Global versus local extinction in a network model of plant–pollinator communities

The loss of a species from an ecological community can trigger a cascade of additional extinctions; the complex interactions that comprise ecological communities make the dynamics and impacts of such a cascade challenging to predict. Previous studies have typically considered global extinctions, where a species cannot re-enter a community once it is lost. However, in some cases a species only becomes locally extinct, and may be able to reinvade from surrounding communities. Here, we use a dynamic, Boolean network model of plant–pollinator community assembly to analyze the differences between global and local extinction events in mutualistic communities. As expected, we find that compared to global extinctions, communities respond to local extinctions with lower biodiversity loss, and less variation in topological network properties. We demonstrate that in the face of global extinctions, larger communities suffer greater biodiversity loss than smaller communities when similar proportions of species are lost. Conversely, smaller communities suffer greater loss in the face of local extinctions. We show that targeting species with the most interacting partners causes more biodiversity loss than random extinctions in the case of global, but not local, extinctions. These results extend our understanding of how mutualistic communities respond to species loss, with implications for community management and conservation efforts.     

Species loss and secondary extinctions in simple and complex model communities

1. The loss of a species from an ecological community can trigger a cascade of secondary extinctions. Here we investigate how the complexity (connectance) of model communities affects their response to species loss. Using dynamic analysis based on a global criterion of persistence (permanence) and topological analysis we investigate the extent of secondary extinctions following the loss of different kinds of species. 2. We show that complex communities are, on average, more resistant to species loss than simple communities: the number of secondary extinctions decreases with increasing connectance. However, complex communities are more vulnerable to loss of top predators than simple communities. 3. The loss of highly connected species (species with many links to other species) and species at low trophic levels triggers, on average, the largest number of secondary extinctions. The effect of the connectivity of a species is strongest in webs with low connectance. 4. Most secondary extinctions are due to direct bottom-up effects: consumers go extinct when their resources are lost. Secondary extinctions due to trophic cascades and disruption of predator-mediated coexistence also occur. Secondary extinctions due to disruption of predator-mediated coexistence are more common in complex communities than in simple communities, while bottom-up and top-down extinction cascades are more common in simple communities. 5. Topological analysis of the response of communities to species loss always predicts a lower number of secondary extinctions than dynamic analysis, especially in food webs with high connectance.     

Early onset of secondary extinctions in ecological communities following the loss of top predators

The large vulnerability of top predators to human-induced disturbances on ecosystems is a matter of growing concern. Because top predators often exert strong influence on their prey populations their extinction can have far-reaching consequences for the structure and functioning of ecosystems. It has, for example, been observed that the local loss of a predator can trigger a cascade of secondary extinctions. However, the time lags involved in such secondary extinctions remain unexplored. Here we show that the loss of a top predator leads to a significantly earlier onset of secondary extinctions in model communities than does the loss of a species from other trophic levels. Moreover, in most cases time to secondary extinction increases with increasing species richness. If local secondary extinctions occur early they are less likely to be balanced by immigration of species from local communities nearby. The implications of these results for community persistence and conservation priorities are discussed.     

Keystone species and vulnerable species in ecological communities: strong or weak interactors?

The loss of a species from an ecological community can trigger a cascade of secondary extinctions. The probability of secondary extinction to take place and the number of secondary extinctions are likely to depend on the characteristics of the species that is lost--the strength of its interactions with other species--as well as on the distribution of interaction strengths in the whole community. Analysing the effects of species loss in model communities we found that removal of the following species categories triggered, on average, the largest number of secondary extinctions: (a) rare species interacting strongly with many consumers, (b) abundant basal species interacting weakly with their consumers and (c) abundant intermediate species interacting strongly with many resources. We also found that the keystone status of a species with given characteristics was context dependent, that is, dependent on the structure of the community where it was embedded. Species vulnerable to secondary extinctions were mainly species interacting weakly with their resources and species interacting strongly with their consumers.     

The local introduction of strongly interacting species and the loss of geographic variation in species and species interactions

Species introductions into nearby communities may seem innocuous, however, these introductions, like long-distance introductions (e.g. trans- and intercontinental), can cause extinctions and alter the evolutionary trajectories of remaining community members. These 'local introductions' can also more cryptically homogenize formerly distinct populations within a species. We focus on several characteristics and the potential consequences of local introductions. First, local introductions are commonly successful because the species being introduced is compatible with existing abiotic and biotic conditions; many nearby communities differ because of historical factors and the absence of certain species is simply the result of barriers to dispersal. Moreover, the species with which they interact most strongly (e.g. prey) may have, for example, lost defences making the establishment even more likely. The loss or absence of defences is especially likely when the absent species is a strongly interacting species, which we argue often includes mammals in terrestrial communities. Second, the effects of the introduction may be difficult to detect because the community is likely to converge onto nearby communities that naturally have the introduced species (hence the perceived innocuousness). This homogenization of formerly distinct populations eliminates the geographic diversity of species interactions and the geographic potential for speciation, and reduces regional species diversity. We illustrate these ideas by focusing on the introduction of tree squirrels into formerly squirrel-less forest patches. Such introductions have eliminated incipient species of crossbills (Loxia spp.) co-evolving in arms races with conifers and will likely have considerable impacts on community structure and ecosystem processes.     

Sixty years of community change in the prairie–savanna–forest mosaic of Wisconsin

Biodiversity loss is a global concern, and maintaining habitat complexity in naturally patchy landscapes can help retain regional diversity. A mosaic of prairie, savanna, and forest historically occurred across central North America but currently is highly fragmented due to human land conversion. It is unclear how each habitat type now contributes to regional diversity. Using legacy data, we resurveyed savanna plant communities originally surveyed in the 1950s to compare change in savannas to that in remnant forests and prairies. Savanna community structure and composition changed substantially over the past 60 years. Tree canopy density nearly doubled and many prairie and savanna specialist species were replaced by forest and non‐native species. All three habitats gained and lost many species since the 1950s, resulting in large changes in community composition from local colonizations and extinctions. Across all three habitats, regional species extinctions matched that of regional colonization resulting in no net change in regional species richness. Synthesis—Despite considerable species turnover within savannas, many species remain within the broader prairie–savanna–forest mosaic. Both regional extinctions and colonizations were high over the past 60 years, and maintaining the presence of all three community types—prairie, savanna and forest—on the landscape is critical to maintaining regional biodiversity.     

Trophically unique species are vulnerable to cascading extinction

Understanding which species might become extinct and the consequences of such loss is critical. One consequence is a cascade of further, secondary extinctions. While a significant amount is known about the types of communities and species that suffer secondary extinctions, little is known about the consequences of secondary extinctions for biodiversity. Here we examine the effect of these secondary extinctions on trophic diversity, the range of trophic roles played by the species in a community. Our analyses of natural and model food webs show that secondary extinctions cause loss of trophic diversity greater than that expected from chance, a result that is robust to variation in food web structure, distribution of interactions strengths, functional response, and adaptive foraging. Greater than expected loss of trophic diversity occurs because more trophically unique species are more vulnerable to secondary extinction. This is not a straightforward consequence of these species having few links with others but is a complex function of how direct and indirect interactions affect species persistence. A positive correlation between a species' extinction probability and the importance of its loss defines high-risk species and should make their conservation a priority.     

Species loss and the structure and functioning of multitrophic aquatic systems

Experiments and theory in single trophic level systems dominate biodiversity and ecosystem functioning research and recent debates. All natural ecosystems contain communities with multiple trophic levels, however, and this can have important effects on ecosystem structure and functioning. Furthermore, many experiments compare assembled communities, rather than examining loss of species directly. We identify three questions around which to organise an investigation of how species loss affects the structure and functioning of multitrophic systems. 1) What is the distribution of species richness among trophic levels; 2) from which trophic levels are species most often lost; and 3) does loss of species from different trophic levels influence ecosystem functioning differently? Our analyses show that: 1) Relatively few high‐quality data are available concerning the distribution of species richness among trophic levels. A new data‐set provides evidence of a decrease in species richness as trophic height increases. 2) Multiple lines of evidence indicate that species are lost from higher trophic levels more frequently than lower trophic levels. 3) A theoretical model suggests that both the structure of food webs (occurrence of omnivory and the distribution of species richness among trophic levels) and the trophic level from which species are lost determines the impact of species loss on ecosystem functioning, which can even vary in the sign of the effect. These results indicate that, at least for aquatic systems, models of single trophic level ecosystems are insufficient for understanding the functional consequences of extinctions. Knowledge is required of food web structure, which species are likely to be lost, and also whether cascading extinctions will occur.     

Reducing horizontal and vertical diversity in a foodweb triggers extinctions and impacts functions

Species loss can result in secondary extinctions and changes in ecosystem functions at distant trophic levels. Such effects of species loss are predicted to be affected by both the number of species lost within a trophic level (horizontal diversity) and the number of trophic levels lost (vertical diversity). We experimentally manipulated horizontal and vertical diversity within an aquatic insect community, and examined responses throughout the food web. Horizontal and vertical diversity both impacted ciliates: reduction of detritivorous insect diversity resulted in secondary extinctions and decreased density of ciliates, but only when an insect predator was simultaneously absent. Horizontal and vertical diversity differed in their effect on other foodweb processes, including detrital processing, predator growth, and densities of rotifers, flagellates and flatworms. These results caution that foodweb effects of multitrophic species loss may not be reliably predicted from manipulations of just one dimension of diversity.     

The structure and stability of model ecosystems assembled in a variable environment

To explore how environmental variability may create non‐random community structure, we simulated the assembly of model communities under varying levels of environmental variability. We assembled communities by creating a large pool of randomly constructed species, and then added species from this pool sequentially, allowing extinctions of invading and resident species to occur until the community became saturated. Because much current research on community structure focuses on single trophic levels, we constructed species pools consisting only of competitors. To compare with more realistic communities, we also created species pools with multiple trophic levels. For both types of communities, following assembly we calculated a variety of metrics of community structure, and five measures of community stability. Communities assembled under high environmental variability had fewer species, fewer and weaker interactions among species, and greater evenness in abundance of persisting species. For single trophic‐level communities, community size was dictated primarily by competitive exclusion. In contrast, for multiple trophic‐level communities, community size was increasingly limited by dynamical instabilities as environmental variability increased. Differences in community structure resulting from assembly under high environmental variability led to differences in community stability. According to two measures of stability related to population variability – the characteristic return rate to equilibrium and the coefficient of variation in individual species densities – stability increased for communities assembled under high environmental variability. In contrast, three additional measures of stability that are not directly related to population variability showed a variety of patterns, either increasing, decreasing, or remaining constant. Thus, communities assembled in highly variable environments are not necessarily generically more stable. Our results demonstrate that environmental variability can structure communities and affect their stability properties in non‐trivial ways. Thus, when making predictions about the response of communities to future extinctions or environmental degradation, account should be given to the forces responsible for community structure.     

Loss of predator species,not intermediate consumers,triggers rapid and dramatic extinction cascades

Ecological networks are tightly interconnected, such that loss of a single species can trigger additional species extinctions. Theory predicts that such secondary extinctions are driven primarily by loss of species from intermediate or basal trophic levels. In contrast, most cases of secondary extinctions from natural systems have been attributed to loss of entire top trophic levels. Here, we show that loss of single predator species in isolation can, irrespective of their identity or the presence of other predators, trigger rapid secondary extinction cascades in natural communities far exceeding those generally predicted by theory. In contrast, we did not find any secondary extinctions caused by intermediate consumer loss. A food web model of our experimental system—a marine rocky shore community—could reproduce these results only when biologically likely and plausible nontrophic interactions, based on competition for space and predator‐avoidance behaviour, were included. These findings call for a reassessment of the scale and nature of extinction cascades, particularly the inclusion of nontrophic interactions, in forecasts of the future of biodiversity.     

The form of direct interspecific competition modifies secondary extinction patterns in multi‐trophic food webs

Forcibly removing species from ecosystems has important consequences for the remaining assemblage, leading to changes in community structure, ecosystem functioning and secondary (cascading) extinctions. One key question that has arisen from single‐ and multi‐trophic ecosystem models is whether the secondary extinctions that occur within competitive communities (guilds) are also important in multi‐trophic ecosystems? The loss of consumer–resource links obviously causes secondary extinction of specialist consumers (topological extinctions), but the importance of secondary extinctions in multi‐trophic food webs driven by direct competitive exclusion remains unknown. Here I disentangle the effects of extinctions driven by basal competitive exclusion from those caused by trophic interactions in a multi‐trophic ecosystem (basal producers, intermediate and top consumers). I compared food webs where basal species either show diffuse (all species compete with each other identically: no within guild extinctions following primary extinction) or asymmetric competition (unequal interspecific competition: within guild extinctions are possible). Basal competitive exclusion drives extra extinction cascades across all trophic levels, with the effect amplified in larger ecosystems, though varying connectance has little impact on results. Secondary extinction patterns based on the relative abundance of the species lost in the primary extinction differ qualitatively between diffuse and asymmetric competition. Removing asymmetric basal species with low (high) abundance triggers fewer (more) secondary extinctions throughout the whole food web than removing diffuse basal species. Rare asymmetric competitors experience less pressure from consumers compared to rare diffuse competitors. Simulations revealed that diffuse basal species are never involved in extinction cascades, regardless of the trophic level of a primary extinction, while asymmetric competitors were. This work highlights important qualitative differences in extinction patterns that arise when different assumptions are made about the form of direct competition in multi‐trophic food webs.     

Ecological communities are vulnerable to realistic extinction sequences

Loss of species will directly change the structure and potentially the dynamics of ecological communities, which in turn may lead to additional species loss (secondary extinctions) due to direct and/or indirect effects (e.g. loss of resources or altered population dynamics). Furthermore, the vulnerability of food webs to repeated species loss is expected to be affected by food web topology, species interactions, as well as the order in which species go extinct. Species traits such as body size, abundance and connectivity might determine a species’ vulnerability to extinction and, thus, the order in which species go primarily extinct. Yet, the sequence of primary extinctions, and their effects on the vulnerability of food webs to secondary extinctions, when species abundances are allowed to respond dynamically, has only recently become the focus of attention. Here, we analyse and compare topological and dynamical robustness to secondary extinctions of model food webs, in the face of 34 extinction sequences based on species traits. Although secondary extinctions are frequent in the dynamical approach and rare in the topological approach, topological and dynamical robustness tends to be correlated for many bottom–up directed, but not for top–down directed deletion sequences. Furthermore, removing species based on traits that are strongly positively correlated to the trophic position of species (such as large body size, low abundance, high net effect) is, under the dynamical approach, found to be as destructive as removing primary producers. Such top–down oriented removal of species are often considered to correspond to realistic extinction scenarios, but earlier studies, based on topological approaches, have found such extinction sequences to have only moderate effects on the remaining community. Thus, our result suggests that the structure of ecological communities, and therefore the integrity of important ecosystem processes could be more vulnerable to realistic extinction sequences than previously believed.     

Using community viability analysis to identify fragile systems and keystone species

Owing to interdependences among species in ecological communities, the loss of one species can trigger a cascade of secondary extinctions with potentially dramatic effects on the functioning and stability of the community. It is, therefore, important to assess the risk and likely extent of secondary extinctions. Community viability analysis is a new technique that can be used to accomplish this goal. The analysis can also be used to identify fragile community structures and keystone species and, hence, to provide guidelines for conservation priorities. Here, we describe the principles underlying community viability analysis and review its contributions to our understanding of the response of ecological communities to species loss.     

The impacts of different management strategies and environmental forcing in ecological communities

Understanding the effects of population management on the community a target species belongs to is of key importance for successful management. It is known that the removal or extinction of a single species in a community may lead to extinctions of other community members. In our study, we assess the impacts of population management on competitive communities, studying the response of both locally stable and unstable communities of varying size (between four and 10 species) to three different management strategies; harvesting of a target species, harvesting with non-targeted catch, and stocking of the target species. We also studied the consequences of selecting target species with different relative abundances, as well as the effects of varying environmental conditions.We show here how the effects of management in competitive communities extend far beyond the target population. A crucial role is played by the underlying stability properties of the community under management. In general, locally unstable communities are more vulnerable to perturbation through management. Furthermore, the community response is shown to be sensitive to the relative density of the target species. Of considerable interest is the result that even a small (2.5%) increase in the population size of the target species through stocking may lead to extinction of other community members. These results emphasize the importance of considering and understanding multi-species interactions in population management.     

Re-assessing current extinction rates

There is a widespread belief that we are experiencing a mass extinction event similar in severity to previous mass extinction events in the last 600 million years where up to 95% of species disappeared. This paper reviews evidence for current extinctions and different methods of assessing extinction rates including species–area relationships and loss of tropical forests, changing threat status of species, co-extinction rates and modelling the impact of climate change. For 30 years some have suggested that extinctions through tropical forest loss are occurring at a rate of up to 100 species a day and yet less than 1,200 extinctions have been recorded in the last 400 years. Reasons for low number of identified global extinctions are suggested here and include success in protecting many endangered species, poor monitoring of most of the rest of species and their level of threat, extinction debt where forests have been lost but species still survive, that regrowth forests may be important in retaining ‘old growth’ species, fewer co-extinctions of species than expected, and large differences in the vulnerability of different taxa to extinction threats. More recently, others have suggested similar rates of extinction to earlier estimates but with the key cause of extinction being climate change, and in particular rising temperatures, rather than deforestation alone. Here I suggest that climate change, rather than deforestation is likely to bring about such high levels of extinction since the impacts of climate change are local to global and that climate change is acting synergistically with a range of other threats to biodiversity including deforestation.     

Habitat-islands in the coastal Atacama Desert: loss of functional redundancy,but not of functional diversity,with decreased precipitation

Background and AimsAridity is increasing in many regions of the world, but microclimatic conditions may buffer plant communities from the direct effects of decreased precipitation, creating habitat islands. However, reduced precipitation can also impact these communities indirectly by decreasing the suitability of the surrounding habitat, thus limiting incoming propagules and increasing the chances of population decline and species loss. We test whether decreased precipitation results in loss of species and functional diversity within habitat islands, evaluating in particular whether declines in species diversity and abundance are less likely to result in loss of functional diversity if species/individual loss is stochastic (i.e. independent of species/individual traits) and communities/populations are functionally redundant.MethodsLomas communities are discrete plant communities embedded in the Atacama Desert, maintained by the microclimatic conditions created by fog. We recorded species and functional diversity in six Lomas communities along a 500 km long precipitation gradient in northern Chile. Functional traits were measured in 20 individuals per species, in those species that accounted for approx. 75 % of the abundance at each site. We calculated functional diversity and functional redundancy of the community, and intraspecific functional variation.Key ResultsDecreased precipitation was associated with lower species diversity and lower species abundances. However, no traits or functional strategies increased or decreased consistently with precipitation, suggesting stochastic species/individual loss. Species with stress-tolerant strategies were predominant in all sites. Although species diversity decreased with decreasing precipitation, functional diversity remained unchanged. Lower functional redundancy in the drier sites suggests that mainly functionally redundant species were lost. Likewise, intraspecific functional variation was similar among communities, despite the lower species abundance in drier sites.ConclusionsDecreased precipitation can impact habitat island communities indirectly by decreasing the suitability of the surrounding habitat. Our results support the idea that a stochastic loss of species/individuals from functionally redundant communities and populations does not result in loss of functional diversity.     

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.     

Natural Shorelines Promote the Stability of Fish Communities in an Urbanized Coastal System

Habitat loss and fragmentation are leading causes of species extinctions in terrestrial, aquatic and marine systems. Along coastlines, natural habitats support high biodiversity and valuable ecosystem services but are often replaced with engineered structures for coastal protection or erosion control. We coupled high-resolution shoreline condition data with an eleven-year time series of fish community structure to examine how coastal protection structures impact community stability. Our analyses revealed that the most stable fish communities were nearest natural shorelines. Structurally complex engineered shorelines appeared to promote greater stability than simpler alternatives as communities nearest vertical walls, which are among the most prevalent structures, were most dissimilar from natural shorelines and had the lowest stability. We conclude that conserving and restoring natural habitats is essential for promoting ecological stability. However, in scenarios when natural habitats are not viable, engineered landscapes designed to mimic the complexity of natural habitats may provide similar ecological functions.     

贵州省普定县不同植被演替阶段的物种组成与群落结构特征

通过对贵州省普定县喀斯特地区不同植被演替阶段群落的调查, 研究了植被演替过程中群落物种组成和群落结构的变化。结果表明, 该地区的植被主要处于5个演替阶段, 即次生乔木林、乔灌过渡林、藤刺灌丛、稀灌草丛以及火烧干扰后的蕨类植物群落。本次调查共记录到植物365种, 隶属89科218属。其中, 蕨类植物31种, 隶属14科23属; 种子植物334种, 隶属75科195属。物种分布较多的科主要有蔷薇科、菊科、禾本科、百合科、忍冬科、唇形科、莎草科、樟科、葡萄科和水龙骨科。随着正向演替的推进, 物种丰富度增加, 群落结构趋于复杂化。藤刺灌丛与乔灌过渡林群落层次不明显, 次生乔木林分层明显。从藤刺灌丛向次生乔木林演替的过程中, 小径级个体所占比例明显降低, 高于1.3 m植物的总密度、乔木密度和藤本密度都先升高后降低, 而灌木密度呈逐渐降低的趋势。对喀斯特地区植被的恢复提出了参考措施。