Species Richness and the Analytic Geometry of Latitudinal and Altitudinal Gradients

Extensive empirical work has shown that species richness decreases roughly exponentially or quadratically with latitude. What appears to be a latitudinal gradient in fact may simply be a negative correlation of latitude with area at that latitude, due to convergence of lines of meridian at the poles. There is simply less area at high latitudes, which means fewer niches and fewer opportunities for speciation, hence diminished biodiversity at high latitudes. Similarly, analytic geometry of a cone shows that species number should decrease linearly with altitude on a conical mountain. Here, I provide an explicit mathematical model of the area hypothesis of species richness along latitude and altitude gradients. I re-analyze a previously published latitudinal gradient dataset and show that species number is a linear function of the predicted area and that species number is more fully explained by predicted area than by a quadratic function of latitude. However, analytic geometry is not needed if precise measures of area are known.     

Landscape Variation in Tree Species Richness in Northern Iran Forests

Mapping landscape variation in tree species richness (SR) is essential to the long term management and conservation of forest ecosystems. The current study examines the prospect of mapping field assessments of SR in a high-elevation, deciduous forest in northern Iran as a function of 16 biophysical variables representative of the area’s unique physiography, including topography and coastal placement, biophysical environment, and forests. Basic to this study is the development of moderate-resolution biophysical surfaces and associated plot-estimates for 202 permanent sampling plots. The biophysical variables include: (i) three topographic variables generated directly from the area’s digital terrain model; (ii) four ecophysiologically-relevant variables derived from process models or from first principles; and (iii) seven variables of Landsat-8-acquired surface reflectance and two, of surface radiance. With symbolic regression, it was shown that only four of the 16 variables were needed to explain 85% of observed plot-level variation in SR (i.e., wind velocity, surface reflectance of blue light, and topographic wetness indices representative of soil water content), yielding mean-absolute and root-mean-squared error of 0.50 and 0.78, respectively. Overall, localised calculations of wind velocity and surface reflectance of blue light explained about 63% of observed variation in SR, with wind velocity accounting for 51% of that variation. The remaining 22% was explained by linear combinations of soil-water-related topographic indices and associated thresholds. In general, SR and diversity tended to be greatest for plots dominated by Carpinus betulus (involving ≥ 33% of all trees in a plot), than by Fagus orientalis (median difference of one species). This study provides a significant step towards describing landscape variation in SR as a function of modelled and satellite-based information and symbolic regression. Methods in this study are sufficiently general to be applicable to the characterisation of SR in other forested regions of the world, providing plot-scale data are available for model generation.     

Species Richness Pattern along Altitudinal Gradient in Central European Beech Forests

The unimodal species richness-altitude distribution pattern seems to be universal. To investigate the validity of this phenomenon in homogeneous substrate and vegetation conditions, we sampled beech-dominated forests in five volcanic mountain ranges in the Western Carpathians. European beech (Fagus sylvatica L.) formed monodominant closed-canopy stands at altitudes from 300 to 1,200 m. Along this gradient, the influence of beech on understory plant species richness was expected to be strong and uniform. The shape of the species richness-altitude relationship was analyzed for three datasets: herb layer, shrub layer, and both layers merged together. Contrary to prediction, the studied species richness-altitude relationship was inversely unimodal, with a minimum at intermediate altitudes. Quadratic regression models were statistically significant for all three datasets (P<0.001) and the explained variability ranged from 12 % to 20 %. The possible explanation for the observed pattern is twofold. In the central part of the altitudinal gradient, low species richness is due to strong competition by monodominant beech with accumulation of leaf litter and uptake soil resources, mainly water. This influence is somewhat released towards the margins of the gradient. Secondly, the species pool from the neighbouring communities increases species richness only in the lower parts of the altitudinal gradient.     

Plant Size as Determinant of Species Richness of Herbivores,Natural Enemies and Pollinators across 21 Brassicaceae Species

Large plants are often more conspicuous and more attractive for associated animals than small plants, e.g. due to their wider range of resources. Therefore, plant size can positively affect species richness of associated animals, as shown for single groups of herbivores, but studies usually consider intraspecific size differences of plants in unstandardised environments. As comprehensive tests of interspecific plant size differences under standardised conditions are missing so far, we investigated effects of plant size on species richness of all associated arthropods using a common garden experiment with 21 Brassicaceae species covering a broad interspecific plant size gradient from 10 to 130 cm height. We recorded plant associated ecto- and endophagous herbivores, their natural enemies and pollinators on and in each aboveground plant organ, i.e. flowers, fruits, leaves and stems. Plant size (measured as height from the ground), the number of different plant organ entities and their biomass were assessed. Increasing plant size led to increased species richness of associated herbivores, natural enemies and pollinating insects. This pattern was found for ectophagous and endophagous herbivores, their natural enemies, as well as for herbivores associated with leaves and fruits and their natural enemies, independently of the additional positive effects of resource availability (i.e. organ biomass or number of entities and, regarding natural enemies, herbivore species richness). We found a lower R² for pollinators compared to herbivores and natural enemies, probably caused by the high importance of flower characteristics for pollinator species richness besides plant size. Overall, the increase in plant height from 10 to 130 cm led to a 2.7-fold increase in predicted total arthropod species richness. In conclusion, plant size is a comprehensive driver of species richness of the plant associated arthropods, including pollinators, herbivores and their natural enemies, whether they are endophagous or ectophagous or associated with leaves or fruits.     

The Influence of Vegetation Height Heterogeneity on Forest and Woodland Bird Species Richness across the United States

Avian diversity is under increasing pressures. It is thus critical to understand the ecological variables that contribute to large scale spatial distribution of avian species diversity. Traditionally, studies have relied primarily on two-dimensional habitat structure to model broad scale species richness. Vegetation vertical structure is increasingly used at local scales. However, the spatial arrangement of vegetation height has never been taken into consideration. Our goal was to examine the efficacies of three-dimensional forest structure, particularly the spatial heterogeneity of vegetation height in improving avian richness models across forested ecoregions in the U.S. We developed novel habitat metrics to characterize the spatial arrangement of vegetation height using the National Biomass and Carbon Dataset for the year 2000 (NBCD). The height-structured metrics were compared with other habitat metrics for statistical association with richness of three forest breeding bird guilds across Breeding Bird Survey (BBS) routes: a broadly grouped woodland guild, and two forest breeding guilds with preferences for forest edge and for interior forest. Parametric and non-parametric models were built to examine the improvement of predictability. Height-structured metrics had the strongest associations with species richness, yielding improved predictive ability for the woodland guild richness models (r² = ∼0.53 for the parametric models, 0.63 the non-parametric models) and the forest edge guild models (r² = ∼0.34 for the parametric models, 0.47 the non-parametric models). All but one of the linear models incorporating height-structured metrics showed significantly higher adjusted-r² values than their counterparts without additional metrics. The interior forest guild richness showed a consistent low association with height-structured metrics. Our results suggest that height heterogeneity, beyond canopy height alone, supplements habitat characterization and richness models of forest bird species. The metrics and models derived in this study demonstrate practical examples of utilizing three-dimensional vegetation data for improved characterization of spatial patterns in species richness.     

An Analysis of Fish Species Richness in Natural Lakes

There is a growing recognition of the need to conserve biodiversity that has been conceptualised in the Convention of Biological Diversity. Maintenance of fish species richness is particularly important, because habitat degradation in inland waters continues to accelerate on a global scale. Here we develop empirical models for predicting fish species richness in natural lakes in various geographical regions of the world. In tropical lakes where fish biodiversity is richer than in temperate lakes, fish species richness can be predicted by a few variables such as lake area and altitude. Low fish species richness in most temperate lakes might be due to the effect of glaciation on colonisation and speciation of fishes. In US, Canadian and northern European lakes, lake acidification is one of the important factors influencing fish species richness. Although limnological characteristics influence fish species richness in temperate lakes, lake area and altitude have greater predictive power. This is in contrast to fish species richness in rivers, which can be reliably predicted by basin area. In the power curves, which describe the relationship between fish species richness and habitat size in lakes and rivers, the exponent is always greater in tropical regions than in temperate regions. Because fish biodiversity is greater in the tropics threats to fish biodiversity through habitat degradation are greater than those in temperate inland waters.     

Patterns of Species Richness and Composition in Re-Created Grassland

The success of many prairie restorations is not well documented. A restoration begun in 1975 at the Fermi National Accelerator Laboratory near Chicago, Illinois allows assessment of restoration efforts as well as changes through time. Data are presented on species richness and composition for 13 restorations planted in successive years between 1975 and 1990 and two remnant prairies. Presence of species was recorded using a stratified random design. Species richness at several scales and non‐metric multidimensional scaling ordination were used to assess trends in the vegetation. Species richness declined through time at all scales examined and was always less in the restored prairies than that found in the remnant prairies. Species composition changed with time but not in the direction of the composition found in the remnants. Our understanding of the maintenance of species richness is not sufficient to allow the re‐creation of patterns of species found in remnant grassland communities.     

Spatial Scaling of Non-Native Fish Richness across the United States

A major goal and challenge of invasion ecology is to describe and interpret spatial and temporal patterns of species invasions. Here, we examined fish invasion patterns at four spatially structured and hierarchically nested scales across the contiguous United States (i.e., from large to small: region, basin, watershed, and sub-watershed). All spatial relationships in both richness and fraction between species groups (e.g., natives vs. exotics) were positive at large scales. However, contrary to predictions using null/neutral models, the patterns at small scales were hump-shaped (unimodal), not simply negative. The fractions of both domestic (introduced among watersheds within the USA) and foreign (introduced from abroad) exotics increased with area across scales but decreased within each scale. The foreign exotics exhibited the highest dominance (lowest evenness) and spatial variation in distribution, followed by domestic exotics and natives, although on average natives still occupy larger areas than domestic and foreign exotics. The results provide new insight into patterns and mechanisms of fish species invasions at multiple spatial scales in the United States.     

The Genetic Content of Chromosomal Inversions across a Wide Latitudinal Gradient

There is increasing evidence regarding the role of chromosomal inversions in relevant biological processes such as local adaptation and speciation. A classic example of the adaptive role of chromosomal polymorphisms is given by the clines of inversion frequencies in Drosophila subobscura, repeatable across continents. Nevertheless, not much is known about the molecular variation associated with these polymorphisms. We characterized the genetic content of ca. 600 individuals from nine European populations following a latitudinal gradient by analysing 19 microsatellite loci from two autosomes (J and U) and the sex chromosome (A), taking into account their chromosomal inversions. Our results clearly demonstrate the molecular genetic uniformity within a given chromosomal inversion across a large latitudinal gradient, particularly from Groningen (Netherlands) in the north to Málaga (Spain) in the south, experiencing highly diverse environmental conditions. This low genetic differentiation within the same gene arrangement across the nine European populations is consistent with the local adaptation hypothesis for th evolutionof chromosomal polymorphisms. We also show the effective role of chromosomal inversions in maintaining different genetic pools within these inverted genomic regions even in the presence of high gene flow. Inversions represent thus an important barrier to gene flux and can help maintain specific allelic combinations with positive effects on fitness. Consistent patterns of microsatellite allele-inversion linkage disequilibrium particularly in loci within inversions were also observed. Finally, we identified areas within inversions presenting clinal variation that might be under selection.     

西藏地衣物种多样性的海拔梯度分析

根据西藏地衣分类和区系的文献资料,对西藏地衣物种多样性的海拔梯度进行了分析。267个分类群按照生长型或者基物被分为六组。每一组地衣的物种丰度随海拔增加呈现单峰曲线形式的变化。多数组物种丰度的峰值出现在海拔3400~3900m之间,对应于山地寒温带针叶林带的上半部分,比尼泊尔对应类群出现极值的海拔要高。此植物带谱内复杂的生态系统可能是物种多样性高的主要原因。壳状地衣物种丰度的峰值出现在5100~5400m区间则可能是因为该区内高大的树木的消失以及具有充裕的阳光。西藏的地衣物种多样性远低于尼泊尔,两地共有的物种数量很少。对西藏地衣物种多样性的调查不充分应当是其主要原因,因此今后中国地衣学工作者应当加强西藏地衣多样性的研究。     

Species Richness and Adaptation of Marine Fungi from Deep-Subseafloor Sediments

The fungal kingdom is replete with unique adaptive capacities that allow fungi to colonize a wide variety of habitats, ranging from marine habitats to freshwater and terrestrial habitats. The diversity, importance, and ecological roles of marine fungi have recently been highlighted in deep-subsurface sediments using molecular methods. Fungi in the deep-marine subsurface may be specifically adapted to life in the deep biosphere, but this can be demonstrated only using culture-based analyses. In this study, we investigated culturable fungal communities from a record-depth sediment core sampled from the Canterbury Basin (New Zealand) with the aim to reveal endemic or ubiquist adapted isolates playing a significant ecological role(s). About 200 filamentous fungi (68%) and yeasts (32%) were isolated. Fungal isolates were affiliated with the phyla Ascomycota and Basidiomycota, including 21 genera. Screening for genes involved in secondary metabolite synthesis also revealed their bioactive compound synthesis potential. Our results provide evidence that deep-subsurface fungal communities are able to survive, adapt, grow, and interact with other microbial communities and highlight that the deep-sediment habitat is another ecological niche for fungi.     

川渝地区马先蒿属物种丰富度空间分布格局及其影响因素

对川渝地区马先蒿属植物的物种多样性分布格局进行了较为系统的研究,川渝地区共有马先蒿属植物181种,分属于11个群70个系,其物种多样性在川渝地区呈现西高东低的分布格局,物种多样性水平在海拔3500～4000m最高。在SPSS软件的支持下,运用因子分析法和相关分析法定量分析川渝地区148个单元格内主要环境因素与马先蒿属物种多样性分布之间的关系,表明熊蜂物种丰富度、海拔高度和≥15℃的积温依次是影响川渝地区马先蒿属物种多样性分布格局最重要的生态环境因子。熊蜂物种丰富度和海拔高度与马先蒿物种丰富度呈正相关,熊蜂对马先蒿属植物的繁殖成功与否起着关键的作用;诸多温度因子中,≥15℃积温对马先蒿属物种多样性分布影响最大,与马先蒿属物种丰富度呈负相关,这与它喜好高海拔地区寒冷的环境相一致。     

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.     

Latitudinal variation in virus-induced mortality of phytoplankton across the North Atlantic Ocean

Viral lysis of phytoplankton constrains marine primary production, food web dynamics and biogeochemical cycles in the ocean. Yet, little is known about the biogeographical distribution of viral lysis rates across the global ocean. To address this, we investigated phytoplankton group-specific viral lysis rates along a latitudinal gradient within the North Atlantic Ocean. The data show large-scale distribution patterns of different virus groups across the North Atlantic that are associated with the biogeographical distributions of their potential microbial hosts. Average virus-mediated lysis rates of the picocyanobacteria Prochlorococcus and Synechococcus were lower than those of the picoeukaryotic and nanoeukaryotic phytoplankton (that is, 0.14 per day compared with 0.19 and 0.23 per day, respectively). Total phytoplankton mortality (virus plus grazer-mediated) was comparable to the gross growth rate, demonstrating high turnover rates of phytoplankton populations. Virus-induced mortality was an important loss process at low and mid latitudes, whereas phytoplankton mortality was dominated by microzooplankton grazing at higher latitudes (>56°N). This shift from a viral-lysis-dominated to a grazing-dominated phytoplankton community was associated with a decrease in temperature and salinity, and the decrease in viral lysis rates was also associated with increased vertical mixing at higher latitudes. Ocean-climate models predict that surface warming will lead to an expansion of the stratified and oligotrophic regions of the world''s oceans. Our findings suggest that these future shifts in the regional climate of the ocean surface layer are likely to increase the contribution of viral lysis to phytoplankton mortality in the higher-latitude waters of the North Atlantic, which may potentially reduce transfer of matter and energy up the food chain and thus affect the capacity of the northern North Atlantic to act as a long-term sink for CO₂.     

Relationship of Productivity to Species Richness in the Xinjiang Temperate Grassland

The relationship between species richness (SR) and aboveground net primary productivity (ANPP) is still a central and debated issue in community ecology. Previous studies have often emphasized the relationship of alpha diversity (number of species identity) to the mean ANPP with respect to the SR-ANPP relationship while neglecting the contribution of beta diversity (dissimilarity in species composition) to the mean ANPP and to the stability of ANPP (coefficient of ANPP: CV of ANPP). In this study, we used alpha and beta diversity, mean ANPP and the CV of ANPP collected from 159 sites and belonging to three vegetation types in the Xinjiang temperate grassland to first examine their trends along climatic factors and among different vegetation types and then test the relationship among alpha (beta) diversity and mean ANPP and the CV of ANPP. Our results showed that in the Xinjiang temperate grasslands, alpha diversity was positively and linearly correlated with MAP but unimodally correlated with MAT. Meanwhile, beta diversity was unimodally correlated with MAP but linearly correlated with MAT. Relative to desert steppe, meadow steppe and typical steppe had the highest alpha and beta diversity, respectively. Except for ANPP exhibiting a quadratic relationship with MAP, no significant relationship was found among ANPP, the CV of ANPP and climatic factors. ANPP and the CV of ANPP also exhibited no apparent patterns in variation among different vegetation types. Our results further showed that mean ANPP was closely associated with alpha diversity. Both linear and unimodal relationships were detected between alpha diversity and mean ANPP, but their particular form was texture-dependent. Meanwhile, the CV of ANPP was positively correlated with beta diversity. Our results indicated that in addition to incorporating alpha diversity and mean ANPP, incorporating beta diversity and the CV of ANPP could expand our understanding of the SR-ANPP relationship.     

Fundamental Causes of the Global Patterns of Species Range and Richness1

Global patterns of species range and richness are a consequence of many interacting factors, including environmental conditions, competition, geographical area, and historical/evolutionary development. Two widely studied global patterns of distribution are the latitudinal and elevation gradients of species range and richness. The fundamental mechanisms by which environment and physiology of the plants themselves interact to generate global-scale correlations between increased species range or decreased species richness and latitude/elevation have not previously been established. This paper develops the hypothesis that the primary climatic variables determining global-scale gradients in ectotherm species range and richness are temperature (T) and temperature variability (T), and that the primary physiological variable defining adaptation of ectotherms to temperature is respiratory energy metabolism. This hypothesis is based on a postulate that adaptation of ectotherms to latitudinal/altitudinal gradients of T and T leads to corresponding gradients in properties of energy metabolism. The gradients of metabolic properties give rise to gradients of species range and richness that are observed on a global scale. We demonstrate that natural selection results in ectotherms with metabolic properties matched to their environment and that energy use efficiency and the temperature range allowing growth are inversely related. Thus, opposing selective pressures to increase metabolic energy-use-efficiency or to increase the probability of surviving climate extremes control adaptation of ectotherms to climate. The principles developed in this paper yield fundamental laws of ecology that allow calculation of the contributions of global temperature patterns to the formation of gradients of species range and diversity. Relative values of richness and range are calculated solely from data on abiotic variables. Predictions agree with known patterns of ectotherm distribution.     

Activity Patterns of Eurasian Lynx Are Modulated by Light Regime and Individual Traits over a Wide Latitudinal Range

The activity patterns of most terrestrial animals are regarded as being primarily influenced by light, although other factors, such as sexual cycle and climatic conditions, can modify the underlying patterns. However, most activity studies have been limited to a single study area, which in turn limit the variability of light conditions and other factors. Here we considered a range of variables that might potentially influence the activity of a large carnivore, the Eurasian lynx, in a network of studies conducted with identical methodology in different areas spanning latitudes from 49°7′N in central Europe to 70°00′N in northern Scandinavia. The variables considered both light conditions, ranging from a day with a complete day–night cycle to polar night and polar day, as well as individual traits of the animals. We analysed activity data of 38 individual free-ranging lynx equipped with GPS-collars with acceleration sensors, covering more than 11,000 lynx days. Mixed linear additive models revealed that the lynx activity level was not influenced by the daily daylight duration and the activity pattern was bimodal, even during polar night and polar day. The duration of the active phase of the activity cycle varied with the widening and narrowing of the photoperiod. Activity varied significantly with moonlight. Among adults, males were more active than females, and subadult lynx were more active than adults. In polar regions, the amplitude of the lynx daily activity pattern was low, likely as a result of the polycyclic activity pattern of their main prey, reindeer. At lower latitudes, the basic lynx activity pattern peaked during twilight, corresponding to the crepuscular activity pattern of the main prey, roe deer. Our results indicated that the basic activity of lynx is independent of light conditions, but is modified by both individual traits and the activity pattern of the locally most important prey.     

Climate and Species Richness Predict the Phylogenetic Structure of African Mammal Communities

We have little knowledge of how climatic variation (and by proxy, habitat variation) influences the phylogenetic structure of tropical communities. Here, we quantified the phylogenetic structure of mammal communities in Africa to investigate how community structure varies with respect to climate and species richness variation across the continent. In addition, we investigated how phylogenetic patterns vary across carnivores, primates, and ungulates. We predicted that climate would differentially affect the structure of communities from different clades due to between-clade biological variation. We examined 203 communities using two metrics, the net relatedness (NRI) and nearest taxon (NTI) indices. We used simultaneous autoregressive models to predict community phylogenetic structure from climate variables and species richness. We found that most individual communities exhibited a phylogenetic structure consistent with a null model, but both climate and species richness significantly predicted variation in community phylogenetic metrics. Using NTI, species rich communities were composed of more distantly related taxa for all mammal communities, as well as for communities of carnivorans or ungulates. Temperature seasonality predicted the phylogenetic structure of mammal, carnivoran, and ungulate communities, and annual rainfall predicted primate community structure. Additional climate variables related to temperature and rainfall also predicted the phylogenetic structure of ungulate communities. We suggest that both past interspecific competition and habitat filtering have shaped variation in tropical mammal communities. The significant effect of climatic factors on community structure has important implications for the diversity of mammal communities given current models of future climate change.