The quest for a null model for macroecological patterns: geometry of species distributions at multiple spatial scales

There have been several attempts to build a unified framework for macroecological patterns. However, these have mostly been based either on questionable assumptions or have had to be parameterized to obtain realistic predictions. Here, we propose a new model explicitly considering patterns of aggregated species distributions on multiple spatial scales, the property which lies behind all spatial macroecological patterns, using the idea we term 'generalized fractals'. Species' spatial distributions were modelled by a random hierarchical process in which the original 'habitat' patches were randomly replaced by sets of smaller patches nested within them, and the statistical properties of modelled species assemblages were compared with macroecological patterns in observed bird data. Without parameterization based on observed patterns, this simple model predicts realistic patterns of species abundance, distribution and diversity, including fractal-like spatial distributions, the frequency distribution of species occupancies/abundances and the species–area relationship. Although observed macroecological patterns may differ in some quantitative properties, our concept of random hierarchical aggregation can be considered as an appropriate null model of fundamental macroecological patterns which can potentially be modified to accommodate ecologically important variables.     

Fifteen important questions in the spatial ecology of diatoms

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Positive relationship between regional distribution and local abundance in stream insects: a consequence of niche breadth or niche position?

A positive relationship between regional distribution and local abundance of species is almost ubiquitous macroecological pattern, yet the mechanisms behind this pattern remain poorly understood. I tested for the relationship between regional distribution and local abundance of stream insect species in a boreal drainage system, with a specific aim to examine if this relationship follows the mechanistic basis of either the niche-based (niche breadth and niche position) or metapopulation models. There was a positive relationship between regional distribution and local abundance of stream insects, and there also were significant relationships between distribution/abundance and niche breadth or niche position. These results thus suggest that widely distributed species tend to be, on average, locally more abundant, have wider niches and lower marginality of niche position with regard to environmental factors than species that have more restricted distributions. However, although significant, there was much unexplained variability around these relationships, suggesting that other mechanisms (e.g. metapopulation dynamics) besides differences in species' niches are likely to affect the distribution and abundance of stream insects, at least within a drainage system. The results thus showed that 1) although niche position was more consistently related to the positive distribution-abundance relationship, ecologists should not abandon niche breadth as a potential mechanism behind this relationship, and 2) that several mechanisms are likely to act in concert in determining the relationship between distribution and abundance of species.     

Climatic niche evolution in turtles is characterized by phylogenetic conservatism for both aquatic and terrestrial species

Understanding how the climatic niche of species evolved has been a topic of high interest in current theoretical and applied macroecological studies. However, little is known regarding how species traits might influence climatic niche evolution. Here, we evaluated patterns of climatic niche evolution in turtles (tortoises and freshwater turtles) and whether species habitat (terrestrial or aquatic) influences these patterns. We used phylogenetic, climatic and distribution data for 261 species to estimate their climatic niches. Then, we compared whether niche overlap between sister species was higher than between random species pairs and evaluated whether niche optima and rates varied between aquatic and terrestrial species. Sister species had higher values of niche overlap than random species pairs, suggesting phylogenetic climatic niche conservatism in turtles. The climatic niche evolution of the group followed an Ornstein–Uhlenbeck model with different optimum values for aquatic and terrestrial species, but we did not find consistent evidence of differences in their rates of climatic niche evolution. We conclude that phylogenetic climatic niche conservatism occurs among turtle species. Furthermore, terrestrial and aquatic species occupy different climatic niches but these seem to have evolved at similar evolutionary rates, reinforcing the importance of habitat in understanding species climatic niches and their evolution.     

Intraspecific range dynamics and niche evolution in Candidula land snail species

The range dynamics of a species can either be governed by the spatial tracing of the fundamental environmental niche or by adaptation that allows to occupy new niches. Therefore, the investigation of spatial variation in the realized environmental niche is central to the understanding of species range limit dynamics. However, the study of intraspecific niche variation has been neglected in most phylogeographical studies. We studied the spatial distribution of the realized environmental niche in three land snail species of the genus Candidula , integrating phylogeographical methods, morphometrics, and spatial biodiversity informatics . The phylogeographical analyses showed significant range expansions in all species. These expansions were accompanied in Candidula gigaxii by a shift in the realized environmental niche, the species Candidula unifasciata followed its ancestral niche during expansion while the climate changed in the area of origin and Candidula rugosiuscula tracked the ancestral environmental conditions. The significant niche shifts were associated with potentially adaptive changes of shell morphology. We propose our presented approach as a practicable framework to test hypotheses on intraspecific niche evolution. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 303–317.     

Of niches and distributions: range size increases with niche breadth both globally and regionally but regional estimates poorly relate to global estimates

The relationship between species’ niche breadth (i.e. the range of environmental conditions under which a species can persist) and range size (i.e. the extent of its spatial distribution) has mostly been tested within geographically restricted areas but rarely at the global extent. Here, we not only tested the relationship between range size (derived from species’ distribution data) and niche breadth (derived from species’ distribution and co‐occurrence data) of 1255 plant species at the regional extent of the European Alps, but also at the global extent and across both spatial scales for a subset of 180 species. Using correlation analyses, linear models and variation partitioning, we found that species’ realized niche breadth estimated at the regional level is a weak predictor of species’ global niche breadth and range size. Against our expectations, distribution‐derived niche breadth was a better predictor for species’ range size than the co‐occurrence‐based estimate, which should, theoretically, account for more than the climatically determined niche dimensions. Our findings highlight that studies focusing on the niche breadth vs range size relationship must explicitly consider spatial mismatches that might have confounded and diminished previously reported relationships.     

Unusual abundance–range size relationship in an Afromontane bird community: the effect of geographical isolation?

Aim To show that the frequently reported positive trend in the abundance–range‐size relationship does not hold true within a montane bird community of Afrotropical highlands; to test possible explanations of the extraordinary shape of this relationship; and to discuss the influence of island effects on patterns of bird abundance in the Cameroon Mountains. Location Bamenda Highlands, Cameroon, Western Africa. Methods We censused birds during the breeding season in November and December 2003 using a point‐count method and mapped habitat structure at these census points. Local habitat requirements of each species detected by point counts were quantified using canonical correspondence analysis, and the size of geographical ranges of species was measured from their distribution maps for sub‐Saharan Africa. We tested differences in abundance, niche breadth and niche position between three species groups: endemic bird species of the Cameroon Mountains, non‐endemic Afromontane species, and widespread species. Results We detected neither a positive nor negative abundance–range‐size relationship in the bird community studied. This pattern was caused by the similar abundance of widespread, endemic and non‐endemic montane bird species. Moreover, endemic and non‐endemic montane species had broader local niches than widespread species. The widespread species also used more atypical habitats, as indicated by the slightly larger values of their niche positions. Main conclusions The relationship detected between abundance and range size does not correspond with that inferred from contemporary macroecological theory. We suggest that island effects are responsible for the observed pattern. Relatively high abundances of montane species are probably caused by their adaptation to local environmental conditions, which was enabled by climatic stability and the isolation of montane forest in the Cameroon Mountains. Such a unique environment provides a less suitable habitat for widespread species. Montane species, which are abundant at present, may also have had large ranges in glacial periods, but their post‐glacial distribution may have become restricted after the retreat of the montane forest. On the basis of comparison of our results with studies describing the abundance structure of bird communities in other montane areas in the Afrotropics, we suggest that the detected patterns may be universal throughout Afromontane forests.     

Co-diversity and co-distribution in phyllostomid bats: Evaluating the relative roles of climate and niche conservatism

Explaining the causes of geographic gradients in biodiversity remains an elusive task. Traditionally, correlative approaches have been used to relate species richness with contemporary climate, without actually explaining the causal factors. Recent approaches propose simulation models as more appropriate tools for assessing potential causes of macroecological patterns. Here we developed stochastic models to assess the relative contribution of climate and niche conservatism in determining compositional similarity among sites (co-diversity) and geographic association among species (co-distribution) in the bat family Phyllostomidae. We used range-diversity plots and variance-ratio tests to describe and evaluate such patterns. Our results supported a strong effect of climate in determining cohesive ranges causing positive co-diversity and co-distribution. We also demonstrated a marginal effect of niche conservatism, as modeled here, among species in shaping these patterns. However, climate and niche conservatism are not sufficient and other processes are still required to explain observed patterns. Our study highlights the importance of historical processes and demonstrates the usefulness of a simulation framework in testing biogeographical hypothesis to understand the relationship between diversity and distribution.     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Niche properties of Central European spiders: shading, moisture and the evolution of the habitat niche

Aim  Niche theory emphasizes the importance of environmental conditions for the distribution and abundance of species. Using a macroecological approach our study aimed at identifying the important environmental gradients for spiders. We generated numerical values of niche position and niche width. We also investigated relationships between these niche properties as well as the degree of phylogenetic conservatism in order to draw conclusions about the evolution of the habitat niche.
Location  Central Europe: lowlands of Austria, Belgium, Germany, Luxembourg, the Netherlands and Switzerland.
Methods  We analysed 244 published spider communities from 70 habitat types by correspondence analysis. The resulting community scores were used to test for correlations with habitat characteristics. Species scores were used to derive niche position (mean scores) and niche width (standard deviation of scores). To test for niche conservatism we estimated variance components across the taxonomic hierarchy.
Results  The first two axes of the correspondence analysis were correlated with shading and moisture, respectively. Niche width had a hump-shaped relationship to both environmental gradients. β-diversity was strikingly higher in open habitats than in forests. Habitat niche conservatism was lower than phylogenetic conservatism in body size.
Main conclusions  Environmental factors are important drivers for the β-diversity of spiders, especially across open habitats. This underlines the importance of preserving the whole range of moisture conditions in open habitats. Narrow niches of species occurring at the ends of both environmental gradients indicate that adaptations to extreme habitats lead to constraints in ecological flexibility. Nevertheless, the habitat niche of species seems to evolve much faster than morphological or physiological traits.     

Comparing macroecological patterns across continents: evolution of climatic niche breadth in varanid lizards

Macroecological analyses often test hypotheses at the global scale, or among more closely related species in a single region (e.g. continent). Here, we test several hypotheses about climatic niche widths among relatively closely related species that occur across multiple continents, and compare patterns within and across continents to see if they differ. We focus on the lizard genus Varanus (monitor lizards), which occurs in diverse environments in Africa, Asia, and Australia. We address three main questions. 1) How do climatic niche breadths of species on a given niche axis change based on the position of species along that niche axis? (E.g. are species that occur in more extreme environments more narrowly specialized for those conditions?) 2) Are there trade‐offs in niche breadths on temperature and precipitation axes among species, or are niche widths on different axes positively related? 3) Is variation in niche breadths among species explained primarily by within‐locality seasonal variation, or by differences in climatic conditions among localities across the species range? We generate a new time‐calibrated phylogeny for Varanus and test these hypotheses within and between continents using climatic data and phylogenetic methods. Our results show that patterns on each continent often parallel each other and global patterns. However, in many other cases, the strength of relationships can change dramatically among closely related species on different continents. Overall, we found that: 1) species in warmer environments have narrower temperature niche breadths, but there is no relationship between precipitation niche breadth and niche position; 2) temperature and precipitation niche breadths tend to be positively related among species, rather than showing trade‐offs; and 3) within‐locality seasonal variation explains most variation in climatic niche breadths. Some of these results are concordant with previous studies (in amphibians and North American lizards), and might represent general macroecological patterns.     

Winter bird communities in the northern Negev: species dispersal patterns, habitat use and implications for habitat conservation

Bird habitat conservation may require different management strategies for different seasonal bird assemblages. We studied habitat use by winter birds in forest and scrubland habitat patches in the northern Negev, Israel. Our goal was to assess whether differences in responses to landscape and habitat structure between breeding and non-breeding seasons require changes in future conservation plans that have been suggested for the Negev breeding bird community. We evaluated habitat and area effects on bird abundance and distribution and tested whether species habitat use during winter involves niche shifts. Compared with breeding birds, a larger proportion of winter bird species occupied both scrubland and forest. As in summer, forest bird species responded to habitat structure, whereas scrubland species were associated with both habitat structure and area. Resident birds disperse into habitats in which they were not present during summer. Consequently, for several species, the correlation between bird densities and environmental factors showed a better fit at the landscape rather than at the habitat scale. In addition, rather than niche shift, birds actually extended their niche breadth. Nest site selection may constrain bird distribution into a realized niche, smaller than their fundamental niche. Despite the scale differences in habitat use, the similar species diversity patterns between seasons suggest that both winter and summer birds would benefit from conservation of scrub patches larger than 50 ha, and enrichment of foliage layers within the planted forests.     

宏生态尺度上景观破碎化对物种丰富度的影响

生物多样性的地理格局及其形成机制是宏生态学与生物地理学的研究热点。大量研究表明,景观尺度上的生境破碎化对物种多样性的分布格局具有重要作用,但目前尚不清楚这种作用是否足以在宏生态尺度上对生物多样性地理格局产生显著影响。利用中国大陆鸟类和哺乳动物的物种分布数据,在100 km×100 km网格的基础上生成了这两个类群生物的物种丰富度地理格局,进一步利用普通最小二乘法模型和空间自回归模型研究了物种丰富度与气候、生境异质性、景观破碎化的相关关系。结果表明,景观破碎化因子与鸟类和哺乳动物的物种丰富度都具有显著的关联关系,其方差贡献率可达约30%—50%(非空间模型)和60%—80%(空间模型),略低于或接近于气候和生境异质性因子。方差分解结果显示,景观破碎化因子与气候和生境异质性因子的方差贡献率的重叠部分达20%—40%。相对鸟类而言,景观破碎化对哺乳动物物种丰富度的地理格局具有更高的解释率。     

Missing the rarest: is the positive interspecific abundance–distribution relationship a truly general macroecological pattern?

Lepidopterists have long acknowledged that many uncommon butterfly species can be extremely abundant in suitable locations. If this is generally true, it contradicts the general macroecological pattern of the positive interspecific relationship between abundance and distribution, i.e. locally abundant species are often geographically more widespread than locally rare species. Indeed, a negative abundance–distribution relationship has been documented for butterflies in Finland. Here we show, using the Finnish butterflies as an example, that a positive abundance–distribution relationship results if the geographically restricted species are missed, as may be the case in studies based on random or restricted sampling protocols, or in studies that are conducted over small spatial scales. In our case, the abundance–distribution relationship becomes negative when approximately 70 per cent of the species are included. This observation suggests that the abundance–distribution relationship may in fact not be linear over the entire range of distributions. This intriguing possibility combined with some taxonomic biases in the literature may undermine the generalization that for a given taxonomic assemblage there is a positive interspecific relationship between local abundance and regional distribution.     

黄河三角洲植物生态位和生态幅对物种分布-多度关系的解释

物种分布与多度间的正相关格局非常普遍,但该格局的生态机制却一直不太明确。研究者提出了很多假说来解释这种分布-多度关系,其中物种的生态幅和生态位（资源可利用性）机制的研究较多。为了验证物种的生态幅和生态位是否能解释物种的多度-分布格局,本文研究了黄河三角洲地区湿地植物分布、多度、生态位和生态幅间的关系,结果表明：该区物种分布与多度呈显著正相关,且均与生态幅显著正相关,物种分布与生态位显著负相关,但物种多度与生态位相关性不显著。这说明物种的分布越广,其多度越高,环境容忍度越大;而可利用资源更多的物种分布更广,环境容忍度越大的物种多度越多,资源可利用性对该区物种多度影响不大。本研究说明物种生态幅能解释物种分布-多度正相关格局,而生态位假说不能很好的解释这一格局;应该还有其他因素一起解释这一物种分布-多度正相关格局。     

Macroecology meets macroevolution: evolutionary niche dynamics in the seaweed Halimeda

Aim Because of their broad distribution in geographical and ecological dimensions, seaweeds (marine macroalgae) offer great potential as models for marine biogeographical inquiry and exploration of the interface between macroecology and macroevolution. This study aims to characterize evolutionary niche dynamics in the common green seaweed genus Halimeda, use the observed insights to gain understanding of the biogeographical history of the genus and predict habitats that can be targeted for the discovery of species of special biogeographical interest. Location Tropical and subtropical coastal waters. Methods The evolutionary history of the genus is characterized using molecular phylogenetics and relaxed molecular clock analysis. Niche modelling is carried out with maximum entropy techniques and uses macroecological data derived from global satellite imagery. Evolutionary niche dynamics are inferred through application of ancestral character state estimation. Results A nearly comprehensive molecular phylogeny of the genus was inferred from a six‐locus dataset. Macroecological niche models showed that species distribution ranges are considerably smaller than their potential ranges. We show strong phylogenetic signal in various macroecological niche features. Main conclusions The evolution of Halimeda is characterized by conservatism for tropical, nutrient‐depleted habitats, yet one section of the genus managed to invade colder habitats multiple times independently. Niche models indicate that the restricted geographical ranges of Halimeda species are not due to habitat unsuitability, strengthening the case for dispersal limitation. Niche models identified hotspots of habitat suitability of Caribbean species in the eastern Pacific Ocean. We propose that these hotspots be targeted for discovery of new species separated from their Caribbean siblings since the Pliocene rise of the Central American Isthmus.     

Mutualistic interactions shape global spatial congruence and climatic niche evolution in Neotropical mimetic butterflies

Understanding the mechanisms underlying species distributions and coexistence is both a priority and a challenge for biodiversity hotspots such as the Neotropics. Here, we highlight that Müllerian mimicry, where defended prey species display similar warning signals, is key to the maintenance of biodiversity in the c. 400 species of the Neotropical butterfly tribe Ithomiini (Nymphalidae: Danainae). We show that mimicry drives large-scale spatial association among phenotypically similar species, providing new empirical evidence for the validity of Müller's model at a macroecological scale. Additionally, we show that mimetic interactions drive the evolutionary convergence of species climatic niche, thereby strengthening the co-occurrence of co-mimetic species. This study provides new insights into the importance of mutualistic interactions in shaping both niche evolution and species assemblages at large spatial scales. Critically, in the context of climate change, our results highlight the vulnerability to extinction cascades of such adaptively assembled communities tied by positive interactions.     

Relationships between survival and habitat suitability of semi‐aquatic mammals

Spatial distribution and habitat selection are integral to the study of animal ecology. Habitat selection may optimize the fitness of individuals. Hutchinsonian niche theory posits the fundamental niche of species would support the persistence or growth of populations. Although niche‐based species distribution models (SDMs) and habitat suitability models (HSMs) such as maximum entropy (Maxent) have demonstrated fair to excellent predictive power, few studies have linked the prediction of HSMs to demographic rates. We aimed to test the prediction of Hutchinsonian niche theory that habitat suitability (i.e., likelihood of occurrence) would be positively related to survival of American beaver (Castor canadensis), a North American semi‐aquatic, herbivorous, habitat generalist. We also tested the prediction of ideal free distribution that animal fitness, or its surrogate, is independent of habitat suitability at the equilibrium. We estimated beaver monthly survival probability using the Barker model and radio telemetry data collected in northern Alabama, United States from January 2011 to April 2012. A habitat suitability map was generated with Maxent for the entire study site using landscape variables derived from the 2011 National Land Cover Database (30‐m resolution). We found an inverse relationship between habitat suitability index and beaver survival, contradicting the predictions of niche theory and ideal free distribution. Furthermore, four landscape variables selected by American beaver did not predict survival. The beaver population on our study site has been established for 20 or more years and, subsequently, may be approaching or have reached the carrying capacity. Maxent‐predicted increases in habitat use and subsequent intraspecific competition may have reduced beaver survival. Habitat suitability‐fitness relationships may be complex and, in part, contingent upon local animal abundance. Future studies of mechanistic SDMs incorporating local abundance and demographic rates are needed.     

Niche evolution in Australian terrestrial mammals? Clarifying scale-dependencies in phylogenetic and functional drivers of co-occurrence

Interactive forces between competition and habitat filtering drive many biogeographic patterns over evolutionary time scales. However, the responsiveness of assemblages to these two forces is highly influenced by spatial scale, forming complex patterns of niche separation. We explored these spatial dependencies by quantifying the influence of phylogeny and functional traits in shaping present day native terrestrial mammal assemblages at multiple scales, principally by identifying the spatial scales at which niche evolution operates. We modelled the distribution of 53 native terrestrial mammal species across New South Wales, Australia. Using predicted distributions, we estimated the range overlap between each pair of species at increasing grain sizes (~0.8, 5.1, 20, 81, 506, 2,025, 8,100 km²). We employed a decision tree to identify how interactions among functional traits and phylogenetic relatedness translated to levels of sympatry at increasing spatial scales. We found that Australian terrestrial mammals displayed phylogenetic over-dispersion that was inversely related to spatial scale, suggesting that ecological processes were more influential than biogeographic sympatry patterns in defining assemblages of species. While the contribution of phylogenetic relatedness to patterns of co-occurrence decreased as spatial scale increased, the reverse was true for habitat preferences. At the same time, functional traits also operated at different scales, as dietary preferences dominated at local spatial scales (<10 km²) while body mass has a stronger effect at larger spatial scales. Our findings show that ecological and evolutionary processes operate at different scales and that Australian terrestrial mammals diverged slower along their micro-scale niche compared to their macro-scale niche. By combining phylogenetic and niche methods through the modelling of species distributions, we assessed whether specific traits were related to a particular niche. More importantly, conducting multi-scale spatial analysis avoids categorical assignment of traits-to-niches, providing a clearer relationship between traits and a species ecological niche and a more precise scaling for the axes of niche evolution.