A global comparative analysis of elevational species richness patterns of ferns

Aim Elevational gradients offer an outstanding opportunity to assess factors determining patterns of species richness, but along single transects potential explanatory factors often covary, making it difficult to distinguish between competing hypotheses. Many previous studies on plants have interpreted their results as supporting the mid‐domain effect (MDE) as a major determinant of species richness, even when climatic factors showed similarly high explanatory power. We compared fern species richness along 20 elevational transects to quantify the relative contribution of climate and MDE as drivers of elevational richness patterns. Location Twenty transects world‐wide. Methods Ferns were sampled in 1039 plots of 400–2500 m² each. Mean annual precipitation and temperature, epiphytic bryophyte cover (as a proxy for air humidity) and MDE predictions were included as independent variables. For each transect, we calculated multiple linear models and partitioned the variance to assess the relative contribution of the independent variables, selecting the most parsimonious models based on Akaike weights and multi‐model inference. Results Along most individual gradients, nearly all variance of fern species richness that could be attributed to either space or MDEs was collinear with climatic factors. Yet, the comparison across transects showed that elevational richness patterns are most parsimoniously accounted for by climatic conditions, especially by low water availability at low elevations and in dry regions in general, and by low temperatures at high elevations and in extra‐tropical regions. Main conclusions Fern species richness is most closely related to climatic factors, and while MDE, surface area and metapopulation processes may somewhat modify the patterns, their importance has been overstated in the past. Future research challenges include determining whether the richness–climate relationship reflects: (1) a direct relationship through the physiological tolerance of the plants, (2) an indirect influence of climate on ecosystem productivity, or (3) an evolutionary legacy of longer or faster diversification processes under certain climatic conditions.     

喜马拉雅山哺乳动物物种多样性垂直分布格局

生物多样性的空间分布及其相关机制一直是生态学、生物地理学和保护生物学研究的热点问题。山地生态系统生境异质性和生物多样性高, 适合研究生物多样性空间分布及其相关机制。喜马拉雅山脉位于青藏高原南缘, 是全球生态热点区域。其地形复杂, 海拔落差大(100-8,844 m), 具有明显的垂直气候带。本研究通过整合野外调查和文献资料, 系统地分析了10目23科160属313种喜马拉雅山地区哺乳动物物种多样性的垂直分布格局, 发现该区域哺乳动物总体及其子集的物种多样性垂直分布格局都为左偏倚的中峰格局, 物种多样性在海拔900-1,400 m之间最高, 不同物种子集的物种多样性垂直分布格局的模式有所不同。UPGMA聚类分析表明, 喜马拉雅山地区哺乳动物群落沿海拔梯度可以划分为5个聚类簇(海拔100-1,500 m、1,500-2,000 m、2,000-3,000 m、3,000-4,200 m以及4,200-6,000 m的地区), 大致与该地区植被的垂直带分布相吻合。喜马拉雅山地区哺乳动物物种多样性在中低海拔最为丰富, 可能跟东洋界与古北界生物群扩散后的交汇地带相关。喜马拉雅山区贯通南北的沟谷是生物扩散和迁移的通道, 沟谷内水热资源较好, 气候稳定性高, 为高山生态系统内各种生物创造了栖息条件。综上, 喜马拉雅山沟谷地区是生物多样性热点地区, 也是生物扩散和交流关键的“生态走廊”, 应加强对喜马拉雅山沟谷地区的保护, 以维系该区域较高的生物多样性。     

The impact of population processes on patterns of species richness: Lessons from elevational gradients

In the last few years, considerable headway has been made towards understanding patterns of species richness along latitudinal and elevational gradients, mostly by focussing on the influences of surface area, climatic factors, evolutionary history, and stochastic processes. However, the potential impact of population-level processes in determining or modifying patterns of species richness has largely been neglected, partly due to the difficulty of gathering such data for numerous species along geographical or ecological gradients. Based on two empirical examples, I here show that dispersal and the resulting source-sink effects modify patterns of plant species richness along elevation gradients, and that the inclusion or exclusion of such sink populations alters the perception of the diversity patterns and hence our interpretation of them. I argue that population processes should be taken into account when studying patterns of species richness, especially at scales at which dispersal is common in the taxon under consideration.     

The effect of area on local and regional elevational patterns of species richness

Aim To calculate the degree to which differences between local and regional elevational species richness patterns can be accounted for by the effects of regional area. Location Five elevational transects in Costa Rica, Ecuador, La Réunion, Mexico and Tanzania. Methods We sampled ferns in standardized field plots and collated regional species lists based on herbarium and literature data. We then used the Arrhenius function S = cA^z to correct regional species richness (S) for the effect of area (A) using three slightly different approaches, and compared the concordance of local and regional patterns prior to and after accounting for the effect of area on regional richness using linear regression analyses. Results We found a better concordance between local and regional elevational species richness after including the effect of area in the majority of cases. In several cases, local and regional patterns are very similar after accounting for area. In most of the cases, the maximum regional richness shifted to a higher elevation after accounting for area. Different approaches to correct for area resulted in qualitatively similar results. Main conclusions The differences between local and regional elevational richness patterns can at least partly be accounted for by area effects, suggesting that the underlying causes of elevational richness patterns might be the same at both spatial scales. Values used to account for the effect of area differ among the different study locations, showing that there is no generally applicable elevational species–area relationship.     

Mapping patterns of ferns species richness through the use of herbarium data

This paper aims to analyse the spatial patterns of sampling effort and species richness of pteridophyte in a well-investigated region as Tuscany, Italy, by using data stored from a geodatabase storing information on the specimens preserved in the main herbaria of the region. A total of 6,905 records about pteridophyte specimens were extracted from the geodatabase, and 5,638 of such specimens were studied through the use of spatial statistical techniques. The data about the sampling effort and species richness were analysed in relation to topographical variables to assess any significant relationship. Specimen-based rarefaction techniques were used to compare areas with different number of detected species. The analysis of the sampling effort data showed a nonhomogeneous distribution of herbarium data, with some areas being intensively sampled and others being almost unsampled. Thus, the geographical distribution of specimens was extremely clustered. The comparison across geographical areas through specimen-based rarefaction curves showed great differences in species richness and sampling completeness. The analysis of the residuals of species–area relationships evidenced that the distance to water bodies was the only significant topographical variable in controlling species diversity.     

Regional and global elevational patterns of microbial species richness and evenness

Although elevational gradients in microbial biodiversity have attracted increasing attention recently, the generality in the patterns and underlying mechanisms are still poorly resolved. Further, previous studies focused mostly on species richness, while left understudied evenness, another important aspect of biodiversity. Here, we studied the elevational patterns in species richness and evenness of stream biofilm bacteria and diatoms in six mountains in Asia and Europe. We also reviewed published results for elevational richness patterns for soil and stream microbes in a literature analysis. Our results revealed that even within the same ecosystem type (that is, stream) or geographical region, bacteria and diatoms showed contrasting patterns in diversity. Stream microbes, including present stream data, tend to show significantly increasing or decreasing elevational patterns in richness, contrasting the findings for soil microbes that typically showed nonsignificant or significantly decreasing patterns. In all six mountains for bacteria and in four mountains for diatoms, species richness and evenness were positively correlated. The variation in bacteria and diatom richness and evenness were substantially explained by anthropogenic driven factors, such as total phosphorus (TP). However, diatom richness and evenness were also related to different main drivers as richness was mostly related to pH, while evenness was most explained by TP. Our results highlight the lack of consistent elevational biodiversity patterns of microbes and further indicate that the two facets of biodiversity may respond differently to environmental gradients.     

Explaining Andean megadiversity: the evolutionary and ecological causes of glassfrog elevational richness patterns

The Tropical Andes are an important global biodiversity hotspot, harbouring extraordinarily high richness and endemism. Although elevational richness and speciation have been studied independently in some Andean groups, the evolutionary and ecological processes that explain elevational richness patterns in the Andes have not been analysed together. Herein, we elucidate the processes underlying Andean richness patterns using glassfrogs (Centrolenidae) as a model system. Glassfrogs show the widespread mid‐elevation diversity peak for both local and regional richness. Remarkably, these patterns are explained by greater time (montane museum) rather than faster speciation at mid‐elevations (montane species pump), despite the recency of the major Andean uplift. We also show for the first time that rates of climatic‐niche evolution and elevational change are related, supporting the hypothesis that climatic‐niche conservatism decelerates species' shifts in elevational distributions and underlies the mid‐elevation richness peak. These results may be relevant to other Andean clades and montane systems globally.     

Saprotrophic and ectomycorrhizal fungi exhibit contrasting richness patterns along elevational gradients in cool-temperate montane forests

Increasing evidence suggests that elevational gradients of soil fungal richness are highly variable, but few studies have examined how diversity components of each guild contribute to overall fungal diversity. Here, we aimed to disentangle the relationships between total, saprotrophic, and ectomycorrhizal fungal richness, and environmental factors across elevational gradients in cool-temperate montane forests. We observed that total and saprotrophic richness decreased but ectomycorrhizal richness increased with increase in elevation. Elevational range size and nestedness analyses illustrated that saprotrophic and ectomycorrhizal source communities were located at lower and higher elevations, respectively. The observed total and saprotrophic richness were directly influenced by soil properties and indirectly influenced by climate and plant communities. Ectomycorrhizal fungal richness was affected by climate and the dominance of ectomycorrhizal trees. We highlight that two directional source–sink dynamics lead to opposite elevational patterns between saprotrophic and ectomycorrhizal fungal richness, shaping the variation in elevational richness gradients.     

The impact of human pressure on landscape patterns and plant species richness in Mediterranean coastal dunes

We analyze the spatial patterns of natural dune cover patches and their plant richness, comparing coastal sites with different levels of human pressure in central Italy. We created a detailed land cover map of dune sites. The spatial pattern of natural dune cover types was characterized by computing a set of patch-based metrics. To quantify patch plant richness, we used 16 m² vegetation plots, randomly distributed on coastal dune cover types. For each patch, the richness of the entire pool of species and of three guilds (i.e., typical dune, ruderal, and alien species) was considered. We compared different levels of human pressure on coastal dunes focusing on pattern metrics and floristic information by using the nonparametric Kruskal–Wallis test. In sites with high human pressure, we have observed a general simplification in the natural dune spatial pattern and a decline of plant richness but with a specific response for each cover type. Alien and ruderal species presented low richness in all patches. In coastal dunes, the harsh ecological conditions and the strong sea–inland gradient shape the distribution of human activities and control the number of ruderal species. The approach effectively describes fragmentation and biodiversity in dune ecosystems.     

神农架自然保护区非飞行哺乳动物的物种丰富度:沿海拔梯度的分布格局

于1999～2001年调查了神农架自然保护区6个地点不同栖息地的非飞行哺乳动物的物种丰富度。栖息地分为8类：原始林、择伐林（采伐枯立木）、次生林、灌木林、草地、常年性河流水溪、农田和人居住区。小型非飞行哺乳动物调查用捕鼠夹;大型非飞行哺乳动物调查主要根据皮张收购资料以及样线法和痕迹法;用10 m×10 m的样方调查林地树种丰富度。调查发现,神农架自然保护区有非飞行哺乳动物59种［不包括引进种梅花鹿（Cervus nippon）］。在同一海拔高度,原始林通常比择伐林和次生林的物种丰富度高,说明采伐严重降低了物种丰富度。对比同一栖息地不同海拔高度的物种丰富度,我们发现,在中海拔地段（800～1700 m）物种丰富度最高：如在原始林和次生林,海拔1700 m的东溪物种丰富度最高;在择伐林,海拔800 m的九冲物种丰富度最高。聚类分析显示,6个地点的哺乳动物物种组成可以分为两组：高海拔组（2100 m以上）和中低海拔组（1700 m以下）。各地点的哺乳动物物种组成与植被的垂直分布是一致的。各地点的物种丰富度与单位面积（100 m²）树种平均丰富度、栖息地类型数和海拔高度相关。3个环境变量间也是相关的：海拔高度对单位面积树种平均丰富度和栖息地类型数有重要影响。根据研究结果提出两点保护建议：第一,保护区的移民迁出和退耕还林工程应首先在物种丰富度最高的九冲进行,而后是东溪和下谷;第二,为了增加个体流和基因流,保护区东西两片相间的非保护区地带应划入保护区,建立栖息地廊道。     

Influences of interpolation of species ranges on elevational species richness gradients

Interpolation of species ranges has been a common approach to compensate for the unevenness or incompleteness in sampling effort in studies of geographic species richness gradients. However, potential biases introduced by this estimation method remain unclear. Here, we presented an explicit examination of the influences of one‐dimensional interpolation on elevational species richness gradients, and discussed potential causes and processes of these influences. We conducted intensive surveys of birds along the elevational gradients of the Ailao Mountains, southwestern China, and compared richness patterns based on interpolation with raw data as well as estimated data from rarefaction and Chao1 non‐parametric estimator; we also compared results of multiple linear regressions and hierarchical partitioning analyses explaining these four measures of richness. Actual evapotranspiration (AET) and the mid‐domain effect (MDE) were highly correlated and separately provided a good potential explanation for the unimodal richness pattern in the Ailao Mountains, with modifying and suppressive effects of other variables such as area. Interpolation consistently and significantly increased the effects of AET/MDE, while it reduced contributions of area and human disturbance. Our results demonstrated that while compensating for biases in sampling effort, interpolation may also spuriously fill genuine distribution gaps, and tend to underestimate the effects of the non‐monotonic or discontinuous influencing factors that are responsible for these gaps, and overestimate the effects of other factors actually suppressed by these factors. These influences were most strong for species with relatively medium elevational ranges. We conclude that at the regional scale, interpolation method is a potential source of bias in identifying and explaining species richness gradients and should be used with careful consideration. It may be advantageous to adopt other robust estimation methods besides interpolation to gain a more accurate assessment of species richness and a more objective understanding of their underlying mechanisms.     

Could temperature and water availability drive elevational species richness patterns? A global case study for bats

Aim A global meta‐analysis was used to elucidate a mechanistic understanding of elevational species richness patterns of bats by examining both regional and local climatic factors, spatial constraints, sampling and interpolation. Based on these results, I propose the first climatic model for elevational gradients in species richness, and test it using preliminary bat data for two previously unexamined mountains. Location Global data set of bat species richness along elevational gradients from Old and New World mountains spanning 12.5° S to 38° N latitude. Methods Bat elevational studies were found through an extensive literature search. Use was made only of studies sampling  70% of the elevational gradient without significant sampling biases or strong anthropogenic disturbance. Undersampling and interpolation were explicitly examined with three levels of error analyses. The influence of spatial constraints was tested with a Monte Carlo simulation program, Mid‐Domain Null. Preliminary bat species richness data sets for two test mountains were compiled from specimen records from 12 US museum collections. Results Equal support was found for decreasing species richness with elevation and mid‐elevation peaks. Patterns were robust to substantial amounts of error, and did not appear to be a consequence of spatial constraints. Bat elevational richness patterns were related to local climatic gradients. Species richness was highest where both temperature and water availability were high, and declined as temperature and water availability decreased. Mid‐elevational peaks occurred on mountains with dry, arid bases, and decreasing species richness occurred on mountains with wet, warm bases. A preliminary analysis of bat richness patterns on elevational gradients in western Peru (dry base) and the Olympic Mountains, WA (wet base), supported the predictions of the climate model. Main conclusions The relationship between species richness and combined temperature and water availability may be due to both direct (thermoregulatory constraints) and indirect (food resources) factors. Abundance was positively correlated with species richness, suggesting that bat species richness may also be related to productivity. The climatic model may be applicable to other taxonomic groups with similar ecological constraints, for instance certain bird, insect and amphibian clades.     

From lowlands to highlands: searching for elevational patterns of species richness and distribution of scarab beetles in Costa Rica

Aim Understanding the heterogeneous distribution of species on mountains is an important aim in ecology. Altitudinal gradients have enormous potential for improving our knowledge of trends in biodiversity and conservation. In this study, we investigated the variation in scarab beetle diversity (Dynastinae, Rutelinae and Melolonthinae) along an elevational tropical forest gradient. Location The Atlantic slope of the Guanacaste mountain range in Costa Rica. Methods Ultraviolet light traps placed in six forests situated from 100 to 1510 m were used. Changes in species composition and richness among elevations were investigated. Differences in the altitudinal patterns using different groups of species were examined: the whole assemblage, each separate subfamily and two different trophic habits (phytophagous or saproxylic). The effects of temperature, humidity and elevation on scarab distribution were tested using canonical correspondence analyses. The relationship between the community similarity of the studied forests and the altitudinal distance among them was also analysed. Results Species composition and richness changed along the gradient. The peak in species richness varied depending on the species group considered and in all cases occurred 500 or 800 m. Forests at these altitudes were also the richest in exclusive species. Species composition turnover among elevations appeared with a clear separation between lowland and highland fauna. The latter was lower in richness but also had exclusive species. Temperature, humidity and altitude affected species distribution, with altitude being the most important factor for all the subfamilies studied. Main conclusions Our results showed that species distribution fits a hump‐shaped pattern. The peak of this pattern varied depending on the taxonomic group and mountain analysed, highlighting the importance of evolutionary processes as species distribution drivers. The fact that species richness peaked at elevations where human impact is currently important underlines the value of the development of conservation strategies for these areas.     

Mammals on mountainsides: elevational patterns of diversity

The four major papers in this special feature present and interpret data from field studies on the distributions and diversity of small mammals in elevational gradients on mountains in the Philippines, Borneo, southern Mexico and western United States. In the introductory paper, Lomolino places these studies in the context of historical, methodological and conceptual themes in contemporary biogeography. In this final paper, I focus on some important similarities and interesting differences among the four case studies. All of the studies provide evidence for the influence of ecological factors, such as climate, productivity and habitat heterogeneity, on mammalian diversity. All also provide evidence for the influence of historical dispersal, extinction, and speciation events. Perhaps the most interesting result is the documentation of a frequent, but not universal, peak in species diversity at some elevation intermediate between the base and peak of a mountain. Efforts to understand the mechanistic basis for this pattern — and why it differs from the continuous decrease in diversity from the equator to the poles — promise to contribute to developing a general theoretical explanation for the major patterns of biodiversity on earth.