Neutral community dynamics, the mid-domain effect and spatial patterns in species richness

The mid‐domain effect (MDE) aims to explain spatial patterns in species richness invoking only stochasticity and geometrical constraints. In this paper, we used simulations to show that its main qualitative prediction, a hump‐shaped pattern in species richness, converges to the expectation of a spatially bounded neutral model when communities are linked by short‐distance migration. As these two models can be linked under specific situations, neutral theory may provide a mechanistic population level basis for MDE. This link also allows establishing in which situations MDE patterns are more likely to be found. Also, in this situation, MDE models could be used as a first approximation to understand the role of both stochastic (ecological drift and migration) and deterministic (adaptation to environmental conditions) processes driving the spatial structure of species richness.     

Geometric constraints and spatial pattern of species richness: critique of range-based null models

Abstract. Does the shape of a biogeographical region influence its spatial patterns of species richness? A complete answer must include careful distinction between the distribution of a species, which is a complex geometric object, and the range of a species, which is relatively simple, especially when reduced to one dimension. We consider range‐based models of species richness, in particular range overlap counts in one dimension, for which we give a unified mathematical treatment via the joint probability P(m,l) of midpoints and lengths of ranges. We discuss a number of difficulties, in practice and in principle, using range‐based models, and show that the so‐called mid‐domain effect, a proposed null model for the effect of geometric constraint, is qualitatively a property of all biologically realistic models based on range overlap counts. As such, range‐based models provide little insight into understanding or explaining biogeographical patterns in species richness. We characterize the quantitative null model for range overlap counts in one dimension, for which we give a simple and direct field test based on P(m,l). We apply this test to a large clade in a complete bioregion (the Proteaceae of the Cape Floristic Region): geometric constraint does not explain the spatial pattern in this case. We show that any geometric constraint on species richness, including range overlap counts, must act via edge effects. Thus, to understand biogeographical patterns, an understanding of the effects and consequences of edges is fundamental.     

A strong Madagascan rainforest MDE and no equatorward increase in species richness: re-analysis of 'The missing Madagascan mid-domain effect', by Kerr J.T., Perring M. & Currie D.J. (Ecology Letters 9:149-159, 2006)

By reanalysing inaccurately presented data of Kerr et al. (2006) , we refute their claims that area-corrected species richness of endemic Madagascan birds and mammals increases toward the Equator and is best explained by environmental factors, and that the rainforest mid-domain effect (MDE) Lees et al. (1999) demonstrated is artefactual.     

Toward a better understanding of the regional causes of local community richness

Despite widespread acknowledgement that local ecological communities are profoundly shaped by regional-scale influences, including evolutionary and biogeographic processes, this perspective has yet to be widely incorporated into ecological research. Drawing on recent research, we propose four steps towards making regional influences a stronger part of research on the richness of local communities: (1) identifying the regional-scale causes of variation in species richness in the systems ecologists study; (2) testing for effects of regional richness on local richness, using improved observational and experimental analyses to overcome earlier problems; (3) simultaneously analysing environmental influences on regional and local species richness as well as the influence of regional richness on local richness and (4) considering the potential reciprocal effects of local processes on regional richness. In conclusion, we suggest some ways that similar approaches could be applied to other aspects of community structure beyond species richness.     

Trait-based species richness: ecology and macroevolution

Understanding the origins of species richness patterns is a fundamental goal in ecology and evolutionary biology. Much research has focused on explaining two kinds of species richness patterns: (i) spatial species richness patterns (e.g. the latitudinal diversity gradient), and (ii) clade-based species richness patterns (e.g. the predominance of angiosperm species among plants). Here, I highlight a third kind of richness pattern: trait-based species richness (e.g. the number of species with each state of a character, such as diet or body size). Trait-based richness patterns are relevant to many topics in ecology and evolution, from ecosystem function to adaptive radiation to the paradox of sex. Although many studies have described particular trait-based richness patterns, the origins of these patterns remain far less understood, and trait-based richness has not been emphasised as a general category of richness patterns. Here, I describe a conceptual framework for how trait-based richness patterns arise compared to other richness patterns. A systematic review suggests that trait-based richness patterns are most often explained by when each state originates within a group (i.e. older states generally have higher richness), and not by differences in transition rates among states or faster diversification of species with certain states. This latter result contrasts with the widespread emphasis on diversification rates in species-richness research. I show that many recent studies of spatial richness patterns are actually studies of trait-based richness patterns, potentially confounding the causes of these patterns. Finally, I describe a plethora of unanswered questions related to trait-based richness patterns.     

神农架海拔梯度上的植物种域分布特征及Rapoport法则检验

根据Rapoport法则, 动物、植物物种的纬度或海拔分布宽度存在着从高纬度或高海拔地区向低纬度或低海拔地区逐渐变窄的现象。本文基于物种的海拔分布数据, 分析了神农架维管束植物及不同种域宽度组的物种丰富度海拔格局; 采用已有的4种方法和本文改进的种域分组中点法, 分析了海拔梯度上种域宽度与种域中点的关系, 并检验其是否符合Rapoport法则。结果表明, 神农架地区维管植物丰富度的海拔梯度分布格局呈单峰型, 峰值在1,000–1,500 m; 不同种域组的物种丰富度分布具有类似的单峰格局, 但随着种域宽度减小, 其物种丰富度的峰值逐渐偏向低海拔。对于神农架的物种海拔分布数据, Stevens方法、Pagel方法和逐种方法的结果都支持Rapoport法则, 而中点法的结果主要反映中域效应的影响, 种域分组中点法可有效控制中域效应的影响, 但不支持Rapoport法则。上述结果表明, 对于Rapoport法则的检验亟待研究方法的改进; 而种域的海拔格局及其形成机制, 还需要更多案例的比较研究, 才能形成普遍性的认识。     

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Aim  Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host–parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites.
Methods  We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores.
Results  We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites.
Main conclusions  Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.     

Invasibility and species richness of island endemic arthropods: a general model of endemic vs. exotic species

Aim This paper has two objectives. First, we examine how a variety of biotic, abiotic and anthropogenic factors influence the endemic and introduced arthropod richness on an oceanic island. Second, we look at the relationship between the endemic and introduced arthropod richness, to ask whether areas with high levels of endemic species richness deter invasions. Location The work was carried out on a young volcanic island, Terceira, in the Azores. Methods We used standard techniques to collect data on arthropod species richness. Environmental data were obtained from the CIELO climatic model and using GIS. The explanatory value of environmental variables on a small‐scale gradient of endemic and exotic arthropod species richness was examined with generalized linear models (GLMs). In addition, the impact of both endemic and exotic species richness in the communities was assessed by entering them after the environmental variable(s) to see if they contributed significantly to the final model (the hierarchical method). Results Abiotic (climatic and geomorphological) variables gave a better explanation of the variation in endemic species richness, whereas anthropogenic variables explained most of the variation in introduced species richness. Furthermore, after accounting for all environmental variables, part of the unexplained variance in the endemic species richness is explained by the introduced species richness and vice‐versa. That is, areas with high levels of endemic species richness had many introduced species. There is evidence of a somewhat inverse spatial distribution between a group of oceanic‐type, forest‐dwelling, endemic, relict arthropods and a group of more generalist endemic arthropods that are able to survive in disturbed marginal sites particularly rich in non‐indigenous species. Main conclusions Richness of endemic species is mainly driven by abiotic factors such as a climatic axis (oceanic‐type localities with lower temperatures and summer precipitations) and a binary variable CALD (location of sites in caldeiras or ravines), whereas richness of introduced species depends on disturbance related factors. However, after factoring out these major influences, there is a correlation between endemic and introduced richness, suggesting that – independent of the environmental and geographical factors that affect the distribution of endemic or introduced species – the richest endemic assemblages are more prone to invasion, due probably to a facilitation process. Inconclusive evidence suggests that non‐indigenous species are limited to those sites under anthropogenic influence located mainly near forest edges, but the rate of expansion of those species to high‐altitude, core pristine sites has still to be tested.     

The mid-domain effect applied to elevational gradients: species richness of small mammals in Costa Rica

Aim The objective of this study was to comprehensively document and examine the alpha and gamma patterns of species richness in non-volant, small mammals (rodents, shrews and mouse opossums) along a tropical elevational gradient. These data were used to determine the support for existing hypotheses of species richness encompassing mid-domain null models, as well as climatic, and community overlap hypotheses. Location Field studies were conducted along a Caribbean slope of the Río Peñas Blancas watershed in the north-eastern region of Costa Rica between 750 and 1850 m at 10 sampling sites. Methods Species richness and abundances of small mammals were surveyed for four seasons including three temporal replicates at each of five elevational sites: late wet season (2000), early wet season (2001), and dry season (2002), and one spatial replicate at five different sites within the same elevations during the late wet season (2001). Species richness at elevations below 700 m was compiled from specimen records from 23 US national and international collections. Predictions of a null model based solely on geometric constraints were examined using a Monte Carlo simulation program, Mid-Domain Null. Results In 16,900 trap nights, 1561 individuals from 16 species were captured. Both alpha and gamma species richness peaked at mid-elevation between 1000 and 1300 m, with richness declining both at higher and lower elevations. Most of the empirical curves of species richness occur within 95% prediction curves of the mid-domain model, although deviations from the null model exist. Regression of the empirical richness on the null model predictions explained nearly half of the variation observed (r² = 0.45, P = 0.002). Main conclusions The geometric constraints of montane topography appear to influence the diversity pattern of small mammals, although climatic conditions including an intermediate rainfall and temperature regime, and distance from the persistent cloud cap also are correlated with the pattern of species richness. The predictions of productivity, and community overlap hypotheses are not supported with the empirical data.     

Spatial and environmental determinants of vascular plant species richness distribution in the Iberian Peninsula and Balearic Islands

Using an exhaustive data compilation, Iberian vascular plant species richness in 50 times 50 UTM grid cells was regressed against 24 explanatory variables (spatial, geographical, topographical, geological, climatic, land use and environmental diversity variables) using Generalized Linear Models and partial regression analysis in order to ascertain the relative contribution of primary, heterogeneous and spatially structured variables. The species richness variation accounted for by these variables is reasonably high (65% of total deviance). Little less than half of this variation is accounted for spatially structured variables. A purely spatial component of variation is hardly significant. The most significant variables are those related to altitude, and particularly maximum altitude, whose cubic response reflects the occurrence of the maximum number of species at the highest altitudes. This result highlighted the importance of Iberian mountains as hotspots of diversity and the relevance of large and small scale historical factors in contemporary plant distribution patterns. Climatic or energy-related variables contributed little, whereas geological (calcareous and acid rocks) and, to a lesser extent, environmental heterogeneity variables (land use diversity and altitude range) seem to be more important.     

Patterns of species richness and turnover in endemic amphibians of the Guineo-Congolian rain forest

Aim

The African Guineo-Congolian (GC) region is a global biodiversity hotspot with high species endemism, bioclimatic heterogeneity, complex landscape features, and multiple biogeographic barriers. Bioclimatic and geographic variables influence global patterns of species richness and endemism, but their relative importance varies across taxa and regions and is poorly understood for many faunas. Here, we test the hypothesis that turnover in endemic amphibians of the GC biodiversity hotspot is influenced mainly by the geographic distance between grid cells and secondarily by rainfall- and temperature-related variables.

Location

West and Central Africa.

Major Taxa Studied

Amphibians.

Methods

We compiled species-occurrence records via field sampling, online databases, and taxonomic literature. Our study used 1205 unique georeferenced records of 222 amphibian species endemic to the GC region. Patterns of species richness were mapped onto a grid with a spatial resolution of 0.5° × 0.5°. We estimated weighted endemism and tested whether endemism was higher than the expected species richness (randomization test). We quantified species turnover using generalized dissimilarity modelling to evaluate the processes underlying observed patterns of species richness in GC endemic amphibians. We explored bioregionalization using agglomerative hierarchical clustering based on the unweighted pair group method with arithmetic averages.

Results

We identified seven areas within the lower GC region – forests in Cameroon, Gabon, Southern Nigeria, Equatorial Guinea, Republic of Congo, Democratic Republic of Congo, and Cote d'Ivoire – as having high species richness of endemic amphibians. The randomization test returned four major areas of significant weighted endemism: Nigeria-Cameroon mountains, forest regions of the Democratic Republic of Congo, Cote d'Ivoire, and Ghana. Our analysis revealed five bioregions for amphibian endemism, four of which were located within the lower Guineo-Congolian forest. Species turnover was strongly related to the geographic distance between grid cells; contributing bioclimatic variables included precipitation of the warmest quarter, mean temperature of the wettest quarter, and mean diurnal temperature range.

Main Conclusions

Our results indicate that geographic distance between grid cells is the primary determinant of turnover in GC endemic amphibians, with secondary but significant effects of rainfall- and temperature-related variables. Our study identifies key areas of endemic amphibian richness that could be prioritized for conservation actions.     

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。     

集合种群模拟模型及其在竞争共存机制与物种多样性研究中的应用

以荒漠区人工植被的恢复与重建为背景,从宏观尺度研究了很集合种群的空间分布新模式,建立了基于Levins集合种群模型的数值模拟方法。对两物种的模拟结果表明：在适当选择参数下,模拟植被区的集合种群可以形成“海藻式”稳定的时空分布结构,在理论上表明相同生态特征的物种在空间生境中可以达成共存。为了达到物种丰富度和生产力最佳,实现持续发展,对多物种集合种群进行了模拟。模拟结果显示当物种的种数为5时,空间上随机播种的模拟种群覆盖率达到最大,因而可发挥最大的治沙作用。另外,模拟还显示在播种时应采取集聚式的空间播种模式,以使种群具有较高的防沙能力。该结果可为生物防沙治沙领域提供理论依据。     

Geographical expansion and increased prevalence of common species in avian assemblages: implications for large-scale patterns of species richness

Aim The assumption that ecological patterns at large spatial scales originate exclusively from non‐anthropogenic processes is growing more questionable with the increasing domination of the biosphere by humans. Because common and rare species are known to respond differently to anthropogenic activities at local scales these differential responses could, over time, be reflected in distributional patterns of species richness at larger spatial scales. This work tests the hypothesis that modern processes have played a role in shaping these patterns, by examining recent changes in the structure and composition of assemblages of breeding avifauna over a large geographical extent. Location The portion of North America containing the contiguous United States and southern Canada. Methods Changes in the geographical range structure of breeding avifauna in North America from 1968 to 2003 were analysed in regions containing historically moderate levels of anthropogenic activities. Two geographical measures, extent of occurrence and area of occupancy, were used to identify the level of rarity or commonality of individual species and to estimate, based on a vector analysis, patterns of change in geographical range structure for individual species and avian assemblages. Results More species experienced patterns of geographical range expansion (51%) than contraction (28%). The majority of avian assemblages (43%) displayed patterns of geographical range expansion: common species increased in number and proportion (6%) in association with reciprocal losses in rare and moderately rare species, resulting in a constant level of species richness. The minority of avian assemblages (21%) displayed patterns of geographical range contraction: gains occurred for common species as well as for rare and moderately rare species, resulting in substantial increases in species richness and a decline in the proportion of common species (4%). The remaining avian assemblages presented equivocal patterns characterized by gains in the number and proportion (2%) of common species and gains in species richness. Main conclusions Modern processes have played a role in shaping the distribution patterns of species richness at large spatial scales based on the composition of common and rare species. This suggests that anthropogenic activities cannot be ignored as a possible causal factor when considering ecological patterns at large spatial scales.     

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.     

Dependence of broad-scale geographical variation in fleshy-fruited plant species richness on disperser bird species richness

Aim We analysed the interdependence of avian frugivore‐ and fruited plant‐species richness at the scale of major river basins across Europe, taking into account several environmental factors along different spatial gradients. Location Continental Europe and the British Isles. Methods We focused on wintering birds and autumn/winter fruiting plants, and used major river basins as geographical units and Structural Equation Modelling as the principal analytical tool. Results The statistical influence of disperser species richness on fleshy‐fruited plant species richness is roughly double that of the reverse. Broad‐scale variation in frugivore richness is more dependent on environmental factors than on fruited plant richness. However, the influence of disperser richness on plant richness is four times higher than the influence of environmental factors. Environmental influences on both birds and plants are greater than purely spatial influences. Main conclusions Our results are interpreted as indicating that biotic dispersal of fruits strongly affects broad‐scale geographical trends of fleshy‐fruited plant species richness, whereas richness of fruited plants moderately affects frugivore richness.