Original Articles: Plant Attribute Diversity, Resilience, and Ecosystem Function: The Nature and Significance of Dominant and Minor Species

This study tested an hypothesis concerning patterns in species abundance in ecological communities. Why do the majority of species occur in low abundance, with just a few making up the bulk of the biomass? We propose that many of the minor species are analogues of the dominants in terms of the ecosystem functions they perform, but differ in terms of their capabilities to respond to environmental stresses and disturbance. They thereby confer resilience on the community with respect to ecosystem function. Under changing conditions, ecosystem function is maintained when dominants decline or are lost because functionally equivalent minor species are able to substitute for them. We have tested this hypothesis with respect to ecosystem functions relating to global change. In particular, we identified five plant functional attributes—height, biomass, specific leaf area, longevity, and leaf litter quality—that determine carbon and water fluxes. We assigned values for these functional attributes to each of the graminoid species in a lightly grazed site and in a heavily grazed site in an Australian rangeland. Our resilience proposition was cast in the form of three specific hypotheses in relation to expected similarities and dissimilarities between dominant and minor species, within and between sites. Functional similarity—or ecological distance—was determined as the euclidean distance between species in functional attribute space. The analyses provide evidence in support of the resilience hypothesis. Specifically, within the lightly grazed community, dominant species were functionally more dissimilar to one another, and functionally similar species more widely separated in abundance rank, than would be expected on the basis of average ecological distances in the community. Between communities, depending on the test used, two of three, or three of four minor species in the lightly grazed community that were predicted to increase in the heavily grazed community did in fact do so. Although there has been emphasis on the importance of functional diversity in supporting the flow of ecosystem goods and services, the evidence from this study indicates that functional similarity (between dominant and minor species, and among minor species) may be equally important in ensuring persistence (resilience) of ecosystem function under changing environmental conditions.     

Effects Of Vegetation and Soil on Species Diversity of Soil Dwelling Diptera in a Heathland Ecosystem

The role of vegetation and soil factors on the biodiversity of two soil dwelling, saprophagous, fly families (Sphaeroceridae and Lonchopteridae) in a heathland ecosystem were investigated. The fly community is primarily affected by soil humidity and the amount of organic matter while the vegetation structure and species composition only indirectly influence the fly communities. There was no correlation between plant species richness and the fly diversity indices. Based on our results and literature data, we hypothesise that the direct effects of the vegetation is more evident for herbivorous insects than for species that do not feed on plants. The investigated families show a clear response to microhabitat differences in the soil factors, which makes them promising indicators for soil health and as tool for monitoring environmental changes.     

Role of Nematodes in Soil Health and Their Use as Indicators

The composition of nematode communities (plant-parasitic and free-living) may be used as bioindicators of soil health or condition because composition correlates well with nitrogen cycling and decomposition, two critical ecological processes in soil. Maturity and trophic diversity indices withstand statistical rigor better than do abundances, proportions, or ratios of trophic groups. Maturity indices respond to a variety of land-management practices, based largely on inferred life history characteristics of families. Similarity indices may be more useful than diversity indices because they reflect taxon composition. Improving existing indices or developing alternative indices refined by a greater understanding of the biology of key taxa may enhance the utility of nematodes as bioindicators.     

Nematode Population Fluctuations during Decomposition of Specific Organic Amendments

Population densities of nematodes in field soil without plants were monitored for 10 months following application of organic amendments to pots in a greenhouse. The four treatments consisted of three different kinds of organic amendments: homogeneous crop residues of maize (Zea mays, C:N = 48.0:1), Texas panicum (Panicum texanum, C:N = 32.9:1), or velvetbean (Mucuna pruriens, C:N = 18.6:1), plus a control without any amendment. Plant-parasitic nematodes declined in all treatments due to absence of a food source. Bacterivore numbers increased following amendment application and remained greater than initial population levels until 4 months after application. Fungivore numbers were higher than initial levels until 6 months after amendment application and did not decline below the initial numbers during the course of the experiment. On several sampling dates, the bacterivorous genera Cervidellus and Eucephalobus were most abundant in pots with maize residues. Among the fungivores, Aphelenchoides numbers early in the experiment were greatest in pots amended with velvetbean, whereas numbers of Aphelenchus, Nothotylenchus, and Tylenchidae (mainly Filenchus) were greatest during the latter half of the experiment following the maize amendment. Omnivorous nematodes, particularly Eudorylaimus, showed two peaks in abundance during the course of the experiment. Results provided some evidence that population levels of some genera of bacterivores and fungivores may be affected by specific organic amendments.     

Nematode Genera in Forest Soil Respond Differentially to Elevated CO2

Previous reports suggest that fungivorous nematodes are the only trophic group in forest soils affected by elevated CO₂. However, there can be ambiguity within trophic groups, and we examined data at a genus level to determine whether the conclusion remains similar. Nematodes were extracted from roots and soil of loblolly pine (Pinus taeda) and sweet gum (Liquidambar styraciflua) forests fumigated with either ambient air or CO₂-enriched air. Root length and nematode biomass were estimated using video image analysis. Most common genera included Acrobeloides, Aphelenchoides, Cephalobus, Ditylenchus, Ecphyadorphora, Filenchus, Plectus, Prismatolaimus, and Tylencholaimus. Maturity Index values and diversity increased with elevated CO₂ in loblolly pine but decreased with elevated CO₂ in sweet gum forests. Elevated CO₂ treatment affected the occurrence of more nematode genera in sweet gum than loblolly pine forests. Numbers were similar but size of Xiphinema decreased in elevated CO₂. Abundance, but not biomass, of Aphelenchoides was reduced by elevated CO₂. Treatment effects were apparent at the genus levels that were masked at the trophic level. For example, bacterivores were unaffected by elevated CO₂, but abundance of Cephalobus was affected by CO₂ treatment in both forests.     

Changes in Individual Allometry Can Lead to Species Coexistence without Niche Separation

The principle of competitive exclusion is a fundamental tenet of ecology. Commonly used competition models predict that at most only one species per limiting resource can coexist in the same environment at steady state; hence, the upper limit to species diversity depends only on the number of limiting resources and not on the rates of resource supply. We demonstrate that such model behavior is the result of both the growth and biomass turnover functions being proportional to the population biomass. We argue that at least the growth function should be a nonlinear, concave downward function of biomass. This form for the growth function should arise simply because of changes in the allometry of individuals in the population. With this change in model structure, we show that any number of species can coexist at an asymptotically stable steady state, even where there is only one limiting resource. Furthermore, if growth increases nonlinearly with biomass, the steady-state resource concentration and hence the potential for biodiversity increases as the resource supply rate increases. Received 31 August 2001; accepted 10 April 2002.     

Trophic Dependence of Ecosystem Resistance and Species Compensation in Experimentally Acidified Lake 302S (Canada)

Ecosystem resistance to the impacts of diverse human insults depends on the replacement of sensitive species by ones more tolerant of the stressor. Here we present evidence from a whole-lake acidification experiment (Lake 302S, Experimental Lakes Area, Canada) that resistance and species compensation decline with increasing trophic level. Diverse and fast-growing algal and rotifer assemblages with high dispersal potentials showed significant compensatory species dynamics, resulting in the maintenance of total biomass despite 30%–80% declines in species richness. Canonical correspondence analysis showed that significant compensatory algal and rotifer dynamics were best explained by differential species tolerances of acidified chemical conditions coupled with release from resource limitation and predation. However, less diverse cladoceran, copepod, and fish assemblages showed significant declines in total biomass and weak species compensation with loss of species during acidification. In comparison, algal and zooplankton species dynamics remained relatively synchronized in a nearby unperturbed reference lake (Lake 239) during the experiment. As a result, Lake 302S showed limited ecosystem resistance to anthropogenic acidification. Therefore, we hypothesize that lost species will increase the susceptibility of acidified lakes to the adverse impacts of other environmental stressors (for example, climate warming, stratospheric ozone depletion, invasive species). Consequently, the ecosystem stability of boreal lakes is expected to decline as global change proceeds. Received 2 January 2001; accepted 12 July 2002.     

Grazing in a porous environment. 2. Nematode community structure

The influence of soil matric potential on nematode community composition and grazing associations were examined. Undisturbed cores (5 cm diameter, 10 cm depth) were collected in an old field dominated by perennial grasses on a Hinckley sandy loam at Peckham Farm near Kingston, Rhode Island. Ten pairs of cores were incubated at −3, −10, −20 and −50 kPa matric potential after saturation for 21–28 or 42–58 days. Nematodes were extracted using Cobb's decanting and sieving method followed by sucrose centrifugal-flotation and identified to family or genus. Collembola and enchytraeids present were also enumerated because they are grazers that reside in air-filled spaces. Direct counts of bacteria and fungi were made to estimate biovolume using fluorescein isothiocyanate and fluorescein diacetate stains, respectively. Trophic diversity and maturity indices were calculated for nematode communities. Three patterns of matric potential effect were observed for nematode taxa. One, there was a consistent effect of matric potential for all seasons for Alaimus, Monhysteridae, Prismatolaimus, Paraxonchium and Dorylaimoides. Two, some effects of matric potential were consistent among seasons and other effects were inconsistent for Aphelenchoides, Aphelenchus, Cephalobidae, Coomansus, Eudorylaimus, Huntaphelenchoides, Panagrolaimidae, Paraphelenchus, Sectonema, and Tripyla. Third, effects of matric potential were always inconsistent among seasons for Aphanolaimus, Aporcelaimellus, Bunonema, Rhabditidae, and Tylencholaimus. As predicted, fungal and bacterial biomass responded oppositely to matric potential. Total bacterial biomass was greater at −3 kPa than −10, −20 and −50 kPa (P=0.0095). Total fungal biomass was greater at −50, −20 and −10 kPa than −3 kPa (P=0.0095). Neither bacterial-feeding, fungal-feeding nor predacious nematodes correlated significantly with bacterial or fungal biomass. Omnivorous and predacious nematodes correlated positively with number of bacterial-feeding nematodes; predacious nematodes also correlated positively with fungal-feeding nematodes. Numbers of Collembola and enchytraeids were more often correlated positively with microbial-grazing nematode numbers in drier than moist soils. From this study, we propose two mechanisms that may explain nematode community structure changes with matric potential: differential anhydrobiosis and/or enclosure hypotheses. The later suggests that drying of soil generates pockets of moisture in aggregates that become isolated from one another enclosing nematodes and their food in relatively high concentrations creating patches of activity separated by larger areas of inactivity. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mobile Link Organisms and Ecosystem Functioning: Implications for Ecosystem Resilience and Management

Current natural resource management seldom takes the ecosystem functions performed by organisms that move between systems into consideration. Organisms that actively move in the landscape and connect habitats in space and time are here termed “mobile links.” They are essential components in the dynamics of ecosystem development and ecosystem resilience (that is, buffer capacity and opportunity for reorganization) that provide ecological memory (that is, sources for reorganization after disturbance). We investigated the effects of such mobile links on ecosystem functions in aquatic as well as terrestrial environments. We identify three main functional categories: resource, genetic, and process linkers and suggest that the diversity within functional groups of mobile links is a central component of ecosystem resilience. As the planet becomes increasingly dominated by humans, the magnitude, frequency, timing, spatial extent, rate, and quality of such organism-mediated linkages are being altered. We argue that global environmental change can lead to (a) the decline of essential links in functional groups providing pollination, seed dispersal, and pest control; (b) the linking of previously disconnected areas, for example, the spread of vector-borne diseases and invasive species; and (c) the potential for existing links to become carriers of toxic substances, such as persistent organic compounds. We conclude that knowledge of interspatial exchange via mobile links needs to be incorporated into management and policy-making decisions in order to maintain ecosystem resilience and hence secure the capacity of ecosystems to supply the goods and services essential to society. Received 23 April 2001; accepted 17 June 2002.     

Taxonomical vs. functional responses of bee communities to fire in two contrasting climatic regions

1. Valuable insights into mechanisms of community responses to environmental change can be gained by analysing in tandem the variation in functional and taxonomic composition along environmental gradients. 2. We assess the changes in species and functional trait composition (i.e. dominant traits and functional diversity) of diverse bee communities in contrasting fire-driven systems in two climatic regions: Mediterranean (scrub habitats in Israel) and temperate (chestnut forests in southern Switzerland). 3. In both climatic regions, there were shifts in species diversity and composition related to post-fire age. In the temperate region, functional composition responded markedly to fire; however, in the Mediterranean, the taxonomic response to fire was not matched by functional replacement. 4. These results suggest that greater functional stability to fire in the Mediterranean is achieved by replacement of functionally similar species (i.e. functional redundancy) which dominate under different environmental conditions in the heterogeneous landscapes of the region. In contrast, the greater functional response in the temperate region was attributed to a more rapid post-fire vegetation recovery and shorter time-window when favourable habitat was available relative to the Mediterranean. 5. Bee traits can be used to predict the functional responses of bee communities to environmental changes in habitats of conservation importance in different regions with distinct disturbance regimes. However, predictions cannot be generalized from one climatic region to another where distinct habitat configurations occur.     

Palaeolimnological insights for biodiversity science: an emerging field

1. Decreases in biodiversity are so widespread that they are now considered a form of global change in their own right. Given the grave nature of this issue, rapid advances in understanding are needed to mitigate further impacts. In this Opinion paper, we argue that palaeolimnological studies have important contributions to make to biodiversity science. 2. Given that long‐term community data are sparse in their geographic coverage and tend to span no more than 5 years, greater insight into biodiversity dynamics can be obtained from palaeoecological analyses. One such approach is palaeolimnology, which is a field that can provide long‐term data on changes in both physico‐chemical and biological components of lake ecosystems. 3. To date, a handful of quantitative palaeolimnological studies have addressed biodiversity questions, focussing primarily on defining the drivers of change in species richness or identifying functional traits that best capture ecosystem processes. Several studies have also quantified the role of spatial variables in determining assemblage structure, a necessary first step in addressing how metacommunity interactions influence biodiversity–ecosystem processes. Overall, these early studies show that palaeolimnological approaches can address both similar and novel questions compared with contemporary ecological studies. However, palaeolimnology allows for a great expansion of the temporal scale of investigation, the quantification of rates of change to stressors and the possibility of conducting experiments by applying resurrection techniques. 4. As an emerging field, there are numerous exciting applications of palaeolimnology to biodiversity science. It is an opportune time to create synergy between contemporary aquatic ecologists and palaeolimnologists.     

Soil Nematode Responses to Crop Management and Conversion to Native Grasses

Soil nematode community response to treatments of three, four-year crop rotations (spring wheat-pea-spring wheat-flax, spring wheat-green manure-spring wheat-flax, and spring wheat-alfalfa-alfalfa-flax) under conventional and organic management, and native tall grass restoration (restored prairie) were assessed in June 2003, and July and August 2004. The research site was the Glenlea Long-term Rotation and Crop Management Study, in the Red River Valley, Manitoba, established in 1992. The nematode community varied more with sample occasion than management and rotation. The restored prairie favored high colonizer-persister (c-p) value omnivores and carnivores, and fungivores but less bacterivores. The restored prairie soil food web was highly structured, mature and low-to-moderately enriched as indicated by structure (SI), maturity (MI) and enrichment (EI) index values, respectively. Higher abundance of fungivores and channel index (CI) values suggested fungal-dominated decomposition. Nematode diversity was low even after more than a decade of restoration. A longer time may be required to attain higher diversity for this restored fragmented prairie site distant from native prairies. No consistent differences were found between organic and conventional management for nematode trophic abundance, with the exception of enrichment opportunists of the c-p 1 group which were favored by conventional management. Although EI was lower and SI was higher for organic than conventional their absolute values suggested decomposition channels to be primarily bacterial, and fewer trophic links with both management scenarios. A high abundance of fungivores in the rotation including the green manure indicates greater fungal decomposition.     

乌鲁木齐市水磨沟公园土壤动物群落多样性研究

为了研究水磨沟公园土壤动物群落结构,于2006年4月15日至9月21日在水磨沟公园不同高度选择3个取样区,每个取样区设立两个不同的取样点,采集了0～5、5～10、10～15cm3个不同层次的大中型土壤动物.共获得土壤动物1153个,分别属于3门8纲18目,其中鞘翅目、中腹足目、小引类为优势类群,占全捕量的61%.蜘蛛目、伪蝎目、半翅目为稀有类群,仅占总捕获量的2.34%,其余为常见类群.分析表明,在不同高度土壤动物群落物种组成和多样性之间有明显的区别;山顶土壤动物个体数量和群落物种多样性指数明显的高于山地土壤动物个体数量和群落物种多样性指数.同时随着土壤深度的增加土壤动物物种数量逐步减少,表现出明显的垂直分布特征.     

Plant Community Composition as a Predictor of Regional Soil Carbon Storage in Alaskan Boreal Black Spruce Ecosystems

The boreal forest is the largest terrestrial biome in North America and holds a large portion of the world’s reactive soil carbon. Therefore, understanding soil carbon accumulation on a landscape or regional scale across the boreal forest is useful for predicting future soil carbon storage. Here, we examined the relationship between floristic composition and ecosystem parameters, such as soil carbon pools, the carbon-to-nitrogen (C/N) ratio of live black spruce needles, and normalized basal area increment (NBAI) of trees in black spruce communities, the most widespread forest type in the boreal forest of Alaska. Variability in ecosystem properties among black spruce stands was as large as that which had previously been documented among all forest types in the central interior of Alaska; we found an eightfold range in NBAI and fivefold range in mineral soil carbon and nitrogen pools. Acidic black spruce communities had significantly more carbon in the organic soil horizon than did nonacidic black spruce communities, but did not differ in any other measured ecosystem parameter. We explained 48% of the variation in total soil carbon with a combination of plant community indices and abiotic and biotic factors. Plant community composition was at least as effective as any single environmental factor or stand characteristic in predicting soil C pools in Alaskan black spruce ecosystems. We conclude that among the community properties analyzed, the presence of key groups of species, overall species composition, and diversity of certain functional types, especially Sphagnum moss species, are important predictors of soil carbon sequestration in the black spruce forest type.