Trait‐based ecology of terrestrial arthropods

In focusing on how organisms' generalizable functional properties (traits) interact mechanistically with environments across spatial scales and levels of biological organization, trait‐based approaches provide a powerful framework for attaining synthesis, generality and prediction. Trait‐based research has considerably improved understanding of the assembly, structure and functioning of plant communities. Further advances in ecology may be achieved by exploring the trait–environment relationships of non‐sessile, heterotrophic organisms such as terrestrial arthropods, which are geographically ubiquitous, ecologically diverse, and often important functional components of ecosystems. Trait‐based studies and trait databases have recently been compiled for groups such as ants, bees, beetles, butterflies, spiders and many others; however, the explicit justification, conceptual framework, and primary‐evidence base for the burgeoning field of ‘terrestrial arthropod trait‐based ecology’ have not been well established. Consequently, there is some confusion over the scope and relevance of this field, as well as a tendency for studies to overlook important assumptions of the trait‐based approach. Here we aim to provide a broad and accessible overview of the trait‐based ecology of terrestrial arthropods. We first define and illustrate foundational concepts in trait‐based ecology with respect to terrestrial arthropods, and justify the application of trait‐based approaches to the study of their ecology. Next, we review studies in community ecology where trait‐based approaches have been used to elucidate how assembly processes for terrestrial arthropod communities are influenced by niche filtering along environmental gradients (e.g. climatic, structural, and land‐use gradients) and by abiotic and biotic disturbances (e.g. fire, floods, and biological invasions). We also review studies in ecosystem ecology where trait‐based approaches have been used to investigate biodiversity–ecosystem function relationships: how the functional diversity of arthropod communities relates to a host of ecosystem functions and services that they mediate, such as decomposition, pollination and predation. We then suggest how future work can address fundamental assumptions and limitations by investigating trait functionality and the effects of intraspecific variation, assessing the potential for sampling methods to bias the traits and trait values observed, and enhancing the quality and consolidation of trait information in databases. A roadmap to guide observational trait‐based studies is also presented. Lastly, we highlight new areas where trait‐based studies on terrestrial arthropods are well positioned to advance ecological understanding and application. These include examining the roles of competitive, non‐competitive and (multi‐)trophic interactions in shaping coexistence, and macro‐scaling trait–environment relationships to explain and predict patterns in biodiversity and ecosystem functions across space and time. We hope this review will spur and guide future applications of the trait‐based framework to advance ecological insights from the most diverse eukaryotic organisms on Earth.     

Emergence of community structure in terrestrial mammal-dominated ecosystems

We present a paper that combines empirical and theoretical research about the trophic organization of biological communities. Some regularities are observed in the analysis of the relationship between the trophic structure (how the species are distributed among a set of feeding groups) of a number of African large mammal communities and the type of ecosystem. Different types of ecosystems are characterized by specific patterns in the trophic structure of the mammal community. In order to explain the origin of these patterns, we propose a model defining the underlying dynamic of mammal-dominated ecosystems. The main aim of this study is to show that it is possible to obtain a dynamic explanation of those patterns. The model is shown to spontaneously define different types of structures in community organization, related to those observed. We propose a model that could help to explain the correlation between different environmental factors and the abundance or diversity of herbivores, and which establishes a general mechanism that makes it possible to understand how some rules constrain the assembly of the communities. In addition, the proposed model leads us to see how biological communities can operate in an integrated way, which allows for the acceptance of their changes on large time-scales as evolutionary. In summary, we suggest that communities are unitary structures with coherent properties that result from the self-organizing dynamic of the whole system.     

Pulsed resources and community dynamics of consumers in terrestrial ecosystems

Many terrestrial ecosystems are characterized by intermittent production of abundant resources for consumers, such as mast seeding and pulses of primary production following unusually heavy rains. Recent research is revealing patterns in the ways that consumer communities respond to these pulsed resources. Studies of the ramifying effects of pulsed resources on consumer communities integrate 'top-down' and 'bottom-up' approaches to community dynamics, and illustrate how the strength of species interactions can change dramatically through time.     

Trophic interactions in the methanogenic microbial community of low-temperature terrestrial ecosystems

The formation of methane in various ecosystems is due to the functioning of an anaerobic community, which combines trophically different groups of microorganisms. The methanogenic microbial community is a complex biological system, which responds to low temperatures by changes in its trophic structure resulting in redistributing matter flows. The enhanced activity of homoacetogenic bacteria at low temperature plays a significant role in this redistribution. Due to their relatively high growth rates and metabolic versatility, homoacetogens can successfully compete with fermenting bacteria and hydrogenotrophic methanogenic archaea for common substrates. The concentration of hydrogen is an important regulatory factor in the psychroactive methanogenic community. At low temperature methanogenic archaea possessing a higher affinity for hydrogen than homoacetogens provide for interspecies H2 transport in syntrophic reactions of fatty acid decomposition. The formation of a balanced community at low temperature is a longtime process. Cold terrestrial ecosystems are dominated by psychroactive (psychrotolerant) microorganisms, which can grow over a wide range of ambient temperatures.     

Linking human impacts to community processes in terrestrial and freshwater ecosystems

Human impacts such as habitat loss, climate change and biological invasions are radically altering biodiversity, with greater effects projected into the future. Evidence suggests human impacts may differ substantially between terrestrial and freshwater ecosystems, but the reasons for these differences are poorly understood. We propose an integrative approach to explain these differences by linking impacts to four fundamental processes that structure communities: dispersal, speciation, species-level selection and ecological drift. Our goal is to provide process-based insights into why human impacts, and responses to impacts, may differ across ecosystem types using a mechanistic, eco-evolutionary comparative framework. To enable these insights, we review and synthesise (i) how the four processes influence diversity and dynamics in terrestrial versus freshwater communities, specifically whether the relative importance of each process differs among ecosystems, and (ii) the pathways by which human impacts can produce divergent responses across ecosystems, due to differences in the strength of processes among ecosystems we identify. Finally, we highlight research gaps and next steps, and discuss how this approach can provide new insights for conservation. By focusing on the processes that shape diversity in communities, we aim to mechanistically link human impacts to ongoing and future changes in ecosystems.     

Survival strategies in polar terrestrial arthropods

Three components of the survival strategy of a terrestrial Antarctic mite, Alaskozetes antarcticus (Acari: Cryptostigmata) are considered: overwintering survival, energetics and life history. Supercooling is an important feature of its cold tolerance, whilst elevation of standard metabolism allows activity at low temperatures, both of which contribute tcTa long development and maximum survival of individuals in the population. These are facets of the overall survival strategy evolved by such a species in response to the Antarctic terrestrial environment, but which may be widespread in polar invertebrates.     

The incidence of bacterial endosymbionts in terrestrial arthropods

Intracellular endosymbiotic bacteria are found in many terrestrial arthropods and have a profound influence on host biology. A basic question about these symbionts is why they infect the hosts that they do, but estimating symbiont incidence (the proportion of potential host species that are actually infected) is complicated by dynamic or low prevalence infections. We develop a maximum-likelihood approach to estimating incidence, and testing hypotheses about its variation. We apply our method to a database of screens for bacterial symbionts, containing more than 3600 distinct arthropod species and more than 150 000 individual arthropods. After accounting for sampling bias, we estimate that 52% (CIs: 48–57) of arthropod species are infected with Wolbachia, 24% (CIs: 20–42) with Rickettsia and 13% (CIs: 13–55) with Cardinium. We then show that these differences stem from the significantly reduced incidence of Rickettsia and Cardinium in most hexapod orders, which might be explained by evolutionary differences in the arthropod immune response. Finally, we test the prediction that symbiont incidence should be higher in speciose host clades. But while some groups do show a trend for more infection in species-rich families, the correlations are generally weak and inconsistent. These results argue against a major role for parasitic symbionts in driving arthropod diversification.     

Speciation and phylogeography of Hawaiian terrestrial arthropods

The Hawaiian archipelago is arguably the world's finest natural laboratory for the study of evolution and patterns of speciation. Arthropods comprise over 75% of the endemic biota of the Hawaiian Islands and a large proportion belongs to species radiations. We classify patterns of speciation within Hawaiian arthropod lineages into three categories: (i) single representatives of a lineage throughout the islands; (ii) species radiations with either (a) single endemic species on different volcanoes or islands, or (b) multiple species on each volcano or island; and (iii) single widespread species within a radiation of species that exhibits local endemism. A common pattern of phylogeography is that of repeated colonization of new island groups, such that lineages progress down the island chain, with the most ancestral groups (populations or species) on the oldest islands. While great dispersal ability and its subsequent loss are features of many of these taxa, there are a number of mechanisms that underlie diversification. These mechanisms may be genetic, including repeated founder events, hybridization, and sexual selection, or ecological, including shifts in habitat and/or host affiliation. The majority of studies reviewed suggest that natural selection is a primary force of change during the initial diversification of taxa.     

Estimating co-extinction threats in terrestrial ecosystems

The biosphere is changing rapidly due to human endeavour. Because ecological communities underlie networks of interacting species, changes that directly affect some species can have indirect effects on others. Accurate tools to predict these direct and indirect effects are therefore required to guide conservation strategies. However, most extinction-risk studies only consider the direct effects of global change—such as predicting which species will breach their thermal limits under different warming scenarios—with predictions of trophic cascades and co-extinction risks remaining mostly speculative. To predict the potential indirect effects of primary extinctions, data describing community interactions and network modelling can estimate how extinctions cascade through communities. While theoretical studies have demonstrated the usefulness of models in predicting how communities react to threats like climate change, few have applied such methods to real-world communities. This gap partly reflects challenges in constructing trophic network models of real-world food webs, highlighting the need to develop approaches for quantifying co-extinction risk more accurately. We propose a framework for constructing ecological network models representing real-world food webs in terrestrial ecosystems and subjecting these models to co-extinction scenarios triggered by probable future environmental perturbations. Adopting our framework will improve estimates of how environmental perturbations affect whole ecological communities. Identifying species at risk of co-extinction (or those that might trigger co-extinctions) will also guide conservation interventions aiming to reduce the probability of co-extinction cascades and additional species losses.     

Soil nematode biodiversity in terrestrial ecosystems

A review of the literature on nematode diversity (=number of species identified) of soil inhabiting nematodes was undertaken and analysed with regard to distance from the equator, vegetation type and sampling effort. After applying a correction factor for sampling effort the results indicated that species richness was greatest in temperate broadleaf forest (61.7 species per sample) followed by cultivated soil, grassland, tropical rainforest, temperate coniferous forests and polar vegetation. The maintenance of high biodiversity in cultivated soils is unexpected but may reflect the impact of dominance in calculating many indices. Species richness was greatest between latitudes 30–40° (93.9 species per sample) and least above 70°, the mean richness near the equator (i.e. 0–10°) was 80.6 species per sample. While these data would suggest that nematode diversity is not necessarily greatest at the equator, and evidence to support a 'humped back' theory of species richness is not conclusive, they contradict the suggestion that nematode diversity increases with increased latitude.     

Desiccation stress at sub-zero temperatures in polar terrestrial arthropods

Cold tolerant polar terrestrial arthropods have evolved a range of survival strategies which enable them to survive the most extreme environmental conditions (cold and drought) they are likely to encounter. Some species are classified as being freeze tolerant but the majority of those found in the Antarctic survive sub-zero temperatures by avoiding freezing by supercooling. For many arthropods, not just polar species, survival of desiccating conditions is equally important to survival of low temperatures. At sub-zero temperatures freeze avoiding arthropods are susceptible to desiccation and may lose water due to a vapour diffusion gradient between their supercooled body fluids and ice in their surroundings. This process ceases once the body fluids are frozen and so is not a problem for freeze tolerant species. This paper compares five polar arthropods, which have evolved different low temperature survival strategies, and the effects of exposure to sub-zero temperatures on their supercooling points (SCP) and water contents. The Antarctic oribatid mite (Alaskozetes antarcticus) reduced its supercooling point temperature from -6 to -30 degrees C, when exposed to decreasing sub-zero temperatures (cooled from 5 to -10 degrees C over 42 days) with little loss of body water during that period. However, Cryptopygus antarcticus, a springtail which occupies similar habitats in the Antarctic, showed a decrease in both water content and supercooling ability when exposed to the same experimental protocol. Both these Antarctic arthropods have evolved a freeze avoiding survival strategy. The Arctic springtail (Onychiurus arcticus), which is also freeze avoiding, dehydrated (from 2.4 to 0.7 g water g(-1) dry weight) at sub-zero temperatures and its SCP was lowered from c. -3 to below -15 degrees C in direct response to temperature (5 to -5.5 degrees C). In contrast, the freeze tolerant larvae of an Arctic fly (Heleomyza borealis) froze at c. -7 degrees C with little change in water content or SCP during further cold exposure and survived frozen to -60 degrees C. The partially freeze tolerant sub-Antarctic beetle Hydromedion sparsutum froze at c. -2 degrees C and is known to survive frozen to -8 degrees C. During the sub-zero temperature treatment, its water content reduced until it froze and then remained constant. The survival strategies of such freeze tolerant and freeze avoiding arthropods are discussed in relation to desiccation at sub-zero temperatures and the evolution of strategies of cold tolerance.     

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.     

Diversity of non-native terrestrial arthropods on woody plants in Canada

A list of non-native phytophagous insects and mites on woody plants (trees, shrubs, vines) in Canada was compiled using information from literature and input from taxonomists. The 419 recorded species include Hemiptera (53% of species), Lepidoptera (22%), Coleoptera (13%) and Hymenoptera (9%). Almost all species originate from the Palearctic, especially Europe, reflecting historical trade patterns. About 41% of species were directly introduced to Canada from countries of origin, and the remainder spread from the United States of America (USA) after initial establishment there. Major ports on the east and west coasts, on Lake Erie and Lake Ontario are the main points of entry for exotic species directly introduced, and southern British Columbia (BC), Ontario (ON) and Quebec (QC) are the major points of entry for species spreading from the USA. Consequently, BC, ON, QC and Nova Scotia have the highest diversity of non-native species, and the prairie provinces and northern territories have the lowest. The extent of the distribution of individual species is related to length of time in Canada, number of introductions and dispersal abilities. Almost all native woody plant genera in Canada have been invaded by exotic phytophages. The large majority of phytophages occur on angiosperms. Woody plant genera with the largest distribution, highest species diversity and highest local abundances tend to host the greatest number of non-native species, including Picea, Pinus, Malus, Prunus, Salix, Betula, Quercus, Pyrus and Populus. The arrival rate of species in Canada increased from the late nineteenth century until about 1960, and declined rapidly thereafter. Quarantine legislation enacted in the USA in 1912 and in Canada in 1976 seems to have reduced the rate of insect invasion. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Global classification of natural terrestrial ecosystems

Summary A global classification system of natural terrestrial ecosystems (including systematic notation), based on the climate zones of Walter, is presented. The basic units of the system are the ecological units biome and biogeocoene. The zonobiomes, which are climate zones corresponding to the largest vegetation units, are subdivided into subzonobiomes and these into individual biomes. The biomes are thus natural, geographical units within the climate zones. They are in turn subdivided into individual biogeocoenes and their constituent synusiae. In addition, the coordinate concepts of pedobiome and orobiome are introduced. These are distinguished from the zonobiomes as follows:1. the pedobiomes by extreme edaphic conditions which cause azonal vegetation.2. the orobiomes, as mountain ranges, by their vertical climate zonation and the altitudinal belts of vegetation.These relationships are explained, and two subseries of pedo-and oro-subunits are established. Transitional zones (zono-ecotones) between individual zonobiomes are also distinguished. The classification system is summarized in a schematic, and a world map of zonobiomes and zono-ecotones is included. More details are presented in Walter (1976).

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Zusammenfassung Ein globales Gliederungssystem der natürlichen terrestrischen Ökosysteme (einschließlich systematischen Bezeichnungen) wird in Beziehung zu den Walter'schen Klimazonen gesetzt. Grundeinheiten des Systems sind die ökologischen Einheiten Biom und Biogeozön. Die Zonobiome werden unterteilt in Subzonobiome und diese in Biome. Die Zonobiome sind Klimazonen und entsprechen den größten Vegetationseinheiten. Die Biome sind natürliche, geographische Einheiten innerhalb der Klimazonen. Sie werden bis zu einzelnen Biogeozönen und ihren Synusien (Teilsytemen) unterteilt. Parallel dazu werden die Begriffe Pedobiom und Orobiom eingeführt. Diese heben sich aus den Zonobiomen heraus: die Pedobiomen durch extreme Böden, die eine azonale Vegetation bedingen, die Orobiome als Gebirge durch die vertikale Klimagliederung und die Höhenstufen der Vegetation. Diese Beziehungen werden erklärt, und zwei Nebenreihen der Pedo- bzw. Orobiom-Untereinheiten werden aufgestellt. Zwischen den einzelnen Zonobiomen werden Übergangszonen (Zonoökotone) unterschieden. Das Gliederungssytem wird bereits in einem Schema zusammengefaßt, und eine Weltkarte der Zonobiome und Zonoökotone wird beigefügt. Ausführlich werden alle diese Fragen bei WALTER (1976) behandelt.

     

Evolution of freezing susceptibility and freezing tolerance in terrestrial arthropods

Arthropods have evolved various adaptations to survive adverse seasons and it has long been discussed why some arthropods are freezing-susceptible and some are freezing-tolerant. However, which mode of frost resistance came first during the course of evolution? A commonly held opinion is that no choice of strategy has been offered in evolution, because each species of arthropod may have its own evolutionary and natural history, leading to cold-hardiness. Freezing tolerance is more frequent in holometabolous insect orders and partially used by certain vertebrates, like some terrestrially hibernating amphibians and reptiles. Supported by phylogenetic, ontogenetic and ecological arguments, we suggest here that freezing tolerance is more recent than freezing susceptibility in the course of arthropods evolution. In addition, we observe that three basic modes of freezing resistance in insect species exist in the field: (i) permanent or year-round freezing-susceptible species, (ii) alternative or seasonal freezing-susceptible/freezing-tolerant species, (iii) permanent or year-round freezing tolerant species.     

Evolutionary regime shifts in simulated ecosystems

Catastrophic regime shifts in ecosystems occur when the system is tipped into a new attractor state under some external forcing. Here we consider whether evolutionary adaptations within ecosystems can trigger similar transitions. We use an individual‐based, evolutionary model of interconnected ecosystems to analyze nonlinear changes in global state resulting from local adaptations. Transitions between periods of stability occur when new traits arise that allow exploitation of under‐utilized resources. Subsequent rapid growth of the population carrying the new trait causes abrupt environmental change that drives incumbent species extinct. We call these transitions ‘evolutionary regime shifts’. These internally generated perturbations can result in ecosystem collapse, followed by recovery to an alternate stable state, or occasionally system‐wide extinction. While these disruptions may have a negative impact on ecosystem productivity in individual simulation runs, mean results over many simulations show a trend for increasing ecosystem productivity and stability over time. Feedback between life and the abiotic environment in the model creates a ‘long‐tailed’ distribution of extinction sizes without any external trigger for large extinction events.