Niche width impacts vertebrate diversification

Aim The size of the climatic niche of a species is a major factor determining its distribution and evolution. In particular, it has been proposed that niche width should be associated with the rate of species diversification. Here, we test whether species niche width affects the speciation and extinction rates of three main clades of vertebrates: amphibians, mammals and birds. Location Global. Methods We obtained the time‐calibrated phylogenies, IUCN conservation status, species distribution maps and climatic data for 2340 species of amphibians, 4563 species of mammals and 9823 species of birds. We computed the niche width for each species as the mean annual temperature across the species range. We estimated speciation, extinction and transition rates associated with lineages with either narrow (specialist) or wide (generalist) niches using phylogeny‐based birth–death models. We also tested if current conservation status was correlated with the niche width of species. Results We found higher net diversification rates in specialist species than in generalist species. This result was explained by both higher speciation rates (for the three taxonomic groups) and lower extinction rates (for mammals and birds only) in specialist than in generalist species. In contrast, current specialist species tended to be more threatened than generalist species. Main conclusions Our diversification analysis shows that the width of the climatic niche is strongly associated with diversification rates and may thus be a crucial factor for understanding the emergence of diversity patterns in vertebrates. The striking difference between our diversification results and current conservation status suggests that the current extinction process may be different from extinction rates estimated from the whole history of the group.     

Low dispersal ability and habitat specificity promote extinctions in rare but not in widespread species: the Orthoptera of Germany

Local extinctions are often non‐randomly associated with range size, dispersal ability and habitat specificity, as well as body size, sexual dimorphism and phylogeny. We used a large data set of the Orthoptera species (bush crickets, crickets, grasshoppers) occurring in Germany and compared the number of occupied grid cells before 1980 to those occupied after 1980, corrected for monitoring intensity. The number of grid cells in which a species went extinct was non‐linearly related to the number of occupied grid cells per species. Using generalized linear modelling we analysed extinction in relation to national distribution (the number of occupied grid cells before 1980), dispersal ability (derived from a large body of literature concerning wing development, colonization dynamics and within‐habitat mobility), habitat specificity (moisture specialists versus generalists), potential reproduction (the number of ovarioles), the degree of sexual size dimorphism and phylogeny (twelve clades). Species with a large global range size also had a large national range size. Species with a large range experienced more total extinction events than species with smaller ranges but relatively fewer compared to range size. The latter relationship was largely shaped by the dispersal ability of the species: the interactions of range size×dispersal ability and range size×habitat specificity explained almost one third of the variation in the number of extinction events. Species with high dispersal ability went extinct in a similar number of grid cells irrespective of their range size. By contrast, species with low dispersal ability went extinct in proportion to their range size. Therefore, comparing the speed of extinction across species in the conventional way of extinction rates (that is the percentage of range contraction) might be flawed because it only applies to species with low dispersal ability. Sexual size dimorphism was not a significant predictor of extinction. Extinction was not concentrated on particular clades.     

Unprotecting the rare species: a niche‐based gap analysis for odonates in a core Cerrado area

Aim We evaluated Odonata distribution data and predicted the compositional resemblance based on niche‐based species distribution models to analyse the following questions: (1) How is estimated species richness distributed, and how can it be preserved under the actual network of conservation units (a gap analysis approach)? (2) How is the estimated odonate beta diversity distributed, and is there a better distribution of conservation units (a priority setting approach)? (3) Is the probability of being under protection a function of the potential species range size? and (4) Will the current conservation network proposals protect odonate taxa? Location Central Brazil in a core Cerrado area. Methods We generated odonate species distribution predictions based on MaxEnt and maps derived from estimated species richness, beta diversity and gap analysis for all species predicted to occur in the study area. Then, we compared these maps with current conservation units, land‐use patterns and proposals for the establishment of conservation units. Results Raw odonate species records provided limited utility for setting conservation priorities without the use of niche‐based models. However, area under the receiver operating curve (AUC) values were characterized by substantial variation that was related to the number of records. No current conservation units overlapped the areas with higher predicted richness and beta diversity, and in general, conservation units were not preserving restricted/rare species. There was a direct linear correlation between species range size and the proportion of its range protected in the current network of conservation units. Finally, we identified three areas with high odonate beta diversity where conservationist actions should be implemented. Main conclusions Current conservation units and future suggested areas do not overlap regions with high conservation values for odonates. Conservation units protect species at random, and the level of protection has a direct relationship with species range size; thus, wide‐range species are expected to be more protected than restricted or threatened species.     

Species richness, species range size and ecological specialisation among African primates: geographical patterns and conservation implications

The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.     

Nested distribution of amphibians in the Zhoushan archipelago, China: can selective extinction cause nested subsets of species?

Because of their poor dispersal ability, amphibians are well suited for testing the selective extinction theory on islands. Amphibian fauna in the Zhoushan archipelago, China, exhibit a high level of nestedness (C = 0.893), and the species number is lower on islands than on similar sized areas on the mainland. No correlation was found between island-specific species richness and the nearest distance from a larger island, distance from the mainland or density of human population. These results suggest that no amphibian colonisation has occurred in the archipelago since island isolation 7000–9000 years ago. Furthermore, the results imply that selective extinction contributes to the nestedness of amphibians in the Zhoushan archipelago. The incidence of a species on the islands is significantly correlated with log area of the smallest island occupied by the species and the number of provinces on the Chinese mainland in which the species occur. However, there is no correlation with average body length of adults and island occurrence. It is concluded that (1) the area of the smallest island occupied by a species is a good estimate of the minimum area for a viable population of the species and a good predictor of species incidence on islands, (2) species with a restricted distribution range are more vulnerable to extinction from islands than those with a wide distribution range and (3) the effect of body size on occurrence on the islands is uncertain, and may be specific to the archipelago and taxa studied. The observed nestedness of amphibian assemblages has two implications for conservation: (1) not only can all the species found in several small reserves be found on a large reserve of the same total size, but additional species can be found on the single large reserve; (2) for a reserve to maintain viable populations of all species in a region it should be at least as large as the smallest island occupied by the most vulnerable species. Received: 16 December 1996 / Accepted: 22 September 1997     

Patterns of species richness hotspots and estimates of their protection are sensitive to spatial resolution

Aim

Species richness is a measure of biodiversity often used in spatial conservation assessments and mapped by summing species distribution maps. Commission errors inherent those maps influence richness patterns and conservation assessments. We sought to further the understanding of the sensitivity of hotspot delineation methods and conservation assessments to commission errors, and choice of threshold for hotspot delineation.

Location

United States.

Methods

We created range maps and 30‐m and 1‐km resolution habitat maps for terrestrial vertebrates in the United States and generated species richness maps with each dataset. With the richness maps and the GAP Protected Areas Dataset, we created species richness hotspot maps and calculated the proportion of hotspots within protected areas; calculating protection under a range of thresholds for defining hotspots. Our method allowed us to identify the influence of commission errors by comparing hotspot maps.

Results

Commission errors from coarse spatial grain data and lack of porosity in the range data inflated richness estimates and altered their spatial patterns. Coincidence of hotspots from different data types was low. The 30‐m hotspots were spatially dispersed, and some were very long distances from the hotspots mapped with coarser data. Estimates of protection were low for each of the taxa. The relationship between estimates of hotspot protection and threshold choice was nonlinear and inconsistent among data types (habitat and range) and grain size (30‐m and 1‐km).

Main conclusions

Coarse mapping methods and grain sizes can introduce commission errors into species distribution data that could result in misidentifications of the regions where hotspots occur and affect estimates of hotspot protection. Hotspot conservation assessments are also sensitive to choice of threshold for hotspot delineation. There is value in developing species distribution maps with high resolution and low rates of commission error for conservation assessments.     

Global patterns of geographic range size in birds

Large-scale patterns of spatial variation in species geographic range size are central to many fundamental questions in macroecology and conservation biology. However, the global nature of these patterns has remained contentious, since previous studies have been geographically restricted and/or based on small taxonomic groups. Here, using a database on the breeding distributions of birds, we report the first (to our knowledge) global maps of variation in species range sizes for an entire taxonomic class. We show that range area does not follow a simple latitudinal pattern. Instead, the smallest range areas are attained on islands, in mountainous areas, and largely in the southern hemisphere. In contrast, bird species richness peaks around the equator, and towards higher latitudes. Despite these profoundly different latitudinal patterns, spatially explicit models reveal a weak tendency for areas with high species richness to house species with significantly smaller median range area. Taken together, these results show that for birds many spatial patterns in range size described in geographically restricted analyses do not reflect global rules. It remains to be discovered whether global patterns in geographic range size are best interpreted in terms of geographical variation in species assemblage packing, or in the rates of speciation, extinction, and dispersal that ultimately underlie biodiversity.     

Dynamics of a northern fulmar (Fulmarus glacialis) population at the southern limit of its range in Europe

In the context of global changes, defining the source–sink dynamics of populations of emblematic species, such as seabirds, within the limits of their distribution range is often crucial to optimize the priorities of surveys and conservation management, especially in protected areas. However, ringing is often not possible and only simple survey methods, such as the ‘apparently occupied sites’ method, can be utilized by managers of protected areas and threatened species. Using data collected between 1997 and 2005, we investigated the population dynamics of the northern fulmar (Fulmarus glacialis) at the southern limit of its range on the western French coast, which hosts increasing populations. Using a robust design spatial occupancy model, we estimated the proportion of occupied nests, the rates of nest colonization, nest extinction and the population growth rates of four colonies of the largest population (Ouessant Island). The estimated annual growth rate was high (average 1.049). A deterministic population dynamics model indicated that the observed rapid increase of Ouessant populations cannot be explained by their intrinsic dynamics, which suggested an important role for immigration. Different demographic scenarios provide several lines of evidence that the large northern fulmar population in this Man and Biosphere Reserve is a sink population. The strong increase in a population located at the limits of the species distributional range implies that it functions as a population sink. Inexpensive effective survey methods could allow investigation of the demographic status of seabird populations and provide relevant information for the hierarchization of conservation priorities.     

Extinction risk and diversification are linked in a plant biodiversity hotspot

It is widely recognized that we are entering an extinction event on a scale approaching the mass extinctions seen in the fossil record. Present-day rates of extinction are estimated to be several orders of magnitude greater than background rates and are projected to increase further if current trends continue. In vertebrates, species traits, such as body size, fecundity, and geographic range, are important predictors of vulnerability. Although plants are the basis for life on Earth, our knowledge of plant extinctions and vulnerabilities is lagging. Here, we disentangle the underlying drivers of extinction risk in plants, focusing on the Cape of South Africa, a global biodiversity hotspot. By comparing Red List data for the British and South African floras, we demonstrate that the taxonomic distribution of extinction risk differs significantly between regions, inconsistent with a simple, trait-based model of extinction. Using a comprehensive phylogenetic tree for the Cape, we reveal a phylogenetic signal in the distribution of plant extinction risks but show that the most threatened species cluster within short branches at the tips of the phylogeny--opposite to trends in mammals. From analyzing the distribution of threatened species across 11 exemplar clades, we suggest that mode of speciation best explains the unusual phylogenetic structure of extinction risks in plants of the Cape. Our results demonstrate that explanations for elevated extinction risk in plants of the Cape flora differ dramatically from those recognized for vertebrates. In the Cape, extinction risk is higher for young and fast-evolving plant lineages and cannot be explained by correlations with simple biological traits. Critically, we find that the most vulnerable plant species are nonetheless marching towards extinction at a more rapid pace but, surprisingly, independently from anthropogenic effects. Our results have important implications for conservation priorities and cast doubts on the utility of current Red List criteria for plants in regions such as the Cape, where speciation has been rapid, if our aim is to maximize the preservation of the tree-of-life.     

A multidimensional characterization of rarity applied to the Aegean tenebrionid beetles (Coleoptera Tenebrionidae)

This paper attempts to use museum collection data to estimate measures of species rarity and then to relate these measures to extinction risk. For this purpose, 170 taxa (138 species and 32 subspecies) of tenebrionid beetles from 32 Aegean Islands (Greece) were considered. For each taxon, rarity was evaluated as geographic distribution (mean incidence on islands in the archipelago), potential habitat exploitation (total area of the islands occupied on the total area of the study system) and contactability (number of decades of taxon’s records on the total number of decades of assumed persistence from 1870 to 2000). All of these indices were correlated to each other. Whether expressed in terms of range size or habitat exploitation rarity was a major determinant of a species’ risk of extinction (evaluated as extinction decade). Thus, the designation of rarity provides a good basis for identifying species that are most in need of conservation at a particular scale.     

The form of the curves: a direct evaluation of MacArthur & Wilson's classic theory

1. We calculate the yearly numbers of bird species immigrating to – and becoming extinct on – 13 small islands of the British Isles, using a long and relatively complete data record.
2. We estimate the size of the colonist pool for each island using four methods.
3. We assume that immigrations and extinctions are distributed binomially, and use a maximum likelihood method to fit concave immigration and extinction functions to the data, utilizing all four species pool estimates.
4. Extinction rates increase significantly and consistently with increasing numbers of breeding species on each island. For nine of the 13 islands the extinction functions are significantly concave.
5. Immigration rates decrease consistently with increasing numbers of breeding species on each island. Seven islands have significantly concave immigration functions.
6. Immigration rates and extinction rates decline consistently, but not significantly, with island distance and island size, respectively. The number of breeding species does not always reflect the number of species likely to have reached an island. Moreover, some species may choose not to breed when their chance of extinction is high. These factors, plus the modest range of island areas and distances in our database, reduce our chances of finding the theoretically predicted effects of area and distance on extinction and immigration rates.     

5. NOTES ON A FEW S. RHODESIAN BIRDS

The Aquatic Warbler is a threatened Afro-Palaearctic migrant with a largely unknown distribution in the winter (non-breeding) season. Protection of wintering sites may be crucial for the conservation of the species. Previous studies have identified extensive areas of north-western sub-Saharan Africa that could potentially be occupied by the species during winter. However, these studies have not necessarily differentiated between potentially suitable and unsuitable habitat types at a spatial resolution appropriate for targeting field surveys. To identify specific sites that could be occupied by non-breeding Aquatic Warblers at a scale appropriate for targeted field surveys, we adopted a modelling approach that combined recent sightings from Djoudj, Senegal, with land cover and climate data. We produced maps of potential distribution using three maximum entropy models. While a paucity of sightings prevented a full test of these maps on independent records, the areas that were predicted to be potentially occupied included areas around which there are historical records. We suggest field surveys should be targeted towards sites in the Inner Niger Delta and a number of marshes along and away from the Niger River in Mali, and to sites in southern Mauritania.     

Adaptive management as a tool to improve the conservation of endemic floras: the case of Sicily, Malta and their satellite islands

The presence of endemic species is among the fundamental criteria for characterizing the biodiversity of a territory. Analyzing species richness, extinction level and distribution drivers is an important preliminary step to set conservation priorities and test environmental policies. This study applied the concept of adaptive management to develop strategies for the conservation of endemic floras by considering, as a case study, Sicily, Malta and their neighboring small islands. Adaptive management can be defined as the systematic acquisition and application of reliable information to improve management over time. The development of adaptive conservation strategies aimed preliminary: (1) to quantify endemic plant diversity; (2) to assess the current IUCN knowledge; (3) to analyze the spatial patterns of species distribution in relation to number of colonized habitats, preferential habitats, altitudinal range, and bedrock; (4) to assess whether Natura 2000 network contributed significantly to increase the overlap between endemic distributional areas and protected surface. Strictly Sicilian endemics were 202 taxa amounting to 7.0 % of the whole native flora (c. 2900 taxa). The current picture of extinction risk is still incomplete because over 50 % of endemics were never assessed or assessed with old IUCN criteria. The spatial range size of endemics depended by 40 % on bedrock, and altitudinal and niche breadth. Habitat type did not influence the range size of endemics. The overlap between endemic distributional areas and protected surface increased from 41.3 to 63.3 % with Natura 2000. Adaptive management needs measurable goals to test the progressive improvement of conservation strategies over time, and the reduction in threatened species may be considered as an indicator of successful conservation outcomes. Feedback plays an important role in the periodic revision of biodiversity assessment, distribution modeling, and environmental management, which are fundamental to predict conservation outcomes in the face of extreme uncertainty. In particular, the exhaustive knowledge of the IUCN status is a primary step to implement adaptive measures of conservation, especially as regards endemic floras that are potentially more vulnerable to large-scale or unpredictable and stochastic threats.     

The conservation of declining butterfly populations in Britain and Europe: priorities, problems and successes

The status, ecology and conservation of butterflies in Europe and Britain are reviewed, as a background to the National Trust's past and future contribution to British conservation. Britain has a poor butterfly fauna by European standards, the main areas of endemism and species richness being in the Alps and southern Europe. To date, the main declines among European butterfly populations have occurred across central-northern Europe, with slightly higher extinction rates in mainland countries than in Britain. The main causes of decline are biotope destruction, the loss of certain species' habitats within surviving semi-natural biotopes due to changed land management, and a failure by several species to track the patches of their habitat that are still being generated in modern fragmented landscapes. Until recently, most conservation programmes failed to take account of the latter two factors, resulting in many local extinctions of rare butterfly species even in conservation areas. Recent measures have been much more successful; many were first tested on National Trust properties.     

Life-history traits associated with fragmentation vulnerability of lizards in the Thousand Island Lake, China

Following habitat fragmentation, the remnant faunal community will undergo a period of species loss or 'relaxation.' Theory predicts that species with particular life-history traits, such as a small population size, small geographical range, low fecundity and large body size, should be more vulnerable to fragmentation. In this study, we investigated the relationships between the above life-history traits and the fragmentation vulnerability index (the number of islands occupied) of five lizard species inhabiting recently isolated land-bridge islands in the Thousand Island Lake, China. Data on life-history traits were collected from field surveys (population density) and from the literature (body size, clutch size and geographical range size). The species–area relationships for lizards sampled from the mainland versus on the islands differed significantly (i.e. the number of species inhabiting islands was decreased relative to similar-sized areas on the mainland), indicating that species extinction has occurred on all of the study islands following isolation. For the fragmentation vulnerability index, model selection based on Akaike's information criterion identified natural density at mainland sites as the best correlate of vulnerability to fragmentation, supporting the hypothesis that rare species are most vulnerable to local extinction and will be lost first from fragmented landscapes. In contrast, there was little evidence for an effect of lizards' snout–vent length, clutch size or geographical range size on fragmentation vulnerability. Identification of species traits that render some species more vulnerable to fragmentation than others has important implications for conservation and can be used to aid direct management efforts.     

Macroecology and extinction risk correlates of frogs

Aim  Our aim was to test whether extinction risk of frog species could be predicted from their body size, fecundity or geographical range size. Because small geographical range size is a correlate of extinction risk in many taxa, we also tested hypotheses about correlates of range size in frogs.
Location  Global.
Methods  Using a large comparative data set ( n  = 527 species) compiled from the literature, we performed bivariate and multiple regressions through the origin of independent contrasts to test proposed macroecological patterns and correlates of extinction risk in frogs. We also created minimum adequate models to predict snout–vent length, clutch size, geographical range size and IUCN Red List status in frogs. Parallel non-phylogenetic analyses were also conducted. We verified the results of the phylogenetic analyses using gridded data accounting for spatial autocorrelation.
Results  The most threatened frog species tend to have small geographical ranges, although the relationship between range and extinction risk is not linear. In addition, tropical frogs with small clutches have the smallest ranges. Clutch size was strongly positively correlated with geographical range size ( r ² = 0.22) and body size ( r ² = 0.28).
Main conclusions  Our results suggest that body size and fecundity only affect extinction risk indirectly through their effect on geographical range size. Thus, although large frogs with small clutches tend to be endangered, there is no comparative evidence that this relationship is direct. If correct, this inference has consequences for conservation strategy: it would be inefficient to allocate conservation resources on the basis of low fecundity or large body size; instead it would be better to protect areas that contain many frog species with small geographical ranges.     

Extinction debt and the role of static and dynamical fragmentation on biodiversity

The mass-extinction events caused by human-driven habitat loss are a current concern in conservation science. However, the observed number of extinctions is considerably smaller than predicted. The overestimation of extinction rates comes from the time-delay which depends on the species sensitivity to habitat changes. The standard method of predicting the effect of habitat loss on biodiversity is to use the species–area relationship and progressively following it backwards to smaller areas. The difference between the actual number of species and the one provided by the backwards species–area relationship is dubbed extinction debt. Previous studies in general adopt a static view for the spatial distribution of species. Nonetheless, a precise understanding of the problem urges us to adopt a dynamic framework to this issue since the time between disturbances of the landscape plays an active role in influencing the strength of the extinction debt. In this context, here we address two distinct approaches for this question: a static and a dynamic view of fragmentation. In the former we quantify the extinction debt in a quenched spatial distribution of species, whereas in the latter the community is let to evolve between disturbance events of the landscape. Here we show that the size of the extinction debt depends on the pattern of the fragmentation. It is found that random distributions of destroyed habitats provide larger extinction debts than those obtained for contiguous areas of fragmentation. Furthermore, in the dynamic approach it is observed that dispersal can lead to unexpected outcomes such as lower biodiversity levels than ones inferred from the backwards species–area relationship.     

厦门市重点保护植物空间优先保护格局研究

生物多样性保护对维持城市生态系统功能具有重要意义。以39种厦门市重点保护植物为对象,通过物种分布模型MaxENT获得物种潜在分布栅格图,利用空间保护优先化定量工具Zonation软件识别理论上既适宜重点保护植物生存又能够保证景观连通性的区域,获得本地重点保护植物景观保护等级。根据2020年全球生物多样性目标,将景观保护等级最高的17%区域视为多物种空间优先保护区,结合Zonation模型生成的随景观丧失物种加权灭绝风险曲线,将保护等级最高的8%区域划为一级保护区,保护等级在8%-17%范围内的区域划为二级保护区。利用MaxENT模型中的jackknife刀切法发现海拔是对本地重点保护植物分布影响最大的环境因子,优先保护区集中分布于海拔较低的海岸带区域。将优先保护区与自然保护地建设现状、厦门市生态功能区规划、土地利用规划、城市总体规划对比发现厦门市岛外西部、北部的优先保护区得到了较好保护;岛外的西南部及东南部、岛内的东部及南部海岸带的优先保护区被建设用地大规模占用,已纳入自然保护地范围的区域较少,存在大量的海岸带优先保护区保护空缺;岛外东南部的部分优先保护区虽未被占用,但规划中属发展备用地,缺乏生态保护。为避免优先保护区面积的进一步萎缩,应重点关注海岸带区域优先保护区的生态保护,将目前属于发展备用地的优先保护区转划为生态留白空间,针对一级、二级优先保护区分别实施刚性和弹性的生态保育措施,在保护生物多样性的同时,严控对海岸带区域优先保护区的进一步开发利用,协调优先保护区内保护与开发利用间的关系。     

A primate at risk in Northeast Brazil: local extinctions of Coimbra Filho’s titi (<Emphasis Type="Italic">Callicebus coimbrai</Emphasis>)

Identifying the factors that determine local extinction of populations is crucial to ensure species conservation. Forest-dwelling primates are especially vulnerable to habitat fragmentation, although few studies have provided systematic evidence of local extinctions. Over an 11-year period, approximately 100 remnant populations of the endangered Coimbra Filho’s titi monkey (Callicebus coimbrai) have been found within the geographic range of the species in Bahia and Sergipe, Northeast Brazil. During the present study, extinction of 13 of these populations was recorded through intensive surveys. These extinctions were detected from evidence of intensive logging and clear-cutting, interviews with local residents and systematic searches of the sites where occurrence of the species had been confirmed in previous surveys. These local extinctions represent approximately 10 % of the known populations of C. coimbrai and up to 28.3 % of the area occupied by the species. Comparison of the vegetation structure in fragments where extinction was recorded and where the species still occurs indicated that sparser understorey may be a correlate of extinction, combined with the fact that extinctions occurred within fragments characterised by relatively high levels of anthropogenic disturbance. These findings reinforce the Endangered status of the species and the urgent need for intensification of conservation measures within the most impacted areas of the geographic distribution of C. coimbrai.