Measuring climatic niche position and breadth may help to determine where species can occur over space and time. Using GIS-based and phylogenetic comparative methods, we investigated global patterns of variation in cl...Measuring climatic niche position and breadth may help to determine where species can occur over space and time. Using GIS-based and phylogenetic comparative methods, we investigated global patterns of variation in climatic niche breadth in lacertid lizards to test the following three hypotheses about climatic niche widths. First, does a species' temperature or precipitation niche breadth relate to its temperature or precipitation niche position(the mean value of annual mean temperature or annual precipitation across sampled localities in the range of each species)? Second, are there trade-offs between a species' temperature niche breadth and precipitation niche breadth? Third, does a species' temperature or precipitation niche breadth relate to altitude or latitude? We expect that:(1) species distributed in cold regions are specialized for low-temperature environments(i.e. narrow niche breadth center around low temperatures);(2) a negative relationship between species niche breadth on temperature and precipitation axes according to the tradeoff hypothesis(i.e. species that tolerate a broad range of precipitation regimes cannot also tolerate a broad range of temperatures);(3) precipitation niche breadth decreases with altitude or latitude, whereas temperature climatic niche breadth increases with altitude or latitude. Based on the analytical results we found that:(1) temperature niche breadth and position are negatively related, while precipitation niche breadth and position are positively related;(2) there is no trade-off between temperature and precipitation niche breadths; and (3) temperature niche breadth and latitude/altitude are positively related, but precipitation niche breadth and latitude/altitude are not significantly related. Our results show many similarities with previous studies on climatic niche widths reported for amphibians and lizards, which provide further evidence that such macroecological patterns of variation in climatic niche breadths may be widespread.展开更多
最大熵原理(the principle of maximum entropy)起源于信息论和统计力学,是基于有限的已知信息对未知分布进行无偏推断的一种数学方法。这一方法在很多领域都有成功应用,但只是近几年才被应用到生态学研究中,并且还存在很多争论。我们...最大熵原理(the principle of maximum entropy)起源于信息论和统计力学,是基于有限的已知信息对未知分布进行无偏推断的一种数学方法。这一方法在很多领域都有成功应用,但只是近几年才被应用到生态学研究中,并且还存在很多争论。我们从基本概念和方法出发,用掷骰子的例子阐明了最大熵原理的概念,并提出运用最大熵原理解决问题需要遵从的步骤。最大熵原理在生态学中的应用主要包括以下方面:(1)用群落水平功能性状的平均值作为约束条件来预测群落物种相对多度的模型;(2)基于气候、海拔、植被等环境因子构建物种地理分布的生态位模型;(3)对物种多度分布、种-面积关系等宏生态学格局的推断;(4)对物种相互作用的推断;(5)对食物网度分布的研究等等。最后我们综合分析了最大熵原理在生态学应用中所存在的争议,包括相应模型的有效性、可靠性等方面,介绍了一些对最大熵原理预测能力及其局限性的检验结果,强调了生态学家应用最大熵原理需要注意的问题,比如先验分布的选择、约束条件的设置等等。在物种相互作用、宏生态学格局等方面对最大熵原理更广泛的讨论与应用可能会给生态学带来新的发展机会。展开更多
Aims Several studies have shown that plant height changes along environmen-tal gradients.However,altitudinal patterns of plant height across species are still unclear,especially in regions sensitive to climate change....Aims Several studies have shown that plant height changes along environmen-tal gradients.However,altitudinal patterns of plant height across species are still unclear,especially in regions sensitive to climate change.As canopy height decreases dramatically near the tree line in alpine areas,we hypothesize that plant height across all species also decreases with increasing altitude,and distinct thresholds exist along this gradient.Methods Using a large dataset of maximum plant height and elevation range(400 to 6000 m a.s.l.)of 4295 angiosperms from the regional flora of the Tibetan Plateau,we regressed plant height for every 100 m belt against elevation to explore the relationships.To identify the approximate boundaries where dramatic changes in plant height occurs for herbaceous plants,shrubs,trees,woody plants and all angiosperms,we used piecewise linear regression.Phylogenetically independent contrast was used to test the potential evolutionary influences on altitudinal patterns at the family level.Important Findings Results showed that for herbaceous plants,shrubs,trees,woody plants and all angiosperms,plant height decreases significantly as altitude increases.In addition,we found that altitude,a proxy for many environmental factors,had obvious thresholds(breakpoints)dictating patterns of plant height.The results of phylogenetically independent contrast also emphasized the importance of evolution-ary history in determining the altitudinal patterns of plant height for some growth forms.Our results highlight the relative intense filter-ing effect of environmental factors in shaping patterns of functional traits and how this could vary for different ranges of environmental variables.展开更多
Background: Although the species-urban green area relationship (SARu) has been analyzed worldwide, the global consistency of its parameters, such as the fit and the slope of models, remains unexplored. Moreover,the SA...Background: Although the species-urban green area relationship (SARu) has been analyzed worldwide, the global consistency of its parameters, such as the fit and the slope of models, remains unexplored. Moreover,the SARu can be explained by 20 different models. Therefore, our objective was to evaluate which models provide a better explanation of SARus and, focusing on the power model,to evaluate the global heterogeneity in its fit and slope. Methods: We tested the performance of multiple statistical models in accounting for the way in which species richness increases with area, and examined whether variability in model form was associated with various methodological and environmental factors. Focusing on the power model, we analyzed the global heterogeneity in the fit and slope of the models through a meta-analysis. Results: Among 20 analyzed models, the linear model provided the best fit to the most datasets, was the top ranked model according to our efficiency criterion, and was the top overall ranked model.The Kobayashi and power models were the second and third overall ranked models, respectively.The number of green areas and the minimum number of species within a green area were the only significant variables explaining the variation in model form and performance, accounting for less than 10% of the variation. Based on the power model,there was a consistent overall fit (r2=0.50) and positive slope of 0.20 for the species richness increase with area worldwide.Conclusions:The good fit of the linear model to our SARu datasets contrasts with the non-linear SAR frequently found in true and non-urban habitat island systems;however, this finding may be a result of the small sample size of many SARu datasets. The overall power model slope of 0.20 suggests low levels of isolation among urban green patches, or alternatively that habitat specialist and area sensitive species have already been extirpated from urban green areas.展开更多
The timing of flowering is an important driver of species distribution and community assembly patterns.However,we still have much to learn about the factors that shape flowering diversity(i.e.,number of species flower...The timing of flowering is an important driver of species distribution and community assembly patterns.However,we still have much to learn about the factors that shape flowering diversity(i.e.,number of species flowering per period) in plant communities.One potential explanation of flowering diversity is the mid-domain effect,which states that geometric constraints on species ranges within a bounded domain(space or time) will yield a mid-domain peak in diversity regardless of ecological factors.Here,we determine whether the mid-domain effect explains peak flowering time(i.e.,when most species of communities are flowering) across China.We used phenological data of 16,267 herbaceous and woody species from the provincial Flora in China and species distribution data from the Chinese Vascular Plant Distribution Database to determine relationships between the observed number of species flowering and the number of species flowering as predicted by the mid-domain effect model,as well as between three climatic variables(mean minimum monthly temperature,mean monthly precipitation,and mean monthly sunshine duration).We found that the mid-domain effect explained a significant proportion of the temporal variation in flowering diversity across all species in China.Further,the mid-domain effect explained a greater proportion of variance in flowering diversity at higher latitudes than at lower latitudes.The patterns of flowering diversity for both herbaceous and woody species were related to both the mid-domain effect and environmental variables.Our findings indicate that including geometric constraints in conjunction with abiotic and biotic predictors will improve predictions of flowering diversity patterns.展开更多
In this brief report, we analyzed ecological correlates of risk of extinction for mammals endemic to China using phylogenetic eigenvector methods to control for the effect of phylogenetic inertia. Extinction risks wer...In this brief report, we analyzed ecological correlates of risk of extinction for mammals endemic to China using phylogenetic eigenvector methods to control for the effect of phylogenetic inertia. Extinction risks were based on the International Union for Conservation of Nature (IUCN) Red List and ecological explanatory attributes that include range size and climatic variables. When the effect of phylogenetic inertia were controlled, climate became the best predictor for quantifying and evaluating extinction risks of endemic mammals in China, accounting for 13% of the total variation. Range size seems to play a trivial role, explaining -1% of total variation; however, when non-phylogenetic variation partitioning analysis was done, the role of range size then explained 7.4% of total variation. Consequently, phylogenetic inertia plays a substantial role in increasing the explanatory power of range size on the extinction risks of mammals endemic to China. Limitations of the present study are discussed, with a focus on under-represented sampling of endemic mammalian species.展开更多
Background:Studies of the African savannas have used national parks to test ecological theories of natural ecosystems,including equilibrium,non-equilibrium,complex adaptive systems,and the role of top-down and bottom-...Background:Studies of the African savannas have used national parks to test ecological theories of natural ecosystems,including equilibrium,non-equilibrium,complex adaptive systems,and the role of top-down and bottom-up physical and biotic forces.Most such studies have excluded the impact of pastoralists in shaping grassland ecosystems and,over the last half century,the growing human impact on the world’s rangelands.The mounting human impact calls for selecting indicators and integrated monitoring methods able to track ecosystem changes and the role of natural and human agencies.Our study draws on five decades of monitoring the Amboseli landscape in southern Kenya to document the declining role of natural agencies in shaping plant ecology with rising human impact.Results:We show that plant diversity and productivity have declined,biomass turnover has increased in response to a downsizing of mean plant size,and that ecological resilience has declined with the rising probability of extreme shortfalls in pasture production.The signature of rainfall and physical agencies in driving ecosystem properties has decreased sharply with growing human impact.We compare the Amboseli findings to the long-term studies of Kruger and Serengeti national parks to show that the human influence,whether by design or default,is increasingly shaping the ecology of savanna ecosystems.We look at the findings in the larger perspective of human impact on African grasslands and the world rangelands,in general,and discuss the implications for ecosystem theory and conservation policy and management.Conclusions:The Amboseli study shows the value of using long-term integrated ecological monitoring to track the spatial and temporal changes in the species composition,structure,and function of rangeland ecosystems and the role of natural and human agencies in the process of change.The study echoes the widespread changes underway across African savannas and world’s rangelands,concluding that some level of ecosystem management is needed to prevent展开更多
Trees are spread worldwide,as the watchmen that experience the intricate ecological effects caused by various environmental factors.In order to better understand such effects,it is preferential to achieve finely and f...Trees are spread worldwide,as the watchmen that experience the intricate ecological effects caused by various environmental factors.In order to better understand such effects,it is preferential to achieve finely and fully mapped global trees and their environments.For this task,aerial and satellite-based remote sensing(RS)methods have been developed.However,a critical branch regarding the apparent forms of trees has significantly fallen behind due to the technical deficiency found within their globalscale surveying methods.Now,terrestrial laser scanning(TLS),a state-of-the-art RS technology,is useful for the in situ three-dimensional(3D)mapping of trees and their environments.Thus,we proposed co-developing an international TLS network as a macroscale ecotechnology to increase the 3D ecological understanding of global trees.First,we generated the system architecture and tested the available RS models to deepen its ground stakes.Then,we verified the ecotechnology regarding the identification of its theoretical feasibility,a review of its technical preparations,and a case testification based on a prototype we designed.Next,we conducted its functional prospects by previewing its scientific and technical potentials and its functional extensibility.Finally,we summarized its technical and scientific challenges,which can be used as the cutting points to promote the improvement of this technology in future studies.Overall,with the implication of establishing a novel cornerstone-sense ecotechnology,the co-development of an international TLS network can revolutionize the 3D ecological understanding of global trees and create new fields of research from 3D global tree structural ecology to 3D macroecology.展开更多
Two key features of human sociality are anatomically complex brains with neuron-dense cerebral cortices,and the propensity to form complex social networks with non-kin.Complex brains and complex social networks facili...Two key features of human sociality are anatomically complex brains with neuron-dense cerebral cortices,and the propensity to form complex social networks with non-kin.Complex brains and complex social networks facilitate flows of fitness-enhancing energy and information at multiple scales of social organization.Here,we consider how these flows interact to shape the emergence of macroscopic regularities in hunter-gatherer macroecology relative to other mammals and non-human primates.Collective computation is the processing of information by complex adaptive systems to generate inferences in order to solve adaptive problems.In hunter-gatherer societies the adaptive problem is to resolve uncertainty in generative models used to predict complex environments in order to maximize inclusive fitness.The macroecological solution is to link complex brains in social networks to form collective brains that perform collective computations.By developing theory and analyzing data,the author shows hunter-gatherers bands of~16 people,or~4 co-residing families,form the largest collective brains of any social mammal.Moreover,because individuals,families,and bands interact at multiple time scales,these fission-fusion dynamics lead to the emergence of the macroscopic regularities in hunter-gatherer macroecology we observe in cross-cultural data.These results show how computation is distributed across spatially-extended social networks forming decentralized knowledge systems characteristic of hunter-gatherer societies.The flow of information at scales far beyond daily interactions leads to the emergence of small-worlds where highly clustered local interactions are embedded within much larger,but sparsely connected multilevel metapopulations.展开更多
基金Financial support was provided by grants from Zhejiang Provincial Natural Science Foundation (LY17C030003)Natural Science Foundation of China (31270571)Science and Technology Bureau of Sanya (2013YD08)
文摘Measuring climatic niche position and breadth may help to determine where species can occur over space and time. Using GIS-based and phylogenetic comparative methods, we investigated global patterns of variation in climatic niche breadth in lacertid lizards to test the following three hypotheses about climatic niche widths. First, does a species' temperature or precipitation niche breadth relate to its temperature or precipitation niche position(the mean value of annual mean temperature or annual precipitation across sampled localities in the range of each species)? Second, are there trade-offs between a species' temperature niche breadth and precipitation niche breadth? Third, does a species' temperature or precipitation niche breadth relate to altitude or latitude? We expect that:(1) species distributed in cold regions are specialized for low-temperature environments(i.e. narrow niche breadth center around low temperatures);(2) a negative relationship between species niche breadth on temperature and precipitation axes according to the tradeoff hypothesis(i.e. species that tolerate a broad range of precipitation regimes cannot also tolerate a broad range of temperatures);(3) precipitation niche breadth decreases with altitude or latitude, whereas temperature climatic niche breadth increases with altitude or latitude. Based on the analytical results we found that:(1) temperature niche breadth and position are negatively related, while precipitation niche breadth and position are positively related;(2) there is no trade-off between temperature and precipitation niche breadths; and (3) temperature niche breadth and latitude/altitude are positively related, but precipitation niche breadth and latitude/altitude are not significantly related. Our results show many similarities with previous studies on climatic niche widths reported for amphibians and lizards, which provide further evidence that such macroecological patterns of variation in climatic niche breadths may be widespread.
文摘最大熵原理(the principle of maximum entropy)起源于信息论和统计力学,是基于有限的已知信息对未知分布进行无偏推断的一种数学方法。这一方法在很多领域都有成功应用,但只是近几年才被应用到生态学研究中,并且还存在很多争论。我们从基本概念和方法出发,用掷骰子的例子阐明了最大熵原理的概念,并提出运用最大熵原理解决问题需要遵从的步骤。最大熵原理在生态学中的应用主要包括以下方面:(1)用群落水平功能性状的平均值作为约束条件来预测群落物种相对多度的模型;(2)基于气候、海拔、植被等环境因子构建物种地理分布的生态位模型;(3)对物种多度分布、种-面积关系等宏生态学格局的推断;(4)对物种相互作用的推断;(5)对食物网度分布的研究等等。最后我们综合分析了最大熵原理在生态学应用中所存在的争议,包括相应模型的有效性、可靠性等方面,介绍了一些对最大熵原理预测能力及其局限性的检验结果,强调了生态学家应用最大熵原理需要注意的问题,比如先验分布的选择、约束条件的设置等等。在物种相互作用、宏生态学格局等方面对最大熵原理更广泛的讨论与应用可能会给生态学带来新的发展机会。
文摘Aims Several studies have shown that plant height changes along environmen-tal gradients.However,altitudinal patterns of plant height across species are still unclear,especially in regions sensitive to climate change.As canopy height decreases dramatically near the tree line in alpine areas,we hypothesize that plant height across all species also decreases with increasing altitude,and distinct thresholds exist along this gradient.Methods Using a large dataset of maximum plant height and elevation range(400 to 6000 m a.s.l.)of 4295 angiosperms from the regional flora of the Tibetan Plateau,we regressed plant height for every 100 m belt against elevation to explore the relationships.To identify the approximate boundaries where dramatic changes in plant height occurs for herbaceous plants,shrubs,trees,woody plants and all angiosperms,we used piecewise linear regression.Phylogenetically independent contrast was used to test the potential evolutionary influences on altitudinal patterns at the family level.Important Findings Results showed that for herbaceous plants,shrubs,trees,woody plants and all angiosperms,plant height decreases significantly as altitude increases.In addition,we found that altitude,a proxy for many environmental factors,had obvious thresholds(breakpoints)dictating patterns of plant height.The results of phylogenetically independent contrast also emphasized the importance of evolution-ary history in determining the altitudinal patterns of plant height for some growth forms.Our results highlight the relative intense filter-ing effect of environmental factors in shaping patterns of functional traits and how this could vary for different ranges of environmental variables.
基金funded by the Consejo Nacional de Investigaciones Científicas y Técnicas and the Universidad de Buenos Aires (Argentina)
文摘Background: Although the species-urban green area relationship (SARu) has been analyzed worldwide, the global consistency of its parameters, such as the fit and the slope of models, remains unexplored. Moreover,the SARu can be explained by 20 different models. Therefore, our objective was to evaluate which models provide a better explanation of SARus and, focusing on the power model,to evaluate the global heterogeneity in its fit and slope. Methods: We tested the performance of multiple statistical models in accounting for the way in which species richness increases with area, and examined whether variability in model form was associated with various methodological and environmental factors. Focusing on the power model, we analyzed the global heterogeneity in the fit and slope of the models through a meta-analysis. Results: Among 20 analyzed models, the linear model provided the best fit to the most datasets, was the top ranked model according to our efficiency criterion, and was the top overall ranked model.The Kobayashi and power models were the second and third overall ranked models, respectively.The number of green areas and the minimum number of species within a green area were the only significant variables explaining the variation in model form and performance, accounting for less than 10% of the variation. Based on the power model,there was a consistent overall fit (r2=0.50) and positive slope of 0.20 for the species richness increase with area worldwide.Conclusions:The good fit of the linear model to our SARu datasets contrasts with the non-linear SAR frequently found in true and non-urban habitat island systems;however, this finding may be a result of the small sample size of many SARu datasets. The overall power model slope of 0.20 suggests low levels of isolation among urban green patches, or alternatively that habitat specialist and area sensitive species have already been extirpated from urban green areas.
基金supported by the National Natural Science Foundation of China (Grant number 42261004)supported by the Jiangsu Social Development Project (BE2022792)。
文摘The timing of flowering is an important driver of species distribution and community assembly patterns.However,we still have much to learn about the factors that shape flowering diversity(i.e.,number of species flowering per period) in plant communities.One potential explanation of flowering diversity is the mid-domain effect,which states that geometric constraints on species ranges within a bounded domain(space or time) will yield a mid-domain peak in diversity regardless of ecological factors.Here,we determine whether the mid-domain effect explains peak flowering time(i.e.,when most species of communities are flowering) across China.We used phenological data of 16,267 herbaceous and woody species from the provincial Flora in China and species distribution data from the Chinese Vascular Plant Distribution Database to determine relationships between the observed number of species flowering and the number of species flowering as predicted by the mid-domain effect model,as well as between three climatic variables(mean minimum monthly temperature,mean monthly precipitation,and mean monthly sunshine duration).We found that the mid-domain effect explained a significant proportion of the temporal variation in flowering diversity across all species in China.Further,the mid-domain effect explained a greater proportion of variance in flowering diversity at higher latitudes than at lower latitudes.The patterns of flowering diversity for both herbaceous and woody species were related to both the mid-domain effect and environmental variables.Our findings indicate that including geometric constraints in conjunction with abiotic and biotic predictors will improve predictions of flowering diversity patterns.
基金This work was supported by the China Scholarship Council (CSC)(201308180004) Acknowledgements: I would like to thank two anonymous reviewers for constructive comments on this manuscript.
文摘In this brief report, we analyzed ecological correlates of risk of extinction for mammals endemic to China using phylogenetic eigenvector methods to control for the effect of phylogenetic inertia. Extinction risks were based on the International Union for Conservation of Nature (IUCN) Red List and ecological explanatory attributes that include range size and climatic variables. When the effect of phylogenetic inertia were controlled, climate became the best predictor for quantifying and evaluating extinction risks of endemic mammals in China, accounting for 13% of the total variation. Range size seems to play a trivial role, explaining -1% of total variation; however, when non-phylogenetic variation partitioning analysis was done, the role of range size then explained 7.4% of total variation. Consequently, phylogenetic inertia plays a substantial role in increasing the explanatory power of range size on the extinction risks of mammals endemic to China. Limitations of the present study are discussed, with a focus on under-represented sampling of endemic mammalian species.
基金funding ACP over the years include the Wildlife Conservation Society and the Liz Claiborne Art Ortenberg Foundation.
文摘Background:Studies of the African savannas have used national parks to test ecological theories of natural ecosystems,including equilibrium,non-equilibrium,complex adaptive systems,and the role of top-down and bottom-up physical and biotic forces.Most such studies have excluded the impact of pastoralists in shaping grassland ecosystems and,over the last half century,the growing human impact on the world’s rangelands.The mounting human impact calls for selecting indicators and integrated monitoring methods able to track ecosystem changes and the role of natural and human agencies.Our study draws on five decades of monitoring the Amboseli landscape in southern Kenya to document the declining role of natural agencies in shaping plant ecology with rising human impact.Results:We show that plant diversity and productivity have declined,biomass turnover has increased in response to a downsizing of mean plant size,and that ecological resilience has declined with the rising probability of extreme shortfalls in pasture production.The signature of rainfall and physical agencies in driving ecosystem properties has decreased sharply with growing human impact.We compare the Amboseli findings to the long-term studies of Kruger and Serengeti national parks to show that the human influence,whether by design or default,is increasingly shaping the ecology of savanna ecosystems.We look at the findings in the larger perspective of human impact on African grasslands and the world rangelands,in general,and discuss the implications for ecosystem theory and conservation policy and management.Conclusions:The Amboseli study shows the value of using long-term integrated ecological monitoring to track the spatial and temporal changes in the species composition,structure,and function of rangeland ecosystems and the role of natural and human agencies in the process of change.The study echoes the widespread changes underway across African savannas and world’s rangelands,concluding that some level of ecosystem management is needed to prevent
基金The work was financially supported by the National Key Research and Development Program of China(No.2022YFE0112700)the National Natural Science Foundation of China(No.32171782 and 31870531).
文摘Trees are spread worldwide,as the watchmen that experience the intricate ecological effects caused by various environmental factors.In order to better understand such effects,it is preferential to achieve finely and fully mapped global trees and their environments.For this task,aerial and satellite-based remote sensing(RS)methods have been developed.However,a critical branch regarding the apparent forms of trees has significantly fallen behind due to the technical deficiency found within their globalscale surveying methods.Now,terrestrial laser scanning(TLS),a state-of-the-art RS technology,is useful for the in situ three-dimensional(3D)mapping of trees and their environments.Thus,we proposed co-developing an international TLS network as a macroscale ecotechnology to increase the 3D ecological understanding of global trees.First,we generated the system architecture and tested the available RS models to deepen its ground stakes.Then,we verified the ecotechnology regarding the identification of its theoretical feasibility,a review of its technical preparations,and a case testification based on a prototype we designed.Next,we conducted its functional prospects by previewing its scientific and technical potentials and its functional extensibility.Finally,we summarized its technical and scientific challenges,which can be used as the cutting points to promote the improvement of this technology in future studies.Overall,with the implication of establishing a novel cornerstone-sense ecotechnology,the co-development of an international TLS network can revolutionize the 3D ecological understanding of global trees and create new fields of research from 3D global tree structural ecology to 3D macroecology.
文摘Two key features of human sociality are anatomically complex brains with neuron-dense cerebral cortices,and the propensity to form complex social networks with non-kin.Complex brains and complex social networks facilitate flows of fitness-enhancing energy and information at multiple scales of social organization.Here,we consider how these flows interact to shape the emergence of macroscopic regularities in hunter-gatherer macroecology relative to other mammals and non-human primates.Collective computation is the processing of information by complex adaptive systems to generate inferences in order to solve adaptive problems.In hunter-gatherer societies the adaptive problem is to resolve uncertainty in generative models used to predict complex environments in order to maximize inclusive fitness.The macroecological solution is to link complex brains in social networks to form collective brains that perform collective computations.By developing theory and analyzing data,the author shows hunter-gatherers bands of~16 people,or~4 co-residing families,form the largest collective brains of any social mammal.Moreover,because individuals,families,and bands interact at multiple time scales,these fission-fusion dynamics lead to the emergence of the macroscopic regularities in hunter-gatherer macroecology we observe in cross-cultural data.These results show how computation is distributed across spatially-extended social networks forming decentralized knowledge systems characteristic of hunter-gatherer societies.The flow of information at scales far beyond daily interactions leads to the emergence of small-worlds where highly clustered local interactions are embedded within much larger,but sparsely connected multilevel metapopulations.