Cascading extinctions and community collapse in model food webs

Species loss in ecosystems can lead to secondary extinctions as a result of consumer–resource relationships and other species interactions. We compare levels of secondary extinctions in communities generated by four structural food-web models and a fifth null model in response to sequential primary species removals. We focus on various aspects of food-web structural integrity including robustness, community collapse and threshold periods, and how these features relate to assumptions underlying different models, different species loss sequences and simple measures of diversity and complexity. Hierarchical feeding, a fundamental characteristic of food-web structure, appears to impose a cost in terms of robustness and other aspects of structural integrity. However, exponential-type link distributions, also characteristic of more realistic models, generally confer greater structural robustness than the less skewed link distributions of less realistic models. In most cases for the more realistic models, increased robustness and decreased levels of web collapse are associated with increased diversity, measured as species richness S, and increased complexity, measured as connectance C. These and other results, including a surprising sensitivity of more realistic model food webs to loss of species with few links to other species, are compared with prior work based on empirical food-web data.     

Using food web dominator trees to catch secondary extinctions in action

In ecosystems, a single extinction event can give rise to multiple ‘secondary’ extinctions. Conservation effort would benefit from tools that help forecast the consequences of species removal. One such tool is the dominator tree, a graph-theoretic algorithm that when applied to food webs unfolds their complex architecture, yielding a simpler topology made of linear pathways that are essential for energy delivery. Each species along these chains is responsible for passing energy to the taxa that follow it and, as such, it is indispensable for their survival. To assess the predictive potential of the dominator tree, we compare its predictions with the effects that followed the collapse of the capelin (Mallotus villosus) in the Barents Sea ecosystem. To this end, we first compiled a food web for this ecosystem, then we built the corresponding dominator tree and, finally, we observed whether model predictions matched the empirical observations. This analysis shows the potential and the drawbacks of the dominator trees as a tool for understanding the causes and consequences of extinctions in food webs.     

Species loss leads to community closure

Global extinction of a species is sadly irreversible. At a local scale, however, extinctions may be followed by re-invasion. We here show that this is not necessarily the case and that an ecological community may close its doors for re-invasion of species lost from it. Previous studies of how communities are assembled have shown that there may be rules for that process and that limitations are set to the order by which species are introduced and put together. Instead of focusing on the assembly process we randomly generated simple competitive model communities that were stable and allowed for two to 10 coexisting species. When a randomly selected single species was removed from the community, the cascading species loss was recorded and frequently the resulting community was more than halved. Cascading extinctions have previously been recorded, but we here show that the relative magnitude of the cascade is dependent on community size (and not only trophic structure) and that the reintroduction of the original species lost often is impossible. Hence, species loss does not simply leave a void potentially refilled, but permanently alters the entire community structure and consequently the adaptive landscape for potential re-invaders.     

Species-rich ecosystems are vulnerable to cascading extinctions in an increasingly variable world

Global warming leads to increased intensity and frequency of weather extremes. Such increased environmental variability might in turn result in increased variation in the demographic rates of interacting species with potentially important consequences for the dynamics of food webs. Using a theoretical approach, we here explore the response of food webs to a highly variable environment. We investigate how species richness and correlation in the responses of species to environmental fluctuations affect the risk of extinction cascades. We find that the risk of extinction cascades increases with increasing species richness, especially when correlation among species is low. Initial extinctions of primary producer species unleash bottom-up extinction cascades, especially in webs with specialist consumers. In this sense, species-rich ecosystems are less robust to increasing levels of environmental variability than species-poor ones. Our study thus suggests that highly species-rich ecosystems such as coral reefs and tropical rainforests might be particularly vulnerable to increased climate variability.     

Keystone species and food webs

Different species are of different importance in maintaining ecosystem functions in natural communities. Quantitative approaches are needed to identify unusually important or influential, ‘keystone’ species particularly for conservation purposes. Since the importance of some species may largely be the consequence of their rich interaction structure, one possible quantitative approach to identify the most influential species is to study their position in the network of interspecific interactions. In this paper, I discuss the role of network analysis (and centrality indices in particular) in this process and present a new and simple approach to characterizing the interaction structures of each species in a complex network. Understanding the linkage between structure and dynamics is a condition to test the results of topological studies, I briefly overview our current knowledge on this issue. The study of key nodes in networks has become an increasingly general interest in several disciplines: I will discuss some parallels. Finally, I will argue that conservation biology needs to devote more attention to identify and conserve keystone species and relatively less attention to rarity.     

Parasites reduce food web robustness because they are sensitive to secondary extinction as illustrated by an invasive estuarine snail

A robust food web is one in which few secondary extinctions occur after removing species. We investigated how parasites affected the robustness of the Carpinteria Salt Marsh food web by conducting random species removals and a hypothetical, but plausible, species invasion. Parasites were much more likely than free-living species to suffer secondary extinctions following the removal of a free-living species from the food web. For this reason, the food web was less robust with the inclusion of parasites. Removal of the horn snail, Cerithidea californica, resulted in a disproportionate number of secondary parasite extinctions. The exotic Japanese mud snail, Batillaria attramentaria, is the ecological analogue of the native California horn snail and can completely replace it following invasion. Owing to the similarities between the two snail species, the invasion had no effect on predator–prey interactions. However, because the native snail is host for 17 host-specific parasites, and the invader is host to only one, comparison of a food web that includes parasites showed significant effects of invasion on the native community. The hypothetical invasion also significantly reduced the connectance of the web because the loss of 17 native trematode species eliminated many links.     

How global extinctions impact regional biodiversity in mammals

Phylogenetic diversity (PD) represents the evolutionary history of a species assemblage and is a valuable measure of biodiversity because it captures not only species richness but potentially also genetic and functional diversity. Preserving PD could be critical for maintaining the functional integrity of the world's ecosystems, and species extinction will have a large impact on ecosystems in areas where the ecosystem cost per species extinction is high. Here, we show that impacts from global extinctions are linked to spatial location. Using a phylogeny of all mammals, we compare regional losses of PD against a model of random extinction. At regional scales, losses differ dramatically: several biodiversity hotspots in southern Asia and Amazonia will lose an unexpectedly large proportion of PD. Global analyses may therefore underestimate the impacts of extinction on ecosystem processes and function because they occur at finer spatial scales within the context of natural biogeography.     

Ecosystem function and species loss – a microcosm study

There are too many kinds of organisms to be able to study and manage each, yet the loss of a single species can sometimes unravel an ecosystem. Such `fusewire species'– critical in the same sense that an electrical fuse can cut out a whole circuit – would be a rewarding focus for research and management effort. However, this approach can only be effective if these `fusewires' represent but a small proportion of the number of species in the system.  

Aim

To demonstrate methods for measuring what proportion of the species in a system are critical to ecosystem function.  

Methods

The prevalence of fusewire species was measured in manipulative experiments on an aquatic microcosm.  

Results

No single genus deletion caused changes in key characteristics of the system.  

Main conclusions

Comparison of these results with other published studies shows that the proportion of critical fusewire species varies amongst different ecosystems. The oxidation pond microcosms were shown to contain no single species indispensable to system function. They appear to be ill-suited to a management strategy which focuses on priority eukaryote species. However, a single study provides no evidence that this result is general or even typical of other kinds of ecosystems; it is presented here as an empirical model. Other methods of investigation are available; they are less experimentally rigorous but more practical. These could provide important guidance in planning an approach to management in a particular ecosystem.     

Extinctions in simple and complex communities

Disagreement exists between the results of theoretical and empirical exploration into the effect of increasing community complexity on the stability of multispecies ecosystems. A recent return to interest in this area suggests previous results should be re-assessed, from both experimental studies and models, to understand where this discrepancy arises from. Here we propose various simple extensions to a standard multispecies community model that each increase the complexity of the system in a different way. We find that increasing the number of species in a community leads to a decrease in community persistence after the system is perturbed, and go on to show that increasing the dynamical diversity of the community members leads to an increase in stability through a reduction in extinction events, relative to the less complex form of the model. Our results suggest that different forms of complexity lead to different outcomes in the stability properties of the community. While aspects of this work agree with previous empirical findings that more complex communities are more robust to perturbation, we stress that the type of complexity included and the measure of stability used in community models must be properly defined, to allow objective comparisons to be made with previous and future work.     

Linking genes to communities and ecosystems: Daphnia as an ecogenomic model

How do genetic variation and evolutionary change in critical species affect the composition and functioning of populations, communities and ecosystems? Illuminating the links in the causal chain from genes up to ecosystems is a particularly exciting prospect now that the feedbacks between ecological and evolutionary changes are known to be bidirectional. Yet to fully explore phenomena that span multiple levels of the biological hierarchy requires model organisms and systems that feature a comprehensive triad of strong ecological interactions in nature, experimental tractability in diverse contexts and accessibility to modern genomic tools. The water flea Daphnia satisfies these criteria, and genomic approaches capitalizing on the pivotal role Daphnia plays in the functioning of pelagic freshwater food webs will enable investigations of eco-evolutionary dynamics in unprecedented detail. Because its ecology is profoundly influenced by both genetic polymorphism and phenotypic plasticity, Daphnia represents a model system with tremendous potential for developing a mechanistic understanding of the relationship between traits at the genetic, organismal and population levels, and consequences for community and ecosystem dynamics. Here, we highlight the combination of traits and ecological interactions that make Daphnia a definitive model system, focusing on the additional power and capabilities enabled by recent molecular and genomic advances.     

Uncertainty in identifying local extinctions: the distribution of missing data and its effects on biodiversity measures

Identifying local extinctions is integral to estimating species richness and geographic range changes and informing extinction risk assessments. However, the species occurrence records underpinning these estimates are frequently compromised by a lack of recorded species absences making it impossible to distinguish between local extinction and lack of survey effort—for a rigorously compiled database of European and Asian Galliformes, approximately 40% of half-degree cells contain records from before but not after 1980. We investigate the distribution of these cells, finding differences between the Palaearctic (forests, low mean human influence index (HII), outside protected areas (PAs)) and Indo-Malaya (grassland, high mean HII, outside PAs). Such cells also occur more in less peaceful countries. We show that different interpretations of these cells can lead to large over/under-estimations of species richness and extent of occurrences, potentially misleading prioritization and extinction risk assessment schemes. To avoid mistakes, local extinctions inferred from sightings records need to account for the history of survey effort in a locality.     

Species loss of stoneflies (Plecoptera) in the Czech Republic during the 20th century

1. Rapid expansion and intensification of anthropogenic activities in the 20th century has caused profound changes in freshwater assemblages. Unfortunately, knowledge of the extent and causes of species loss (SL) is limited due to the lack of reliable historical data. An unusual data set allows us to compare changes in the most sensitive of aquatic insect orders, the Plecoptera, at some 170 locations in the Czech Republic between two time periods, 1955–1960 and 2006–2010. Historical data (1890–1911) on assemblages of six lowland rivers allow us to infer even earlier changes. 2. Regional stonefly diversity decreased in the first half of the 20th century. Streams at lower altitudes lost a substantial number of species, which were never recovered. In the second half of the century, large‐scale anthropogenic pressure caused SL in all habitats, leading to a dissimilarity of contemporary and previous assemblages. The greatest changes were found at sites affected by organic pollution and a mixture of organic pollution and channelisation or impoundment. Colonisation of new habitats was observed in only three of the 80 species evaluated. 3. Species of moderate habitat specialisation and tolerance to organic pollution were most likely to be lost. Those with narrow specialisations in protected habitats were present in both historical and contemporary collections. 4. Contemporary assemblages are the consequence of more than a 100 years of anthropogenic impacts. In particular, streams at lower altitude and draining intensively exploited landscapes host a mere fragment of the original species complement. Most stonefly species are less frequently present than before, although their assemblages remain almost intact in near‐natural mountain streams. Our analyses demonstrate dramatic restriction of species ranges and, in some cases, apparent changes in altitudinal preference throughout the area.     

Predicting invasion success in complex ecological networks

A central and perhaps insurmountable challenge of invasion ecology is to predict which combinations of species and habitats most effectively promote and prevent biological invasions. Here, we integrate models of network structure and nonlinear population dynamics to search for potential generalities among trophic factors that may drive invasion success and failure. We simulate invasions where 100 different species attempt to invade 150 different food webs with 15–26 species and a wide range (0.06–0.32) of connectance. These simulations yield 11 438 invasion attempts by non-basal species, 47 per cent of which are successful. At the time of introduction, whether or not the invader is a generalist best predicts final invasion success; however, once the invader establishes itself, it is best distinguished from unsuccessful invaders by occupying a lower trophic position and being relatively invulnerable to predation. In general, variables that reflect the interaction between an invading species and its new community, such as generality and trophic position, best predict invasion success; however, for some trophic categories of invaders, fundamental species traits, such as having the centre of the feeding range low on the theoretical niche axis (for non-omnivorous and omnivorous herbivores), or the topology of the food web (for tertiary carnivores), best predict invasion success. Across all invasion scenarios, a discriminant analysis model predicted successful and failed invasions with 76.5 per cent accuracy for properties at the time of introduction or 100 per cent accuracy for properties at the time of establishment. More generally, our results suggest that tackling the challenge of predicting the properties of species and habitats that promote or inhibit invasions from food web perspective may aid ecologists in identifying rules that govern invasions in natural ecosystems.     

Regime shifts in marine communities: a complex systems perspective on food web dynamics

Species composition and habitats are changing at unprecedented rates in the world''s oceans, potentially causing entire food webs to shift to structurally and functionally different regimes. Despite the severity of these regime shifts, elucidating the precise nature of their underlying processes has remained difficult. We address this challenge with a new analytic approach to detect and assess the relative strength of different driving processes in food webs. Our study draws on complexity theory, and integrates the network-centric exponential random graph modelling (ERGM) framework developed within the social sciences with community ecology. In contrast to previous research, this approach makes clear assumptions of direction of causality and accommodates a dynamic perspective on the emergence of food webs. We apply our approach to analysing food webs of the Baltic Sea before and after a previously reported regime shift. Our results show that the dominant food web processes have remained largely the same, although we detect changes in their magnitudes. The results indicate that the reported regime shift may not be a system-wide shift, but instead involve a limited number of species. Our study emphasizes the importance of community-wide analysis on marine regime shifts and introduces a novel approach to examine food webs.     

Paleoecological patterns in molluscan extinctions and recoveries: comparison of the Cretaceous-Paleogene and Eocene-Oligocene extinctions in North America

Exposures across the Cretaceous-Tertiary (K-T) and Eocene-Oligocene (E-O) boundaries, in Texas and Mississippi, respectively, probably represent the most complete and best-preserved fossil molluscan sequences across these boundary intervals in the world. Outcrops from both boundaries contain pristine aragonitic and calcitic molluscan shells, which were deposited in fine-grained sediments from open marine environments. The K-T and the E-O extinctions exhibit very different recovery patterns, probably reflecting very different causes as well as magnitudes of extinction.The K-T sequence contains a molluscan fossil record that is consistent with an abrupt extinction event at the K-T boundary and a prolonged initial recovery in hostile oceanographic conditions. The uppermost 10 m of Upper Cretaceous sediments contain a diverse (approximately 40 species) molluscan fauna dominated by suspension feeders. The earliest Paleocene sediments immediately above the tsunami bed contain an impoverished fauna dominated by deposit feeders. The Paleocene fauna slowly climbs in diversity but remains relatively impoverished and dominated by deposit feeders for several hundred thousand years after the extinction in conjunction with anomalous δ¹³C values that suggest prolonged suppression of marine primary productivity. Diverse suspension-feeder dominated molluscan assemblages reappear with the resumption of normal conditions of primary production. In the long term, early to middle Paleocene gamma diversity includes evolutionary “bloom taxa,” families that exhibit unusual speciation bursts that subside in the Eocene. Total diversity for the Gulf Coast does not approach Cretaceous levels until the Late Eocene representing a total recovery interval of nearly 25 million years.While the E-O event also reflects a molluscan extinction rate of over 90% in the Gulf of Mexico, there are no signs of hostile environmental conditions in the recovery fauna. Early Oligocene molluscan assemblages are diverse and dominated by suspension feeders characteristic of normal marine conditions. The hiatus at the E-O boundary, however, could have obscured a short-term recovery fauna. There is also no sign of long-term perturbation by the E-O extinction. There are no bloom taxa and gamma diversity approaches pre-extinction levels within a few million years. The overall pattern of the E-O extinction is consistent with extinction (and/or migration) associated with long-term cooling.