Factors influencing herpetofauna abundance and diversity in a tropical agricultural landscape mosaic

Agricultural intensification and the associated factors, including land transformation, are among the major global threats affecting biodiversity especially herpetofauna. However, little information is available about how different factors shape herpetofauna species assemblages in agricultural landscape at different spatial scales from patch (125 – 250m) to the landscape (500 – 1000m). We assessed the diversity of amphibians and reptiles in areas under low and high degrees of agricultural intensification and explored different factors regulating diversity at different spatial scales using four sampling methods. Diversity and abundance of amphibians varied significantly between the two zones, but not for reptiles. Agricultural intensification index (AII), calculated based on agrochemical use and area under agriculture at 250m scale, seemed to affect amphibians both at patch as well as at 500m and 1000m landscape scales. The AII influenced reptilian diversity only at patch and 500m scales. Vicinity of natural forest had a stronger influence on reptilian abundance. Seminatural vegetation impacted herpetofauna diversities at larger spatial scales. The extent of water bodies influenced the reptilian abundance at 250m patch scale and amphibian abundance both at 250m and 1000m scale. Fallow lands affected only reptilian diversity at all spatial scales. Plantation affected amphibian at all scales but reptiles only at the landscape scale. Habitat heterogeneity regulated only amphibian diversity. These results highlight the fact that different patch and landscape-scale factors regulate the diversity of reptiles and amphibians differentially. Such scale-specific information will crucially inform future conservation action for the herpetofauna in the agricultural landscape.     

Arthropod biodiversity in olive groves under two soil management systems: presence versus absence of herbaceous cover crop

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湖南壶瓶山国家级自然保护区两栖 爬行动物多样性的时空格局

本研究利用2013至2016年湖南壶瓶山国家级自然保护区巡护监测数据,评估壶瓶山保护区两栖爬行动物多样性水平。壶瓶山保护区在8个监测站点布设了25条两栖爬行动物调查样线,进行常规监测,其中,日间样线14条,夜间样线11条,日间样线平均长度4 km,夜间样线平均长度0.75 km。每个月,每条样线平均调查3次。4年共调查到62种两栖爬行动物,占湖南省两栖爬行动物总种数的38.04%。逐月计算各监测站点的alpha多样性指数和整个保护区的beta多样性指数,按季节比较不同站点间物种组成的Hellinger距离。结果显示,保护区不同位置间、不同的监测站点间以及年际间的两栖爬行动物多样性没有显著差异,提示保护区环境具有相对稳定性。但是,个别位于实验区或邻近实验区的站点,物种组成有着剧烈的波动,表明人为活动影响了两栖爬行动物的分布。另外,保护区两栖爬行动物的物种组成与多样性有着显著的季节差异,这与两栖爬行动物的生物学特性有关。以上结果说明,壶瓶山保护区两栖爬行动物的多样性结构具有稳定性和敏感性的特点,可以考虑用作保护区生物多样性水平的监测指标。     

Exploring the effect of soil management intensity on taxonomic and functional diversity of ants in Mediterranean olive groves

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Amphibian diversity increases in an heterogeneous agricultural landscape

As a group amphibians are the vertebrates most affected by anthropic activity, particularly by agriculture. The rapid advance of the agricultural frontier makes it important to identify the role of agroecosystems as habitat supply for amphibians. We analyzed the differences in amphibian assemblages and populations between habitats with different plant covers and different degrees of human intervention in northwestern Argentina. For three years we conducted 114 high frequency trap samplings to quantify abundance, specific composition and species richness of amphibian assemblages in three habitat types (lemon and sugarcane crops and secondary forest) of a piedmont agroecosystem of Tucumán province. Crops hosted more species and individuals than secondary forests, but the specific composition of forest was different from that of crops suggesting that they could be complementary. Although the assemblage abundance of every observation responded to climate, the strong effect of sampling year was not related to climatic factors suggesting that there might be long term fluctuations that were not analyzed. We also found that responses to agricultural practices were species specific, so no generalizations about these practices should be done. Our study shows that cultivated areas are not hostile environments for amphibians since they can lodge huge amphibian populations and that the occurrence of disturbances associated to agricultural practices is not necessarily reflected in a decrease in the abundance and richness of amphibians in the short term. However, our results also show that forested lands are necessary to lodge some specialist species which are very rare in the croplands. This suggests that environmental heterogeneity generated by the combination of natural and cultivated patches can increase biodiversity at landscape scale because this allows the coexistence of species related to either kind of environment.     

Spatial Biodiversity Patterns of Madagascar's Amphibians and Reptiles

Madagascar has become a model region for testing hypotheses of species diversification and biogeography, and many studies have focused on its diverse and highly endemic herpetofauna. Here we combine species distribution models of a near-complete set of species of reptiles and amphibians known from the island with body size data and a tabulation of herpetofaunal communities from field surveys, compiled up to 2008. Though taxonomic revisions and novel distributional records arose since compilation, we are confident that the data are appropriate for inferring and comparing biogeographic patterns among these groups of organisms. We observed species richness of both amphibians and reptiles was highest in the humid rainforest biome of eastern Madagascar, but reptiles also show areas of high richness in the dry and subarid western biomes. In several amphibian subclades, especially within the Mantellidae, species richness peaks in the central eastern geographic regions while in reptiles different subclades differ distinctly in their richness centers. A high proportion of clades and subclades of both amphibians and reptiles have a peak of local endemism in the topographically and bioclimatically diverse northern geographic regions. This northern area is roughly delimited by a diagonal spanning from 15.5°S on the east coast to ca. 15.0°S on the west coast. Amphibian diversity is highest at altitudes between 800–1200 m above sea-level whereas reptiles have their highest richness at low elevations, probably reflecting the comparatively large number of species specialized to the extended low-elevation areas in the dry and subarid biomes. We found that the range sizes of both amphibians and reptiles strongly correlated with body size, and differences between the two groups are explained by the larger body sizes of reptiles. However, snakes have larger range sizes than lizards which cannot be readily explained by their larger body sizes alone. Range filling, i.e., the amount of suitable habitat occupied by a species, is less expressed in amphibians than in reptiles, possibly reflecting their lower dispersal capacity. Taxonomic composition of communities assessed by field surveys is largely explained by bioclimatic regions, with communities from the dry and especially subarid biomes distinctly differing from humid and subhumid biomes.     

Environmental determinants of amphibian and reptile species richness in China

Understanding the factors that regulate geographical variation in species richness has been one of the fundamental questions in ecology for decades, but our knowledge of the cause of geographical variation in species richness remains poor. This is particularly true for herpetofaunas (including amphibians and reptiles). Here, using correlation and regression analyses, we examine the relationship of herpetofaunal species richness in 245 localities across China with 30 environmental factors, which include nearly all major environmental factors that are considered to explain broad-scale species richness gradients in such theories as ambient energy, water–energy dynamics, productivity, habitat heterogeneity, and climatic stability. We found that the species richness of amphibians and reptiles is moderately to strongly correlated with most of the environmental variables examined, and that the best fit models, which include explanatory variables of temperature, precipitation, net primary productivity, minimum elevation, and range in elevation, explain ca 70% the variance in species richness for both amphibians and reptiles after accounting for sample area. Although water and temperature are important explanatory variables to both amphibians and reptiles, water variables explain more variance in amphibian species richness than in reptile species richness whereas temperature variables explain more variance in reptile species richness than in amphibian species richness, which is consistent with different physiological requirements of the two groups of organisms.     

北京市两栖爬行动物空间分布格局及影响因素

基于遥感和野外调查数据,研究北京市两栖爬行动物空间分布格局及其影响因素。结果显示,北京记录两栖爬行动物2纲3目10科17属22种,其中,保护物种15种,占物种总数的68.19%。物种分布整体呈现由西北向东南递减的空间特征,西北部山区丰富度和相对多度较大,门头沟和延庆物种丰富度最大(14种),占调查物种总数的66.67%,优势种为中国林蛙、黑斑侧褶蛙、中华蟾蜍和山地麻蜥;门头沟、怀柔、房山分布了全市71.54%的种群数量;200-500 m海拔段物种丰富度最高(18种),主要生境为森林灌丛;物种数在6种以上的网格有11个,包括大滩、怀九河、百花山、拒马河、密云水库等。总体来看,物种种类和数量相较之前呈现减少趋势,而土地用途转换具有重要影响。1980-2018年,建设用地表现出以东西城为核心向外围扩张的趋势,侵占的农田占生态系统总转换面积的42.85%;城六区和平原区的主要物种为黑斑侧褶蛙和中华蟾蜍,少有中国林蛙分布。进一步分析两栖动物1500 m的迁徙距离内,森林灌丛和农田为主要的生境类型,城市扩张侵占了部分农田;同时,污染物的排放引起水体水质下降,特别是北运河水系,造成两栖动物的繁殖率降低和栖息地破坏。目前,北京市各类保护地虽覆盖两栖爬行动物种类的90.91%和数量的60.45%,但在怀柔宝山镇白河上段、密云水库来水的潮河上段辛庄桥、平谷洳河大兴庄镇-峪口镇等地,仍然存在较大的保护空缺。对此,建议开展长期定位观测,构建生物多样性观测网络,并探索建立高强度土地利用下的生物多样性保护机制,从而实现生物资源可持续利用。     

Amphibian species and functional diversity as indicators of restoration success in tropical montane forest

Tropical forest restoration is increasingly seen as an activity that may counteract or reduce biodiversity loss. However, few studies monitor fauna or consider measures of functional diversity to assess restoration success. We assessed the effect of a tropical montane forest restoration program on species and functional diversity, using amphibians as the target group. We compared amphibian assemblages in three types of land use: restoration areas, tropical montane cloud forest (TMCF; reference ecosystem) and cattle pastures (degraded ecosystem) in southern Mexico. We also described microclimate, microhabitat heterogeneity, woody vegetation structure and diversity for each type of land use, and their relationship to amphibian species and functional diversity. Compared to TMCF, restoration areas had similar environmental conditions. However, amphibian species richness was similar in the three types of land use and abundance was lower in the restoration areas. In TMCF, the amphibian assemblage was dominated by forest-specialist species, the pastures by generalist species, and the restoration areas by a combination of both species types. Interestingly, functional richness, functional evenness and functional divergence did not vary with land use, though the number of functional groups in restoration areas and TMCF was slightly higher. Overall, the results suggest that after seven years, active restoration provided habitat heterogeneity and recovered woody vegetation capable of maintaining amphibian species and functional groups similar to those inhabiting TMCF. Forest fragments adjacent to restoration areas seem to facilitate fauna recolonization and this emphasizes the importance of the conservation of the reference ecosystems to achieving restoration success.     

Can a large metropolis sustain complex herpetofauna communities? An analysis of the suitability of green space fragments in Rome

Urban areas are primary causes of species' range fragmentation and reduction. However, relatively few studies have attempted to describe the habitat variables influencing the diversity and conservation of amphibians and reptiles, particularly in Mediterranean Europe and in large metropolitan areas. We explored this broad conservation ecology problem by studying the richness and diversity patterns in relation to a suite of six independent habitat variables in Rome, one of the most ancient cities of the world. We considered all the green remnant areas ( n =62) of Rome, ranging 1 to >1000 ha in size, which are interspersed within a sea of urbanized matrix. A total of 10 amphibian and 15 reptile species were studied. Their presence/absence patterns were assessed and the effects of the various habitat variables on each species were predicted by a logistic regression model. A total of 1261 presence records (404 amphibians and 857 reptiles) were analysed. Fragment size and wood size within each fragment did correlate significantly with the species richness of both amphibians and reptiles, and there was a clear threshold effect after 50 ha of wooded surface. The presence of water bodies positively affected the species distribution. One amphibian and three reptiles inhabited exclusively fragments >50 ha. The distance from the centre did not affect fragment species richness. The presence of most species of both amphibians and reptiles was positively influenced by the irregular versus circular shape of the wooded area. The legal protection of a given area did not influence the observed patterns but the total number of sheltered species. Overall, our study suggests that, in order to maintain the current diversity and population viability, it is necessary, in addition to water bodies' maintenance, to (1) preserve the wooded landscapes over 50 ha; (2) promote irregularly shaped increases in the wood surface; (3) maintain ecotonal boundaries.     

Conventional land‐use intensification reduces species richness and increases production: A global meta‐analysis

Most current research on land‐use intensification addresses its potential to either threaten biodiversity or to boost agricultural production. However, little is known about the simultaneous effects of intensification on biodiversity and yield. To determine the responses of species richness and yield to conventional intensification, we conducted a global meta‐analysis synthesizing 115 studies which collected data for both variables at the same locations. We extracted 449 cases that cover a variety of areas used for agricultural (crops, fodder) and silvicultural (wood) production. We found that, across all production systems and species groups, conventional intensification is successful in increasing yield (grand mean + 20.3%), but it also results in a loss of species richness (?8.9%). However, analysis of sub‐groups revealed inconsistent results. For example, small intensification steps within low intensity systems did not affect yield or species richness. Within high‐intensity systems species losses were non‐significant but yield gains were substantial (+15.2%). Conventional intensification within medium intensity systems revealed the highest yield increase (+84.9%) and showed the largest loss in species richness (?22.9%). Production systems differed in their magnitude of richness response, with insignificant changes in silvicultural systems and substantial losses in crop systems (?21.2%). In addition, this meta‐analysis identifies a lack of studies that collect robust biodiversity (i.e. beyond species richness) and yield data at the same sites and that provide quantitative information on land‐use intensity. Our findings suggest that, in many cases, conventional land‐use intensification drives a trade‐off between species richness and production. However, species richness losses were often not significantly different from zero, suggesting even conventional intensification can result in yield increases without coming at the expense of biodiversity loss. These results should guide future research to close existing research gaps and to understand the circumstances required to achieve such win‐win or win‐no‐harm situations in conventional agriculture.     

Climate and landscape explain richness patterns depending on the type of species’ distribution data

Understanding the patterns of species richness and their environmental drivers, remains a central theme in ecological research and especially in the continental scales where many conservation decisions are made. Here, we analyzed the patterns of species richness from amphibians, reptiles and mammals at the EU level. We used two different data sources for each taxon: expert-drawn species range maps, and presence/absence atlases. As environmental drivers, we considered climate and land cover. Land cover is increasingly the focus of research, but there still is no consensus on how to classify land cover to distinct habitat classes, so we analyzed the CORINE land cover data with three different levels of thematic resolution (resolution of classification scheme ˗ less to more detailed). We found that the two types of species richness data explored in this study yielded different richness maps. Although, we expected expert-drawn range based estimates of species richness to exceed those from atlas data (due to the assumption that species are present in all locations throughout their region), we found that in many cases the opposite is true (the extreme case is the reptiles where more than half of the atlas based estimates were greater than the expert-drawn range based estimates). Also, we detected contrasting information on the richness drivers of biodiversity patterns depending on the dataset used. For atlas based richness estimates, landscape attributes played more important role than climate while for expert-drawn range based richness estimates climatic variables were more important (for the ectothermic amphibians and reptiles). Finally we found that the thematic resolution of the land cover classification scheme, also played a role in quantifying the effect of land cover diversity, with more detailed thematic resolution increasing the relative contribution of landscape attributes in predicting species richness.     

Land use and biodiversity congruences at local scale: applications to conservation strategies

The establishment of protected areas is traditionally considered indispensable to preserve biodiversity hotspots or areas inhabited by threatened species. The analysis of correlations between composition or richness of several taxonomic groups in a specific area has emerged as a useful mechanism to quickly identify areas of biological and conservation interest, and is currently used to select and design protected areas. The effect of habitat fragmentation on these correlations at a regional scale has been studied, but the effect of land use on the correlations obtained at local scale is poorly understood. We evaluated the relationships among different taxonomic groups frequently used in biological assessments for reserve design (vascular flora, amphibians, reptiles, birds and mammals), taking into account the effects of land use on these correlations. We compared richness and species composition of these groups in 1 km² quadrants within a total area of 75 km² in a low mountain Mediterranean area. Richness and composition were significantly correlated between several groups, both using the complete data set and also analyzing natural and disturbed areas separately. Species composition and species richness correlations were not congruent at the full landscape approach, nor natural and disturbed quadrants. Factors extrinsic to the communities also varied their influence in the assemblage of the community and species richness or not land uses were taken into account. According to our results, natural and human-disturbed areas should not be combined in cross-taxon congruences analysis at local scale.     

Amphibian and reptile communities of rainforest fragments: minimum patch size to support high richness and abundance

Information on animal communities inhabiting Neotropical fragmented landscapes is important for developing conservation strategies. The structure of amphibian and reptile communities in six tropical rainforest fragments (<20?ha) and two reference areas in continuous forest at Los Tuxtlas, Mexico was studied. A total of 3,481 individuals of 51 species of amphibians and reptiles were recorded across 12 bimonthly surveys during 2 years. Taxonomic composition was different between the smallest fragments and the reference areas. Six species were exclusive to large undisturbed forest and richness was significantly lower in the five smallest fragments (1.4–6.6?ha) compared with the largest patch, one or both of the reference areas. Amphibian abundance tended to be higher in large areas, while reptiles were more abundant in the five smallest fragments. Craugastor loki and Anolis uniformis were the dominant species in all sites, and particularly in the smaller fragments. Amphibian and reptile richness was positively related to larger patch sizes, deeper leaf litter, closed canopy cover, and higher relative humidity and negatively related to linear patch shape and high temperatures. Abundance of reptiles was positively associated to high temperatures, high density of woody debris, and closed canopy cover; it was negatively affected by linear patch shape, low humidity levels, and steeper slopes. While amphibian and reptile communities were strongly affected in vegetation fragments, these patches retained a considerable number of rainforest species. Fragments up to at least 17?ha have the potential for preserving communities with similar structure to those occurring in large tropical rainforests.     

Environmental Correlates for Species Richness among Amphibians and Reptiles in a Climate Transition Area

Correlations between environmental factors and the distribution of amphibian and reptile species richness were investigated in a climate transition area, Peneda-Gerês National Park (PNPG), in North-Western Portugal. Using presence-data at a local-scale (1 × 1 km), Ecological-Niche Factor Analysis (ENFA) identified a mixture of climatic (precipitation and number of days with fog), topographical (altitude and relief) and habitat factors (number of watercourses and water surfaces, the type of the largest water surface and tree diversity cover), as accurate predictors of species occurrence. Three factors were common for both taxonomic groups, and consistently presented a positive relation with species occurrence: precipitation, number of water surfaces, and tree diversity cover; suggesting a strong coincidence in the environmental correlates that influence amphibian and reptile species richness. Distribution patterns of observed and predicted species richness were compared using a Geographical Information System. Overall, three high species richness areas were predicted in common for both taxonomic groups and two additional areas for amphibians only. These areas matched with the observed species richness but suggested larger areas of high species richness. The location of the PNPG in a biogeographic crossroad, between Euro-Siberian and Mediterranean provinces, emphasised species richness of amphibians and reptiles and suggests a high priority conservation status for this protected area. Most of Central-Northern Portugal is located in a climatic transition area; therefore, increased species richness should be expected for other areas. Local scale studies for other protected areas should be planned as a framework for the development of multi-scale conservation planning by Portuguese authorities.     

Evolutionarily distinct amphibians are disproportionately lost from human‐modified ecosystems

Humans continue to alter terrestrial ecosystems, but our understanding of how biodiversity responds is still limited. Anthropogenic habitat conversion has been associated with the loss of evolutionarily distinct bird species at local scales, but whether this evolutionary pattern holds across other clades is unknown. We collate a global dataset on amphibian assemblages in intact forests and nearby human‐modified sites to assess whether evolutionary history influences susceptibility to land conversion. We found that evolutionarily distinct amphibian species are disproportionately lost when forested habitats are converted to alternative land‐uses. We tested the hypothesis that grassland‐associated amphibian lineages have both higher diversification and are pre‐adapted to human landscapes, but found only weak evidence supporting this. The loss of evolutionarily distinct amphibians with land conversion suggests that preserving remnant forests will be vital if we aim to preserve the amphibian tree of life in the face of mounting anthropogenic pressures.     

Butterfly diversity at the ecotone between agricultural and semi‐natural habitats across a climatic gradient

Aim Understanding the response of species to ecotones and habitat edges is essential to designing conservation management, especially in mosaic agricultural landscapes. This study examines how species diversity and composition change with distance from semi‐natural habitats, over ecotones into agricultural fields, and how within‐site patterns of community transition change across a climatic gradient and differ between crop types. Location A total of 19 sites in Israel where semi‐natural habitats border agricultural fields (wheat fields or olive groves) distributed along a sharp climatic gradient ranging between 100 and 800 mm mean annual rainfall. Methods  We performed butterfly surveys in 2006. We analysed species richness (α‐diversity), diversity, community nestedness and species turnover (β‐diversity) within sites and between sites (γ‐diversity). We also assessed where species of conservation concern occurred. Results In wheat sites, richness and diversity declined abruptly from ecotones to fields and remained homogenously poor throughout the fields, regardless of climate. In olive sites, despite the sharp structural boundary, richness and diversity remained high from the semi‐natural habitat to the grove margins and then declined gradually into groves. Species of conservation concern occurred across all habitats at olive sites, but none were found inside wheat fields or at their ecotones. The contrast in community structure between semi‐natural habitats and fields was affected by both climate and field type. Irrigation in arid regions did not augment species diversity. Main conclusions Our results indicate that consideration of crop type, within a climatic context, should receive high priority in biodiversity conservation in agricultural areas. In ‘hostile’ crops, such as wheat, we suggest favouring a combination of high‐intensity management and wide margins over less intensive management without margins, which may merely aid generalist butterfly species. The scarcity of butterflies in arid irrigated fields suggests a need to carefully assess the effects of irrigation and agrochemicals on species’ communities.     

Biological diversity of a temporary pond herpetofauna in north Florida sandhills

From 1985 through 1990, the herpetofauna of a temporary pond in an uplands longleaf pine sandhills community in north-central Florida was monitored. A drift fence completely encircled the pond. Animals were captured in pitfall traps and marked as they entered and exited the pond basin. I captured 16 155 individuals of 42 species (16 amphibians, 26 reptiles). The species richness, diversity (using Margalef's Diversity Index) and dominance (using the Berger-Parker Index) varied among years. Between 62.5% and 87.5% of the amphibian species and 65% to 81% of the reptile species were captured in any one year. Daily amphibian capture was positively correlated with rainfall, whereas reptile capture was either not correlated or weakly negatively correlated with rainfall. Hydroperiod duration was not correlated with the numbers of either amphibians or reptiles captured. Neither the amphibian nor the reptile community showed any trends in diversity or dominance indices during the course of the study, although both communities were dominated by a few species. However, the species responsible for community dominance changed somewhat as the study progressed. Assessing the results of this study is hampered by the lack of comparable studies elsewhere, expected natural fluctuations of amphibian populations, and a prolonged drought, especially during the latter stages of the study. The herpetological community at Breezeway Pond does not appear to follow theoretical predictions of community response to stress. Temporary ponds are important centres of herpetofaunal biodiversity in uplands sandhills communities. Long-term studies are needed to monitor the composition, structure, and functional interactions of their resident species.     

The distribution of amphibian species richness in protected areas of Chiapas,Mexico

Due to the current environmental crisis, many animal species face extinction problems. Amphibian populations have been affected by this crisis. Our goal is to study amphibian species diversity in Chiapas, which has 7.6% of the endemic amphibians in Mexico and 53 protected areas. Only 58% of the protected areas have management plans or information on their resident amphibians. We aim to determine the extent of protection provided by the network of natural areas for the conservation of amphibian species in the state and to discuss the effectiveness of this protection. Therefore, we compiled a georeferenced database of 112 amphibian species in Chiapas to create each distribution model. In addition, we carried out representativeness, beta diversity, and species richness analyses. As a result, we obtained a high degree of representativeness for the records and species distribution models. However, we found a decrease in the richness of amphibians involving 20% of total species, 13% of endemics, 18% threatened according to NOM-059, and 31% threatened according to IUCN between 1800 and 2020 and 1980–2020. We also identified two biodiversity hotspots in the Sierra Madre de Chiapas and the Northern Highlands physiographic regions. Finally, based on potential distributions, we found more endemic and threatened species outside protected natural areas than inside them. Our results give a broader picture of how amphibian richness is distributed in Chiapas. This information can help to prioritize conservation efforts toward those areas rich in threatened or endemic species, such as the Northern Mountains Hotspot we identified in northern Chiapas.     

Disentangling vegetation and climate as drivers of Australian vertebrate richness

Determining drivers of species richness is recognised as highly complex, involving many synergies and interactions. We examine the utility of newly available remote sensing representations of vegetation productivity and vegetation structure to examine drivers of species richness at continental and regional scales. We related richness estimates derived from stacked species distribution models for birds, mammals, amphibians, and reptiles to estimates of actual and potential evapotranspiration (AET and PET), forest structure, and forest productivity across Australia as a whole as well as by bioclimatic zones. We used structural equation modeling to partition correlations between climate energy and vegetation attributes and their subsequent associations with species richness. Continentally, vertebrate richness patterns were strongly related to patterns of energy availability. Richness of amphibians, mammals, and birds were positively associated with AET. However, reptile richness was most strongly associated with PET. Regionally, forest structure and productivity associations with bird, mammal, and amphibian richness were strongest. Again, reptile richness associated most strongly with PET. Our results suggest that a hierarchy of drivers of broad‐scale vertebrate richness patterns exist (reptiles excluded): 1) climate energy is most important at the continental scale; next, 2) vegetation productivity and vegetation structure are most important at the regional scale; except 3) at low extremes of climate energy when energy becomes limiting.