海南岛霸王岭热带天然林景观中主要木本植物关键种的潜在分布

在较大的空间尺度上生态位模型是预测物种潜在分布的有效途径之一。为了探讨在热带天然林景观中木本植物(限于乔木和灌木)主要关键种的潜在分布,在对海南岛霸王岭的热带天然林进行按公里网格样方调查的基础上,采用演替地位和最大潜在高度两个功能性指标对物种进行了功能群划分,并在功能群框架下运用优势度指数法进行了关键种的确定;采用基于地理信息系统(Geographic information system, GIS)的基于规则集合预测的遗传算法(Algorithm for rule-set prediction, GARP)生态位模型对主要关键种的地理分布进行了预测,并应用受试者工作特征分析进行了模型精度验证;应用多元线性回归分析对影响各关键种潜在分布的关键因子进行了确定。结果表明:除了顶极次林层乔木功能群和顶极主林层乔木功能群外,在先锋种功能群、顶极灌木种功能群和顶极超冠层乔木功能群中采用优势度指数法划分出的关键种较为理想;一般来讲,在进行预测的8个关键种中,除了先锋主林层乔木种海南杨桐(Adinandra hainanensis),其它3个先锋种毛稔(Melastoma sanquiueum)、银柴(Aporosa chinensis)和枫香(Liquidambar formosana) 在研究区北部、西部以及西南部均具有较高的发生概率,而顶极种除了顶极超冠层乔木种南亚松(Pinus merkusii)外,九节(Psychotria rubra)、高脚罗伞(Ardisia quinquegona)和海南椎(Castanopsis hainanensis)具有相似的潜在分布格局,在研究区中部、东南部和南部地区具有较高的发生概率;相关分析表明极端最低温、年均温、极端最高温、年均降水量、海拔和坡向6大因子是影响研究区关键种潜在分布的关键因子;精度检验表明,GARP模型对8个关键种的潜在分布预测效果均较好,而其中又以银柴和海南椎的预测精度最高。     

Reduction and recovery of keystone predation pressure after disease‐related mass mortality

Disturbances such as disease can reshape communities through interruption of ecological interactions. Changes to population demographics alter how effectively a species performs its ecological role. While a population may recover in density, this may not translate to recovery of ecological function. In 2013, a sea star wasting syndrome outbreak caused mass mortality of the keystone predator Pisaster ochraceus on the North American Pacific coast. We analyzed sea star counts, biomass, size distributions, and recruitment from long‐term intertidal monitoring sites from San Diego to Alaska to assess regional trends in sea star recovery following the outbreak. Recruitment, an indicator of population recovery, has been spatially patchy and varied within and among regions of the coast. Despite sea star counts approaching predisease numbers, sea star biomass, a measure of predation potential on the mussel Mytilus californianus, has remained low. This indicates that post‐outbreak populations have not regained their full predation pressure. The regional variability in percent of recovering sites suggested differences in factors promoting sea star recovery between regions but did not show consistent patterns in postoutbreak recruitment on a coast‐wide scale. These results shape predictions of where changes in community composition are likely to occur in years following the disease outbreak and provide insight into how populations of keystone species resume their ecological roles following mortality‐inducing disturbances.     

Mistletoe as a keystone resource: an experimental test

Various entities have been designated keystone resources, but few tests have been attempted and we are unaware of any experimental manipulations of purported keystone resources. Mistletoes (Loranthaceae) provide structural and nutritional resources within canopies, and their pervasive influence on diversity led to their designation as keystone resources. We quantified the effect of mistletoe on diversity with a woodland-scale experiment, comparing bird diversities before and after all mistletoe plants were removed from 17 treatment sites, with those of 11 control sites and 12 sites in which mistletoe was naturally absent. Three years after mistletoe removal, treatment woodlands lost, on average, 20.9 per cent of their total species richness, 26.5 per cent of woodland-dependent bird species and 34.8 per cent of their woodland-dependent residents, compared with moderate increases in control sites and no significant changes in mistletoe-free sites. Treatment sites lost greater proportions of birds recorded nesting in mistletoe, but changes in species recorded feeding on mistletoe did not differ from control sites. Having confirmed the status of mistletoe as a keystone resource, we suggest that nutrient enrichment via litter-fall is the main mechanism promoting species richness, driving small-scale heterogeneity in productivity and food availability for woodland animals. This explanation applies to other parasitic plants with high turnover of enriched leaves, and the community-scale influence of these plants is most apparent in low productivity systems.     

Keystone Interactions: Salmon and Bear in Riparian Forests of Alaska

The term “keystone species” is used to describe organisms that exert a disproportionately important influence on the ecosystems in which they live. Analogous concepts such as “keystone mutualism” and “mobile links” illustrate how, in many cases, the interactions of two or more species produce an effect greater than that of any one species individually. Because of their role in transporting nutrients from the ocean to river and riparian ecosystems, Pacific salmon (Oncorhynchus spp.) and brown bear (Ursus arctos) have been described as keystone species and mobile links, although few data are available to quantify the importance of this interaction relative to other nutrient vectors. Application of a mass balance model to data from a southwestern Alaskan stream suggests that nitrogen (N) influx to the riparian forest is significantly increased in the presence of both salmon and bear, but not by either species individually. The interactions of salmon and bear may provide up to 24% of riparian N budgets, but this percentage varies in time and space according to variations in salmon escapement, channel morphology and watershed vegetation characteristics, suggesting interdependence and functional redundancy among N sources. These findings illustrate the complexity of interspecific interactions, the importance of linkages across ecosystem boundaries and the necessity of examining the processes and interactions that shape ecological communities, rather than their specific component parts.     

Identifying ‘climate keystone species’ as a tool for conserving ecological communities under climate change

Aim

Climate change affects ecological communities via impacts on species. The community's response to climate change can be represented as the temporal trend in a climate-related functional property that is quantified using a relevant functional trait. Noteworthy, some species influence this response in the community more strongly than others.

Innovation

Leveraging on the concept of keystone species, we propose that species with a strong effect on the community's functional response to climate change beyond their relative abundance can be considered as ‘climate keystone species’. We develop a stepwise tool to determine species' effects on a community's climate response and identify climate keystone species. We quantify the species-specific effect by measuring the difference in the community's climate response with and without the species. Next, we identify climate keystone species as those with a strong residual effect after weighting with their relative abundances in the community.

Main Conclusions

To illustrate the use of the stepwise tool with empirical data, we identify climate keystone species that have a strong effect on the change in the average temperature niche in North American bird communities over time and find the identification tool ecologically relevant. Identification of climate keystone species can serve as an additional conservation method to efficiently protect ecological communities and, in turn, the ecosystem functions they provide.     

Macrogeographic variation of a pond predator's top-down effects in a common garden environment

Geographic variation in species behavior and life history has been well documented in biology. Species with wide geographic distributions (i.e., across a continent) but small home ranges (i.e., <1 km²) likely experience wide variability in abiotic environments across the entirety of their range, possibly exhibiting strong local adaptation. Understanding variation across a large geographic scale is especially important when considering species that have strong ecological importance, such as keystone species. Yet, few studies have compared the potential cascading ecological effects of a predator with a keystone role in at least part of its range. To understand how keystone ability in pond food webs can vary across a large geographic range, we conducted an artificial pond experiment with a known keystone predator in at least part of its range, the marbled salamander (Ambystoma opacum). To do so, we collected size-matched salamander larvae from three geographically distant populations (>650 km apart) in Ohio, Mississippi, and North Carolina and placed them in mesocosms with a suite of spring breeding amphibian prey species. We observed differential survival of some prey species leading to differences in spring-breeding amphibian diversity among the three predator populations, indicating that keystone predation may vary at a geographic scale. Prey diversity was lowest with predators from northern (Ohio) populations of salamanders. Further understanding of large-scale variability in ecologically important predators and the potential effects of translocating wide-ranging ambystomatid species is needed to direct future conservation efforts and preserve biodiversity.     

The ecological importance of crocodylians: towards evidence‐based justification for their conservation

Large‐bodied predators are well represented among the world's threatened and endangered species. A significant body of literature shows that in terrestrial and marine ecosystems large predators can play important roles in ecosystem structure and functioning. By contrast, the ecological roles and importance of large predators within freshwater ecosystems are poorly understood, constraining the design and implementation of optimal conservation strategies for freshwater ecosystems. Conservationists and environmentalists frequently promulgate ecological roles that crocodylians are assumed to fulfil, but often with limited evidence supporting those claims. Here, we review the available information on the ecological importance of crocodylians, a widely distributed group of predominantly freshwater‐dwelling, large‐bodied predators. We synthesise information regarding the role of crocodylians under five criteria within the context of modern ecological concepts: as indicators of ecological health, as ecosystem engineers, apex predators, keystone species, and as contributors to nutrient and energy translocation across ecosystems. Some crocodylians play a role as indicators of ecosystem health, but this is largely untested across the order Crocodylia. By contrast, the role of crocodylian activities in ecosystem engineering is largely anecdotal, and information supporting their assumed role as apex predators is currently limited to only a few species. Whether crocodylians contribute significantly to nutrient and energy translocation through cross‐ecosystem movements is unknown. We conclude that most claims regarding the importance of crocodylians as apex predators, keystone species, ecosystem engineers, and as contributors to nutrient and energy translocation across ecosystems are mostly unsubstantiated speculation, drawn from anecdotal observations made during research carried out primarily for other purposes. There is a paucity of biological research targeted directly at: understanding population dynamics; trophic interactions within their ecological communities; and quantifying the short‐ and long‐term ecological impacts of crocodylian population declines, extirpations, and recoveries. Conservation practices ideally need evidence‐based planning, decision making and justification. Addressing the knowledge gaps identified here will be important for achieving effective conservation of crocodylians.     

Ecosystem‐level effects of keystone species reintroduction: a literature review

The keystone species concept was introduced in 1969 in reference to top‐down regulation of communities by predators, but has expanded to include myriad species at different trophic levels. Keystone species play disproportionately large, important roles in their ecosystems, but human‐wildlife conflicts often drive population declines. Population declines have resulted in the necessity of keystone species reintroduction; however, studies of such reintroductions are rare. We conducted a literature review and found only 30 peer‐reviewed journal articles that assessed reintroduced populations of keystone species, and only 11 of these assessed ecosystem‐level effects following reintroduction. Nine of 11 publications assessing ecosystem‐level effects found evidence of resumption of keystone roles; however, these publications focus on a narrow range of species. We highlight the deficit of peer‐reviewed literature on keystone species reintroductions, and draw attention to the need for assessment of ecosystem‐level effects so that the presence, extent, and rate of ecosystem restoration driven by keystone species can be better understood.     

Node centrality indices in food webs: Rank orders versus distributions

Network analysis examines the role of species in ecological communities. The most common approach involves measurement of centrality of species or other groups of individuals based on their topological positions in food webs, followed by establishing the rank order of importance of these groups. However, ranking may differ considerably with indices of centrality and therefore comparison of rank orders is essential to obtain more meaningful results on species performance. Since ranking ignores absolute differences between centrality values, species orders may neglect important structural information in food webs. Consequently, simultaneous examination of the distribution of index values is inevitable. Hierarchical clustering and consensus generation revealed that rank orders of centrality exhibit a similar pattern over six example food webs, while distributions differ not only with indices because their relationships are largely inconsistent with food webs as well. Therefore, optimal analysis of networks and the selection of keystone species in any ecological study should rely upon both of these procedures. Similar conclusions are drawn from the detailed evaluation of a sample food web from the Florida Bay.     

Keystone predators (eastern newts, Notophthalmus viridescens) reduce the impacts of an aquatic invasive species

Predation, competition, and their interaction are known to be important factors that influence the structure of ecological communities. In particular, in those cases where a competitive hierarchy exists among prey species, the presence of certain keystone predators can result in enhanced diversity in the prey community. However, little is known regarding the influence of keystone predator presence on invaded prey communities. Given the widespread occurrence of invasive species and substantial concern regarding their ecological impacts, studies on this topic are needed. In this study I used naturalistic replications of an experimental tadpole assemblage to assess the influence of predatory eastern newts, Notophthalmus viridescens, on the outcome of interspecific competition among native and nonindigenous tadpoles. When newts were absent, the presence of the tadpoles of one invasive species, the Cuban treefrog, Osteopilus septentrionalis, resulted in decreased survival and growth rate of the dominant native species, Bufo terrestris, and dominance of the tadpole assemblage by O. septentrionalis. However, the presence of one adult newt generally reduced or eliminated the negative impacts of O. septentrionalis tadpoles, resulting in comparable survival and performance of native species in invaded and noninvaded treatments. Differential mortality among the tadpole species suggests that newts preyed selectively on O. septentrionalis tadpoles, supporting the hypothesis that newts acted as keystone predators in the invaded assemblage. The presence of nonindigenous larval cane toads, Bufo marinus, did not significantly affect native species, and this species was not negatively affected by the presence of newts. Collectively, these results suggest that eastern newts significantly modified the competitive hierarchy of the invaded tadpole assemblage and reduced the impacts of a competitively superior invasive species. If general, these results suggest that the presence of certain species may be an essential factor regulating the ecological impacts of biological invasions.     

Identifying potential keystone species from field data – an example from temporary ponds

Identifying keystone species is essential for understanding community dynamics and preserving species richness. However, few studies have used quantitative, a priori methods to identify potential keystone species. Species known to act as keystones in North Carolina (NC) temporary ponds ( Notophthalmus viridescens , eastern newt, and Siren intermedia , lesser siren) were tested to see whether they played the same role in similar habitats in South Carolina (SC). Newts and sirens had no effect on anuran species richness in SC. Instead, another salamander ( Ambystoma talpoideum , mole salamander) absent from the NC ponds was identified as a strong keystone in SC. It functioned independently of environmental factors and the densities of other predators. Larval dragonflies ( Tramea carolina , Carolina saddlebag) were identified as weak, context-dependent keystones in SC, supporting anuran richness in isolated ponds with very low pH. The results suggest that the identity of keystone species varies, even in similar habitats within a physiographic region.     

The co-evolution of culturally inherited altruistic helping and cultural transmission under random group formation

Limited migration results in kin selective pressure on helping behaviors under a wide range of ecological, demographic and life-history situations. However, such genetically determined altruistic helping can evolve only when migration is not too strong and group size is not too large. Cultural inheritance of helping behaviors may allow altruistic helping to evolve in groups of larger size because cultural transmission has the potential to markedly decrease the variance within groups and augment the variance between groups. Here, we study the co-evolution of culturally inherited altruistic helping behaviors and two alternative cultural transmission rules for such behaviors. We find that conformist transmission, where individuals within groups tend to copy prevalent cultural variants (e.g., beliefs or values), has a strong adverse effect on the evolution of culturally inherited helping traits. This finding is at variance with the commonly held view that conformist transmission is a crucial factor favoring the evolution of altruistic helping in humans. By contrast, we find that under one-to-many transmission, where individuals within groups tend to copy a “leader” (or teacher), altruistic helping can evolve in groups of any size, although the cultural transmission rule itself hitchhikes rather weakly with a selected helping trait. Our results suggest that culturally determined helping behaviors are more likely to be driven by “leaders” than by popularity, but the emergence and stability of the cultural transmission rules themselves should be driven by some extrinsic factors.     

Neuroecology, chemical defense, and the keystone species concept

Neuroecology unifies principles from diverse disciplines, scaling from biophysical properties of nerve and muscle cells to community-wide impacts of trophic interactions. Here, these principles are used as a common fabric, woven from threads of chemosensory physiology, behavior, and population and community ecology. The "keystone species" concept, for example, is seminal in ecological theory. It defines a species whose impacts on communities are far greater than would be predicted from its relative abundance and biomass. Similarly, neurotoxins could function in keystone roles. They are rare within natural habitats but exert strong effects on species interactions at multiple trophic levels. Effects of two guanidine alkaloids, tetrodotoxin (TTX) and saxitoxin (STX), coalesce neurobiological and ecological perspectives. These molecules compose some of the most potent natural poisons ever described, and they are introduced into communities by one, or only a few, host species. Functioning as voltage-gated sodium channel blockers for nerve and muscle cells, TTX and STX serve in chemical defense. When borrowed by resistant consumer species, however, they are used either in chemical defense against higher order predators or for chemical communication as chemosensory excitants. Cascading effects of the compounds profoundly impact community-wide attributes, including species compositions and rates of material exchange. Thus, a diverse array of physiological traits, expressed differentially across many species, renders TTX and STX fully functional as keystone molecules, with vast ecological consequences at multiple trophic levels.     

Keystone species in seed dispersal networks are mainly determined by dietary specialization

A central issue in ecology is the definition and identification of keystone species, i.e. species that are relatively more important than others for maintaining community structure and ecosystem functioning. Network theory has been pointed out as a robust theoretical framework to enhance the operationality of the keystone species concept. We used the concept of centrality as a proxy for a species’ relative importance for the structure of seed dispersal networks composed of either frugivorous bats or birds and their food‐plants. Centrality was expected to be determined mainly by dietary specialization, but also by body mass and geographic range size. Across 15 Neotropical datasets, only specialized frugivore species reached the highest values of centrality. Furthermore, the centrality of specialized frugivores varied widely within and among networks, whereas that of secondary and opportunistic frugivores was consistently low. A mixed‐effects model showed that centrality was best explained by dietary specialization, but not by body mass or range size. Furthermore, the relationship between centrality and those three ecological correlates differed between bat– and bird–fruit networks. Our findings suggest that dietary specialization is key to understand what makes a frugivore species a keystone in seed dispersal networks, and that taxonomic identity also plays a significant role. Specialized frugivores may play a central role in network structuring and ecosystem functioning, which has important implications for conservation and restoration.     

生物入侵对鸟类的生态影响

生物入侵是全球生物多样性面临的最主要威胁之一, 入侵种在改变入侵地环境的同时也使当地的生物受到极大影响。鸟类在生态系统中处于较高的营养级, 生态系统中任何一个环节的变化都可能对鸟类造成一定的影响。本文回顾了哺乳动物、鸟类、无脊椎动物和植物等不同生物类群的入侵对本地鸟类生态影响方面的研究进展。外来生物对鸟类的影响主要表现在以下几方面: (1)外来哺乳动物对成鸟、幼鸟或鸟卵的捕食作用; (2)外来鸟类与本地鸟类竞争栖息地和食物资源, 与当地的近缘种杂交而造成基因流失; (3)外来无脊椎动物改变本地鸟类的栖息环境和食物状况, 甚至直接捕食本地鸟类; (4)外来植物入侵改变入侵地的植物群落组成和结构, 造成本地鸟类的栖息地丧失或破碎化, 并通过改变入侵地生态系统的食物链结构而对高营养级的鸟类产生影响。最后, 作者还提出了该领域有待解决的问题和今后可能的研究方向。     

Functional importance vs keystoneness: Reformulating some questions in theoretical biocenology

Abstract Two concepts relating to the influence of individual species on the biocenoses in which they occur are reviewed. The first, the general functional importance of a species, is denned as the sum, over all species, of the changes (sign ignored) in productivity which would occur on removal of the particular species from the biocenosis. General functional importance is calculated as: where P_j is the productivity of the jth species before (t= 0) and after (t= 1) removal of the particular (ith) species being evaluated. Though I_i values cannot be determined empirically, this concept raises provocative questions for theoretical biocenology. The second concept reviewed is that of the keystone species. Never having been precisely or operationally defined,‘keystone’ has come to mean little more than ‘important for something.’ Moreover, there is no empirical or theoretical foundation for the idea that there exists in any biocenosis a natural dichotomy corresponding to the verbal one of keystone and non-keystone species. Some investigators have implied that such a dichotomy is suggested by the frequency distributions of experimentally determined values of interaction strength. The patterns they refer to are, however, artifacts resulting from small sample sizes and the plotting of frequency distributions on arithmetic rather than logarithmic scales. As a casual metaphor ‘keystone species’ was appealing and harmless; but the pretence that it is a well-defined concept or phenomenon has had a stultifying effect on ecological thought and argument.     

A two level mutation-selection model of cultural evolution and diversity

Cultural evolution is a complex process that can happen at several levels. At the level of individuals in a population, each human bears a set of cultural traits that he or she can transmit to its offspring (vertical transmission) or to other members of his or her society (horizontal transmission). The relative frequency of a cultural trait in a population or society can thus increase or decrease with the relative reproductive success of its bearers (individual’s level) or the relative success of transmission (called the idea’s level). This article presents a mathematical model on the interplay between these two levels. The first aim of this article is to explore when cultural evolution is driven by the idea’s level, when it is driven by the individual’s level and when it is driven by both. These three possibilities are explored in relation to (a) the amount of interchange of cultural traits between individuals, (b) the selective pressure acting on individuals, (c) the rate of production of new cultural traits, (d) the individual’s capacity to remember cultural traits and to the population size. The aim is to explore the conditions in which cultural evolution does not lead to a better adaptation of individuals to the environment. This is to contrast the spread of fitness-enhancing ideas, which make individual bearers better adapted to the environment, to the spread of “selfish” ideas, which spread well simply because they are easy to remember but do not help their individual bearers (and may even hurt them). At the same time this article explores in which conditions the adaptation of individuals is maximal. The second aim is to explore how these factors affect cultural diversity, or the amount of different cultural traits in a population. This study suggests that a larger interchange of cultural traits between populations could lead to cultural evolution not improving the adaptation of individuals to their environment and to a decrease of cultural diversity.     

Mismatches between supply and demand in wildlife tourism: Insights for assessing cultural ecosystem services

Assessing cultural ecosystem services provided by biodiversity requires a combination of ecological and social approaches. In this study, we investigated the capacity of large African mammal species to provide the cultural ecosystem service of wildlife tourism by using a supply and demand framework. First, we tested the relationship between supply and demand for large mammal species in wildlife tourism. Second, we tested whether the trophic level and body size of mammals influenced the mismatch between supply and demand, and whether the patterns of mismatches were consistent among four protected areas (PAs) in three Southern African countries. To quantify supply of species, we counted large mammals along 196 five km road transects within the four PAs; to estimate demand, we gathered 651 face-to-face questionnaires of wildlife tourists and distinguished between their expectation and hope to see specific species. Results show that a higher supply of large mammal species increased the expectation to see a species (linear regression slope β = 0.28, p < 0.01), whereas supply did not affect the hopes to see a specific species (β = −0.04, p = 0.63). Analyses of mismatches revealed that predator species were more demanded in relation to their supply than ungulates. Finally, we found that the demands of wildlife tourists for mammal species in relation to their supply were consistent across the four PAs. Supply-demand analyses reveal that species’ traits, in particular trophic level, shape the hopes of wildlife tourists to see specific mammal species. We propose that the quantification of supply-demand mismatches can be used to identify charismatic species and relevant species’ traits, and can be applied for wildlife tourism assessments within as well as across regions. Supply-demand analyses provide a useful framework and deliver indicators for better assessing cultural ecosystem services involving wildlife and nature-based tourism, and can be used for conservation management.     

基于Ecopath模型的莱州湾中国对虾增殖生态容量

通过增殖放流,增加优质渔业资源、改善种群结构是渔业资源养护的重要手段,而增殖生态容量的研究是科学实施增殖放流的前提.本文根据2009-2010年的渔业资源与生态环境数据,构建了由26个功能群组成的莱州湾生态系统Ecopath模型,利用该模型分析了生态系统的总体特征、营养相互关系与关键种,计算了放流品种中国对虾的增殖生态容量.结果表明：系统的总初级生产量/总呼吸(TPP/TR)为1.53,总初级生产量/总生物量(TPP/B)为24.54,同时具有较低的循环指数(FCI=0.07)、较高的剩余生产量（434.41 t·km^-2·a^-1）和较低的系统连接指数(CI=0.29),该系统目前处于发育的早期阶段.中国对虾目前不是莱州湾生态系统的关键种,当前中国对虾的生物量为0.1143 t·km^-2,有较大的增殖潜力;当生物量增长25.8倍时,仍不会超过增殖生态容量2.9489 t·km^-2.     

Identifying keystone trophic groups in benthic ecosystems: Implications for fisheries management

Many species inhabiting the benthic marine ecosystems of the central and northern Chilean coast have been intensively harvested and this exploitation has increased considerably in recent years. Despite this harvest pressure, few studies have attempted to establish a more holistic, systems-based management plan. On the contrary, research continues to rely on population models in which the species of interest are isolated from their ecological context. This work offers several keystone indices in order to help multispecies fisheries management. The indices used are: (1) functional indices based on steady-state and dynamic trophic models; (2) structural indices based on bottom-up and top-down control mechanisms; and (3) qualitative keystone species indices using loop models (mixed control). The quantitative trophic models were constructed using Ecopath with Ecosim (EwE; v. 5.0) software, and the qualitative model was analysed using Loop Analysis. All models describe the interactions of the most representative species and functional groups inhabiting the benthic ecosystems of Tongoy Bay, La Rinconada Marine Reserve (Antofagasta Bay), and the kelp forest of Mejillones Peninsula (Antofagasta). Even though our results only represent the short-term dynamics of these systems, we have found keystoneness properties of several species and functional groups, including primary producers, herbivores, and top predators. Despite this wide variability of groups, we detected a different core set of species or functional groups, each of which contained prey–predator and plant–herbivore relationships. Because the traditional keystone concept of a single species is difficult to apply, we suggest shifting away from this view towards a more holistic alternative such as that of a keystone species complex. This kind of approach would facilitate the design and assessment of sustainable management strategies for ecological marine ecosystems. Despite the ecological relevance of our results, further experimental studies and modelling using other theoretical frameworks should be performed.