Habitat complexity has been considered a key driver of biodiversity and other ecological phenomena for nearly a century. However, there is still no consensus over the definition of complexity or how ...to measure it. Up‐to‐date and clear guidance on measuring complexity is urgently needed, particularly given the rise of remote sensing and advent of technologies that allow environments to be scanned at unprecedented spatial extents and resolutions. Here we review how complexity is measured in ecology. We provide a framework for metrics of habitat complexity, and for the related concept of spatial heterogeneity. We focus on the two most commonly used complexity metrics in ecology: fractal dimension and rugosity. We discuss the pros and cons of these metrics using practical examples from our own empirical data and from simulations. Fractal dimension is particularly widely used, and we provide a critical examination of it drawing on research from other scientific fields. We also discuss informational metrics of complexity and their potential benefits. We chart a path forward for research on measuring habitat complexity by presenting, as a guide, sets of essential and desirable criteria that a metric of complexity should possess. Lastly, we discuss the applied significance of our review.
Given the rise of remote‐sensing, up‐to‐date and clear guidance on measuring complexity is urgently needed. Here we review how complexity is measured in ecology. We provide a framework for metrics of habitat complexity, and for the related concept of spatial heterogeneity. We demonstrate the pros and cons of various complexity metrics particularly focusing on popular ones; we provide a modern critical examination of fractal dimension and show how and why it is untenable despite its appeal. We discuss informational metrics of complexity, and their potential benefits and chart a path forward for research on measuring complexity in ecology.
•Non-pharmaceutical interventions (NPIs) were enacted in COVID-19 pandemic.•We examine effect of NPIs on total disease burden of a non-target respiratory disease.•To do this, we mathematically ...analyse and simulate four models of varying complexity.•We show that NPIs over about a year have a negative effect on total disease burden.•The negative effect increased in magnitude with the rate at which immunity is lost.
The recent global COVID-19 pandemic resulted in governments enacting non-pharmaceutical interventions (NPIs) targeted at reducing transmission of SARS-CoV-2. But the NPIs also affected the transmission of viruses causing non-target seasonal respiratory diseases, including influenza and respiratory syncytial virus (RSV). In many countries, the NPIs were found to reduce cases of such seasonal respiratory diseases, but there is also evidence that subsequent relaxation of NPIs led to outbreaks of these diseases that were larger than pre-pandemic ones, due to the accumulation of susceptible individuals prior to relaxation. Therefore, the net long-term effects of NPIs on the total disease burden of non-target diseases remain unclear. Knowledge of this is important for infectious disease management and maintenance of public health. In this study, we shed light on this issue for the simplified scenario of a set of NPIs that prevent or reduce transmission of a seasonal respiratory disease for about a year and are then removed, using mathematical analyses and numerical simulations of a suite of four epidemiological models with varying complexity and generality. The model parameters were estimated using empirical data pertaining to seasonal respiratory diseases and covered a wide range. Our results showed that NPIs reduced the total disease burden of a non-target seasonal respiratory disease in the long-term. Expressed as a percentage of population size, the reduction was greater for larger values of the basic reproduction number and the immunity loss rate, reflecting larger outbreaks and hence more infections averted by imposition of NPIs. Our study provides a foundation for exploring the effects of NPIs on total disease burden in more-complex scenarios.
Abstract
Hyperdiverse tropical forest tree communities illustrate a fundamental problem in ecology: How can many species coexist given relatively few limiting resources? Neutral theory provides a ...solution by positing that species have equal fitness and hence drift to extinction slowly. However, neutral theory seriously under‐predicts temporal changes in species abundances. This can be remedied by breaking neutrality and adding temporal environmental stochasticity (TES), but the mechanisms mediating the effects of TES on species richness remain unclear. Here, we make progress by analysing a local community model with species competing for a common resource under TES, to derive formulae partitioning species richness according to different mechanisms.
By applying our formulae to generic parameter sets for tropical forest tree communities, we found that when the autocorrelation time of TES was short, the dominant mechanism driving species richness was non‐linear averaging of the interspecific competition term over time, which reduced the typical strength of interspecific competition and boosted richness relative to the neutral case. However, greater immigration to the community resulted in more species and hence weaker non‐linear averaging due to the law of large numbers. In contrast, when the autocorrelation time of TES was long, the dominant mechanism driving species richness was strong selection between changes in environmental conditions, which increased the typical strength of interspecific competition and reduced richness relative to the neutral case.
By applying our formulae to a specific parameter set for a tropical forest tree community in Panama, we found that TES had minor effects on species richness because (i) the immigration rate was sufficiently large for non‐linear averaging of the interspecific competition term to be weak and (ii) the autocorrelation time was sufficiently short to suppress the effects of selection.
Synthesis
. We provide a novel mechanistic explanation of how TES affects tree species richness in tropical forests, in particular how TES often has minor effects on richness despite having substantial effects on temporal changes in species abundances. This provides a deeper insight into why a neutral model with added TES can accurately capture static and dynamic aspects of tree community diversity in the tropics.
AbstractA goal of ecology is to identify the stabilizing mechanisms that maintain species diversity in the face of competitive exclusion and drift. For tropical forest tree communities, it has been ...hypothesized that high diversity is maintained via Janzen-Connell effects, whereby host-specific natural enemies prevent any one species from becoming too abundant. Here we explore the plausibility of this hypothesis with theoretical models. We confirm a previous result that when added to a model with drift but no competitive exclusion-that is, a neutral model where intrinsic fitnesses are perfectly equalized across species-Janzen-Connell effects maintain very high species richness that scales strongly with community size. However, when competitive exclusion is introduced-that is, when intrinsic fitnesses vary across species-the number of species maintained by Janzen-Connell effects is substantially reduced and scales much less strongly with community size. Because fitness variation is pervasive in nature, we conclude that the potential of Janzen-Connell effects to maintain diversity is probably weak and that the mechanism does not yet provide a sufficient explanation for the observed high diversity of tropical forest tree communities. We also show that, surprisingly, dispersal limitation can further reduce the ability of Janzen-Connell effects to maintain diversity.
Carbon sequestration by afforestation can help mitigate global climate change but may have adverse environmental and economic impacts in some regions. For example, economic incentives for carbon ...sequestration may encourage the expansion of
Pinus radiata timber plantations in the Fynbos biome of South Africa, with negative consequences for water supply and biodiversity. I built a dynamic ecological–economic model to investigate whether afforestation of a Fynbos catchment with
Pinus radiata is economically viable when the potential benefits of carbon sequestration and timber production are balanced against the losses to water supply. I found that afforestation appears viable to the forestry industry under current water tariffs and current carbon accounting legislation, but would appear unviable if the forestry industry were to pay the true cost of water used by the plantations. I also found that under various plausible future economic scenarios, afforestation can be associated with either large future economic gains or losses, suggesting a need for future analyses based on branches of decision theory that deal with severe uncertainty. I conclude with a general recommendation that climate legislation should be explicit about the conditions under which afforestation for carbon sequestration of native vegetation is a legitimate climate mitigation strategy.
Aim
Artificial island habitats such as human‐made wetlands are emerging novel ecosystems. Understanding the drivers of diversity in such artificial systems is essential for balancing the goals of ...biodiversity conservation and human socio‐economic needs.
Location
Telangana state, India.
Methods
We surveyed water birds in a network of 57 artificial wetlands and assessed four macroecological biodiversity patterns: spatial betadiversity, temporal betadiversity, species‐abundance distributions (SADs), and the species–area relationship (SAR). We employed a mix of phenomenological and mechanistic models to examine the four macroecological patterns. We hypothesized that the wetland bird communities are primarily structured by immigration–extinction dynamics and thus that spatial and temporal betadiversity would be high, the within‐wetland SADs would exhibit a large number of rare species and a monotonically declining overall shape, and that the SAR across wetlands would be strongly increasing.
Results
Spatial and temporal betadiversity were both high and mostly attributable to turnover rather than nestedness. While the pooled SAD exhibited an interior mode, the SAD for individual wetlands was generally log‐series distributed, consistent with a model in which immigration among wetlands is high. The SAR exhibited an increasing trend, with the ‘small‐island effect’, which reflects constraints on immigration and is often observed for true island archipelagos, being absent.
Main Conclusions
We tentatively conclude that bird diversity in this network of artificial wetlands is mainly structured by immigration–extinction dynamics, although we acknowledge that some of the patterns are also consistent with niche dynamics and future research should measure relevant biotic and abiotic variables in these wetlands. We encourage future work in which our rich dataset is used to fit dynamic models that permit more‐detailed quantitative inferences about mechanisms structuring diversity in this novel ecosystem, which can ultimately also inform conservation management.
Half a century ago, Janzen and Connell hypothesized that the high tree species diversity in tropical forests is maintained by specialized natural enemies. Along with other mechanisms, these can cause ...conspecific negative density dependence (CNDD) and thus maintain species diversity. Numerous studies have measured proxies of CNDD worldwide, but doubt about its relative importance remains. We find ample evidence for CNDD in local populations, but methodological limitations make it difficult to assess if CNDD scales up to control community diversity and thereby local and global biodiversity patterns. A combination of more robust statistical methods, new study designs, and eco-evolutionary models are needed to provide a more definite evaluation of the importance of CNDD for geographic variation in plant species diversity.
Stabilizing conspecific negative density dependence (CNDD) is a classical explanation for diversity maintenance in species-rich ecosystems, particularly in tropical forests.Fifty years after Janzen and Connell popularized this idea, a plethora of empirical evidence suggests that CNDD can be found at all latitudes and for many tree species.On closer inspection, however, it is still unclear if locally measured density effects are indeed stabilizing community dynamics and, moreover, if they have a causal effect on large-scale diversity and abundance patterns, such as the latitudinal diversity gradient.More robust and comparable CNDD estimates are needed, coupled with a theoretical research program that aims at understanding the role of CNDD for coexistence in stochastic multispecies communities as well as macroecological and macroevolutionary diversity patterns.
A fundamental challenge in ecology is to understand the mechanisms that govern patterns of relative species abundance. Previous numerical simulations have suggested that complex niche-structured ...models produce species abundance distributions (SADs) that are qualitatively similar to those of very simple neutral models that ignore differences between species. However, in the absence of an analytical treatment of niche models, one cannot tell whether the two classes of model produce the same patterns via similar or different mechanisms. We present an analytical proof that, in the limit as diversity becomes large, a strong niche model give rises to exactly the same asymptotic form of SAD as the neutral model, and we verify the analytical predictions for a Panamanian tropical forest data set. Our results strongly suggest that neutral processes drive patterns of relative species abundance in high-diversity ecological communities, even when strong niche structure exists. However, neutral theory cannot explain what generates high diversity in the first place, and it may not be valid in low-diversity communities. Our results also confirm that neutral theory cannot be used to infer an absence of niche structure or to explain ecosystem function.
Ecology lacks a holistic approach that can model phenomena across temporal and spatial scales, largely because of the challenges in modelling systems with a large number of interacting constituents. ...This hampers our understanding of complex ecosystems and the impact that human interventions (e.g., deforestation, wildlife harvesting and climate change) have on them. Here we use density functional theory, a computational method for many-body problems in physics, to develop a computational framework for ecosystem modelling. Our methods accurately fit experimental and synthetic data of interacting multi-species communities across spatial scales and can project to unseen data. As the key concept we establish and validate a cost function that encodes the trade-offs between the various ecosystem components. We show how this single general modelling framework delivers predictions on par with established, but specialised, approaches for systems from predatory microbes to territorial flies to tropical tree communities. Our density functional framework thus provides a promising avenue for advancing our understanding of ecological systems.
LaManna
(Reports, 30 June 2017, p. 1389) found higher conspecific negative density dependence in tree communities at lower latitudes, yielding a possible mechanistic explanation for the latitudinal ...diversity gradient. We show that their results are artifacts of a selective data transformation and a forced zero intercept in their fitted model. A corrected analysis shows no latitudinal trend.
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