Ecological and conservation insights from reconstructive studies of temperate old-growth forests

Reconstructive studies that use paleoecological, dendroecological, historical and other approaches in order to interpret long-term ecosystem dynamics are increasingly generating valuable insights for ecologists, conservationists and foresters who are interested in the ecology, protection and management of old-growth forests. In most cases, the historical context provided by these studies reveals a long-term pattern of change that challenges assumptions about the pristine condition of these systems. Ironically, it is the history of environmental fluctuations, natural disturbance processes, and subtle, often indirect, human impacts that is revealed by reconstructive work that may shape the characteristic structure, composition and ecosystem processes of old-growth forests, and that will certainly provide the greatest challenge to their future conservation and management.     

Linking macroecology and community ecology: refining predictions of species distributions using biotic interaction networks

Macroecological models for predicting species distributions usually only include abiotic environmental conditions as explanatory variables, despite knowledge from community ecology that all species are linked to other species through biotic interactions. This disconnect is largely due to the different spatial scales considered by the two sub‐disciplines: macroecologists study patterns at large extents and coarse resolutions, while community ecologists focus on small extents and fine resolutions. A general framework for including biotic interactions in macroecological models would help bridge this divide, as it would allow for rigorous testing of the role that biotic interactions play in determining species ranges. Here, we present an approach that combines species distribution models with Bayesian networks, which enables the direct and indirect effects of biotic interactions to be modelled as propagating conditional dependencies among species’ presences. We show that including biotic interactions in distribution models for species from a California grassland community results in better range predictions across the western USA. This new approach will be important for improving estimates of species distributions and their dynamics under environmental change.     

Ecosystem functioning in a disturbance-recovery context: Contribution of macrofauna to primary production and nutrient release on intertidal sandflats

Although many ecologists have described relationships between biodiversity and ecosystem functioning, changes in functioning after disturbance and during recovery are not well documented. Diversity-functioning theory predicts a decline in functioning with decreased diversity (e.g., following disturbance) and a gradual increase in functioning coincident with biotic recovery. We tested this theory empirically by measuring primary production and nutrient release (indices of sandflat functioning) at three sites at the beginning, middle and end of a year-long recovery period. Although microphytobenthos stock (sediment chlorophyll a content) recovered quickly, rates of gross photosynthetic oxygen production were reduced in disturbed plots, relative to controls, in 7 of 9 comparisons. Plots with high macrofaunal richness had enhanced ammonium efflux, and greater efflux of ammonium likely increased rates of primary production. The site with the highest bioturbation potential, which was dominated by large bivalves, exhibited the largest decline in functioning immediately after disturbance. This site also had the most persistent treatment-control differences during recovery, probably because the large bivalves remained at low abundance in treated plots throughout the year-long experiment.     

The bog climax hypothesis: fossil arthropod and stratigraphic evidence in peat sections from southeast Alaska, USA

The hypothesis that bogs are climax (structurally and compositionally stable) communities has been proposed for several large regions including southeast Alaska. A logical consequence of the bog climax is the hypothesis that, where there is infrequent large-scale physical disturbance, landscape-level changes from early successional woodland to late successional peatland should have occurred during the Holocene. This hypothesis is tested in southeast Alaska using analyses of fossil arthropods and site stratigraphy, as well as a review of the ecological literature.
Results indicate at least three major landscape types have dominated since deglaciation: deciduous woodland (about 9000 to 6000 yr BP), coniferous woodland (about 6000 to 3000 yr BP), and peatland (about 3000 yr BP to present). Other paleoecological and ecological studies from around the region provide evidence for directional changes, both past and present, from woodland to peatland. Furthermore, there is no clear evidence for peatland to woodland transitions in the absence of large-scale disturbance. These results suggest that the bog climax hypothesis deserves a more careful examination by ecologists studying succession.     

Disturbance and vegetation response in relation to environmental gradients in the Great Smoky Mountains

In constructing models of species and community distributions along environmental gradients in the Great Smoky Mountains, R. H. Whittaker (1956) focused on old-aged, apparently stable, natural communities. More recent studies indicate that disturbance gradients potentially influence and are influenced by the complex environmental gradients of Whittaker's original models. Using primarily fire and exotic species invasion as examples, this paper shows: 1) disturbance parameters vary along the topographic, elevation and moisture gradients in the Great Smoky Mountains in much the same way as temperature, moisture and solar radiation change; 2) species composition at different locations along the major environmental gradients is partially determined by the disturbance parameter; 3) species characteristics such as mode of reproduction are often correlated with specific disturbance parameters; 4) functional aspects of ecosystem response to disturbance vary along environmental gradients; and 5) man-caused disturbance may vary along environmental or biotic gradients. Since disturbance gradients may parallel physical environmental gradients, the two may be difficult to distinguish. Modification of disturbance frequencies along major environmental gradients may result in slow shifts in the distribution of both individual species and whole communities.Botanical nomenclature follows Radford et al. (1968).     

A niche for ecosystem multifunctionality in global change research

Concern about human modification of Earth's ecosystems has recently motivated ecologists to address how global change drivers will impact the simultaneous provisioning of multiple functions, termed ecosystem multifunctionality (EMF). However, metrics of EMF have often been applied in global change studies with little consideration of the information they provide beyond single functions, or how and why EMF may respond to global change drivers. Here, we critically review the current state of this rapidly expanding field and provide a conceptual framework to guide the effective incorporation of EMF in global change research. In particular, we emphasize the need for a priori identification and explicit testing of the biotic and abiotic mechanisms through which global change drivers impact EMF, as well as assessing correlations among multiple single functions because these patterns underlie shifts in EMF. While the role of biodiversity in mediating global change effects on EMF has justifiably received much attention, empirical support for effects via other biotic and physicochemical mechanisms are also needed. Studies also frequently stated the importance of measuring EMF responses to global change drivers to understand the potential consequences for multiple ecosystem services, but explicit links between measured functions and ecosystem services were missing from many such studies. While there is clear potential for EMF to provide novel insights to global change research, predictive understanding will be greatly improved by insuring future research is strongly hypothesis‐driven, is designed to explicitly test multiple abiotic and biotic mechanisms, and assesses how single functions and their covariation drive emergent EMF responses to global change drivers.     

Dispersal and neutral sampling mediate contingent effects of disturbance on plant beta‐diversity: a meta‐analysis

A major challenge in ecology, conservation and global‐change biology is to understand why biodiversity responds differently to similar environmental changes. Contingent biodiversity responses may depend on how disturbance and dispersal interact to alter variation in community composition (β‐diversity) and assembly mechanisms. However, quantitative syntheses of these patterns and processes across studies are lacking. Using null‐models and meta‐analyses of 22 factorial experiments in herbaceous plant communities across Europe and North America, we show that disturbance diversifies communities when dispersal is limited, but homogenises communities when combined with increased immigration from the species pool. In contrast to the hypothesis that disturbance and dispersal mediate the strength of niche assembly, both processes altered β‐diversity through neutral‐sampling effects on numbers of individuals and species in communities. Our synthesis suggests that stochastic effects of disturbance and dispersal on community assembly play an important, but underappreciated, role in mediating biotic homogenisation and biodiversity responses to environmental change.     

陆地植物群落物种多样性维持机制

从空间尺度和特定生物区系两个尺度对物种多样性的维持机制进行了综述.在大的空间尺度,简述了引起物种多样性存在差异的物理和自然因子的作用,包括植物群落的历史和年龄、梯度变化(纬度梯度、水分梯度、海拔梯度、土壤养分梯度)、面积效应和隔离程度;针对特定生物区系,从生物因素(生产力、种间关系、林隙动态)和非生物因素(演替、干扰及空间异质性、人类活动)方面论述其与物种多样性之间的关系.     

Biotic homogenisation and differentiation as directional change in beta diversity: synthesising driver–response relationships to develop conceptual models across ecosystems

Biotic homogenisation is defined as decreasing dissimilarity among ecological assemblages sampled within a given spatial area over time. Biotic differentiation, in turn, is defined as increasing dissimilarity over time. Overall, changes in the spatial dissimilarities among assemblages (termed ‘beta diversity’) is an increasingly recognised feature of broader biodiversity change in the Anthropocene. Empirical evidence of biotic homogenisation and biotic differentiation remains scattered across different ecosystems. Most meta-analyses quantify the prevalence and direction of change in beta diversity, rather than attempting to identify underlying ecological drivers of such changes. By conceptualising the mechanisms that contribute to decreasing or increasing dissimilarity in the composition of ecological assemblages across space, environmental managers and conservation practitioners can make informed decisions about what interventions may be required to sustain biodiversity and can predict potential biodiversity outcomes of future disturbances. We systematically reviewed and synthesised published empirical evidence for ecological drivers of biotic homogenisation and differentiation across terrestrial, marine, and freshwater realms to derive conceptual models that explain changes in spatial beta diversity. We pursued five key themes in our review: (i) temporal environmental change; (ii) disturbance regime; (iii) connectivity alteration and species redistribution; (iv) habitat change; and (v) biotic and trophic interactions. Our first conceptual model highlights how biotic homogenisation and differentiation can occur as a function of changes in local (alpha) diversity or regional (gamma) diversity, independently of species invasions and losses due to changes in species occurrence among assemblages. Second, the direction and magnitude of change in beta diversity depends on the interaction between spatial variation (patchiness) and temporal variation (synchronicity) of disturbance events. Third, in the context of connectivity and species redistribution, divergent beta diversity outcomes occur as different species have different dispersal characteristics, and the magnitude of beta diversity change associated with species invasions also depends strongly on alpha and gamma diversity prior to species invasion. Fourth, beta diversity is positively linked with spatial environmental variability, such that biotic homogenisation and differentiation occur when environmental heterogeneity decreases or increases, respectively. Fifth, species interactions can influence beta diversity via habitat modification, disease, consumption (trophic dynamics), competition, and by altering ecosystem productivity. Our synthesis highlights the multitude of mechanisms that cause assemblages to be more or less spatially similar in composition (taxonomically, functionally, phylogenetically) through time. We consider that future studies should aim to enhance our collective understanding of ecological systems by clarifying the underlying mechanisms driving homogenisation or differentiation, rather than focusing only on reporting the prevalence and direction of change in beta diversity, per se.     

Scaling of the mean and variance of population dynamics under fluctuating regimes

Theoretical ecologists have long sought to understand how the persistence of populations depends on the interactions between exogenous (biotic and abiotic) and endogenous (e.g., demographic and genetic) drivers of population dynamics. Recent work focuses on the autocorrelation structure of environmental perturbations and its effects on the persistence of populations. Accurate estimation of extinction times and especially determination of the mechanisms affecting extinction times is important for biodiversity conservation. Here we examine the interaction between environmental fluctuations and the scaling effect of the mean population size with its variance. We investigate how interactions between environmental and demographic stochasticity can affect the mean time to extinction, change optimal patch size dynamics, and how it can alter the often-assumed linear relationship between the census size and the effective population size. The importance of the correlation between environmental and demographic variation depends on the relative importance of the two types of variation. We found the correlation to be important when the two types of variation were approximately equal; however, the importance of the correlation diminishes as one source of variation dominates. The implications of these findings are discussed from a conservation and eco-evolutionary point of view.     

生态学系统的空间异质性

空间异质性是生态学系统的一个普遍牧场生,生态学家对它在生态学中的重要性已取得了比以往更深刻的认识。试图从空间异质性的含义,空间异质性与尺度和等级的关系,空间异质性的定量描述,空间异质性对生物和非生物过程的影响,以及空间异质性的动态等5个方面综述了有关空间异质性的生态学研究的新进展。     

Bird based Index of Biotic Integrity: Assessing the ecological condition of Atlantic Forest patches in human-modified landscape

Wooded biomes converted to human-modified landscapes (HML) are common throughout the tropics, yielding small and isolated forest patches surrounded by an agricultural matrix. Diverse anthropogenic interventions in HMLs influence patches in complex ways, altering natural dynamics. Assessing current condition or ecological integrity in these patches is a challenging task for ecologists. Taking the Brazilian Atlantic Forest as a case study, we used the conceptual framework of the Index of Biotic Integrity (IBI), a multimetric approach, to assess the ecological integrity of eight small forest patches in a highly disturbed HML with different configurations and histories. The IBI was developed using bird assemblages found in these patches, and its performance was compared with analytical approaches commonly used in environmental assessment, such as general richness and Shannon’s diversity index. As a first step, the IBI procedure identifies an existing gradient of human disturbance in the study region and checks which biotic characteristics (candidate metrics) vary systematically across the gradient. A metric is considered valid when its’ relationship with the gradient provides an ecological interpretation of the environment. Then, the final IBI is elaborated using each valid metric, obtaining a score for each site. Over one year of sampling, 168 bird species were observed, providing 74 different bird candidate metrics to be tested against the disturbance gradient. Seven of them were considered valid:richness of threatened species; richness of species that use both “forest and non-forest” habitats; abundance of endemics, abundance of small understory-midstory insectivores, abundance of exclusively forest species; abundance of non-forest species, and abundance of species that forage exclusively in the midstory stratum. Each metric provided complementary information about the patch’s ecological integrity. The resulting IBI showed a significant linear relationship with the gradient of human disturbance, while total species richness and Shannońs diversity index did not. Application of numerical approaches, such as total species richness and Shannon’s diversity, did not distinguish ecological traits among species. The IBI proved better for assessing and interpreting ecological and environmental condition of small patches in highly disturbed HML. The IBI framework, its multimetric character, and the ease with which it can be adapted to diverse situations, make it an effective approach for assessing environmental conditions in the Atlantic Forest region, and also for many other small forest patches in the tropics.     

A multidimensional framework for measuring biotic novelty: How novel is a community?

Anthropogenic changes in climate, land use, and disturbance regimes, as well as introductions of non‐native species can lead to the transformation of many ecosystems. The resulting novel ecosystems are usually characterized by species assemblages that have not occurred previously in a given area. Quantifying the ecological novelty of communities (i.e., biotic novelty) would enhance the understanding of environmental change. However, quantification remains challenging since current novelty metrics, such as the number and/or proportion of non‐native species in a community, fall short of considering both functional and evolutionary aspects of biotic novelty. Here, we propose the Biotic Novelty Index (BNI), an intuitive and flexible multidimensional measure that combines (a) functional differences between native and non‐native introduced species with (b) temporal dynamics of species introductions. We show that the BNI is an additive partition of Rao's quadratic entropy, capturing the novel interaction component of the community's functional diversity. Simulations show that the index varies predictably with the relative amount of functional novelty added by recently arrived species, and they illustrate the need to provide an additional standardized version of the index. We present a detailed R code and two applications of the BNI by (a) measuring changes of biotic novelty of dry grassland plant communities along an urbanization gradient in a metropolitan region and (b) determining the biotic novelty of plant species assemblages at a national scale. The results illustrate the applicability of the index across scales and its flexibility in the use of data of different quality. Both case studies revealed strong connections between biotic novelty and increasing urbanization, a measure of abiotic novelty. We conclude that the BNI framework may help building a basis for better understanding the ecological and evolutionary consequences of global change.     

Individual to community-level faunal responses to environmental change from a marine fossil record of Early Miocene global warming

Modern climate change has a strong potential to shift earth systems and biological communities into novel states that have no present-day analog, leaving ecologists with no observational basis to predict the likely biotic effects. Fossil records contain long time-series of past environmental changes outside the range of modern observation, which are vital for predicting future ecological responses, and are capable of (a) providing detailed information on rates of ecological change, (b) illuminating the environmental drivers of those changes, and (c) recording the effects of environmental change on individual physiological rates. Outcrops of Early Miocene Newport Member of the Astoria Formation (Oregon) provide one such time series. This record of benthic foraminiferal and molluscan community change from continental shelf depths spans a past interval environmental change (≈ 20.3-16.7 mya) during which the region warmed 2.1-4.5°C, surface productivity and benthic organic carbon flux increased, and benthic oxygenation decreased, perhaps driven by intensified upwelling as on the modern Oregon coast. The Newport Member record shows that (a) ecological responses to natural environmental change can be abrupt, (b) productivity can be the primary driver of faunal change during global warming, (c) molluscs had a threshold response to productivity change while foraminifera changed gradually, and (d) changes in bivalve body size and growth rates parallel changes in taxonomic composition at the community level, indicating that, either directly or indirectly through some other biological parameter, the physiological tolerances of species do influence community change. Ecological studies in modern and fossil records that consider multiple ecological levels, environmental parameters, and taxonomic groups can provide critical information for predicting future ecological change and evaluating species vulnerability.     

Comparative assessment of indices of freshwater habitat conditions using different invertebrate taxon sets

Monitoring changes in population levels of a wide range of species in biodiversity research and conservation requires practical, easy-to-use and efficient assessment and monitoring methods. Dragonflies (Insecta: Odonata) are a valuable tool for assessing aquatic systems and have been used as indicators of ecological health, ecological integrity, and environmental change, including climatic change, as well as indicators of habitat recovery. We field-tested a freshwater ecological integrity index, the Dragonfly Biotic Index (DBI), based on dragonfly assemblages at the local scale, and compared the DBI to a biodiversity index (average taxonomic distinctness, AvTD) as well as to a standard freshwater benthic macroinvertebrate-based freshwater health index (South African Scoring System, using Average Score Per Taxon, ASPT). We sampled 20 river sites, selected a priori. Adult dragonflies and benthic macroinvertebrates were collected using standardized methods. Environmental variables were collected in situ, and water samples taken. Temperature and pH were the most important physical environmental variables in explaining the assemblage structure, and we found significant abiotic–biotic relationships, as well as biotic–biotic relationships. Overall, dragonflies were more sensitive to changes in river condition than were macroinvertebrates, in part because they were responding at the species rather than higher taxonomic level. AvTD scores did not show any significant relationship with changes in river condition. Furthermore, sites with low biotic scores (indicating disturbance) had high AvTD values. In contrast, DBI site value and ASPT scores were highly significantly correlated. We conclude that dragonfly assemblages in the form of a DBI are an excellent tool for environmental assessment and monitoring freshwater biodiversity, with the potential to replace labour-intensive benthic macroinvertebrate-based freshwater quality assessments, such as SASS.     

Ecological theory meets soil ecotoxicology: Challenge and chance

The degradation of soils due to various anthropogenic stress factors is alarming. Although chemicals are a major reason for soil degradation, most ecologists are not interested in studying such effects. We try to wake their interest by addressing a number of unsolved soil ecotoxicological problems that are related to disturbance ecology, biodiversity, ecosystem functioning and modelling. Features distinguishing chemical from natural stress render promising new aspects in disturbance ecology. Ecotoxicological studies are ideal models of disturbance, particularly regarding frequency, intensity or multitude of stress. Patterns of secondary succession after a major chemical damage can directly be related to the intermediate disturbance hypothesis. More knowledge on altered life history patterns following stress could support both evolutionary ecology and risk assessment. We raise the question if inherent resource competition makes communities more vulnerable to stress. Three aspects of ecotoxicological risk assessment are introduced: (1) exposure and bioavailability, which is directly connected to environmental heterogeneity; (2) tests on ecosystem functioning, suffering from major drawbacks; and (3) modelling. Here, promising approaches exist but need substantial input for being applicable to soils. Ecological modelling should put more emphasis on simulating both natural and chemical disturbances, including behavioural aspects and environmental variability. Finally, research needs for ecological risk assessment in soils are derived such as a simple system to assess the impact of chemicals on soil biodiversity, the inclusion of behavioural changes of keystone species or the consideration of density-dependent effects. Common research efforts of basic ecologists and soil ecotoxicologists could render a lot of mutual benefits.     

Three levels of integrating ecology with the conservation of South American temperate forests: the initiative of the Institute of Ecological Research Chiloé, Chile

The diversity of native species assemblages and that of indigenous cultures that once characterized the temperate forests of southern Chile have experienced a process of homogenization ever since the Spanish conquest. Today this process continues to erode both biotic and cultural diversity. With the goal of linking ecological research with actions to conserve the biological and cultural richness of this region, we established the Institute of Ecological Research Chiloé. The Institute's philosophy and activities involve three approaches: (1) participation of professional ecologists in environmental education and decision making, through collaboration with the community at local, regional, and global scales. (2) programs of ecological education, which include planting indigenous trees in urban areas and creating a local botanical garden with representative Chilean forest species. (3) critical analyses of the narrow economic and utilitarian environmental ethics that currently prevail in Chile, and often in other Latin American nations, and examination of traditional or novel alternative ethics and perspectives that address multiple interrelations between biological and cultural dimensions.     

Coevolutionary aesthetics in human and biotic artworlds

This work proposes a coevolutionary theory of aesthetics that encompasses both biotic and human arts. Anthropocentric perspectives in aesthetics prevent the recognition of the ontological complexity of the aesthetics of nature, and the aesthetic agency of many non-human organisms. The process of evaluative coevolution is shared by all biotic advertisements. I propose that art consists of a form of communication that coevolves with its own evaluation. Art and art history are population phenomena. I expand Arthur Danto’s Artworld concept to any aesthetic population of producers and evaluators. Current concepts of art cannot exclusively circumscribe the human arts from many forms of non-human biotic art. Without assuming an arbitrarily anthropocentric perspective, any concept of art will need to engage with biodiversity, and either recognize many instances of biotic advertisements as art, or exclude some instances of human art. Coevolutionary aesthetic theory provides a heuristic account of aesthetic change in both human and biotic artworlds, including the coevolutionary origin of aesthetic properties and aesthetic value within artworlds. Restructuring aesthetics, art criticism, and art history without human beings at the organizing centers of these disciplines stimulate new progress in our understanding of art, and the unique human contributions to aesthetics and aesthetic diversity.     

Interspecific Competition and Species' Distributions: The Ghosts of Theories and Data Past

The idea that biological interactions between species restricttheir distributions beyond limits set by the inorganic environmentwas established by plant ecologists well before the work ofVolterra, Gause, and the development of modern niche theory.Mechanisms of competition between plant species, and the influenceof predation and environmental factors on the outcome of interspecificcompetition, were demonstrated by extensive field experiments.These early botanical achievements were ignored by zoologistswhose subsequent, independently derived ideas about competitionparalleled those of plant ecologists to a remarkable degree.Application of niche theory to the real world was closely tiedto the advent of the "modern synthesis." The primary innovationof niche theory was not simply a new appreciation of the possibleimportance of competition between animals, but the incorporationof an evolutionary perspective to distributional problems largelyabsent among ecologists since Darwin.     

Tropical Islands as Paleoecological Laboratories: Gauging the Consequences of Human Arrival

Inter-island paleoecological comparisons have provided useful information concerning the role of humans vs. background-level disturbance in tropical ecosystems. Major ecological changes have occurred since human arrival in Madagascar, the West Indies, the Hawaiian Islands, and elsewhere. Prehuman vegetation changes and disturbances have also been documented for many islands. Instructive inter-island similarities and differences have been detected in the chronology, distribution, and extent of human activities, vegetation changes, and biotic extinctions. The earliest stratigraphic proxy evidence for initial human impacts (including increased charcoal particle influx to sediments, first appearance of exotic pollen, increase in ruderal pollen, and paleolimnological evidence for cultural eutrophication of lake waters) generally confirm but sometimes predate the earliest conventional archaeological evidence for human activity. Carefully chosen sites permitting the close integration of palynological, paleontological, and archaeological data from a variety of island settings with differing geographic and historical contingencies can enable investigators to more fully evaluate the importance of a range of human and ecological variables in determining the overall character and dynamics of ecosystems.