A near half‐century of temporal change in different facets of avian diversity

Assessments of spatial patterns of biodiversity change are essential to detect a signature of anthropogenic impacts, inform monitoring and conservation programs, and evaluate implications of biodiversity loss to humans. While taxonomic diversity (TD) is the most commonly assessed attribute of biodiversity, it misses the potential functional or phylogenetic implications of species losses or gains for ecosystems. Functional diversity (FD) and phylogenetic diversity (PD) are able to capture these important trait‐based and phylogenetic attributes of species, but their changes have to date only been evaluated over limited spatial and temporal extents. Employing a novel framework for addressing detectability, we here comprehensively assess a near half‐century of changes in local TD, FD, and PD of breeding birds across much of North America to examine levels of congruency in changes among these biodiversity facets and their variation across spatial and environmental gradients. Time‐series analysis showed significant and continuous increases in all three biodiversity attributes until ca. 2000, followed by a slow decline since. Comparison of avian diversity at the beginning and end of the temporal series revealed net increase in TD, FD, and PD, but changes in TD were larger than those in FD and PD, suggesting increasing biotic homogenization of avian assemblages throughout the United States. Changes were greatest at high elevations and latitudes – consistent with purported effects of ongoing climate change on biodiversity. Our findings highlight the potential of combining new types of data with novel statistical models to enable a more integrative monitoring and assessment of the multiple facets of biodiversity.     

Whether including exotic species alters conservation prioritization: a case study in the Min River in southeastern China

Conservation practices from the perspective of functional diversity (FD) and conservation prioritization need to account for the impacts of exotic species in freshwater ecosystems. This work first simulated the influence of exotic species on the values of FD in a schemed mechanistic model, and then a practical case study of conservation prioritization was performed in the Min River, the largest river in southeastern China, to discuss whether including exotic species alters prioritization. The mechanistic model revealed that exotic species significantly altered the expected FD if the number of exotic species occupied 2% of the community. Joint species distribution modelling indicated that the highest FD occurred in the west, northwest and north upstreams of the Min River. Values of FD in 64.69% of the basin decreased after the exotic species were removed from calculation. Conservation prioritization with the Zonation software proved that if first the habitats of exotic species were removed during prioritization, 62.75% of the highest prioritized areas were shifted, average species representation of the endemic species was improved and mean conservation efficiency was increased by 7.53%. Existence of exotic species will significantly alter the metrics of biodiversity and the solution for conservation prioritization, and negatively weighting exotic species in the scope of conservation prioritization is suggested to better protect endemic species. This work advocates a thorough estimate of the impacts of exotic species on FD and conservation prioritization, providing complementary evidence for conservation biology and valuable implications for local freshwater fish conservation.     

Impacts of logging and conversion of rainforest to oil palm on the functional diversity of birds in Sundaland

Commercial selective logging and the conversion of primary and degraded forests to agriculture are the biggest threats to tropical biodiversity. Our understanding of the impacts of these disturbances and the resulting local extinctions on the functional roles performed by the remaining species is limited. We address this issue by examining functional diversity (FD), which quantifies a range of traits that affect a species' ecological role in a community as a single continuous metric. We calculated FD for birds across a gradient of disturbance from primary forest through intensively logged forest to oil palm plantations on previously forested land in Borneo, Southeast Asia, a hotspot of imperilled biodiversity. Logged rainforest retained similar levels of FD to unlogged rainforest, even after two logging rotations, but the conversion of logged forest to oil palm resulted in dramatic reductions in FD. The few remaining species in oil palm filled a disproportionately wide range of functional roles but showed very little clustering in terms of functional traits, suggesting that any further extinctions from oil palm would reduce FD even further. Determining the extent to which the changes we recorded were due to under‐utilization of resources within oil palm or a reduction in the resources present is an important next step. Nonetheless our study improves our understanding of the stability and resilience of functional diversity in these ecosystems and of the implications of land‐use changes for ecosystem functioning.     

Projected impacts of climate warming on the functional and phylogenetic components of coastal Mediterranean fish biodiversity

Climate warming affects biodiversity distribution across all ecosystems. However, beyond changes in species richness, impacts on other biodiversity components are still overlooked, particularly in the marine realm. Here we forecasted the potential effect of climate warming on the phylogenetic and functional components of coastal Mediterranean fish biodiversity. To do so, we used species distribution models to project the potential distribution of 230 coastal fish species by the end of the 21st century based on the IPCC A2 scenario implemented with the Mediterranean climatic model NEMOMED8. From these projections, we assessed the changes in phylogenetic (PD) and functional diversity (FD) of fish assemblages at multiple spatial scales using a dated molecular phylogeny and an extensive functional trait database. At the scale of the entire Mediterranean Sea, the projected extinctions of 40 coastal fish species would lead to a concomitant erosion of PD and FD (13.6 and 3%, respectively). However, a null model revealed that species loss at this scale would not lead to a disproportionate erosion of PD and FD. Similar results were found when considering fish assemblages at the grid cell scale. Indeed, at this scale, the projected changes in species richness would lead to unexpected losses of PD and FD for localized and small areas only. A disproportionate erosion of PD under climate warming was only forecasted when analysing fish assemblages at an intermediate spatial scale, namely the Mediterranean marine ecoregions. Overall, our results emphasize the importance of considering multiple spatial scales when assessing potential impacts of climate warming on the multiple components marine biodiversity.     

Numerical abundance and biomass reveal different temporal trends of functional diversity change in tropical fish assemblages

Understanding how the biodiversity of freshwater fish assemblages changes over time is an important challenge. Until recently most emphasis has been on taxonomic diversity, but it is now clear that measures of functional diversity (FD) can shed new light on the mechanisms that underpin this temporal change. Fish biologists use different currencies, such as numerical abundance and biomass, to measure the abundance of fish species. Nonetheless, because they are not necessarily equivalent, these alternative currencies have the potential to reveal different insights into trends of FD in natural assemblages. In this study, the authors asked how conclusions about temporal trends in FD are influenced by the way in which the abundance of species has been quantified. To do this, the authors computed two informative metrics, for each currency, for 16 freshwater fish assemblages in Trinidad's Northern Range that had been surveyed repeatedly over 5 years. The authors found that numerical abundance and biomass uncover different directional trends in these assemblages for each facet of FD, and as such inform hypotheses about the ways in which these systems are being restructured. On the basis of these results, the authors concluded that a combined approach, in which both currencies are used, contributes to our understanding of the ecological processes that are involved in biodiversity change in freshwater fish assemblages.     

Biodiversity scenarios neglect future land‐use changes

Efficient management of biodiversity requires a forward‐looking approach based on scenarios that explore biodiversity changes under future environmental conditions. A number of ecological models have been proposed over the last decades to develop these biodiversity scenarios. Novel modelling approaches with strong theoretical foundation now offer the possibility to integrate key ecological and evolutionary processes that shape species distribution and community structure. Although biodiversity is affected by multiple threats, most studies addressing the effects of future environmental changes on biodiversity focus on a single threat only. We examined the studies published during the last 25 years that developed scenarios to predict future biodiversity changes based on climate, land‐use and land‐cover change projections. We found that biodiversity scenarios mostly focus on the future impacts of climate change and largely neglect changes in land use and land cover. The emphasis on climate change impacts has increased over time and has now reached a maximum. Yet, the direct destruction and degradation of habitats through land‐use and land‐cover changes are among the most significant and immediate threats to biodiversity. We argue that the current state of integration between ecological and land system sciences is leading to biased estimation of actual risks and therefore constrains the implementation of forward‐looking policy responses to biodiversity decline. We suggest research directions at the crossroads between ecological and environmental sciences to face the challenge of developing interoperable and plausible projections of future environmental changes and to anticipate the full range of their potential impacts on biodiversity. An intergovernmental platform is needed to stimulate such collaborative research efforts and to emphasize the societal and political relevance of taking up this challenge.     

Mismatches between birds' spatial and temporal dynamics reflect their delayed response to global changes

Global changes alter the dynamics of biodiversity, and are forecasted to continue or worsen in the decades to come. Modelling approaches used to anticipate these impacts are mainly based on the equivalence between spatial and temporal response to environmental forcings, generally called space-for-time substitution. However, several processes are known to generate deviations between spatial and temporal responses, potentially undermining the prediction based on space-for-time substitution. We here used high-resolution data from the french breeding bird survey to quantify and map the deviation between spatial and temporal patterns of bird abundances resulting from the dynamics of 124 species monitored in 2133 sites between 2001 and 2012. Using independent empirical data, we then tested specific predictions linked to the determinants (anthropogenic activities) and processes (lagged responses to environmental changes) potentially generating these deviations. We found that deviations between spatial and temporal patterns of abundances were particularly structured in space for bird communities. Following our predictions, these space–time deviations were positively correlated with the human influence on ecosystems, and linked with colonization–extinction ratios and community completeness, two markers of ongoing delayed responses to environmental changes. Our results suggest that the deviations between spatial and temporal patterns are related to recent anthropogenic environmental changes and disequilibrium responses to these changes. Investigating deviations between spatial and temporal patterns of biodiversity might open promising perspectives for a formal quantification of disequilibrium state of biodiversity at large spatial scale.     

Functional diversity changes during tropical forest succession

Functional diversity (FD) ‘those components of biodiversity that influence how an ecosystem operates or functions’ is a promising tool to assess the effect of biodiversity loss on ecosystem functioning. FD has received ample theoretical attention, but empirical studies are limited. We evaluate changes in species richness and FD during tropical secondary forest succession after shifting cultivation in Mexico. We also test whether species richness is a good predictor of FD. FD was calculated based on a combination of nine functional traits, and based on two individual traits important for primary production (specific leaf area) and carbon sequestration (wood density). Stand basal area was a good predictor of successional changes in diversity and FD, in contrast to fallow age. Incidence-based FD indices increased logarithmically with stand basal area, but FD weighted by species’ importance values lacked pattern with succession. Species richness and diversity are strong predictors of FD when all traits were considered; linear relationships indicate that all species are equally functionally complementary, suggesting there is little functional redundancy. In contrast, when FD was calculated for individual traits and weighted for abundances, species richness may underestimate FD.Selection of functional trait(s) critically determines FD, with large consequences for studies relating biodiversity to ecosystem functioning. Careful consideration of the traits required to capture the ecosystem process of interest is thus essential.     

Biodiversity–ecosystem function experiments reveal the mechanisms underlying the consequences of biodiversity change in real world ecosystems

In a recent Forum paper, Wardle (Journal of Vegetation Science, 2016) questions the value of biodiversity–ecosystem function (BEF) experiments with respect to their implications for biodiversity changes in real world communities. The main criticism is that the previous focus of BEF experiments on random species assemblages within each level of diversity has ‘limited the understanding of how natural communities respond to biodiversity loss.’ He concludes that a broader spectrum of approaches considering both non‐random gains and losses of diversity is essential to advance this field of research. Wardle's paper is timely because of recent observations of frequent local and regional biodiversity changes across ecosystems. While we appreciate that new and complementary experimental approaches are required for advancing the field, we question criticisms regarding the validity of BEF experiments. Therefore, we respond by briefly reiterating previous arguments emphasizing the reasoning behind random species composition in BEF experiments. We describe how BEF experiments have identified important mechanisms that play a role in real world ecosystems, advancing our understanding of ecosystem responses to species gains and losses. We discuss recent examples where theory derived from BEF experiments enriched our understanding of the consequences of biodiversity changes in real world ecosystems and where comprehensive analyses and integrative modelling approaches confirmed patterns found in BEF experiments. Finally, we provide some promising directions in BEF research.     

Multifaceted Analysis of Patch-Level Plant Diversity in Response to Landscape Spatial Pattern and History on Mediterranean Dunes

Landscape structure is known to critically affect biodiversity. However, although the multi-facetted character of biodiversity is widely recognized, few studies have linked landscape spatial pattern and history simultaneously to multiple facets (taxonomic, functional, and phylogenetic) and spatial components (α, β, and γ) of plant diversity. We set out to reveal whether landscape parameters have specific effects on the separate diversity facets and components of plant diversity at a patch scale on coastal dune landscapes of Central Italy. For each landscape patch, we computed a set of patch-based metrics relying on multi-temporal land-cover maps. Based on a database of plant community plots, on functional traits from field measurements and on a dated phylogenetic tree, we calculated taxonomic (TD), functional (FD), and phylogenetic diversity (PD) within each patch at α, β, and γ level. Diversity measures were then related to the landscape metrics via linear mixed-effect models. Landscape pattern and transformations affected TD only moderately in coastal dune ecosystems. We found much stronger and contrasted effects on FD and PD. FD increased in patches surrounded by human-dominated habitats; PD was higher in fragmented patches, particularly in the Mediterranean macchia. Moreover, landscape pattern affected differently the single communities, the turnover among communities and the pool of species within the patch (α, β, and γ components). Our results call for the combined inclusion of FD and PD and their partitions into ecological analyses, being TD too crude to capture the comprehensive and contrasted response of plant diversity to landscape spatial pattern.     

Functional diversity,functional composition and functional β diversity drive aboveground biomass across different bioclimatic rangelands

There has been a growing interest in understanding how biodiversity affects the functioning of ecosystems, while, it is still unclear how and to what extent different aspects of biodiversity are related to ecosystems function. We tested the relationship between aboveground biomass (AGB) and various aspects of functional diversity (functional richness (FRic), functional evenness (FEve) and functional divergence (FDiv)), functional β diversity and functional composition. Our study areas were located in four different bioclimatic regions including desert-steppe, steppe, semi-steppe, and Mediterranean rangelands extending along a productivity gradient created by the elevational-rainfall variations in the central Zagros Mountains of Iran.The results showed that FDiv positively predicted AGB in more productive (i.e., Mediterranean) and negatively in less productive rangelands of desert-steppe, steppe and semi-steppe. Functional β diversity predicted AGB only in more productive regions in semi-steppe and Mediterranean rangelands. Functional β diversity decomposed into turnover and nestedness, with turnover being more important in more productive and nestedness being more important in less productive ecosystems. The results also showed that the effects of FD and functional β diversity on AGB are generally directed by animal grazing through homogenizing community-wide trait composition in less productive ecosystems and inducing shifting mosaics across locations in more productive ecosystem.The results indicate that different components of FD were significantly related to AGB and the strength and direction of these effects vary with the different components studied and the bioclimatic regions under the study. In higher productive ecosystems, both functional diversity and functional β diversity are likely to contribute more to AGB through the complementary use of the resources. In less productive ecosystems, however, the functional identity of dominant species and traits would drive AGB.     

Scaling up the diversity–resilience relationship with trait databases and remote sensing data: the recovery of productivity after wildfire

Understanding the mechanisms underlying ecosystem resilience – why some systems have an irreversible response to disturbances while others recover – is critical for conserving biodiversity and ecosystem function in the face of global change. Despite the widespread acceptance of a positive relationship between biodiversity and resilience, empirical evidence for this relationship remains fairly limited in scope and localized in scale. Assessing resilience at the large landscape and regional scales most relevant to land management and conservation practices has been limited by the ability to measure both diversity and resilience over large spatial scales. Here, we combined tools used in large‐scale studies of biodiversity (remote sensing and trait databases) with theoretical advances developed from small‐scale experiments to ask whether the functional diversity within a range of woodland and forest ecosystems influences the recovery of productivity after wildfires across the four‐corner region of the United States. We additionally asked how environmental variation (topography, macroclimate) across this geographic region influences such resilience, either directly or indirectly via changes in functional diversity. Using path analysis, we found that functional diversity in regeneration traits (fire tolerance, fire resistance, resprout ability) was a stronger predictor of the recovery of productivity after wildfire than the functional diversity of seed mass or species richness. Moreover, slope, elevation, and aspect either directly or indirectly influenced the recovery of productivity, likely via their effect on microclimate, while macroclimate had no direct or indirect effects. Our study provides some of the first direct empirical evidence for functional diversity increasing resilience at large spatial scales. Our approach highlights the power of combining theory based on local‐scale studies with tools used in studies at large spatial scales and trait databases to understand pressing environmental issues.     

A review of dystrophic lake and pool habitat in Europe: An Irish perspective

Freshwater lakes and pools contained within peatlands are unique habitats that support rare and specialised species. Despite this, these ecosystems have been overlooked in conservation and management practices. One of these habitats, ‘3160 Natural dystrophic lakes and ponds’, is protected under the European Union (EU) Habitats Directive with a concerning proportion of these habitats having an “unfavourable-bad” or an “unfavourable-inadequate” conservation status across Europe. Our current understanding of the key physico-chemical and ecological features of this habitat is inadequate which is hindering the implementation of effective conservation measures. This review summarises the current knowledge of this protected lake habitat as defined under the EU Habitats Directive. With a focus on Ireland, we demonstrate how the current monitoring and assessment methods used to characterise and assess the structure and function and conservation status of this habitat, which relies largely on the use of macrophyte community composition and surrogate physico-chemical data collected under the EU Water Framework Directive, is ineffective. We propose the incorporation of further or alternative ecological metrics including, but not limited to, algae and macroinvertebrates which are needed to improve our understanding of the structure and function of this priority lake habitat. In addition, application of such data via ecological metrics would allow for the quantification of biodiversity and species rarity metrics which would aid in identifying sites of conservation importance.     

Impacts of invasion by <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas, 1771) on the performance of macroinvertebrate assessment tools for eutrophication pressure in lakes

Aquatic ecosystems are experiencing increasing disturbance from multiple stressors caused by anthropogenic activities. The potential for multiple stressors to modify each others’ impacts is not well understood. Legislation such as the EU Water Framework Directive (WFD) requires the development of tools to assess human impacts in aquatic systems that incorporate ecological elements, such as macroinvertebrates. Nutrient enrichment and invasive species are major threats to freshwater systems. The invasive zebra mussel Dreissena polymorpha is a conspicuous invader in freshwater aquatic systems in Europe and North America, and has been linked to drastic changes in macroinvertebrate communities and lake ecology. In 31 lake sites varying in nutrient pressure and in the presence or absence of D. polymorpha we tested three ecological quality assessment tools based on macroinvertebrate assemblages (% Sensitive Taxa to Total Phosphorus (TP), TP Score and Indicator Taxa Metric) and two basic ecological metrics. There were highly significant changes in macroinvertebrate diversity, structure, and composition associated with the invasion by D. polymorpha. While the three metrics performed consistently well in non-invaded systems, they lost explanatory power for eutrophication pressure in invaded systems. Our results suggest that metrics may need to be developed separately for invaded and non-invaded systems, and that the interaction between alien species and nutrient enrichment requires further investigation.     

Grasshopper （Orthoptera： Acrididae） biodiversity and grassland ecosystems

Interesting results may arise by combining studies on the structure and function of ecosystems with that of biodiversity for certain species. Grasshopper biodiversity is the result of the evolution of grassland ecosystems; however, it also impacts on the structure and the function of those ecosystems. We consider there to be a close relationship between the health of grassland ecosystems and grasshopper biodiversity. The main problems involved in this relationship are likely to include： （i） grasshopper biodiversity and its spatial pattern; （ii） the effect of grasshopper biodiversity on the ecological processes of grassland ecosystems; （iii） the biodiversity threshold of grasshopper population explosions; （iv） the relationship between grasshopper biodiversity and the natural and human factors that affect grassland ecosystems; and （v） grasshopper biodiversity and the health of grassland ecosystems. The solutions to these problems may provide sound bases for controlling disasters caused by grasshoppers and managing grassland ecosystems in the west of China. In this paper, we introduced two concepts for grasshopper biodiversity, that is, ＂spatial pattern＂ and ＂biodiversity threshold＂. It is helpful to understand the action of the spatial pattern of grasshopper biodiversity on the ecological processes of grassland ecosystems and the effect of this spatial pattern on the health of those ecosystems, owing to the fact that, in the west of China, grasslands are vast and grasshoppers are widely distributed. Moreover, we inferred that the change in the level of component richness at each type of grasshopper biodiversity can make an impact on grassland ecosystems, and therefore, there is likely to be a threshold to grasshopper biodiversity for the stability and the sustainability of those ecosystems.     

三江并流区树线生境小型兽类多样性多维度评价及其保护启示

三江并流区具有极高的环境异质性和丰富的生物多样性, 树线以上的高山生境是该地区的核心景观之一。本研究拟通过物种、系统发育和功能多样性对高山小型兽类多样性进行评价, 并从保护生物学的角度讨论生物多样性多维度评价的意义。2013年9月至2018年11月对三江并流区树线区域展开规范化野外调查, 收集小型兽类群落信息。从野外采集的组织样品中提取、扩增、测定线粒体CYTB基因序列, 构建小型兽类物种系统发育树。通过α多样性的不同维度以及功能冗余度评价了三江并流区树线生境小型兽类多样性。结果显示, 使用不同数据类型(出现/缺失数据和多度数据)的结果并不一致, 说明仅使用传统的丰富度指数对于描述多样性分布存在较大局限。物种和系统发育多样性在不同地点间的分布相似, 但功能多样性的分布与这两者有所差异, 从另一角度说明综合评价生物多样性的不同维度对于全面保护演化历史和生态功能的必要性。与高黎贡山、云岭相比较, 怒山山脉在5个样带尺度的多样性指数上呈现最高水平, 且具有最低的功能冗余度, 说明怒山的研究和保护力度均亟待加强。     

基于MODIS EVI时间序列的鄱阳湖湿地植被覆盖和生产力的动态变化

鄱阳湖是我国最大的淡水湖, 是与长江保持自由连通的两个湖泊之一, 也是最为重要的候鸟越冬地之一, 其生境质量对全球的生物多样性保护至关重要。枯水期的鄱阳湖由众多子湖构成, 不同子湖具有不同的水文控制与管理模式, 尤其是位于长江上游的三峡大坝2006年正式运行之后, 不同水文控制模式下的子湖展现出不同的退水机制, 对退水期洲滩出露的时间与湿生植被覆盖和生产力产生了不同的影响。近年来, 遥感和生态模型在研究植被变化中应用广泛。本文以MODIS增强植被指数(enhanced vegetation index, EVI)时间序列表示地表属性, 并利用EVI时间序列模型, 建立了2000-2014年植被覆盖和生产力的时空变化趋势。在研究区内建立的网格中, 随机提取了107个斑块, 采集其每16天间隔的MODIS EVI时间序列(2000年2月至2015年4月), 将自适应Savitzky-Golay平滑算法应用于EVI时间序列分析, 提取了4个关键的植被生长指标, 即生长季开始的日期、生长季长度、生长季EVI峰值和生产力。研究结果表明: (1)具有不同水文控制模式下的湿地植被生长特征表现出显著的差异, 尤其位于自由连通子湖的植被与其他模式的子湖相比: 生长季开始的时间更晚, 生长季较短, EVI峰值较低, 并且生长季节的初级生产力较低; (2)由于水文情势的改变, 自由连通子湖2006年前后的双生长周期湿地植被的生长特征差异明显, 秋季生长季提前, 导致了生物量的过度积累, 降低了越冬雁类食源的适口性; 但位于局部水文控制子湖的湿地植被不存在这种差异。(3)自由连通与局部水文控制的子湖对鄱阳湖越冬候鸟的保护均具有十分重要的意义, 需要保证这两种类型子湖的面积, 为越冬候鸟提供更广阔的食源; 当水文情势发生改变时, 局部的水文人为控制可在一定程度上减缓鄱阳湖水情变化对湿地植被生长带来的影响。     

Imperfect detection distorts depth‐related trends in marine macrofaunal species richness

Ecology and biodiversity research are underpinned by species richness patterns and their environmental drivers. However, a key topic in this discussion is the accuracy of these patterns which are greatly dependent on species detection probabilities. Due to variations in detection of species, true ecological patterns may be distorted. This is particularly true for subtidal macro‐infaunal communities. We tested three hypothesized relationships between marine benthic macrofaunal diversity and depth using species richness per site estimated with a capture–recapture heterogeneity model that accounts for variable detection probabilities. These metrics were based on data from 42 replicated sites across the continental shelf of the Southern Benguela. Average detection probability decreased with greater depth but species richness increased along the same depth gradient. The conflation of these trends in observed diversity data resulted in a positively near‐linear depth–diversity relationship, while accounting for variable species detection revealed a much stronger relationship. Ignoring species detection in ecosystems with imperfect detection could therefore distort species richness patterns, which has implications for ecological theory, management and conservation.     

Region‐wide changes in marine ecosystem dynamics: state‐space models to distinguish trends from step changes

Regime shifts are sudden changes in ecosystem structure that can be detected across several ecosystem components. The concept that regime shifts are common in marine ecosystems has gained popularity in recent years. Many studies have searched for the step‐like changes in ecosystem state expected under a simple interpretation of this idea. However, other kinds of change, such as pervasive trends, have often been ignored. We assembled over 300 ecological time series from seven UK marine regions, covering two to three decades. We developed state‐space models for the first principal component of the time series in each region, a common measure of ecosystem state. Our models allowed both trends and step changes, possibly in combination. We found trends in three of seven regions and step changes in two of seven regions. Gradual and sudden changes are therefore important trajectories to consider in marine ecosystems.