Endemic vertebrates are the most effective surrogates for identifying conservation priorities among Brazilian ecoregions

Many studies have tested the performance of terrestrial vertebrates as surrogates for overall species diversity, because these are commonly used in priority‐setting conservation appraisals. Using a database of 3663 vertebrate species in 38 Brazilian ecoregions, we evaluated the effectiveness of various subsets for representing diversity of the entire vertebrate assemblage. Because ecoregions are established incorporating information on biotic assemblages, they are potentially more amenable to regional comparison than are national or state lists. We used 10 potential indicator groups (all species; all mammals, birds, reptiles, or amphibians; all endemic species; and endemic species within each class) to find priority sets of ecoregions that best represent the entire terrestrial vertebrate fauna. This is the first time such tests are employed to assess the effectiveness of indicator groups at the ecoregion level in Brazil. We show that patterns of species richness are highly correlated among mammals, birds, amphibians, and reptiles. Furthermore, we demonstrate that ecoregion sets selected according to endemic species richness captured more vertebrate species per unit area than sets based on overall vertebrate richness itself, or than those selected at random. Ecoregion sets based on endemic bird, endemic reptile, or endemic amphibian richness also performed well, capturing more species overall than random sets, or than those selected based on species richness of one or all vertebrate classes within ecoregions. Our results highlight the importance of evaluating biodiversity concordance and the use of indicator groups as well as aggregate species richness. We conclude that priority sets based on indicator groups provide a basis for a first assessment of priorities for conservation at an infracontinental scale. Areas with high endemism have long been highlighted for conservation of species. Our findings provide evidence that endemism is not only a worthwhile conservation goal, but also an effective surrogate for the conservation of all terrestrial vertebrates in Brazil.     

燕辽生物群脊椎动物的多样性及与其他生物群的对比分析*

燕辽生物群已发现脊椎动物54属58种, 包括鱼类、两栖类、爬行类、哺乳类等, 但其脊椎动物多样性及其成因机制还未有详细研究。本文对该生物群脊椎动物进行统计分析, 并与同时代的其他生物群脊椎动物类型进行对比, 这为认识燕辽生物群脊椎动物的多样性及其成因提供了重要的证据。早期代表道虎沟生物群与晚期代表玲珑塔生物群虽存在时代上的传承关系, 但生物组合特征明显不同。对比燕辽生物群与相近时代的新疆五彩湾动物群和四川大山铺恐龙动物群, 脊椎动物组合特征差异显著。燕辽生物群恐龙类群主要以小型兽脚类恐龙为主, 还包括一些小型鸟臀类恐龙。另外还具有非常丰富的翼龙和哺乳动物。脊椎动物生态多样性高, 适应飞行、树栖、水生、穴居等多种生活方式, 但是脊椎动物的类型与同时代的相近地区明显不同。翼龙、恐龙和哺乳动物等类群都展现出独特的生物组合特征。有证据表明该时期东亚地区与其他地区可能存在一定程度的地理隔离, 结合陆生脊椎动物组合特征推测燕辽生物群脊椎动物与外界可能存在一定的交流障碍。     

Global concordance in diversity patterns of vascular plants and terrestrial vertebrates

The factors that determine large-scale patterns of species richness are poorly understood. In particular, biologists have not determined the relative roles of taxon-specific characteristics that influence diversification and distribution, and region-specific features that promote and constrain diversity. We show that the numbers of species of vascular plants and of four terrestrial vertebrate taxa (mammals, birds, reptiles and amphibians) vary in parallel across 296 geographic areas covering most of the globe, even after accounting for sample area, climate, topographic heterogeneity and differences between continents. Thus, a common set of regional characteristics and processes appears to shape patterns of species richness in a diverse set of taxa, despite substantial differences in their biological traits.     

Global associations between terrestrial producer and vertebrate consumer diversity

In both ecology and conservation, often a strong positive association is assumed between the diversity of plants as primary producers and that of animals, specifically primary consumers. Such a relationship has been observed at small spatial scales, and a begetting of diversity by diversity is expected under various scenarios of co-evolution and co-adaptation. But positive producer-consumer richness relationships may also arise from similar associations with past opportunities for diversification or contemporary environmental conditions, or from emerging properties of plant diversity such as vegetation complexity or productivity. Here we assess whether the producer-consumer richness relationship generalizes from plot to regional scale and provide a first global test of its strength for vascular plants and endothermic vertebrates. We find strong positive richness associations, but only limited congruence of the most diverse regions. The richness of both primary and higher-level consumers increases with plant richness at similar strength and rate. Environmental conditions emerge as much stronger predictors of consumer richness, and after accounting for environmental differences little variation is explained by plant diversity. We conclude that biotic interactions and strong local associations between plants and consumers only relatively weakly scale up to broad geographical scales and to functionally diverse taxa, for which environmental constraints on richness dominate.     

Partitioning sources of variation in vertebrate species richness

Aim To explore biogeographic patterns of terrestrial vertebrates in Maine, USA using techniques that would describe local and spatial correlations with the environment. Location Maine, USA. Methods We delineated the ranges within Maine (86,156 km²) of 275 species using literature and expert review. Ranges were combined into species richness maps, and compared to geomorphology, climate, and woody plant distributions. Methods were adapted that compared richness of all vertebrate classes to each environmental correlate, rather than assessing a single explanatory theory. We partitioned variation in species richness into components using tree and multiple linear regression. Methods were used that allowed for useful comparisons between tree and linear regression results. For both methods we partitioned variation into broad‐scale (spatially autocorrelated) and fine‐scale (spatially uncorrelated) explained and unexplained components. By partitioning variance, and using both tree and linear regression in analyses, we explored the degree of variation in species richness for each vertebrate group that could be explained by the relative contribution of each environmental variable. Results In tree regression, climate variation explained richness better (92% of mean deviance explained for all species) than woody plant variation (87%) and geomorphology (86%). Reptiles were highly correlated with environmental variation (93%), followed by mammals, amphibians, and birds (each with 84–82% deviance explained). In multiple linear regression, climate was most closely associated with total vertebrate richness (78%), followed by woody plants (67%) and geomorphology (56%). Again, reptiles were closely correlated with the environment (95%), followed by mammals (73%), amphibians (63%) and birds (57%). Main conclusions Comparing variation explained using tree and multiple linear regression quantified the importance of nonlinear relationships and local interactions between species richness and environmental variation, identifying the importance of linear relationships between reptiles and the environment, and nonlinear relationships between birds and woody plants, for example. Conservation planners should capture climatic variation in broad‐scale designs; temperatures may shift during climate change, but the underlying correlations between the environment and species richness will presumably remain.     

Global comparisons of beta diversity among mammals, birds, reptiles, and amphibians across spatial scales and taxonomic ranks

Beta diversity is the change in species composition among areas in a geographic region. The proportion of species shared between two areas often decreases when the distance separating them increases, leading to an increase in beta diversity. This study compares beta diversity among four classes of terrestrial vertebrates （mammals, birds, reptiles, and amphibians） at both regional （biogeographic realm） and global extents, using the same sets of faunal sample units for all four groups in each comparison. Beta diversity is lower for the two endothermic taxa （birds and mammals） than for the two ectothermic taxa （reptiles and amphibians） in all six biogeographic realms examined. When the four taxa in the six biogeographic realms are combined, beta diversity at the species rank is higher than that of the genus rank by a factor of 1.24, and is higher than that of the family rank by a factor of 1.85. The ratio of beta diversity at the genus rank to that at the family rank is 1.50. Beta diversity is slightly higher for ecoregions of 5000-99,999 km^2 than for ecoregions of 100,000-5,000,000 km^2.     

Global tests of regional effect on species richness of vascular plants and terrestrial vertebrates

Determining the effects of regional and contemporary factors on large-scale patterns in species richness has been a fundamental question in modern ecology and biogeography. However, few studies have examined effects of historical and regional factors on species richness at the global scale, and conclusions are often inconsistent or controversial. Here, I use a comprehensive dataset to examine regional effects on species richness of vascular plants and four taxa of terrestrial vertebrates (mammals, birds, reptiles, and amphibians) in the same set of sample units (ecoregions) in seven biogeographic realms across the globe. The same spatial scale and the same set of environmental variables, which are thought to influence large-scale patterns in species richness of vascular plants and terrestrial vertebrates, are used for all the five taxa. Species richness of each taxon is compared across biogeographic realms. Regional effect on species richness has been found for all the five taxa. Of the 90 realm-pair comparisons for the five taxa between observed richness of a region and the richness of the region predicted by the richness–environment relationship derived from the data of another region, 74 (82.2%) showed significant differences between observed and predicted species richness, indicating that a species richness–environment relationship developed for one region cannot accurately predict species richness in other regions of similar environments.     

Hot,dry and different: Australian lizard richness is unlike that of mammals,amphibians and birds

Aims (1) To map the species richness of Australian lizards and describe patterns of range size and species turnover that underlie them. (2) To assess the congruence in the species richness of lizards and other vertebrate groups. (3) To search for commonalities in the drivers of species richness in Australian vertebrates. Location Australia. Methods We digitized lizard distribution data to generate gridded maps of species richness and β‐diversity. Using similar maps for amphibians, mammals and birds, we explored the relationship between species richness and temperature, actual evapotranspiration, elevation and local elevation range. We used spatial eigenvector filtering and geographically weighted regression to explore geographical patterns and take spatial autocorrelation into account. We explored congruence between the species richness of vertebrate groups whilst controlling for environmental effects. Results Lizard richness peaks in the central deserts (where β‐diversity is low) and tropical north‐east (where β‐diversity is high). The intervening lowlands have low species richness and β‐diversity. Generally, lizard richness is uncorrelated with that of other vertebrates but this low congruence is strongly spatially structured. Environmental models for all groups also show strong spatial heterogeneity. Lizard richness is predicted by different environmental factors from other vertebrates, being highest in dry and hot regions. Accounting for environmental drivers, lizard richness is weakly positively related to richness of other vertebrates, both at global and local scales. Main conclusions Lizard species richness differs from that of other vertebrates. This difference is probably caused by differential responses to environmental gradients and different centres of diversification; there is little evidence for inter‐taxon competition limiting lizard richness. Local variation in habitat diversity or evolutionary radiations may explain weak associations between taxa, after controlling for environmental variables. We strongly recommend that studies of variation in species richness examine and account for non‐stationarity.     

The contribution of contemporary climate to ectothermic and endothermic vertebrate distributions in a glacial refuge

Aim Climatic factors are known to influence species distributions. However, elucidating the underlying mechanisms is challenging because direct and indirect effects of climatic and non‐climatic factors are correlated. In the absence of this covariation and at fine‐grain resolutions the direct effect of climate via physiological constraints should be stronger on the distributions of ectothermic organisms. So far, no comprehensive study has explicitly tested the influence of climate on species distributions by quantitatively comparing ectothermic and endothermic vertebrates. Location Peninsular Spain. Methods Presence–absence data of native terrestrial vertebrates in Peninsular Spain were modelled using generalized additive models to disentangle the influence of climate and other contemporary correlated factors (topography and plant cover). We performed partial regressions to partition the deviance explained by climatic and non‐climatic effects into independent and shared components. We compared the independent contributions of climatic and non‐climatic effects between ectothermic and endothermic vertebrates, and among mammals, birds, reptiles and amphibians. Results After reducing the covariation with non‐climatic factors, climate explained a greater proportion of deviance in ectotherms than in endotherms. Also, the contribution of temperature was highest for reptiles, and the contribution of precipitation was highest for amphibians, after extracting their overlaps with precipitation and temperature, respectively. The contribution of topography and plant cover remained high for birds after extracting the overlap with climate. Main conclusions Our results are consistent with the prediction that, at fine resolutions, the direct influence of climate (via physiological constraints) on range distributions is stronger in ectothermic vertebrates. Also, at least for birds, indirect effects of climate (via plant productivity) and other habitat characteristics remained relatively important once their covariation with climate was reduced. This study shows that controlling the direct effects of climate by their indirect effects and/or other correlated factors, combined with comparisons among functional groups, can be a useful approach to elucidate causal links with the spatial patterns of organisms.     

The role of fire in terrestrial vertebrate richness patterns

Productivity is strongly associated with terrestrial species richness patterns, although the mechanisms underpinning such patterns have long been debated. Despite considerable consumption of primary productivity by fire, its influence on global diversity has received relatively little study. Here we examine the sensitivity of terrestrial vertebrate biodiversity (amphibians, birds and mammals) to fire, while accounting for other drivers. We analyse global data on terrestrial vertebrate richness, net primary productivity, fire occurrence (fraction of productivity consumed) and additional influences unrelated to productivity (i.e., historical phylogenetic and area effects) on species richness. For birds, fire is associated with higher diversity, rivalling the effects of productivity on richness, and for mammals, fire's positive association with diversity is even stronger than productivity; for amphibians, in contrast, there are few clear associations. Our findings suggest an underappreciated role for fire in the generation of animal species richness and the conservation of global biodiversity.     

Diversity in isochore structure among cold-blooded vertebrates based on GC content of coding and non-coding sequences

To review the general consideration about the different compositional structure of warm and cold-blooded vertebrates genomes, we used of the increasing number of genetic sequences, including coding (exons) and non-coding (introns) regions, that have been deposited on the databases throughout last years. The nucleotide distributions of the third codon positions (GC3) have been analyzed in 1510 coding sequences (CDS) of fish, 1414 CDS of amphibians and 320 CDS of reptiles. Also, the relationship between GC content of 74, 56 and 25 CDS of fish, amphibians and reptiles, respectively and that of their corresponding introns (GCI) have been considerated. In accordance with recent data, sequence analysis showed the presence of very GC3-rich CDS in these poikilotherm vertebrates. However, very high diversity in compositional patterns among different orders of fish, amphibians and reptiles was found. Significant positive correlations between GC3 and GCI was also confirmed for the genes analyzed. Nevertheless, introns resulted to be poorer in GC than their corresponding CDS, this difference being larger than in human genome. Because the limited number of available sequences including exons and introns we must be cautious about the results derived from them. However, the indicious of higher GC richness of coding sequences than of their corresponding introns could aid to understand the discrepancy of sequence analysis with the ultracentrifugation studies in cold-blooded vertebrates that did not predict the existence of GC-rich isochores.     

黄河流域被子植物和陆栖脊椎动物丰富度格局及其影响因子

生物多样性的大尺度空间分布格局及其形成机制一直是生态学和生物地理学的核心内容。黄河流域是我国重要的生态屏障, 明确该区域动植物多样性分布格局及其影响因素, 对我国黄河流域生态保护和高质量发展具有重要意义。本研究通过收集黄河流域被子植物和陆栖脊椎动物分布数据, 结合气候、环境异质性和人类活动等信息, 探讨了黄河流域被子植物和陆栖脊椎动物物种丰富度格局及其主要影响因素。结果表明, 黄河流域被子植物和陆栖脊椎动物物种丰富度在区域尺度具有相似的分布格局: 南部山地动植物物种丰富度最高, 而东部高寒区和北部干旱区物种丰富度最低。回归树模型表明, 冠层高度范围和净初级生产力范围分别是黄河流域被子植物和陆栖脊椎动物物种丰富度最重要的预测因子; 当移除空间自相关影响后, 环境异质性和气候因子依然对区域尺度的动植物物种丰富度具有较高且相似的解释度。表明环境异质性和气候共同决定了黄河流域被子植物和陆栖脊椎动物物种丰富度格局, 而人类使用土地面积并不是影响黄河流域动植物物种丰富度格局的主要因子。因此, 在未来的研究中若针对不同区域筛选出更精准的环境驱动因子或选用更多不同类别的环境异质性因子进行分析, 将有助于更深入理解物种多样性格局的成因。     

Microdissectional investigation of the nephrons in some fishes,amphibians, and reptiles inhabiting different environment

The structure of nephrons in 83 species of fishes, amphibians, and reptiles was investigated by microdissection. Glomerular diameter, nephron length, length of nephron segments, and the ratios between glomerular size and nephron length are presented. In diifferent groups of fishes (Elasmobranchii, Acipenseridae, Teleostei), the adaptation to freshwater or seawater environment may lead to diverse changes of nephron structure. The kidneys of euryhaline teleosts capable of living in fresh water may be aglomerular, as are those of some marine fishes. In contrast, the diameter of glomeruli in some marine fishes is larger than in true freshwater fishes. In amphibians, the adaptation to freshwater environment, as in teleost fishes, has led to different changes of nephron structure. The size of glomeruli in freshwater reptiles is larger in comparison to terrestrial animals, and the distal tubule in desert and freshwater reptiles is longer than in nondesert species. This probably reflects the adaptive changes of the reptilian nephron to freshwater and desert environments. The results of this study show that the nephron structure of lower vertebrates is predominantly determined by the different environments they occupy rather than by progressive changes within the vertebrate sequence. © 1994 Wiley-Liss, Inc.     

Life stage and taxonomy the most important factors determining vertebrate stoichiometry: A meta‐analysis

Whole‐body elemental composition is a key trait for determining how organisms influence their ecosystems. Using mass‐balance, ecological stoichiometry predicts that animals with higher concentrations of element X will selectively retain more X and will recycle less X in their waste than animals with lower X concentrations. These animals will also store high quantities of X during their lives and after their deaths (prior to full decomposition). Vertebrates may uniquely impact nutrient cycling because they store high quantities of phosphorus (P) in their bones. However, vertebrates have diverse body forms and invest variably in bone. Current analyses of vertebrate elemental content predominately evaluate fishes, typically neglecting other vertebrates and leaving much of the diversity unexplored. We performed a systematic review and identified 179 measurements of whole‐body percent phosphorus (%P), percent nitrogen (%N), and N to P ratio (N:P) from 129 unique species of non‐fish vertebrates (amphibians: 39 species; reptiles: 19 species; birds: 27 species; mammals: 46 species). We found that %P (mean: 1.94%; SD [standard deviation] = 0.77) and N:P (mean: 12.52) varied with taxonomy and life stage, while %N (mean: 10.51%; SD = 3.25) varied primarily with taxonomy. Habitat, diet, and size had small and inconsistent effects in different groups. Our study highlights two research gaps. Life stage, which is frequently neglected in stoichiometric studies, is an important factor determining vertebrate %P. Furthermore, amphibians dominate our dataset, while other vertebrate taxa are poorly represented in the current literature. Further research into these neglected vertebrate taxa is essential.     

Does herbivore diversity depend on plant diversity? The case of California butterflies

It is widely believed that the diversity of plants influences the diversity of animals, and this should be particularly true of herbivores. We examine this supposition at a moderate spatial extent by comparing the richness patterns of the 217 butterfly species resident in California to those of plants, including all 5,902 vascular plant species and the 552 species known to be fed on by caterpillars. We also examine the relationships between plant/butterfly richness and 20 environmental variables. We found that although plant and butterfly diversities are positively correlated, multiple regression, path models, and spatial analysis indicate that once primary productivity (estimated by a water-energy variable, actual evapotranspiration) and topographical variability are incorporated into models, neither measure of plant richness has any relationship with butterfly richness. To examine whether butterflies with the most specialized diets follow the pattern found across all butterflies, we repeated the analyses for 37 species of strict monophages and their food plants and found that plant and butterfly richness were similarly weakly associated after incorporating the environmental variables. We condude that plant diversity does not directly influence butterfly diversity but that both are probably responding to similar environmental factors.     

Links between global taxonomic diversity,ecological diversity and the expansion of vertebrates on land

Tetrapod biodiversity today is great; over the past 400 Myr since vertebrates moved onto land, global tetrapod diversity has risen exponentially, punctuated by losses during major extinctions. There are links between the total global diversity of tetrapods and the diversity of their ecological roles, yet no one fully understands the interplay of these two aspects of biodiversity and a numerical analysis of this relationship has not so far been undertaken. Here we show that the global taxonomic and ecological diversity of tetrapods are closely linked. Throughout geological time, patterns of global diversity of tetrapod families show 97 per cent correlation with ecological modes. Global taxonomic and ecological diversity of this group correlates closely with the dominant classes of tetrapods (amphibians in the Palaeozoic, reptiles in the Mesozoic, birds and mammals in the Cenozoic). These groups have driven ecological diversity by expansion and contraction of occupied ecospace, rather than by direct competition within existing ecospace and each group has used ecospace at a greater rate than their predecessors.     

Evolution of oestrogen functions in vertebrates

Steroidal oestrogens have been isolated from marine and terrestrial animals representative of all major classes of vertebrates including fish, amphibians, reptiles, birds and mammals. In general, oestrogens are responsible for most features characteristic of the female sex of a species, such as metabolic, behavioural and morphological changes during the stages of reproduction; they also support several processes in males. The evolution of the hormonal system always involves both the ligand and its sites of interaction. In the case of oestrogens, the steroid producing enzymes, mainly the aromatase complex, and the oestrogen receptor belong together within their co-evolution. The finding of oestrogenic steroids, the more recent identification of aromatase and receptor genes and their expression fit together, thereby confirming the importance for all vertebrates. Within the present paper, the evolution of the physiological functions of oestrogens from oviparous vertebrates to Eutherian mammals, oestrogen biosynthesis, metabolization and signalling pathways will be reviewed in detail.     

Disentangling the effects of geographic distance and environmental dissimilarity on global patterns of species turnover

Aim To distinguish the effects of geographic distance and environmental dissimilarity on global patterns of species turnover in four classes of terrestrial vertebrates (mammals, birds, reptiles and amphibians). Location Six hundred and sixty terrestrial ecoregions across the globe. Methods We calculated species turnover between each pair of ecoregions, using the Jaccard index (J). We selected seven variables to quantify environment in each ecoregion, and subjected the environmental values to a principal components analysis. For each realm, we applied multiple regression analysis relating the natural logarithm of the Jaccard index (lnJ) to geographic distance alone and in combination with differences in the environment variables measured as principal components (PC). We used partial correlations to partition variance in lnJ between unique contributions of distance and environmental PC scores, the covariation between distance and environment, and unexplained variance. To examine the latitude and species turnover relationship, we regressed lnJ on latitude with distance between ecoregions being included as a covariate. Results The natural logarithm of the Jaccard index (lnJ) decreased significantly with increasing geographic distance for all vertebrate classes in each zoogeographic realm, and the slopes of the relationships per 1000 km ranged from ?0.251 to ?1.043. With environmental differences included in the analysis, both geographic distance and environmental differences were substantial predictors of lnJ for every combination of taxon and realm. On average, the unique contribution of geographic distance to variation in species turnover between ecoregions was about 1.4 times that of the environmental differences between ecoregions. Species turnover generally decreased with increasing latitude when controlling for geographic distance. The value of lnJ for each vertebrate class was highly and positively correlated with those of the other vertebrate classes. Main conclusions Our analyses suggest that both dispersal‐based and niche‐based processes have played important roles in determining faunal similarities among vertebrate assemblages at the spatial scale examined. Furthermore, reptiles and amphibians exhibited greater distance‐independent faunal heterogeneity among ecoregions and greater turnover among ecoregions with respect to geographic and environmental distance than birds and mammals.     

Global bioregions of reptiles confirm the consistency of bioregionalization processes across vertebrate clades

Aim

The identification of biogeographical zones has been fundamental in broadscale biodiversity analyses over the last 150 years. If processes underlying bioregionalization, such as climatic differences, tectonics and physical barriers, are consistent across vertebrate clades, we expect that groups with more similar ecological characteristics would show more similar bioregions. Lack of data has so far hampered the delineation of global bioregions for reptiles. Therefore, we integrated comprehensive geographic distribution and phylogenetic data of lepidosaurian reptiles to delineate global reptile bioregions, compare determinants of biogeographical boundaries across terrestrial vertebrates and test whether clades showing similar responses to environmental factors also show more similar bioregions.

Location

Global.

Time Period

Present.

Major Taxa Studied

Reptiles, amphibians, birds, mammals.

Methods

For reptiles, we used phylogenetic beta diversity to quantify changes in community composition, and hierarchical clustering to identify biogeographic ‘realms’ and ‘regions’. Then, we assessed the determinants of biogeographical boundaries using spatially explicit regression models, testing the effect of climatic factors, physical barriers and tectonics. Bioregions of reptiles were compared to those of other vertebrate clades by testing the overall similarity of the spatial structure of bioregions, and the match of the position of biogeographical boundaries.

Results

For reptiles, we identified 24 evolutionarily unique regions, nested within 14 realms. Biogeographical boundaries of reptiles were related to both climatic factors and past tectonic movements. Bioregions were very consistent across vertebrate clades. Bioregions of reptiles and mammals showed the highest similarity, followed by reptiles/birds and mammals/birds while amphibian bioregions were less similar to those of the other clades.

Main Conclusions

The overall high similarity among bioregions suggests that bioregionalization was affected by similar underlying processes across terrestrial vertebrates. Nevertheless, clades with different eco-physiological characteristics respond somewhat differently to the same environmental factors, resulting in similar but not identical regionalizations across vertebrate clades.