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Understanding ecosystem dynamics for conservation of biota.

1. Ecosystems have higher-order emerging properties that can affect the conservation of species. We identify some of these properties in order to facilitate a better understanding of them. 2. Nonlinear, indirect effects of food web interactions among species can produce counterintuitive changes in populations. 3. Species differ in their roles and linkages with other species in the system. These roles are a property of the system. Such differences in roles influence how we conserve individual species. 4. Ecosystems operate at a multitude of interacting spatial and temporal scales, which together structure the system and affect the dynamics of individual populations. 5. Disturbance also structures an ecosystem, producing both long-term slow changes and sudden shifts in ecosystem dynamics. 6. Ecosystems therefore can have multiple states, determined both by disturbance regimes and biotic interactions. Conservation should recognize a possible multiplicity of natural states while avoiding aberrant (human-induced) states. 7. Ecosystem processes are influenced by the composition of the biota they contain. Disturbances to the biota can distort processes and functions, which in turn can endanger individual species. 8. The goal of ecosystem conservation is the long-term persistence of the biota in the system. There are two paradigms: community-based conservation (CBC) and protected area conservation. Both have their advantages but neither is sufficient to protect the biota on its own. 9. CBC is required to conserve the majority of the world's biota not included in protected areas. However, current CBC methods favour a few idiosyncratic species, distort the species complex, and ignore the majority. More comprehensive methods are required for this approach to meet the goal of ecosystem conservation. 10. Protected areas are essential to conserve species unable to coexist with humans. They also function as ecological baselines to monitor the effects of humans on their own ecosystems. 11. However, protected areas suffer from loss of habitat through attrition of critical areas. Thus, renewal (addition) of habitat is required in order to achieve the long-term persistence of biota in functioning ecosystems. Identification of minimum habitat areas and restoration of ecosystems become two major priorities for future research.

Animals↗

Effects of interspecific competition, predation, and their interaction on survival and development time of immature Anopheles quadrimaculatus.

We examined the effect of predation by the backswimmer (Notonecta undulata; Hemiptera: Notonectidae), competition by zooplankton and snails, and both predation and competition on the survival and development time of larval Anopheles quadrimaculatus mosquitoes in experimental mesocosms. We found that both predation and interspecific competition greatly reduced the survivorship of larvae and the number of larvae emerging into adulthood. Treatments with both predators and competitors had fewer larvae transitioning among instars and into adulthood but not in an additive way. In addition, mosquito larvae in the presence of predators and competitors took two days longer to emerge than where predators and competitions were absent. Our work provides evidence that biotic interactions, such as predation and competition, can strongly regulate the number of mosquito larvae by reducing the number of larvae that survive through instars and to emergence and by increasing the generation time.

Animals↗

Recurrent seasonal variations in abundance and composition of filamentous SOL cluster bacteria (Saprospiraceae, Bacteroidetes) in oligomesotrophic Lake Mondsee (Austria).

The spatial and temporal variation of SOL cluster bacteria was assessed in oligomesotrophic Lake Mondsee and adjacent lakes by fluorescence in situ hybridization over two annual cycles. The filamentous SOL bacteria were present in Lake Mondsee throughout the study period, and the seasonal dynamics of the SOL community were remarkably similar with respect to both abundance and composition in the two consecutive years. Only two of the three SOL subclusters were detected in Lake Mondsee and four connected lakes. These two populations significantly differed in size distribution and demonstrated pronounced but recurrent differences in seasonality and length of period of appearance in Lake Mondsee. Extensive sampling of the lakes in September 2003 revealed low horizontal variation in the composition of the SOL community within Lake Mondsee but marked variations with depth. Between connected habitats pronounced differences in the composition and abundance of the SOL community were detected. The interaction of SOL bacteria with bacterivorous protists, mesozooplankton, and phytoplankton was investigated in order to reveal variables controlling the structure and dynamics of SOL communities. No strong indication for a bottom-up influence of phytoplankton was found, while the estimated community grazing rates of mesozooplankton on SOL bacteria indicated a top-down control of SOL abundance during mesozooplankton peaks in spring and early autumn. Furthermore, species-specific differences in grazing of mesozooplankton on SOL bacteria were observed. In general, the overall composition of SOL communities was controlled by abiotic factors (water chemistry), while their dynamics seemed to be controlled by abiotic and biotic interactions.

Animals↗

H. A. Gleason's 'individualistic concept' and theory of animal communities: a continuing controversy.

A tradition of natural history and of the lore of early twentieth-century ecology was that organisms lived together and interacted to form natural entities or communities. Before there was a recognizable science of ecology, Mobius (1877) had provided a name 'biocoenosis' for such entities. This concept persisted in the early decades of ecological science; at an extreme it was maintained that the community had integrating capabilities and organization like those of an individual organism, hence the term organismic community. In the 1950s-1970s an alternative individualist concept, derived from the ideas of H. A. Gleason (1939), gained credence which held that communities were largely a coincidence of individualistic species characteristics, continuously varying environments and different probabilities of a species arriving on a given site. During the same period, however, a body of population based theory of animal communities became dominant which perpetuated the idea of patterns in nature based on biotic interactions among species resulting in integrated communities. This theory introduced an extended terminology and mathematical models to explain the organization of species into groups of compatible species governed by rules. In the late 1970s the premises and methods of the theory came under attack and a vigorous debate ensued. The alternatives proposed were, at an extreme, null models of random aggregations of species or stochastic, individualistic aggregations of species, sensu Gleason. Extended research and debate ensued during the 1980s resulting in an explosion of studies of animal communities and a plethora of symposia and volumes of collected works concerning the nature of animal communities. The inherent complexity of communities and the traditional differences among animal ecologists about how they should be defined and delimited, at what scale of taxa, space and time to study them, and appropriate methods of study and analysis have resulted in extended and as yet inconclusive discussions. Recent differences and discussions are considered under five general categories, evolution and community theory, individualistic concept, community definition, questions from community ecology and empirical studies. Communities are seen by some ecologists as entities of coevolving species and, in any case, it is necessary to integrate evolutionary ideas with the varied concepts of community.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Ecological impacts of atmospheric CO2 enrichment on terrestrial ecosystems.

Global change has many facets, of which land use and the change of atmospheric chemistry are unquestioned primary agents, which induce a suite of secondary effects, including climatic changes. The largest single contribution to the compositional change of the atmosphere, CO(2) enrichment, has (besides its influence on climate) immediate and direct effects on plants. Quantitatively, CO(2) is the plant 'food' number one, and the rate of photosynthetic CO(2) uptake by leaves is not yet CO(2)-saturated. This abrupt change of the biosphere's diet does and will affect all aspects of life, including our food. However, the plant and ecosystem responses are more subtle than had been assumed from the results of responses of isolated, well-fertilized and well-watered plants in greenhouses during the early days of CO(2)-enrichment research. In this article, I discuss potential responses of complex natural grassland and diverse forests, and address three key themes: CO(2) and nutrients; CO(2) and water; CO(2) and plant-animal interactions. Examples from a suite of climatic regions emphasize that the most important ecosystem level responses to elevated CO(2) will be introduced by differential responses of species. Atmospheric CO(2) enrichment is a biodiversity issue. Classical physiological baseline responses of leaves to elevated CO(2) can be overrun by biodiversity effects to such an extent that some of the traditional predictions may even become reversed. For instance, biodiversity effects may cause humid tropical forests (those which avoid destruction) to become more dynamic and store less, rather than more, carbon as CO(2) enrichment continues. The abundance of certain life forms and species and their lifespans exert major controls over the half-life of carbon stored in forest biomass, and there is evidence that elevated CO(2) can affect these controls and most likely does so already. Also, long-term hydrological consequences of atmospheric CO(2) enrichment will be driven by biodiversity effects, given that some, but not all, species reduce their rate of water loss when exposed to a CO(2)-rich atmosphere. Such insights call for much more realistic experimental conditions and larger-scale test units, which permit biotic interactions across taxa and trophic levels to occur while simulating our CO(2) future. The evidence currently available suggests that ecosystem processes reflect the composition of their biological inventory and this will be affected by a shift in carbon supply.

Animals↗

Disturbance frequency and community structure in a twenty-five year intervention study.

Models of community regulation commonly incorporate gradients of disturbance inversely related to the role of biotic interactions in regulating intermediate trophic levels. Higher trophic-level organisms are predicted to be more strongly limited by intermediate levels of disturbance than are the organisms they consume. We used a manipulation of the frequency of hydrological disturbance in an intervention analysis to examine its effects on small-fish communities in the Everglades, USA. From 1978 to 2002, we monitored fishes at one long-hydroperiod (average 350 days) and at one short-hydroperiod (average 259 days; monitoring started here in 1985) site. At a third site, managers intervened in 1985 to diminish the frequency and duration of marsh drying. By the late 1990s, the successional dynamics of density and relative abundance at the intervention site converged on those of the long-hydroperiod site. Community change was manifested over 3 to 5 years following a dry-down if a site remained inundated; the number of days since the most recent drying event and length of the preceding dry period were useful for predicting population dynamics. Community dissimilarity was positively correlated with the time since last dry. Community dynamics resulted from change in the relative abundance of three groups of species linked by life-history responses to drought. Drought frequency and intensity covaried in response to hydrological manipulation at the landscape scale; community-level successional dynamics converged on a relatively small range of species compositions when drought return-time extended beyond 4 years. The density of small fishes increased with diminution of drought frequency, consistent with disturbance-limited community structure; less-frequent drying than experienced in this study (i.e., longer return times) yields predator-dominated regulation of small-fish communities in some parts of the Everglades.

Animals↗

Global production increased by spatial heterogeneity in a population dynamics model.

Spatial and temporal heterogeneity are often described as important factors having a strong impact on biodiversity. The effect of heterogeneity is in most cases analyzed by the response of biotic interactions such as competition of predation. It may also modify intrinsic population properties such as growth rate. Most of the studies are theoretic since it is often difficult to manipulate spatial heterogeneity in practice. Despite the large number of studies dealing with this topics, it is still difficult to understand how the heterogeneity affects populations dynamics. On the basis of a very simple model, this paper aims to explicitly provide a simple mechanism which can explain why spatial heterogeneity may be a favorable factor for production. We consider a two patch model and a logistic growth is assumed on each patch. A general condition on the migration rates and the local subpopulation growth rates is provided under which the total carrying capacity is higher than the sum of the local carrying capacities, which is not intuitive. As we illustrate, this result is robust under stochastic perturbations.

Models, Theoretical↗

A multilevel modeling approach to assessing regional and local landscape features for lake classification and assessment of fish growth rates.

The ecoregion and watershed frameworks are landscape-based classifications that have been used to group waterbodies with respect to measures of community structure; however, they have yet to be evaluated for grouping lakes for demographic characteristics of fish populations. We used a multilevel modeling approach to determine if variability in mean fish length at age could be partitioned by ecoregions and watersheds. For the ecoregions analysis, we then examined if within-ecoregion variability could be explained by local water quality and lake morphometry characteristics. We used data from agency surveys conducted during 1974-1984 for age 2 and 3 fish of seven common warm and coolwater fish species. Variance in mean length at age between ecoregions for all species was not significant, and between-watershed variance estimates were only significant in 3 out of 14 analyses; however, the total amount of variation between watersheds was very small (ranging from 1.8% to 3.7% of the total variance), indicating that ecoregions and watersheds were ineffective in partitioning variability in mean length at age. Within ecoregions, water quality and lake morphometric characteristics accounted for 2%-23% of the variation in mean length at age. Measures of lake productivity were the most common significant covariates, with mean length at age increasing with increasing lake productivity. Much of the variability in mean length at age was not accounted for, suggesting that other local factors such as biotic interactions, fish density, and exploitation are important. The results indicate that the development of an effective regional framework for managing inland lakes will require a substantial effort to understand sources of demographic variability and that managers should not rely solely on ecoregions or watersheds for grouping lakes with similar growth rates.

Animals↗

Temporal variation in the production of four secondary metabolites in a colonial ascidian.

The ecological and applied importance of ascidian secondary metabolites is well reported, but little is known about temporal variation in the production of these compounds. Here, we examined seasonal variation in the production of the four main pyridoacridine alkaloids (shermilamine B, kuanoniamine D, and their deacetylated forms) of the purple morph of the ascidian Cystodytes sp. Five replicate samples were taken monthly from July 2004 to February 2006, and analyzed by high-performance liquid chromatography. Our results showed a high degree of intercolony variability in secondary metabolite concentration and a lack of statistically significant seasonal variation in the production of the four compounds. Minimum values, however, were consistently recorded over the studied years in late summer, after the reproductive period. These minima may be attributable to sexual exhaustion and seasonally varying biotic interactions or abiotic parameters.

Acridines↗

The response of stream-dwelling fish to liming.

The fish fauna in 22 limed and seven unlimed small streams was monitored using yearly electrofishing to assess the effects of liming on species occurrence and abundance. The liming techniques were divided into three main methods, lake liming, doser liming and wetland liming, to evaluate whether different strategies had different effects on the fish fauna. The predominant species at the investigated stations were salmonids, mainly brown trout (Salmo trutta) and Atlantic salmon (Salmo salar). Other species generally occurred in low numbers. The average number of fish species did not increase significantly after liming, which was probably due to recolonization difficulties caused by the frequent occurrence of migration obstacles in the streams. After liming the density of salmonids increased significantly, irrespective of the liming method. Other fish species showed no general increase, but in individual streams significant increases of European minnow (Phoxinus phoxinus) and bullhead (Cottus gobio) occurred. Some unlimed streams had acid spates with a pH below 6, which immediately lowered the numbers of salmonid parr. Other unlimed streams lost fish species progressively due to increasing acidification. pH, and probably increased levels of metals, were the major factors regulating the fish fauna. Few examples of biotic interactions were observed, but with an increase of Atlantic salmon parr after liming, brown trout abundances decreased in three streams.

Journal Article↗

Biological responses to the chemical recovery of acidified fresh waters in the UK.

We report biological changes at several UK Acid Waters Monitoring Network lakes and streams that are spatially consistent with the recovery of water chemistry induced by reductions in acid deposition. These include trends toward more acid-sensitive epilithic diatom and macroinvertebrate assemblages, an increasing proportional abundance of macroinvertebrate predators, an increasing occurrence of acid-sensitive aquatic macrophyte species, and the recent appearance of juvenile (<1 year old) brown trout in some of the more acidic flowing waters. Changes are often shown to be directly linked to annual variations in acidity. Although indicative of biological improvement in response to improving water chemistry, "recovery" in most cases is modest and very gradual. While specific ecological recovery endpoints are uncertain, it is likely that physical and biotic interactions are influencing the rate of recovery of certain groups of organisms at particular sites.

Acid Rain↗

Host manipulation of a freshwater crustacean (Gammarus roeseli) by an acanthocephalan parasite (Polymorphus minutus) in a biological invasion context.

Several gammarid species serve as intermediate hosts for the acanthocephalan parasite Polymorphus minutus. This parasite influences gammarid behaviour in order to favour transmission to its ultimate host, generally a bird. We investigated this host manipulation in Gammarus roeseli, a gammarid species introduced in France 150 years ago which now coexists with several exotic species from different origins. In the field, vertical distribution of G. roeseli revealed a higher proportion of infected individuals close to the water's surface and the size distribution of infected gammarids revealed predation pressure on infected individuals. However, under laboratory conditions both infected and non-infected individuals remained benthic. The addition of a second gammarid, Dikerogammarus villosus, to the experimental device involved a vertical displacement of infected G. roeseli towards the water's surface. Dikerogammarus villosus, originating from the Ponto-Caspian basin, can be considered as an aggressive predator. The substitution of D. villosus with Atyaephyra desmarestii, a planktivore decapod, did not alter the gammarids' distribution, with both infected and uninfected G. roeseli staying benthic. Thus, biotic interactions between D. villosus and G. roeseli represent selective pressure encouraging the expression of manipulated behaviour in infected amphipods. Through manipulation, P. minutus was found to increase the survival of infected G. roeseli when faced with non-host predators and to make it more vulnerable to predation by the parasite's definitive host.

Acanthocephala↗

Relaxed molecular clocks for dating historical plant dispersal events.

Age estimation from molecular sequences has emerged as a powerful tool for inferring when a plant lineage arrived in a particular area. Knowing the tenure of lineages within a region is key to understanding the evolution of traits, the evolution of biotic interactions, and the evolution of floras. New analytical methods model change in substitution rates along individual branches of a phylogenetic tree by combining molecular data with time constraints, usually from fossils. These "relaxed clock" approaches can be applied to several gene regions that need not all have the same substitution rates, and they can also incorporate multiple simultaneous fossil calibrations. Since 1995, at least 100 plant biogeographic studies have used molecular-clock dating, and about a fifth has used relaxed clocks. Many of these report evidence of long-distance dispersal. Meta-analyses of studies from the same geographic region can identify directional biases because of prevailing wind or water currents and the relative position and size of landmasses.

Geography↗

Species coexistence in temperate, mixed deciduous forests.

The response of tree life-history traits to community profiles (horizontal and vertical heterogeneity, disturbances and biotic interactions) determines community assembly rules, which are currently a hot issue in community ecology. Important mechanisms of coexistence differ throughout the developing stages of tree life history. Many processes of niche partitioning and tradeoffs that potentially enable tree coexistence have been reported to be present in temperate forests, although some of these life-history traits are either correlated with each other or are not independent. Not all of the proposed mechanisms explain coexistence equally well; some could predominate in determining the community organization of forest communities. Population studies need to concentrate more on the component species of a target community to detect the ecological assembly rule. These approaches can also address how chance factors contribute to the composition of temperate tree communities, which might be less dependent on chance than are tropical ones.

Journal Article↗

Compound-specific stable isotope analysis of soil mesofauna using thermally assisted hydrolysis and methylation for ecological investigations.

Stable isotope mass spectrometric approaches are proving to be valuable tools in unravelling biotic interactions in complex ecosystems, yielding information on trophic preferences and functional roles of individual species. Gas chromatography/combustion/isotope ratio mass spectrometry (GC/C/IRMS) provides considerable opportunities to assist in studies concerned with ecosystem processes mediated by soil invertebrates and microorganisms by determination of delta(13)C values of individual compounds, for example, lipids, amino acids etc. However, techniques conventionally adopted for "wet" chemical extractions and derivatizations necessary for compound-specific stable isotope determinations restrict the size of soil organism that can be studied and can limit investigations of individuals or even parts of individuals. We demonstrate here that individual soil mesofauna can be probed directly for their fatty acid stable isotope signatures by pyrolysis-GC/C/IRMS. A thermally assisted hydrolysis and methylation (THM) reaction is described for the determination of delta(13)C fatty acid values using trimethylsulfonium hydroxide (TMSH). Authentic fatty acids, acyl lipids, and individual Collembola (Folsomia candida) raised on C(3) and C(4) isotopically labeled yeast were analyzed initially by py-GC/MS with TMSH and then by py-GC/C/IRMS. A kinetic isotope effect (KIE) observed with the THM reaction prevents direct calculation of the fatty acid delta(13)C values by simple mass balance equations. However, the KIE is shown to be both reproducible and robust and can therefore be accounted for by the use of correction factors. The fatty acid methyl ester compositions of individual F. candida and their respective delta(13)C values were determined and shown to agree with those obtained by conventional "wet" chemical procedures applied to much larger numbers of Collembola, thus enhancing the scope to which stable isotopes can be applied to the study of invertebrates in complex food webs in any environment.

Animals↗

Transformation of chloropicrin and 1,3-dichloropropene by metam sodium in a combined application of fumigants.

Combined application of fumigants is a potential strategy to replace methyl bromide in the control of soil-borne pests. Unfortunately, abiotic and biotic interactions among fumigants restrict some combined application approaches. In this study, the kinetics and mechanisms of reaction between metam sodium (sodium methyldithiocarbamate) and the halogenated fumigants chloropicrin (trichloronitromethane) and 1,3-dichloropropene (1,3-D) were investigated in aqueous solution. For chloropicrin, an extremely rapid oxidation-reduction process occurred in the presence of metam sodium. The second-order rate constant for the reaction between chloropicrin and metam sodium was approximately 2 orders of magnitude greater than that for the reaction between 1,3-D isomers and metam sodium. Transformation of 1,3-D by metam sodium was associated with an aliphatic S(N)2 nucleophilic substitution process. The nucleophilic reaction of cis-1,3-D with metam sodium was significantly faster than that of the trans isomer and was correlated with a lower reaction activation energy for the cis isomer in the transition state. Combining Telone C-35 (65% 1,3-D and 35% chloropicrin) and metam sodium in solution might yield some nucleophilic sulfur species, which played an important role in the dissipation of 1,3-D. The incompatibility of chloropicrin and 1,3-D with metam sodium was also examined in soil under different application scenarios. Simultaneous application of metam sodium with chloropicrin or 1,3-D accelerated the transformation of the two halogenated fumigants, reducing their availability in soil. A sequential strategy for multiple fumigants was developed, which could be applied without the loss of active ingredient that occurs due to the reaction between fumigants. The proposed methodology may enhance pest control while maintaining environmental protection.

Allyl Compounds↗

Elevational trends in defense chemistry, vegetation, and reproduction in Sanguinaria canadensis.

Evaluation of biotic interactions along geographic gradients reveals that pressure on plant populations by herbivores and pathogens increases as latitude decreases, and is accompanied by a parallel increase in the number and toxicity of alkaloid-bearing plants. We compared rhizome alkaloid content with plant reproductive and vegetative characters in Sanguinaria canadensis (Papaveraceae) along an elevational gradient over two growing seasons to ascertain 1) if alkaloid production in bloodroot varies among populations and systematically with elevation, and 2) if there exists a correlation between isoquinoline alkaloid, vegetative and reproductive production. In general, alkaloid content in bloodroot rhizomes declines with elevation, increases with rhizome water content, varies by site, and fluctuates seasonally with plant growth and reproduction. Alkaloid content was positively correlated with vegetative and reproductive effort with few exceptions. Analysis of total protopine and benzophenanthridine alkaloid concentrations revealed generally similar patterns as those of individual alkaloid concentrations, although significant differences did appear between individual alkaloid concentrations.

Adaptation, Physiological↗

General patterns of taxonomic and biomass partitioning in extant and fossil plant communities.

A central goal of evolutionary ecology is to identify the general features maintaining the diversity of species assemblages. Understanding the taxonomic and ecological characteristics of ecological communities provides a means to develop and test theories about the processes that regulate species coexistence and diversity. Here, using data from woody plant communities from different biogeographic regions, continents and geologic time periods, we show that the number of higher taxa is a general power-function of species richness that is significantly different from randomized assemblages. In general, we find that local communities are characterized by fewer higher taxa than would be expected by chance. The degree of taxonomic diversity is influenced by modes of dispersal and potential biotic interactions. Further, changes in local diversity are accompanied by regular changes in the partitioning of community biomass between taxa that are also described by a power function. Our results indicate that local and regional processes have consistently regulated community diversity and biomass partitioning for millions of years.

Animals↗