The land flora: a phototroph-fungus partnership?

Numerous mutualistic associations between phototrophs and fungi exist in the extant land biota. Some are widespread, such as lichens and mycorrhizae, but some are less well known or restricted to special ecological conditions, such as endophytes in plants and algae. Recent molecular data and fossils suggest that associations arose repeatedly and that some of them are ancient, and even ancestral in the case of land plants. Mutualism, that provides various adaptations to terrestrial constraints, may have played a crucial role during terrestrialization and evolution of land phototrophs.     

The quest for a unified view of bacterial land colonization

Exploring molecular mechanisms underlying bacterial water-to-land transition represents a critical start toward a better understanding of the functioning and stability of the terrestrial ecosystems. Here, we perform comprehensive analyses based on a large variety of bacteria by integrating taxonomic, phylogenetic and metagenomic data, in the quest for a unified view that elucidates genomic, evolutionary and ecological dynamics of the marine progenitors in adapting to nonaquatic environments. We hypothesize that bacterial land colonization is dominated by a single-gene sweep, that is, the emergence of dnaE2 derived from an early duplication event of the primordial dnaE, followed by a series of niche-specific genomic adaptations, including GC content increase, intensive horizontal gene transfer and constant genome expansion. In addition, early bacterial radiation may be stimulated by an explosion of land-borne hosts (for example, plants and animals) after initial land colonization events.     

Plant–microbe interactions that have impacted plant terrestrializations

Plants display a tremendous diversity of developmental and physiological features, resulting from gains and losses of functional innovations across the plant phylogeny. Among those, the most impactful have been undoubtedly the ones that allowed plant terrestrializations, the transitions from an aquatic to a terrestrial environment. Although the embryophyte terrestrialization has been particularly scrutinized, others occurred across the plant phylogeny with the involvement of mutualistic symbioses as a common theme. Here, we review the current pieces of evidence supporting that the repeated colonization of land by plants has been facilitated by interactions with mutualistic symbionts. In that context, we detail two of these mutualistic symbioses: the arbuscular mycorrhizal symbiosis in embryophytes and the lichen symbiosis in chlorophyte algae. We suggest that associations with bacteria should be revisited in that context, and we propose that overlooked symbioses might have facilitated the emergence of other land plant clades.

Diverse plant lineages have independently colonized emerged lands over the last 450 million years and were helped in this process by mutualistic associations with fungi and potentially bacteria.     

The development of early terrestrial ecosystems

Summary

In this review of terrestrialization by plants and animals in the early Phanerozoic, the classical idea of a major mid-Palaeozoic event is discarded in favour of gradual colonization over a long time period. Four phases of colonization of the land by plants are recognized but their limits are often difficult to define. The first, of microbial mats comprising prokaryotes and later photosynthesizing protists (algae) but with no direct fossil evidence, extends from the Precambrian and may persist in environments unsuitable for colonization by higher plants and animals today. The second, based on microfossils (spores and cuticles) possibly from plants of bryophyte aspect (if not affinity) started in the Ordovician (c. 460 Ma ago) and ended in the Lower Devonian, but was overlapped by the third phase beginning early in the Silurian (c. 430 Ma). This consisted of small plants of axial organization with terminal sporangia probably allied to the tracheophytes. The advent of taller vascular plants of varied organization around the Silurian — Devonian boundary (c. 420–400 Ma) signalled the final pioneering phase — that of major adaptative radiations on land, culminating in the appearance of extant groups, in changes in reproductive strategy and in the development of complex vegetation structure. The animal record is sparser than that of the plants, but suggests an early land fauna in the mid-Palaeozoic which differed from later terrestrial assemblages in lacking herbivores, with the first direct fossil evidence for land animals in the late Silurian.     

A comparison of the colonization of the planktic realm and the land

Metazoans evolved in the sea and quickly invaded benthic niches. It was from these niches that other large ecosystems were colonized through the later Precambrian and the Phanerozoic. Entering these new environments, the colonists encountered new sets of macroevolutionary controls. Narratives of each colonization are important for historical ecology, while comparison of different events can throw light on these underlying controls. Recent narrative work on the evolution of plankton has suggested that this series of events was unique in respect of its duration and method. Animals have been migrating into the plankton since the Proterozoic, and all the main planktic groups have a benthic origin. Migration was usually by single species acting independently of their native community. A comparison with the other major Phanerozoic colonization, that of the land, highlights these unusual features. Colonization of the land began in the Ordovician and seems to have occurred sporadically. Several of the major groups of terrestrial animals originated on land. The invasion of land is usually related to the evolution of land plants and is perhaps best regarded as migration at the level of the community. The major differences between the two colonizations may be attributable to the dominant control of the environment over plankton versus that of biotic interaction over land animals.     

Azospirillum genomes reveal transition of bacteria from aquatic to terrestrial environments

Fossil records indicate that life appeared in marine environments ～3.5 billion years ago (Gyr) and transitioned to terrestrial ecosystems nearly 2.5 Gyr. Sequence analysis suggests that "hydrobacteria" and "terrabacteria" might have diverged as early as 3 Gyr. Bacteria of the genus Azospirillum are associated with roots of terrestrial plants; however, virtually all their close relatives are aquatic. We obtained genome sequences of two Azospirillum species and analyzed their gene origins. While most Azospirillum house-keeping genes have orthologs in its close aquatic relatives, this lineage has obtained nearly half of its genome from terrestrial organisms. The majority of genes encoding functions critical for association with plants are among horizontally transferred genes. Our results show that transition of some aquatic bacteria to terrestrial habitats occurred much later than the suggested initial divergence of hydro- and terrabacterial clades. The birth of the genus Azospirillum approximately coincided with the emergence of vascular plants on land.     

Origin and evolution of green plants in the light of key evolutionary events

Green plants (Viridiplantae) are ancient photosynthetic organisms that thrive both in aquatic and terrestrial ecosystems, greatly contributing to the changes in global climates and ecosystems. Significant progress has been made toward understanding the origin and evolution of green plants, and plant biologists have arrived at the consensus that green plants first originated in marine deep-water environments and later colonized fresh water and dry land. The origin of green plants, colonization of land by plants and rapid radiation of angiosperms are three key evolutionary events during the long history of green plants. However, the comprehensive understanding of evolutionary features and molecular innovations that enabled green plants to adapt to complex and changeable environments are still limited. Here, we review current knowledge of phylogenetic relationships and divergence times of green plants, and discuss key morphological innovations and distinct drivers in the evolution of green plants. Ultimately, we highlight fundamental questions to advance our understanding of the phenotypic novelty, environmental adaptation, and domestication of green plants.     

Seasonal dynamics of arbuscular mycorrhizal fungi in differing wetland habitats

The dynamics and role of arbuscular mycorrhizal fungi (AMF) have been well described in terrestrial ecosystems; however, little is known about how the dynamics of AMF are related to the ecology of wetland ecosystems. The seasonal dynamics of arbuscular mycorrhizal (AM) colonization within different wetland habitats were examined in this study to determine the factors that influence AM associations and to further assess the ecological role of AMF in wetlands. Fen and marsh habitats of four wetlands in west central Ohio were sampled monthly from March to September. AMF were found at all four sites for each month sampled and were present in all of the dominant plant species. A significant effect of month (P<0.001) on AM colonization did occur and was attributable to maximum colonization levels in the spring and minimum levels in late summer. This trend existed in all four wetlands in both fen and marsh habitats, regardless of variation in water levels, percent soil moisture, or available phosphorus levels. Because abiotic factors had minimal influence on AM colonization variation and the level of AM colonization paralleled plant growth patterns, we conclude that the AM seasonal dynamic was in response to plant phenology. Our data suggest that AM associations in temperate fen and marsh habitats are prevalent in the spring during new root and vegetative growth, even for plants experiencing flooded conditions. Evidence of an overriding AM seasonal trend indicates that future studies should include a seasonal component to better assess the role and distribution of AMF in wetland ecosystems.     

Microplastics as an emerging threat to terrestrial ecosystems

Microplastics (plastics <5 mm, including nanoplastics which are <0.1 μm) originate from the fragmentation of large plastic litter or from direct environmental emission. Their potential impacts in terrestrial ecosystems remain largely unexplored despite numerous reported effects on marine organisms. Most plastics arriving in the oceans were produced, used, and often disposed on land. Hence, it is within terrestrial systems that microplastics might first interact with biota eliciting ecologically relevant impacts. This article introduces the pervasive microplastic contamination as a potential agent of global change in terrestrial systems, highlights the physical and chemical nature of the respective observed effects, and discusses the broad toxicity of nanoplastics derived from plastic breakdown. Making relevant links to the fate of microplastics in aquatic continental systems, we here present new insights into the mechanisms of impacts on terrestrial geochemistry, the biophysical environment, and ecotoxicology. Broad changes in continental environments are possible even in particle‐rich habitats such as soils. Furthermore, there is a growing body of evidence indicating that microplastics interact with terrestrial organisms that mediate essential ecosystem services and functions, such as soil dwelling invertebrates, terrestrial fungi, and plant‐pollinators. Therefore, research is needed to clarify the terrestrial fate and effects of microplastics. We suggest that due to the widespread presence, environmental persistence, and various interactions with continental biota, microplastic pollution might represent an emerging global change threat to terrestrial ecosystems.     

Diverse adaptations of an ancestral gill: a common evolutionary origin for wings,breathing organs,and spinnerets

Changing conditions of life impose new requirements on the morphology and physiology of an organism. One of these changes is the evolutionary transition from aquatic to terrestrial life, leading to adaptations in locomotion, breathing, reproduction, and mechanisms for food capture. We have shown previously that insects' wings most likely originated from one of the gills of ancestral aquatic arthropods during their transition to life on land. Here we investigate the fate of these ancestral gills during the evolution of another major arthropod group, the chelicerates. We examine the expression of two developmental genes, pdm/nubbin and apterous, that participate in the specification of insects' wings and are expressed in particular crustacean epipods/gills. In the horseshoe crab, a primitively aquatic chelicerate, pdm/nubbin is specifically expressed in opisthosomal appendages that give rise to respiratory organs called book gills. In spiders (terrestrial chelicerates), pdm/nubbin and apterous are expressed in successive segmental primordia that give rise to book lungs, lateral tubular tracheae, and spinnerets, novel structures that are used by spiders to breathe on land and to spin their webs. Combined with morphological and palaeontological evidence, these observations suggest that fundamentally different new organs (wings, air-breathing organs, and spinnerets) evolved from the same ancestral structure (gills) in parallel instances of terrestrialization.     

Paradigm shifts in soil organic matter research affect interpretations of aquatic carbon cycling: transcending disciplinary and ecosystem boundaries

New conceptual models that highlight the importance of environmental, rather than molecular, controls on soil organic matter affect interpretations of organic matter (OM) persistence across terrestrial and aquatic boundaries. We propose that changing paradigms in our thinking about OM decomposition explain some of the uncertainties surrounding the fate of land-derived carbon (C) in marine environments. Terrestrial OM, which historically has been thought to be chemically recalcitrant to decay in soil and aquatic environments, dominates inputs to rivers yet is found in trace amounts in the ocean. We discuss three major transformations in our understanding of OM persistence that influence interpretations of the fate of aquatic OM: (1) a shift away from an emphasis on chemical recalcitrance as a primary predictor of turnover; (2) new interpretations of radiocarbon ages, which affect predictions of reactivity; and (3) the recognition that most OM leaving soils in dissolved form has been microbially processed. The first two explain rapid turnover for terrigenous OM in aquatic ecosystems once it leaves the soil matrix. The third suggests that the presence of terrestrial OM in aquatic ecosystems may be underestimated by the use of plant biomarkers. Whether these mechanisms occur in isolation of each other or in combination, they provide insight into the missing terrestrial C signature in the ocean. Spatially and temporally varying transformations of OM along land–water networks require that common terrestrial source indicators be interpreted within specific environmental contexts. We identify areas of research where collaborations between aquatic and terrestrial scientists will enhance quantification of C transfer from soils to inland water bodies, the ocean, and the atmosphere. Accurate estimates of OM processing are essential for improving predictions of the response of vulnerable C pools at the interface of soil and water to changes in climate and land use.     

大尺度陆地生态系统科学研究的理论基础及其技术体系之探讨

大尺度的陆地生态系统科学研究是在生物多样性保护、应对全球气候变化、区域生态环境治理及全球社会经济可持续发展的科技需求背景下应运而生的生态学及生态系统科学前沿领域,并在我国的生态文明建设社会实践中快速发展.本文在系统论述大尺度陆地生态系统科学研究的科技使命基础上,重点梳理和探讨了区域及大陆尺度的生态系统科学研究理论基础和...     

Conservation and restoration of New Zealand Island ecosystems

An ecological collapse has precipitated pioneering conservation initiatives in New Zealand. Many terrestrial communities in t he New Zealand archipelago have been devastated by over-exploitation, introduced mammals and habitat destruction. More recently, marine ecosystems have been depleted by over-harvesting. To mitigate against these losses, conservation in terrestrial environments has focused on protection of species and habitats. A similar approach is now under way in marine environments with the establishment of ‘no-take’ marine reserves. On land, conservation is now reaching beyond protection t o the eradication of pests from islands and restoration of their terrestrial ecosystems. Restoration on islands not only reduces threats to rare species; it also raises opportunities to investigate how species interact. In the sea, marine reserves not only enhance the diversity of depleted marine communities; they may also augment stocks of commercially harvested species. These initiatives provide many lessons that could be applied to degraded habitats elsewhere.     

基于陆地-水生态系统耦合的海河流域水生态功能分区

水生态功能分区是针对水生态系统特征的陆地生态系统划分,是为流域水生态管理提供生态背景和基本单元。陆地-水生态系统的耦合是水生态功能分区的核心,但多停留在个别小流域进行理论探讨,大型流域的实际案例较少。针对海河流域独特的气候、地貌、水文和人类活动特征,提出了水生态功能分区的三级指标体系。一级二级区针对气候、地貌、水文背景进行"自上而下"的分区,三级区针对人类活动对水资源、水环境、生境影响,采用"自下而上"的分区方法。最终,海河流域划分了6个一级区、16个二级区和73个三级区。研究充分体现了"以水定陆、以陆控水"的基本原则,以及"自下而上"和"自上而下"分区方法的优点,结果可为海河流域水生态管理提供科学依据,为水资源空间调配与合理利用、产业结构布局与区域协调等服务。     

Resource subsidies between stream and terrestrial ecosystems under global change

Streams and adjacent terrestrial ecosystems are characterized by permeable boundaries that are crossed by resource subsidies. Although the importance of these subsidies for riverine ecosystems is increasingly recognized, little is known about how they may be influenced by global environmental change. Drawing from available evidence, in this review we propose a conceptual framework to evaluate the effects of global change on the quality and spatiotemporal dynamics of stream–terrestrial subsidies. We illustrate how changes to hydrological and temperature regimes, atmospheric CO₂ concentration, land use and the distribution of nonindigenous species can influence subsidy fluxes by affecting the biology and ecology of donor and recipient systems and the physical characteristics of stream–riparian boundaries. Climate‐driven changes in the physiology and phenology of organisms with complex life cycles will influence their development time, body size and emergence patterns, with consequences for adjacent terrestrial consumers. Also, novel species interactions can modify subsidy dynamics via complex bottom‐up and top‐down effects. Given the seasonality and pulsed nature of subsidies, alterations of the temporal and spatial synchrony of resource availability to consumers across ecosystems are likely to result in ecological mismatches that can scale up from individual responses, to communities, to ecosystems. Similarly, altered hydrology, temperature, CO₂ concentration and land use will modify the recruitment and quality of riparian vegetation, the timing of leaf abscission and the establishment of invasive riparian species. Along with morphological changes to stream–terrestrial boundaries, these will alter the use and fluxes of allochthonous subsidies associated with stream ecosystems. Future research should aim to understand how subsidy dynamics will be affected by key drivers of global change, including agricultural intensification, increasing water use and biotic homogenization. Our conceptual framework based on the match–mismatch between donor and recipient organisms may facilitate understanding of the multiple effects of global change and aid in the development of future research questions.     

Living in a fungal world: impact of fungi on soil bacterial niche development

The colonization of land by plants appears to have coincided with the appearance of mycorrhiza-like fungi. Over evolutionary time, fungi have maintained their prominent role in the formation of mycorrhizal associations. In addition, however, they have been able to occupy other terrestrial niches of which the decomposition of recalcitrant organic matter is perhaps the most remarkable. This implies that, in contrast to that of aquatic organic matter decomposition, bacteria have not been able to monopolize decomposition processes in terrestrial ecosystems. The emergence of fungi in terrestrial ecosystems must have had a strong impact on the evolution of terrestrial bacteria. On the one hand, potential decomposition niches, e.g. lignin degradation, have been lost for bacteria, whereas on the other hand the presence of fungi has itself created new bacterial niches. Confrontation between bacteria and fungi is ongoing, and from studying contemporary interactions, we can learn about the impact that fungi presently have, and have had in the past, on the ecology and evolution of terrestrial bacteria. In the first part of this review, the focus is on niche differentiation between soil bacteria and fungi involved in the decomposition of plant-derived organic matter. Bacteria and fungi are seen to compete for simple plant-derived substrates and have developed antagonistic strategies. For more recalcitrant organic substrates, e.g. cellulose and lignin, both competitive and mutualistic strategies appear to have evolved. In the second part of the review, bacterial niches with respect to the utilization of fungal-derived substrates are considered. Here, several lines of development can be recognized, ranging from mutualistic exudate-consuming bacteria that are associated with fungal surfaces to endosymbiotic and mycophagous bacteria. In some cases, there are indications of fungal specific selection in fungus-associated bacteria, and possible mechanisms for such selection are discussed.     

Effects of climate legacies on above‐ and belowground community assembly

The role of climatic legacies in regulating community assembly of above‐ and belowground species in terrestrial ecosystems remains largely unexplored and poorly understood. Here, we report on two separate regional and continental empirical studies, including >500 locations, aiming to identify the relative importance of climatic legacies (climatic anomaly over the last 20,000 years) compared to current climates in predicting the relative abundance of ecological clusters formed by species strongly co‐occurring within two independent above‐ and belowground networks. Climatic legacies explained a significant portion of the variation in the current community assembly of terrestrial ecosystems (up to 15.4%) that could not be accounted for by current climate, soil properties, and management. Changes in the relative abundance of ecological clusters linked to climatic legacies (e.g., past temperature) showed the potential to indirectly alter other clusters, suggesting cascading effects. Our work illustrates the role of climatic legacies in regulating ecosystem community assembly and provides further insights into possible winner and loser community assemblies under global change scenarios.