面积、温度及分布区限制对物种丰富度海拔格局的影响: 以秦岭太白山为例

 物种丰富度的分布格局及其形成机制是生态学研究的热点。以往的研究主要描述丰富度的格局, 而对其形成机制研究较少, 且主要集中于探讨单个因子或过程的影响。物种丰富度同时受到多个因子和过程的综合作用, 面积、温度及物种分布区限制被认为是控制山地物种丰富度海拔格局的主要因素, 三者同时沿海拔梯度而变化, 同时作用于丰富度的海拔格局。幂函数种-面积关系(SAR)、生态学代谢理论(MTE)及中域效应假说(MDE)分别基于以上3个因素, 从机制上解释了物种丰富度 的海拔格局。探讨这些假说的相对影响对研究物种丰富度的大尺度格局及其形成机制具有重要意义。方差分离方法有利于分解不同因素的影响, 为此, 该文以秦岭太白山的植物物种丰富度为例, 采用方差分离和逐步回归方法, 分析了SAR、MTE及MDE对物种丰富度海拔格局的影响。结果表明, 太白山的植物物种丰富度沿海拔梯度呈单峰分布格局, 但丰富度峰值存在类群差异; 对太白山所有植物物种丰富度的垂直格局而言, SAR、MTE及MDE分别解释了其物种丰富度随海拔变化的66.4%、19.8%和37.9%, 共同解释了84.6%, 在消除其他因素的影响后, SAR和MTE的独立影响较高(分别为25.5%和17.7%), 而MDE的独立影响不显著; 分类群研究则发现, 苔藓植物丰富度的海拔格局主要受MDE的影响, 蕨类植物丰富度的海拔格局同时受到SAR、MTE以及MDE的影响, 而种子植物物种丰富度的海拔格局主要受SAR和MTE影响。     

Small mammal species richness and turnover along elevational gradient in Yulong Mountain,Yunnan, Southwest China

Understanding the species diversity patterns along elevational gradients is critical for biodiversity conservation in mountainous regions. We examined the elevational patterns of species richness and turnover, and evaluated the effects of spatial and environmental factors on nonvolant small mammals (hereafter “small mammal”) predicted a priori by alternative hypotheses (mid‐domain effect [MDE], species–area relationship [SAR], energy, environmental stability, and habitat complexity]) proposed to explain the variation of diversity. We designed a standardized sampling scheme to trap small mammals at ten elevational bands across the entire elevational gradient on Yulong Mountain, southwest China. A total of 1,808 small mammals representing 23 species were trapped. We observed the hump‐shaped distribution pattern of the overall species richness along elevational gradient. Insectivores, rodents, large‐ranged species, and endemic species richness showed the general hump‐shaped pattern but peaked at different elevations, whereas the small‐ranged species and endemic species favored the decreasing richness pattern. The MDE and the energy hypothesis were supported, whereas little support was found for the SAR, the environmental stability hypothesis, and the habitat complexity. However, the primary driver(s) for richness patterns differed among the partitioning groups, with NDVI (the normalized difference vegetation index) and MDE being the most important variables for the total richness pattern. Species turnover for all small mammal groups increased with elevation, and it supported a decrease in community similarity with elevational distance. Our results emphasized for increased conservation efforts in the higher elevation regions of the Yulong Mountain.     

西双版纳种子植物物种多样性的垂直格局及机制

物种多样性海拔分布格局及其形成机制的研究是生物地理学和宏观生态学的重要议题之一。本文利用西双版纳植物专著资料, 结合高分辨率的地形和气候等数据, 探讨了面积、边界限制和现代气候对西双版纳野生种子植物物种丰富度及物种密度海拔分布格局的影响。结果表明: (1)物种丰富度呈单峰分布格局, 面积(81.9%)、边界限制(17.5%)和气候(60.0-69.3%)都不同程度地解释了物种丰富度的单峰格局; (2)利用幂函数种-面积关系计算的物种密度沿海拔大致呈减小的分布趋势, 气候的解释率降低为32.6-40.6%, 与边界限制无显著相关关系; (3)利用等面积高度带划分得到的物种密度沿海拔呈单峰变化趋势, 物种密度与边界限制无显著相关性, 但气候对物种密度的解释率为81.6-89.9%。研究结果有助于准确全面地理解物种多样性的海拔分布格局及其成因机制, 为西双版纳生物多样性保护提供理论支撑和实践指导。     

The mid‐domain effect and habitat complexity applied to elevational gradients: Moss species richness in a temperate semihumid monsoon climate mountain of China

The utility of elevational gradients as tools to test either ecological hypotheses and delineate elevation‐associated environmental factors that explain the species diversity patterns is critical for moss species conservation. We examined the elevational patterns of species richness and evaluated the effects of spatial and environmental factors on moss species predicted a priori by alternative hypotheses, including mid‐domain effect (MDE), habitat complexity, energy, and environment proposed to explain the variation of diversity. Last, we assessed the contribution of elevation toward explaining the heterogeneity among sampling sites. We observed the hump‐shaped distribution pattern of species richness along elevational gradient. The MDE and the habitat complexity hypothesis were supported with MDE being the primary driver for richness patterns, whereas little support was found for the energy and the environmental factors.     

When does diversity fit null model predictions? Scale and range size mediate the mid‐domain effect

Aim  Recently, a flurry of studies have focused on the extent to which geographical patterns of diversity fit mid-domain effect (MDE) null models. While some studies find strong support for MDE null models, others find little. We test two hypotheses that might explain this variation among studies: small-ranged groups of species are less likely than large-ranged species to show mid-domain peaks in species richness, and mid-domain null model predictions are less robust for smaller spatial extents than for larger spatial extents.
Location  We analyse data sets from elevational, riverine, continental and other domains from around the world.
Methods  We use a combination of Spearman rank correlations and binomial tests to examine whether differences within and among studies and domains in the predictive power of MDE null models vary with spatial scale and range size.
Results  Small-ranged groups of species are less likely to fit mid-domain predictions than large-ranged groups of species. At large spatial extents, diversity patterns of taxonomic groups with large mean range sizes fit MDE null model predictions better than did diversity patterns of groups with small mean range sizes. MDE predictions were more explanatory at larger spatial extents than at smaller extents. Diversity patterns at smaller spatial extents fit MDE predictions poorly across all range sizes. Thus, MDE predictions should be expected to explain patterns of species richness when ranges and the scale of analysis are both large.
Main conclusions  Taken together, the support for these hypotheses offers a more sophisticated model of when MDE predictions should be expected to explain patterns of species richness, namely when ranges and the scale of analysis are both large. Thus the circumstances in which the MDE is important are finite and apparently predictable.     

The mid-domain effect in ectomycorrhizal fungi: range overlap along an elevation gradient on Mount Fuji,Japan

Mid-domain effect (MDE) models predict that the random placement of species'' ranges within a bounded geographical area leads to increased range overlap and species richness in the center of the bounded area. These models are frequently applied to study species-richness patterns of macroorganisms, but the MDE in relation to microorganisms is poorly understood. In this study, we examined the characteristics of the MDE in richness patterns of ectomycorrhizal (EM) fungi, an ecologically important group of soil symbionts. We conducted intensive soil sampling to investigate overlap among species ranges and the applicability of the MDE to EM fungi in four temperate forest stands along an elevation gradient on Mount Fuji, Japan. Molecular analyses using direct sequencing revealed 302 EM fungal species. Of 73 EM fungal species found in multiple stands, 72 inhabited a continuous range along the elevation gradient. The maximum overlap in species range and the highest species richness occurred at elevations in the middle of the gradient. The observed richness pattern also fit within the 95% confidence interval of the mid-domain null model, supporting the role of the MDE in EM fungal richness. Deviation in observed richness from the mean of the mid-domain null estimation was negatively correlated with some environmental factors, including precipitation and soil C/N, indicating that unexplained richness patterns could be driven by these environmental factors. Our results clearly support the existence of microbial species'' ranges along environmental gradients and the potential applicability of the MDE to better understand microbial diversity patterns.     

Elevational gradients in ant species richness: area, geometry, and Rapoport's rule

Studying the distributions of plants and animals along environmental gradients can illuminate the factors governing and maintaining species diversity. There are two general predictions of how species richness and elevation are related: either species richness decreases monotonically with increasing elevation or richness peaks at mid-elevations. Several processes might contribute to this pattern. In this paper, I examine patterns in ant species richness along elevational gradients in three states in the western US: Colorado, Nevada, and Utah. I test for the effects of available area and the geometric constraints model on species richness patterns. I also test Rapoport's rescue hypothesis, which relates the extent of species' elevational ranges to patterns in species richness. In each state, species richness peaked at mid-elevations. Area explained more variation in species richness than the geometric constraints model in Colorado and Utah, but not in Nevada. Area and geometric constraints together explained 90%, 99%, and 57% of the variation in species richness in Colorado, Nevada, and Utah, respectively. Even though there were peaks at mid-elevations, I still found a strong Rapoport effect. This work suggests that the influences of area and geometric constraints cannot be overlooked when examining patterns in species richness along environmental gradients.     

Altitudinal patterns of seed plant richness in the Gaoligong Mountains, south-east Tibet, China

Elevational patterns of species richness and their underlying mechanisms have long been a controversial issue in biodiversity and biogeographical research, and several hypotheses have been proposed in the past decades. Local and regional studies have suggested that area and geometric constraint are two of major factors affecting the elevational pattern of species richness. In this study, using data of seed plants and their distribution ranges and a Digital Elevation Model data set, we explored altitudinal patterns of seed plant richness and quantified the effects of area and the mid-domain effect (MDE) on the richness patterns in a high mountain area, Gaoligong Mountains (ranging from 215 m to 5791 m a.s.l.) located in south-eastern Tibet, China. The results showed that richness and density (richness/log-transformed area) of seed plants at species, genus, and family levels all showed hump-shaped patterns along the altitudinal gradient. The altitudinal changes in richness of species with three different range sizes (< 500 m, 500–1500 m, and > 1500 m), species of different plant life-forms (trees, shrubs, and herbs), and endemic species further confirmed this finding. Analysis of Generalized Linear Model depicted that although the area of each elevational band was always in high correlation with the species richness, the MDE could explain 84.9%, 33.8%, 83.8%, and 84.5% of the total variation in richness for all species and the three species groups with different range sizes, respectively. This suggests that the MDE significantly influences the patterns of species richness and is likely be stronger for broad-ranged species than for narrow-ranged ones in the Gaoligong Mountains.     

Orchid Species Richness along Elevational and Environmental Gradients in Yunnan,China

The family Orchidaceae is not only one of the most diverse families of flowering plants, but also one of the most endangered plant taxa. Therefore, understanding how its species richness varies along geographical and environmental gradients is essential for conservation efforts. However, such knowledge is rarely available, especially on a large scale. We used a database extracted from herbarium records to investigate the relationships between orchid species richness and elevation, and to examine how elevational diversity in Yunnan Province, China, might be explained by mid-domain effect (MDE), species–area relationship (SAR), water–energy dynamics (WED), Rapoport’s Rule, and climatic variables. This particular location was selected because it is one of the primary centers of distribution for orchids. We recorded 691 species that span 127 genera and account for 88.59% of all confirmed orchid species in Yunnan. Species richness, estimated at 200-m intervals along a slope, was closely correlated with elevation, peaking at 1395 to 1723 m. The elevational pattern of orchid richness was considerably shaped by MDE, SAR, WED, and climate. Among those four predictors, climate was the strongest while MDE was the weakest for predicting the elevational pattern of orchid richness. Species richness showed parabolic responses to mean annual temperature (MAT) and mean annual precipitation (MAP), with maximum richness values recorded at 13.7 to 17.7°C for MAT and 1237 to 1414 mm for MAP. Rapoport’s Rule also helped to explain the elevational pattern of species richness in Yunnan, but those influences were not entirely uniform across all methods. These results suggested that the elevational pattern of orchid species richness in Yunnan is collectively shaped by several mechanisms related to geometric constraints, size of the land area, and environments. Because of the dominant role of climate in determining orchid richness, our findings may contribute to a better understanding of the potential effects of climate change on orchid diversity, and the development of conservation strategies for orchids.     

Terrestrial gastropod diversity in an alpine region: disentangling effects of elevation,area, geometric constraints,habitat type and land‐use intensity

Elevational gradients have proven to be useful to examine key factors shaping species diversity patterns. This study examines the effects of elevation, area, geometric constraints, habitat type, environmental factors and land‐use intensity on terrestrial gastropod diversity patterns in Val Müstair, an alpine region influenced by different types of agricultural land use in the eastern Alps, Switzerland. Gastropods were sampled using a standardized method in 180 sites spanning an elevational range from 1215 to 2770 m and covering 11 different habitat types. A total of 11 102 specimens representing 70 species were recorded. Observed species richness, statistically estimated true richness (Chao) and geographically interpolated observed richness were used as measures of local species richness. The comparison of three alternative models (environmental, geometric constraints and gastropod abundance models) revealed that the environmental model explained most of the variation in all measures of local diversity. The best model combining the predictors of all three models showed that elevation, soil pH and habitat type affected all measures of local species richness. Similar analyses conducted at the level of 150‐m elevational bands showed that elevation was again the best predictor of species richness, while the area of the elevational band did not have any influence. However, in one out of the two measures of band species richness, the best model indicated that geometric constraints may also contribute to the observed pattern. At both spatial scales, all measures of species richness decreased with increasing elevation. An analysis of species‐specific life‐history traits showed that adult shell size of land snails decreased with increasing elevation. Most species with large shells were confined to lower elevations. The results indicate that environmental factors might be most important in shaping the observed patterns.     

Geographical patterns in species richness of the benthic polychaetes in the continental shelf of the Gulf of California, Mexican Pacific

The present study is the first attempt to describe meso-scale patterns in the species richness of polychaetes along the Gulf of California, which stretches from about 23°N to 31°N. We examine herein the spatial changes in species distribution and explore the overlapping of species’ ranges towards the centre of the Gulf, to test whether the mid-domain effect (MDE) could explain an expected mid-domain peak in species richness. The faunal composition and the latitudinal range of 244 species of polychaetes recorded along the continental shelf of the Gulf of California were analysed in latitude bands of 1°. The species composition changes around the Gulf’s archipelago (~29°N), and the highest values of species richness are found at the 25° (197 species) and 26° (193 species) of latitude. Although the species richness pattern could be described by a parabolic shape, the regional trend was not strongly consistent with the peak of diversity at 27°N (176–191 species) predicted by the mid-domain effect: the random sorting of species’ ranges within spatial domain does not explain satisfactorily the geographical patterns of diversity. Nevertheless, a partial contribution of MDE to these natural patterns of diversity could be detected, and the increase in species richness towards middle latitudes was basically determined by species with distribution ranges larger than 6°. The low level of significance between the empirical species richness pattern and the mid-domain model prediction for polychaetes in the Gulf does not restrict their use as a model for exploring the randomness of the diversity patterns.     

Elevational Gradient of Vascular Plant Species Richness and Endemism in Crete – The Effect of Post-Isolation Mountain Uplift on a Continental Island System

Understanding diversity patterns along environmental gradients and their underlying mechanisms is a major topic in current biodiversity research. In this study, we investigate for the first time elevational patterns of vascular plant species richness and endemism on a long-isolated continental island (Crete) that has experienced extensive post-isolation mountain uplift. We used all available data on distribution and elevational ranges of the Cretan plants to interpolate their presence between minimum and maximum elevations in 100-m elevational intervals, along the entire elevational gradient of Crete (0–2400 m). We evaluate the influence of elevation, area, mid-domain effect, elevational Rapoport effect and the post-isolation mountain uplift on plant species richness and endemism elevational patterns. Furthermore, we test the influence of the island condition and the post-isolation mountain uplift to the elevational range sizes of the Cretan plants, using the Peloponnese as a continental control area. Total species richness monotonically decreases with increasing elevation, while endemic species richness has a unimodal response to elevation showing a peak at mid-elevation intervals. Area alone explains a significant amount of variation in species richness along the elevational gradient. Mid-domain effect is not the underlying mechanism of the elevational gradient of plant species richness in Crete, and Rapoport''s rule only partly explains the observed patterns. Our results are largely congruent with the post-isolation uplift of the Cretan mountains and their colonization mainly by the available lowland vascular plant species, as high-elevation specialists are almost lacking from the Cretan flora. The increase in the proportion of Cretan endemics with increasing elevation can only be regarded as a result of diversification processes towards Cretan mountains (especially mid-elevation areas), supported by elevation-driven ecological isolation. Cretan plants have experienced elevational range expansion compared to the continental control area, as a result of ecological release triggered by increased species impoverishment with increasing elevation.     

Different responses of avian feeding guilds to spatial and environmental factors across an elevation gradient in the central Himalaya

Although elevational patterns of species richness have been well documented, how the drivers of richness gradients vary across ecological guilds has rarely been reported. Here, we examined the effects of spatial factors (area and mid‐domain effect; MDE) and environmental factors, including metrics of climate, productivity, and plant species richness on the richness of breeding birds across different ecological guilds defined by diet and foraging strategy. We surveyed 12 elevation bands at intervals of 300 m between 1,800 and 5,400 m a.s.l using line‐transect methods throughout the wet season in the central Himalaya, China. Multiple regression models and hierarchical partitioning were used to assess the relative importance of spatial and environmental factors on overall bird richness and guild richness (i.e., the richness of species within each guild). Our results showed that richness for all birds and most guilds displayed hump‐shaped elevational trends, which peaked at an elevation of 3,300–3,600 m, although richness of ground‐feeding birds peaked at a higher elevation band (4,200–4,500 m). The Normalized Difference Vegetation Index (NDVI)—an index of primary productivity—and habitat heterogeneity were important factors in explaining overall bird richness as well as that of insectivores and omnivores, with geometric constraints (i.e., the MDE) of secondary importance. Granivore richness was not related to primary production but rather to open habitats (granivores were negatively influenced by habitat heterogeneity), where seeds might be abundant. Our findings provide direct evidence that the richness–environment relationship is often guild‐specific. Taken together, our study highlights the importance of considering how the effects of environmental and spatial factors on patterns of species richness may differ across ecological guilds, potentially leading to a deeper understanding of elevational diversity gradients and their implications for biodiversity conservation.     

Elevational Gradient in Species Richness Pattern of Epigaeic Beetles and Underlying Mechanisms at East Slope of Balang Mountain in Southwestern China

We report on the species richness patterns of epigaeic beetles (Coleoptera: Carabidae and Staphylinidae) along a subtropical elevational gradient of Balang Mountain, southwestern China. We tested the roles of environmental factors (e.g. temperature, area and litter cover) and direct biotic interactions (e.g. foods and antagonists) that shape elevational diversity gradients. Beetles were sampled at 19 sites using pitfall traps along the studied elevational gradient ranging from 1500 m–4000 m during the 2004 growing season. A total of 74416 specimens representing 260 species were recorded. Species richness of epigaeic beetles and two families showed unimodal patterns along the elevational gradient, peaking at mid-elevations (c. 2535 m), and the ranges of most beetle species were narrow along the gradient. The potential correlates of both species richness and environmental variables were examined using linear and second order polynomial regressions. The results showed that temperature, area and litter cover had strong explanatory power of beetle species richness for nearly all richness patterns, of beetles as a whole and of Carabidae and Staphylinidae, but the density of antagonists was associated with species richness of Carabidae only. Multiple regression analyses suggested that the three environmental factors combined contributed most to richness patterns for most taxa. The results suggest that environmental factors associated with temperature, area and habitat heterogeneity could account for most variation in richness pattern of epigaeic beetles. Additionally, the mid-elevation peaks and the small range size of most species indicate that conservation efforts should give attention to the entire gradient rather than just mid-elevations.     

What drives elevational patterns of diversity? A test of geometric constraints,climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcán Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid‐domain effect, MDE). Results The species richness of the 484 recorded species showed a hump‐shaped pattern with elevation with a richness peak at mid‐elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large‐ and small‐ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid‐elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.     

River fish assemblages along an elevation gradient in the eastern extremity of Europe

Studies on assemblages of freshwater fishes along elevational gradients of rivers are lacking, even in Europe. In this paper we have explored the entire range of elevational gradients existing in the European part of Russia. We analyzed how fish biodiversity (species richness, abundance, diversity indices) at 435 river sites differed by elevation. The impact of elevation on the distribution of freshwater fish species was analyzed using regression and ordination methods. For the first time for a large area of Eastern Europe, optimum points and niche breadth for fish species along altitude gradients were estimated. Our analyses showed: (1) species richness and Shannon index decreased in the upper part of the gradient; fish abundance showed a unimodal response to elevation; highest numbers were found at elevations between 250 and 500 m; (2) ordination analysis demonstrated an upstream-downstream gradient of the fish assemblages; (3) regression analysis showed significant preferences for elevation by 19 species, all of which were monotonic; (4) optimum and niche breadth (tolerance) were highly variable between species; only five species (brown trout, grayling, common minnow, bullhead and stone loach) were encountered at elevations above 650 m; and (5) in our region, the habitat of grayling was higher in the mountains, and its abundance (numbers) at extreme elevations was greater, than brown trout. These results show how fish assemblages differ with elevation. Our findings identify the data that can be used for regional environmental monitoring of the state of small rivers and for aquatic conservation.     

Contrasting patterns of elevational zonation for birds and mammals in the Andes of southeastern Peru

Abstract. To determine the generality of avian diversity patterns, we investigated patterns of elevational zonation shown by birds and mammals along the eastern slope of the Andes Mountains in southeastern Peru. The strong environmental gradient sampled, entirely within Peru's Manu National Park and Biosphere Reserve, supports highly diverse faunas. Elevational distributions of 901 bird species, 129 bat species, and twenty-eight species of native mice exhibit contrasting patterns in species richness, species composition, and species turnover. Birds and bats showed smooth declines of species richness with elevation, whereas the richness of mouse assemblages was unrelated to elevation. For all three groups, the greatest differences were between lowland and highland faunas, although cutoff points for this contrast varied among groups (≈ 500 m for birds, 750 m for bats, and 1000 m for mice). Differences in composition also separated bird and bat faunas on either side of c. 1400 m (the boundary between montance forest and cloud forest); for mice, this faunal transition may take place nearer to 2000 m. Bird and bat faunas lacked the more discrete zonations suggested for mouse assemblages, as indicated by elevational range profiles and nested subset analyses. Distinct highland assemblages are apparent in two-dimensional histograms of range limits of birds and mice, but not for bats. Highland bat species occupy broader elevational ranges than lowland bat species, but for both birds and mice, species at intermediate elevations had the broadest amplitudes. Finally, clumping of range maxima and minima along the gradient identified zones of pronounced species turnover in each group, but these were generally not strongly associated with the locations of ecotones. Differences in zonation of these groups appear to reflect their different biological attributes and phylogenetic histories. Such differences obviously complicate discussions of ‘general’ diversity patterns, and limit the usefulness of birds to forecast or predict diversity patterns in other more poorly known groups—other groups may show elevated diversity and endemism in areas where avian diversity patterns appear unremarkable. The pronounced contrasts between bats and mice, and the generally intermediate character of avian patterns, suggest that future analyses might profitably partition birds into finer, more homogeneous groups of historically and/or ecologically similar species. Group differences in zonation may ultimately prove explicable with information on both species-abundance patterns and resource distributions.     

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.     

The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient

Aim The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored. Location The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas. Methods Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance‐based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity. Results A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump‐shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction‐based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r² in Geometridae. The potential effects of geometric constraints (mid‐domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small‐ranged species showed stronger deviations from model predictions than large‐ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small‐ranged species. We hypothesize that small‐ranged species (at least of the Ennominae) may tend to be host specialists, whereas large‐ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern. Conclusions The results clearly show that moth diversity shows a hump‐shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.     

Elevational patterns of fern species assemblages and richness in central Japan

We conducted field surveys in 807 quadrats to evaluate the elevational belts, boundary and richness patterns of ferns and lycophytes in the temperate region of central Japan. We analysed fern species assemblages at 100 m elevational steps by cluster analysis and tested the number of upper and lower boundaries for elevational intervals against a null model of random distribution of elevational limits. We compared the pattern of fern species richness along the elevational gradients in central Japan with patterns in several locations to evaluate the fern flora in central Japan in relation to the rest of the world. We recorded 261 ferns species in total, which is one-third of the Japanese ferns. We found clear elevational boundaries of fern assemblages at 900 and 1,800 m and three fern elevational zones, which corresponded well to the elevational limits of forest types in central Japan. The pattern of fern species richness in central Japan was an asymmetric hump-shaped pattern that peaked close to the sea level, with the peak of local richness at lower elevations than that of regional richness. We found that the peak of fern species richness along the elevational gradient in Japan was located at lower elevations than that of fern elevational patterns in several locations around the world.