how extinction/colonization rates could be predicted by regional occupancy described at five nested spatial scales. Results show that extinction and colonization rates depend on regional occupancy, revealing existence a rescue effect. We also find that these effects are scale dependent and their absolute contribution to colonization and extinction tends to decrease from river section to larger basin scales.

Conclusions

In terms of management, we show that regional occupancy quantification allows the evaluation of local species extinction/colonization dynamics and reduction of local extinction risks for freshwater fish species implies the preservation of suitable habitats at both local and drainage basin scales.     

Range maps and species richness patterns: errors of commission and estimates of uncertainty

Range maps are often combined into “range overlap maps” to estimate spatial variation in species richness. Range maps are, in most cases, designed to represent a species’ maximum geographical extent and not patterns of occupancy within the range. As a consequence, range maps overestimate occupancy by presenting false occupancy (errors of commission) within the interior of the range. To assess the implications of errors of commission when developing and applying range overlap maps, we used neutral landscapes to simulate range maps and patterns of occupancy within ranges. We explored several scenarios based on combinations of six parameters defining biogeographical and cartographic factors typically encountered by investigators. Our results suggest that, in general, uncertainty is lowest when map resolutions are moderately fine, the majority of species have geographically restricted ranges, species occur throughout their range, patterns of occupancy within the range are not correlated among species, and geographically local and widespread species tend to occupy different regions. Several of these outcomes are associated with broad geographical extents, the scale at which range overlap maps are typically applied. Thus, under most circumstances, reasonably accurate and precise representation of species richness patterns can be achieved. However, these representations can be improved by enhancing occupancy data for widespread species – a primary source of uncertainty – and selecting a map resolution that captures relevant biological and environmental heterogeneity. Hence, by determining where a study is situated within the scenarios examined in our simulations, uncertainty and its sources and implications can be ascertained. With this knowledge, research goals, methods, and data sources can be reassessed and refined and, in the end, conclusions and recommendations can be qualified. Alternatively, unique regional, taxonomic, or cartographic factors could be included in future simulations to provide direct assessments of uncertainty.