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.     

Macroscopic network properties and short-term dynamic simulations in coastal ecological systems at Fildes Bay (King George Island,Antarctica)

A trophic mass-balanced of the benthic/pelagic system dominated by large brown macroalgae in Fildes Bay (Antarctica) was constructed by integrating biomass, production, food spectrum, and consumption related information. The resulting trophic model was used to determine the macroscopic (emergent) properties, overall health and propagation of dynamical higher order effects within this complex Antarctic ecological system in response to simulated impacts. The magnitude of the Relative Ascendency, Relative Overhead, and Redundancy values indicates that the coastal benthic/pelagic Fildes Bay system is likely to remain less developed and therefore more resistant to perturbations than other ecological systems dominated by brown macroalgae. In terms of model component contributions to the Ascendency, detritus accounted for ∼33% of the value, followed by the phyto-zooplankton complex (∼26%), macroalgae (∼19%), filter-feeders (∼7%), small epifauna (5%), and top predators (2%). Short-term or transient Ecosim dynamical responses to increase the total mortality of each model component-given mixed and top-down vulnerabilities-revealed that changes in macroalgae levels had a limited impact on the other components of the system. The filter feeder, small epifauna and benthic fishés functional groups had the greatest effects on the remaining Fildes Bay system components. The magnitude of the System Recovery Time indicated that the Nacella concinna and small epifauna components would take the longest time to return to their initial state. Based on the outcomes obtained from the model, we suggest that this preliminary trophic model, including simulated impacts, provides promising possibilities for the determination of macroscopic baseline conditions and the most sensitive components of the Fildes Bay ecological system.     

Cascading effects of artificial light at night: resource-mediated control of herbivores in a grassland ecosystem

Artificial light at night has a wide range of biological effects on both plants and animals. Here, we review mechanisms by which artificial light at night may restructure ecological communities by modifying the interactions between species. Such mechanisms may be top-down (predator, parasite or grazer controlled), bottom-up (resource-controlled) or involve non-trophic processes, such as pollination, seed dispersal or competition. We present results from an experiment investigating both top-down and bottom-up effects of artificial light at night on the population density of pea aphids Acyrthosiphon pisum in a diverse artificial grassland community in the presence and absence of predators and under low-level light of different spectral composition. We found no evidence for top-down control of A. pisum in this system, but did find evidence for bottom-up effects mediated through the impact of light on flower head density in a leguminous food plant. These results suggest that physiological effects of light on a plant species within a diverse plant community can have detectable demographic effects on a specialist herbivore.     

Top-down and bottom-up control on cougar and its prey in a central Mexican natural reserve

Top-down and bottom-up controls are hypothesized to regulate population structures in many ecosystems. However, few studies have had the opportunity to analyze both processes in the natural environment, especially on large carnivores like the cougar (Puma concolor). Previously, studies show that cougar diet in the Sierra Nanchititla Natural Reserve (SNNR), central Mexico, is mainly armadillo, coati, and white-tailed deer. We assess whether top-down and/or bottom-up control regulate this endangered food web: (a) we predicted that seasonal per capita changes in abundance (pca) of cougar will be positively affected by the abundance of their main prey; (b) primary productivity in SNNR will affect the pca of prey species, driving bottom-up control; and (c) armadillo, coati, and white-tailed deer pca will be affected by the abundance of cougar, generating top-down control. Using 15 camera traps for 6 years in the SNNR, we calculated a relative abundance index (RAI) and pca for cougar and each of the focal prey, and we used the normalized difference vegetation index (NDVI) as a proxy of primary productivity. We constructed multiple regression models and selected the best linear models based on ranking the AIC_c values. Our analysis suggests that P. concolor pca is best explained by bottom-up control and intraspecific feedback. White-tailed deer and armadillo pca were both significantly affected by cougar abundance, indicating top-down control for these prey species, but NDVI was not retained in any of the models selected for prey pca. Our results indicate that both bottom-up and top-down control are involved in regulating this endangered food web in the SNNR, Mexico.     

Climate-induced changes in bottom-up and top-down processes independently alter a marine ecosystem

Climate change has complex structural impacts on coastal ecosystems. Global warming is linked to a widespread decline in body size, whereas increased flood frequency can amplify nutrient enrichment through enhanced run-off. Altered population body-size structure represents a disruption in top-down control, whereas eutrophication embodies a change in bottom-up forcing. These processes are typically studied in isolation and little is known about their potential interactive effects. Here, we present the results of an in situ experiment examining the combined effects of top-down and bottom-up forces on the structure of a coastal marine community. Reduced average body mass of the top predator (the shore crab, Carcinus maenas) and nutrient enrichment combined additively to alter mean community body mass. Nutrient enrichment increased species richness and overall density of organisms. Reduced top-predator body mass increased community biomass. Additionally, we found evidence for an allometrically induced trophic cascade. Here, the reduction in top-predator body mass enabled greater biomass of intermediate fish predators within the mesocosms. This, in turn, suppressed key micrograzers, which led to an overall increase in microalgal biomass. This response highlights the possibility for climate-induced trophic cascades, driven by altered size structure of populations, rather than species extinction.     

Inter-hemispheric comparison of bottom-up effects on community structure: Insights revealed using the comparative-experimental approach

The comparative-experimental approach uses identically designed, replicated experiments at different sites along environmental gradients in order to gain insight into the changing dynamics of communities with changing environmental conditions. Such studies reveal how ecological processes vary in intensity and interact to produce community structure. Early emphases were on the community consequences of shifting top-down impacts, competition and disturbance with environmental stress. Recent advances include the more precise quantification of gradients and thus a better understanding of species responses to the environment, and the revelation that bottom-up forces can vary significantly on within-region scales, with major consequences for the impact of top-down forces and thus community dynamics. Here the use of the method to examine the role of geographic location (coastal ecosystems in different hemispheres) and oceanographic conditions (upwelling vs downwelling) on these bottom-up/top-down linkages is advanced. We show that a bottom-up factor (prey recruitment) and a top-down effect (predation rate) vary consistently with oceanographic conditions within each coastal ecosystem, and also between geographic locations (New Zealand, Oregon). In general, both recruitment and predation rates are higher in Oregon. It is postulated that these differences are common responses to oceanographic variation, and that between-hemisphere differences result from the stronger and more persistent upwelling in the California Current ecosystem.     

Centrality measures and the importance of generalist species in pollination networks

Studies of complex networks show that nodes with high centrality scores are important to network structure and stability. Following this rationale, centrality measures can be used to (i) identify keystone species in ecological networks, a major issue in community ecology, and (ii) differentiate the keystone species concept, e.g. species may play a key role in a network for different topological reasons. In 34 pollination communities we examine the relationship between the generalization level of species (ND) and two complementary centrality indices: closeness (CC) and betweenness centrality (BC). CC measures the proximity of a species to all other species in the community, while BC describes the importance of a species as a connector. Most networks had a linear ND–CC relationship with a minimum CC value of 0.41. Hence, species were close to each and will be likely to be rapidly affected by disturbances. Contrarily, in most networks, the ND–BC relationships were power-law distributed with exponents larger than one. Only 59% of the species were connectors (BC > 0). In particular, there was a connector threshold value of ND = 0.46. Species above this threshold represent ～40%, almost all of which were connectors. These results indicate that in pollination systems the most generalized species are usually network keystone species, playing at least two roles: (i) interact closely with most other species (high CC) and (ii) connect otherwise unconnected subnetworks (high BC). We discuss the implications of centrality measures to community-based conservation ecology.     

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.     

海州湾鱼类群落功能多样性的时空变化

功能多样性是联系物种、生态环境和生态系统功能的基础.从功能多样性的角度研究群落结构,有助于更好地理解生物多样性与生态系统功能之间的关系.本研究根据2011—2017年(除2012年外)春、秋季海州湾渔业资源底拖网调查数据,选取反映鱼类摄食、运动、生态适应性、繁殖行为和种群动态特征的13个功能性状,利用功能丰富度指数、功能均匀度指数、功能离散度指数和群落特征加权平均数指数(CWM),研究了海州湾鱼类群落功能多样性的季节、年际和空间的变化.结果表明: 海州湾鱼类群落功能多样性指数具有显著的季节差异,其中秋季功能丰富度指数显著高于春季,春季功能离散度指数显著高于秋季,鱼类洄游是造成功能多样性指数季节变化的主要原因;CWM分析表明,春季鱼类群落优势种为营养级较高、运动能力较强、生长系数、恢复力和脆弱性较高的冷/暖温性鱼类,秋季则相反,春、秋季鱼类群落均以产浮性卵鱼类为主;鱼类群落功能多样性指数呈现一定的年际变化,其中春、秋季各功能多样性指数在不同年际均呈波动趋势,表明海州湾鱼类群落稳定性较低;鱼类群落功能多样性指数呈现显著的空间差异,其中20 m以深远岸水域功能离散度指数较高.海州湾鱼类群落功能多样性具有明显的时空变化特征,鱼类生态位与其对资源的利用均随季节、年际和空间而变化.     

Terrestrial food web of the Malpelo Fauna and Flora Sanctuary,Colombia: An analysis from a topological approach

Malpelo Island is the largest marine protected area (MPA) in the Colombian Pacific; however, the lack of information regarding its ecological dynamics suggests that management and conservation strategies are developed from an individual approach (species or groups of species) and not from an ecosystem one. This study analyzes the terrestrial food web of Malpelo Island, Colombia, based on topological analysis (e.g., centrality). The food web was constructed from 27 nodes representing the main trophogroups, which consist of species or functional groups. Malpelo Island had a food web of four steps with a maximum separation among all trophogroups and trophic pathways, with two steps in average length. Furthermore, the food web was divided into three food web compartments, with a recurrence of connectivity patterns dominated by apparent and exploitative competition, followed by tri-trophic chains and omnivory. Five key trophogroups control the energy flow throughout the food web (detritus, the land crab Johngarthia malpilensis, the lizard Anolis agassizi, the Malpelo dotted galliwasp Diploglossus millepunctatus, and the Nazca booby Sula granti). The high importance of detritus suggests that bottom-up processes act as a control and regulation mechanism of trophic flows. The low number of food web compartments and a high recurrence of specific connectivity patterns in the Malpelo Island terrestrial ecosystem evidence different ecological processes centered on five trophogroups, allowing stability against disturbances. In addition, the simulation of trophogroup removal (randomly or directed) suggests that food web can be vulnerable to structural alterations in their properties, which may have consequences on the resilience of this ecosystem. This study contributes to the knowledge of the trophic dynamics of Malpelo Island, providing a potential tool for management and conservation measures from an ecosystemic approach.     

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.     

Microscale methods to assemble mammalian cells into tissue-like structures

Different cell types make up tissues and organs hierarchically and communicate within a complex, three-dimensional (3D) environment. The in vitro recapitulation of tissue-like structures is meaningful, not only for fundamental cell biology research, but also for tissue engineering (TE). Currently, TE research adopts either the top-down or bottom-up approach. The top-down approach involves defining the macroscopic tissue features using biomaterial scaffolds and seeding cells into these scaffolds. Conversely, the bottom-up approach aims at crafting small tissue building blocks with precision-engineered structural and functional microscale features, using physical and/or chemical approaches. The bottom-up strategy takes advantage of the repeating structural and functional units that facilitate cell-cell interactions and cultures multiple cells together as a functional unit of tissue. In this review, we focus on currently available microscale methods that can control mammalian cells to assemble into 3D tissue-like structures.     

Deterministic diversity changes in freshwater phytoplankton in the Yunnan–Guizhou Plateau lakes in China

Diversity measures reflect different aspects of a community, which are determined by different ecological processes. However, information is still limited on the ecological processes that are represented by different measures of species diversity. In this study, the primary driving factors for richness and diversity indices were tested. The possible ecological processes represented by each index were analyzed. First, the type of ecological process that governed the phytoplankton community in the Yunnan–Guizhou Plateau lakes, either deterministic or stochastic, was identified by Caswell's neutral model. The results indicate that a deterministic process governs the phytoplankton community. Second, the driving factors of richness and diversity indices were screened with mixed models. The results suggest that the variation of phytoplankton richness in different lakes or sites was primarily related to bottom-up factors. The variations in evenness and other measures based on the relative abundance were driven by both top-down and bottom-up factors, such as zooplankton biomass, and pH and mean light, respectively. Finally, although the different measures of diversity may respond to specific bottom-up or top-down processes, the responses to the two processes were not independent of each other. These findings will increase our understanding of the relationships between ecological processes and diversity measures for freshwater phytoplankton.     

胶州湾近岸浅水区鱼类群落结构及多样性

根据2009年3月-2010年2月在胶州湾西北部近岸浅水区进行的鱼类逐月定置网调查数据,采用生态多样性指数和多元统计分析等方法研究了该海域的鱼类群落结构及多样性特征.结果表明:调查共鉴定出44种鱼类,其中海洋性27种,河口性鱼类17种.鱼类种类组成与底层水温和盐度有密切关系.胶州湾近岸浅水区鱼类群落Margalef种类丰富度指数(R)变化范围为0.33-3.50,Shannon-Wiener多样性指数(H’)为0.42-2.25,Pielou均匀度指数(J’)为0.23-0.93.聚类分析(CLUSTER)和多维标度排序(MDS)分析表明,不同鱼类种类组成的各月份样品可划分为3个组分:水温较低的冬季组G1(12月-翌年4月)、水温次高的春秋季组G2(5-6月、10-11月)和水温最高的夏季组G3(7-9月).其中G1的典型种主要为尖海龙(Syngnathus acus)、方氏云鳚(Enedrias fangi)和玉筋鱼(Ammodytes personatus)等冷温性常栖类群种类,G2的主要典型种为鳀鱼(Engraulis japonicus)和尖海龙,G3的主要典型种为鲐鱼(Scomber japonicus)、细条天竺鱼(Apogonichthys lineatus)和普氏缰虾虎鱼(Amoya pflaumi)等洄游性鱼类.单因子相似性分析(ANOSIM)表明,3个月份组间鱼类群落结构差异显著,不同月份组间两两差异均显著.生物-环境分析(BIOENV)表明,底层水温是影响胶州湾近岸浅水区鱼类群落结构时间变化的主要环境因子.在胶州湾内近岸浅水区发现40余种幼鱼,但占优势的主要为低值鱼类,表明胶州湾的产卵、肥育场功能有所下降,应加强胶州湾近岸水域生态环境和渔业资源的保护及修复.     

The impact of human trampling on a threatened coastal Mediterranean plant: The case of Anchusa littorea Moris (Boraginaceae)

In a 5-period study (2007–2011) we examined the effects of human trampling on the last remaining population of Anchusa littorea on its sandy coastal habitat in Sardinia. This species, considered extinct in the wild for several years, was casually rediscovered in a small population at Is Arenas (SW Sardinia). We monthly monitored six trampled and six un-trampled permanent plots, mapping individuals and recording their size and reproductive variables. Trampled and un-trampled plots showed significant differences with respect to plant density, plant size and reproductive performance (flowers and fruits production) of A. littorea. This study demonstrated that human trampling represents a severe threat to the conservation of this species that can be appreciated as a keystone species with concern to human trampling effects on coastal dune plants. In case of A. littorea urgent measures should be undertaken to protect this unique remnant population in the dune system of Is Arenas. In particular, tourist paths should be redirected and confined to others areas in order to promote the natural expansion of A. littorea in its original habitat. A possible integrated strategy for the conservation and management of the species consists in a combination of in situ and ex situ measures.     

海岸带区域陆海统筹生态安全研究

海岸带既是区域经济发展的黄金地带,亦是陆海系统交互胁迫的敏感区域,快速城镇化与工业化导致海岸带关键生态系统服务受损、生态网络格局紊乱、珍稀物种安全保障空间萎缩等重大生态安全问题。以闽三角海岸带珍稀物种-栖息地系统为研究对象,从“多物种-多生境-多尺度”视角出发,基于陆海统筹理念系统评估了闽三角海岸带珍稀物种-栖息地系统的脆弱性,建立了基于物种水平的海岸带生态保护与修复网络,构建了海岸带陆海统筹生态安全“一张图”。研究结果表明：(1)高强度围填海驱动下,闽三角红树林与水鸟栖息地脆弱性显著增加,高脆弱区域主要集中于九龙江河口、厦门同安湾和泉州湾;(2)闽三角生态安全网络由12条水鸟迁徙廊道和11个生态节点组成,其中影响陆海统筹生态安全的生态节点4个,预警生态节点3个;(3)闽三角海岸带未来安全韧性生境(水鸟栖息地与文昌鱼、中国鲎低脆弱生境较密集的区域)分布于云霄红树林保护区、东山岛近岸、九龙江河口、厦门大嶝海域、围头湾、深沪湾。本研究能够为设计规划未来韧性海岸带“受损海水-滨海湿地-珍稀物种”一体化修复与保护的生态安全格局,保障陆海统筹生态安全提供理论支撑和一体化解决方案。     

Adaptive genetic variation mediates bottom-up and top-down control in an aquatic ecosystem

Research in eco-evolutionary dynamics and community genetics has demonstrated that variation within a species can have strong impacts on associated communities and ecosystem processes. Yet, these studies have centred around individual focal species and at single trophic levels, ignoring the role of phenotypic variation in multiple taxa within an ecosystem. Given the ubiquitous nature of local adaptation, and thus intraspecific variation, we sought to understand how combinations of intraspecific variation in multiple species within an ecosystem impacts its ecology. Using two species that co-occur and demonstrate adaptation to their natal environments, black cottonwood (Populus trichocarpa) and three-spined stickleback (Gasterosteus aculeatus), we investigated the effects of intraspecific phenotypic variation on both top-down and bottom-up forces using a large-scale aquatic mesocosm experiment. Black cottonwood genotypes exhibit genetic variation in their productivity and consequently their leaf litter subsidies to the aquatic system, which mediates the strength of top-down effects from stickleback on prey abundances. Abundances of four common invertebrate prey species and available phosphorous, the most critically limiting nutrient in freshwater systems, are dictated by the interaction between genetic variation in cottonwood productivity and stickleback morphology. These interactive effects fit with ecological theory on the relationship between productivity and top-down control and are comparable in strength to the effects of predator addition. Our results illustrate that intraspecific variation, which can evolve rapidly, is an under-appreciated driver of community structure and ecosystem function, demonstrating that a multi-trophic perspective is essential to understanding the role of evolution in structuring ecological patterns.     

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。