Environmental fluctuations have important consequences in the organization of ecological communities, and understanding how such a variability influences the biodiversity of an ecosystem is a major question in ecology. In this paper, we analyze the case of two species competing for the resources within the framework of the neutral theory in the presence of environmental noise, devoting special attention on how such a variability modulates species fitness. The environment is dichotomous and stochastically alternates between periods favoring one of the species while disfavoring the other one, preserving neutrality on the long term. We study two different scenarios: in the first one species fitness varies linearly with the environment, and in the second one the effective fitness is re-scaled by the total fitness of the individuals competing for the same resource. We find that, in the former case environmental fluctuations always reduce the time of species coexistence, whereas such a time can be enhanced or reduced in the latter case, depending on the correlation time of the environment. This phenomenon can be understood as a direct consequence of Chesson's storage effect.
The savanna biome is characterised by a continuous vegetation cover, comprised of herbaceous and woody plants. The coexistence of species in arid savannas, where water availability is the main limiting resource for plant growth, provides an apparent contradiction to the classical principle of competitive exclusion. Previous theoretical work using nonspatial models has focussed on the development of an understanding of coexistence mechanisms through the consideration of resource niche separation and ecosystem disturbances. In this paper, we propose that a spatial self-organisation principle, ca…
Vegetation patterns are a ubiquitous feature of water-deprived ecosystems. Despite the competition for the same limiting resource, coexistence of several plant species is commonly observed. We propose a two-species reaction-diffusion model based on the single-species Klausmeier model, to analytically investigate the existence of states in which both species coexist. Ecologically, the study finds that coexistence is supported if there is a small difference in the plant species' average fitness, measured by the ratio of a species' capabilities to convert water into new biomass to its mortality r…
Resource competition theory predicts coexistence and exclusion patterns based on species R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesize that coexistence under preemption requires an R*-preemption tradeoff, i.e., the species with the priority access should ha…
Cyclic dominance between species may yield spiral waves that are known to provide a mechanism enabling persistent species coexistence. This observation holds true even in presence of spatial heterogeneity in the form of quenched disorder. In this work we study the effects on spatio-temporal patterns and species coexistence of structured spatial heterogeneity in the form of habitats that locally provide one of the species with an advantage. Performing extensive numerical simulations of systems with three and six species we show that these structured habitats destabilize spiral waves. Analyzing …
Explaining coexistence in species-rich communities of primary producers remains a challenge for ecologists because of their likely competition for shared resources. Following Hutchinson's seminal suggestion, many theoreticians have tried to create diversity through a fluctuating environment, which impairs or slows down competitive exclusion. However, fluctuating-environment models often only produce a dozen of coexisting species at best. Here, we investigate how to create richer communities in fluctuating environments, using an empirically parameterized model. Building on the forced Lotka-Volt…
Local coexistence of species in large ecosystems is traditionally explained within the broad framework of niche theory. However, its rationale hardly justifies rich biodiversity observed in nearly homogeneous environments. Here we consider a consumer-resource model in which a coarse-graining procedure accounts for a variety of ecological mechanisms and leads to effective spatial effects which favour species coexistence. Herein, we provide conditions for several species to live in an environment with very few resources. In fact, the model displays two different phases depending on whether the n…
Patterned vegetation is a characteristic feature of many dryland ecosystems. While plant densities on the ecosystem-wide scale are typically low, a spatial self-organisation principle leads to the occurrence of alternating patches of high biomass and patches of bare soil. Nevertheless, intraspecific competition dynamics other than competition for water over long spatial scales are commonly ignored in mathematical models for vegetation patterns. In this chapter, I address the impact of local intraspecific competition on a modelling framework for banded vegetation patterns. Firstly, I show that …
We introduce an asymmetric noisy voter model to study the joint effect of immigration and a competition-dispersal tradeoff in the dynamics of two species competing for space in regular lattices. Individuals of one species can invade a nearest-neighbor site in the lattice, while individuals of the other species are able to invade sites at any distance but are less competitive locally, i.e., they establish with a probability $g \le 1$. The model also accounts for immigration, modeled as an external noise that may spontaneously replace an individual at a lattice site by another individual of the …
How do species coexist? A framework known as Modern Coexistence Theory measures mechanisms of coexistence by comparing a species perturbed to low density (the invader) to other species that remain at their typical densities (the residents); this invader-resident comparison measures a rare-species advantage that results from specialization. However, there are several reasonable ways (i.e., methods) to compare invaders and residents, each differing in practicality and biological interpretation. Here, using theoretical arguments and case studies, we compare four such methods for calculating coexi…
The paradox of the plankton highlights the apparent contradiction between Gause's law of competitive exclusion and the observed diversity of phytoplankton. It is well known that phytoplankton dynamics depend heavily on two main resources: light and nutrients. Here we treat light as a continuum of resources rather than a single resource by considering the visible light spectrum. We propose a spatially explicit reaction-diffusion-advection model to explore under what circumstance coexistence is possible from mathematical and biological perspectives. Furthermore, we provide biological context as …
How large ecosystems can create and maintain the remarkable biodiversity we see in nature is probably one of the biggest open questions in science, attracting attention from different fields, from Theoretical Ecology to Mathematics and Physics. In this context, modeling the stable coexistence of species competing for limited resources is a particularly challenging task. From a mathematical point of view, coexistence in competitive dynamics can be achieved when dominance among species forms intransitive loops. However, these relationships usually lead to species' relative abundances neutrally c…
We study a class of the stochastic May-Leonard models, with three species dominating each other in a cyclic nonhierarchical way, according to the rock-paper-scissors game. We introduce an unevenness in the system, by considering that one of the species is weaker because of a lower selection probability. The simulation results show that the pattern formation is drastically affected by the presence of the weaker species, with no spiral waves arising immediately from random initial conditions. Instead, single-species spatial domains cyclically dominate the entire territory until a region occupied…
To understand the mechanisms underlying species coexistence, ecologists often study invasion growth rates of theoretical and data-driven models. These growth rates correspond to average per-capita growth rates of one species with respect to an ergodic measure supporting other species. In the ecological literature, coexistence often is equated with the invasion growth rates being positive. Intuitively, positive invasion growth rates ensure that species recover from being rare. To provide a mathematically rigorous framework for this approach, we prove theorems that answer two questions: (i) When…
Reaction-diffusion systems with a Lotka-Volterra-type reaction term, also known as competition-diffusion systems, have been used to investigate the dynamics of the competition among $m$ ecological species for a limited resource necessary to their survival and growth. Notwithstanding their rather simple mathematical structure, such systems may display quite interesting behaviours. In particular, while for $m=2$ no coexistence of the two species is usually possible, if $m \ge 3$ we may observe coexistence of all or a subset of the species, sensitively depending on the parameter values. Such coex…
Selection Rules for Species Coexistence in a Hierarchical May-Leonard Model
Raja Rammohun Roy Mahavidyalaya（拉贾·拉姆莫汉·罗伊大学）；Center for Computational Natural Sciences and Bioinformatics, International Institute of Information Technology Hyderabad（海德拉巴信息技术学院计算自然科学和生物信息中心）；Sister Nibedita Govt. General Degree College for Girls（西比尔·尼贝迪塔女子政府综合学位学院）；School of Science, Constructor University（Constructor大学理学院）；Department of Mathematics, Nalanda University（那烂陀大学数学系）
Frequency-dependent selection reflects the interaction between different species as they battle for limited resources in their environment. In a stochastic evolutionary game the species relative fitnesses guides the evolutionary dynamics which fluctuate due to random drift. Dependence of species selection advantages on the environment introduces additional possibilities for the evolutionary dynamics. We analyse a simple model in which a random environment allows competing species to coexist for a long time before a fixation of a single species happens. In our analysis we use stability in a lin…
Statistical Physics has proved essential to analyze multi-agent environments. Motivated by the empirical observation of various non-equilibrium features in Barro Colorado and other ecological systems, we analyze a plant-species abundance model, presenting analytical evidence of scale-invariant plant clusters and non-trivial emergent modular correlations. Such first theoretical confirmation of a scale-invariant region, based on percolation processes, reproduces the key features in actual ecological ecosystems and can confer the most stable equilibrium for ecosystems with vast biodiversity.
Disentangling the interactive effects of anthropogenic disturbances on biodiversity
RIKEN Wako Campus, Division of Fundamental Mathematical Science（理化学研究所和光园区基础数学科学部）；Tokyo Metropolitan University, Department of Biological Sciences（东京都立大学生物科学系）
We introduce and analyze a spatial Lotka-Volterra competition model with local and nonlocal interactions. We study two alternative classes of nonlocal competition that differ in how each species' characteristics determine the range of the nonlocal interactions. In both cases, nonlocal interactions can create spatial patterns of population densities in which highly populated clumps alternate with unpopulated regions. This non-populated regions provide spatial niches for a weaker competitor to establish in the community and persist in conditions in which local models predict competitive exclusio…
Towards a less spherical cow: Species differences dilute the stabilizing effect of higher-order interactions
École Polytechnique Fédérale de Lausanne (EPFL)（洛桑联邦理工学院）；Institute for Cross-Disciplinary Physics and Complex Systems (IFISC), CSIC-UIB（跨学科物理与复杂系统研究所（CSIC-UIB））；Centro Studi e Ricerche “Enrico Fermi” (CREF)（“恩里科·费米”研究与学习中心）；Institute for Applied Mathematics Mauro Picone (IAC) CNR（马乌罗·皮科内应用数学研究所（CNR））；Estación Biológica de Doñana (CSIC)（多尼亚纳生物站（CSIC））；University of Oxford（牛津大学）
Neutral species facilitate coexistence among cyclically competing species under birth and death processes
School of Statistics and Mathematics, Yunnan University of Finance and Economics（云南财经大学统计与数学学院）；School of Mathematics and Statistics, Xidian University（西安电子科技大学数学与统计学院）；Department of Applied Mathematics, Colleage of Applied Sciences, Kyung Hee University（韩国庆熙大学应用科学学院应用数学系）；G-LAMP NEXUS Institute, Kyung Hee University（韩国庆熙大学G-LAMP NEXUS研究所）
We analyze the long term behavior of interacting populations which can be controlled through harvesting. The dynamics is assumed to be discrete in time and stochastic due to the effect of environmental fluctuations. We present extinction and coexistence criteria when there are one or two interacting species. We then use these tools in order to see when harvesting leads to extinction or persistence of species, as well as what the optimal harvesting strategies, which maximize the expected long term yield, look like. For single species systems, we show under certain conditions that the optimal ha…
Decoding species coexistence: A reinforcement learning perspective
School of Physics and Information Technology, Shaanxi Normal University（陕西师范大学物理与信息技术学院）；School of Physics and Electronic Science, East China Normal University（华东师范大学物理与电子科学学院）；School of Computer Science, Soochow University（苏州大学计算机科学与技术学院）；School of Physics, Ningxia University（宁夏大学物理学院）
