Co-variation and indicators of species diversity: Can richness of forest-dwelling species be predicted in northern boreal forests?

Design and establishment of ecologically good networks of conservation areas often requires quick assessments of their biodiversity. Reliable indicators would be useful when doing such assessments. In order to explore the potential indicators for species richness in boreal forests, we studied (1) the co-variation of species richness and composition of species assemblages among beetles, polypores, birds and vascular plants, (2) the relationships between species richness and four boreal forest site types, (3) the relationship between species richness and forest physical structure and (4) the suitability of potential indicator groups within the four taxa to predict the species richness generally. The data show that there are probably not a single taxonomic or forest structural characteristic to be used as a general biodiversity indicator or surrogate for all the species. The correlations in species richness among the four taxa studied were low. However, group-specific indicators were obvious: forest site type was a good surrogate for vascular plant richness, and quantity and quality of dead wood predicted the species richness of polypores. The results support the view that different indicators shall be used for different forest types and taxonomic groups. These indicators should facilitate relatively rapid methods to assess biodiversity patterns at the forest stand level.     

Host-specialization and species diversity in fish parasites: phylogenetic conservatism?

The pattern of parasite species diversification and specialization, appreciated by host range, is investigated in fish parasites. We test whether host range is linked with phylogeny at a high taxonomic level, and if there is a relationship between host range and host species diversification. For this purpose we used two sets of data, one on macro-parasites of marine fishes of the Mediterranean Sea and the other on macro-parasites of marine and freshwater fishes of Canada. Similar patterns of host range among parasitic groups were found. Our findings suggest that habitat (marine vs freshwater) and geographic localization (Canada vs Mediterranean region) play little role in determining the observed patterns of host range. We highlight the potential influence of phylogeny (high-taxonomic level) on the level host range in parasites. We find that parasites with free-swimming larval stages and with direct life cycles have a narrower range of host species than do parasites with indirect life cycle, even if we cannot control for phylogenetic effects because of the lack of variation of life cycles within each parasitic group. Finally, a positive relationship was found between the number of known hosts and parasite species diversity in the case of Mediterranean parasite species. The relationship between host range and species diversification should be related to the mechanism of cospeciation.     

Invasion-resistance in experimental grassland communities: species richness or species identity?

The question as to why some communities are more invasible than others has pro-found implications for conservation biology and land management. The theoretical issues involved go right to the heart of our understanding of species coexistence and community assembly. The experiment reported here indicates that for productive, small-scale grassland plots, species identity matters more than species richness in determining both the number of invading species and the total biomass of invasives.     

Nutrient limitation of plant productivity in scrubby flatwoods: does fire shift nitrogen versus phosphorus limitation?

Differences in the biogeochemistry of nitrogen (N) and phosphorus (P) lead to differential losses and inputs during and over time after fire such that fire may affect nutrient limitation of primary productivity. We conducted a nutrient addition experiment in scrubby flatwoods, a Florida scrub community type, to test the hypothesis that nutrient limitation of primary productivity shifts from N limitation in recently burned sites to P limitation in longer unburned sites. We added three levels of N, P, and N and P together to sites 6 weeks, 8 years, and 20 years postfire and assessed the effects of nutrient addition on above- and belowground productivity and nutrient concentrations. At the community level, nutrient addition did not affect aboveground biomass, but root productivity increased with high N?+?P addition in sites 8 and 20 years after fire. At the species level, N addition increased leaf biomass of saw palmetto (Serenoa repens) in sites 6 weeks and 20 years postfire, while P addition increased foliar %P and apical shoot growth of scrub oak (Quercus inopina) in sites 8 and 20 years postfire, respectively. Contrary to our hypothesis, nutrient limitation does not appear to shift with time after fire; recently burned sites show little evidence of nutrient limitation, while increased belowground productivity indicates that scrubby flatwoods are co-limited by N and P at intermediate and longer times after fire.     

Nitrogen limitation on land and in the sea: How can it occur?

The widespread occurrence of nitrogen limitation to net primary production in terrestrial and marine ecosystems is something of a puzzle; it would seem that nitrogen fixers should have a substantial competitive advantage wherever nitrogen is limiting, and that their activity in turn should reverse limitation. Nevertheless, there is substantial evidence that nitrogen limits net primary production much of the time in most terrestrial biomes and many marine ecosystems. We examine both how the biogeochemistry of the nitrogen cycle could cause limitation to develop, and how nitrogen limitation could persist as a consequence of processes that prevent or reduce nitrogen fixation. Biogeochemical mechansism that favor nitrogen limitation include:

the substantial mobility of nitrogen across ecosystem boundaries, which favors nitogen limitation in the “source” ecosystem — especially where denitrification is important in sediments and soils, or in terrestrial ecosystems where fire is frequent;

differences in the biochemistry of nitrogen as opposed to phosphorus (with detrital N mostly carbon-bonded and detrital P mostly ester-bonded), which favor the development of nitrogen limitation where decomposition is slow, and allow the development of a positive feedback from nitrogen limitation to producers, to reduced decomposition of their detritus, and on to reduced nitrogen availability; and

other more specialized, but perhaps no less important, processes.

A number of mechanisms could keep nitrogen fixation from reversing nitrogen limitation. These include:

energetic constraints on the colonization or activity of nitrogen fixers;

limitation of nitrogen fixers or fixation by another nutrient (phosphorus, molybdenum, or iron) — which would then represent the ultimate factor limiting net primary production;

other physical and ecological mechanisms.

The possible importance of these and other processes is discussed for a wide range of terrestrial, freshwater, and marine ecosystems.     

How can we preserve and restore species richness of pollinating insects on agricultural land?

During recent decades, concern about the loss of biodiversity on agricultural land has increased, and semi‐natural grasslands have been highlighted as critical habitats. Temperate European agricultural landscapes require distinct and appropriate management to prevent further impoverishment of the flora and fauna. This is especially urgent for pollinating insects that provide important ecosystem services. Our aim was to examine how species richness of three important groups of pollinating insects; solitary bees, butterflies and burnet moths are related to different farm characteristics, and if there are any differences between these three groups. A further aim was to test if red‐listed species are related to any farm characteristics. Species richness of solitary bees, butterflies and burnets was measured on all semi‐natural grasslands at 16 farms in a forest‐dominated area of 50 km² in southern Sweden, using systematic transect walks in April to September 2003 (only butterflies and burnets) and 2005. Species richness of solitary bees and butterflies was intercorrelated, both before and after controlling for the area of semi‐natural grassland. Species richness of solitary bees increased with the area of semi‐natural grassland. After controlling for the effect of the area of semi‐natural grassland species richness was strongly positively related with the density of the plant Knautia arvensis and negatively related with the proportion of grazed grassland. The results were similar for solitary bees and butterflies. The number of red‐listed solitary bees was positively related to the proportion of meadows with late harvest (after mid‐July) and decreased with increased farm isolation. The number of burnet species (all red‐listed) was positively related to vegetation height, flower density and the proportion of meadows with late harvest on a farm. Areas with a high density of K. arvensis and with traditional hay‐meadow with late harvest present, harbour most species. Promoting traditional hay‐meadows, late extensive grazing and the herb K. arvensis, people managing agricultural biodiversity can encompass high species richness of pollinating insects and support red‐listed species. Further, we suggest that the density of K. arvensis at a farm can be used as a biodiversity indicator, at least for pollinating insects.     

Overyielding and species diversity: what should we expect?

Recent empirical studies have found evidence of increased biomass production ('overyielding') in species mixtures relative to monoculture, but the interpretation of these results remains controversial, in part, because of the lack of a theoretical expectation. Here, we examined the expected frequency and stability of overyielding species mixtures using Lotka-Volterra models of species dynamics in two- and four-species systems in conjunction with community, population, and specific rate of biomass production (SRP) definitions of overyielding. Overyielding plant mixtures represented > 55% of potential species assemblages under community definitions and approximately 100% of species were either overyielding or underyielding under the population definition. Our species simulations approached their equilibria in 1-2 yr, supporting the relevancy of an equilibrial analysis. The range of parameter space that we explored produced realistic values of plot biomass, supporting their biological relevance. We show that overyielding is expected to be common under community definitions and population definitions. Overyielding, under community or population definitions, does not imply an actual increase in the specific rate of biomass production. In addition, assemblages of overyielding and underyielding species under all three definitions can be stable over time with underyielding species persisting in the presence of overyielding species.     

Does coevolution promote species richness in parasitic cuckoos?

Why some lineages have diversified into larger numbers of species than others is a fundamental but still relatively poorly understood aspect of the evolutionary process. Coevolution has been recognized as a potentially important engine of speciation, but has rarely been tested in a comparative framework. We use a comparative approach based on a complete phylogeny of all living cuckoos to test whether parasite–host coevolution is associated with patterns of cuckoo species richness. There are no clear differences between parental and parasitic cuckoos in the number of species per genus. However, a cladogenesis test shows that brood parasitism is associated with both significantly higher speciation and extinction rates. Furthermore, subspecies diversification rate estimates were over twice as high in parasitic cuckoos as in parental cuckoos. Among parasitic cuckoos, there is marked variation in the severity of the detrimental effects on host fitness; chicks of some cuckoo species are raised alongside the young of the host and others are more virulent, with the cuckoo chick ejecting or killing the eggs/young of the host. We show that cuckoos with a more virulent parasitic strategy have more recognized subspecies. In addition, cuckoo species with more recognized subspecies have more hosts. These results hold after controlling for confounding geographical effects such as range size and isolation in archipelagos. Although the power of our analyses is limited by the fact that brood parasitism evolved independently only three times in cuckoos, our results suggest that coevolutionary arms races with hosts have contributed to higher speciation and extinction rates in parasitic cuckoos.     

Nitrogen limitation of decomposition and decay: How can it occur?

The availability of nitrogen (N) is a critical control on the cycling and storage of soil carbon (C). Yet, there are conflicting conceptual models to explain how N availability influences the decomposition of organic matter by soil microbial communities. Several lines of evidence suggest that N availability limits decomposition; the earliest stages of leaf litter decay are associated with a net import of N from the soil environment, and both observations and models show that high N organic matter decomposes more rapidly. In direct contrast to these findings, experimental additions of inorganic N to soils broadly show a suppression of microbial activity, which is inconsistent with N limitation of decomposition. Resolving this apparent contradiction is critical to representing nutrient dynamics in predictive ecosystem models under a multitude of global change factors that alter soil N availability. Here, we propose a new conceptual framework, the Carbon, Acidity, and Mineral Protection hypothesis, to understand the effects of N availability on soil C cycling and storage and explore the predictions of this framework with a mathematical model. Our model simulations demonstrate that N addition can have opposing effects on separate soil C pools (particulate and mineral‐protected carbon) because they are differentially affected by microbial biomass growth. Moreover, changes in N availability are frequently linked to shifts in soil pH or osmotic stress, which can independently affect microbial biomass dynamics and mask N stimulation of microbial activity. Thus, the net effect of N addition on soil C is dependent upon interactions among microbial physiology, soil mineralogy, and soil acidity. We believe that our synthesis provides a broadly applicable conceptual framework to understand and predict the effect of changes in soil N availability on ecosystem C cycling under global change.     

Biotic resistance in freshwater fish communities: species richness,saturation or species identity?

Some communities are susceptible to invasions and some are not. Why? Elton suggested in 1958 that the ability of the community to withstand invading species – its biotic resistance – depends on the number of resident species. Later contributors have emphasized the habitat's ability to support species, as well as the contribution of individual species to the resistance. In this study we use information from 184 introductions of Arctic char into Swedish lakes to study both abiotic and biotic aspects of the resident community's ability to resist introductions. We find that the best model included the proportion of forest cover and the proportion of agricultural land cover in the watershed in combination with the presence versus absence of northern pike. Thus, the most important biotic factor to explain the outcome of introductions of Arctic char is the presence of northern pike, a large piscivore. This means that one single species explains the outcome of the introductions better than does the species richness or the saturation level of the community.     

Elevational gradients and species richness: do methods change pattern perception?

Aim Species richness patterns along elevational gradients have been documented extensively. Yet, the implications of differences in how the data are compiled are seldom explored. We investigate the effect of grain size on the richness–elevation relationship. Grain size varies among the principal methods used to collect or aggregate species occurrences: localized sites, elevational ‘bins’ and interpolation of species ranges. Assumptions of sampling and species distributions also vary among these methods. Methodology can influence the pattern that is perceived and comparability of results. We compare patterns from all three methods explicitly using the same suite of observations, based on museum records and field surveys of non‐flying small mammals. Our assessment is enhanced by comparing patterns resulting from each method for each of six adjacent mountain ranges. Location Utah, North America. Methods We document elevational species richness patterns using generalized linear models (GLMs), comparing the general shape of the trend as well as curvature, location and magnitude of peak richness across methods, both within and among gradients. We also introduce a new procedure to test for richness peaks using site‐based occurrences. Results We find a general congruence of the richness–elevation relationship, depicting a hump‐shaped pattern with a second‐order polynomial GLM showing a significant fit to nearly all gradient‐methodology combinations. However, underlying characteristics of the trend may vary with grain size. As grain size coarsens, maximum species richness increases and elevation of the mode slightly decreases. Results for curvature vary, but degree of curvature tends to increase as grain size coarsens. The richness–elevation patterns are independent of sampling effects. Main conclusions The perceived elevational diversity pattern for small mammals along these mountain ranges is not scale‐dependent. Differences in how the data are compiled are not reflected in major differences in patterns, even when local samples are neither uniformly spaced nor sampled with the same intensity. This result lends confidence to the assertion that patterns documented in similar studies with different methodologies and for which sampling is sufficiently comprehensive are good indicators of diversity. However, consistency of results from more than one compilation method may help to address issues of scale‐dependence, more so when these comparisons are made explicit.     

What does species richness tell us about functional trait diversity? Predictions and evidence for responses of species and functional trait diversity to land‐use change

In the conservation literature on land‐use change, it is often assumed that land‐use intensification drives species loss, driving a loss of functional trait diversity and ecosystem function. Modern research, however, does not support this cascade of loss for all natural systems. In this paper we explore the errors in this assumption and present a conceptual model taking a more mechanistic approach to the species–functional trait association in a context of land‐use change. We provide empirical support for our model's predictions demonstrating that the association of species and functional trait diversity follows various trajectories in response to land‐use change. The central premise of our model is that land‐use change impacts upon processes of community assembly, not species per se. From the model, it is clear that community context (i.e. type of disturbance, species pool size) will affect the response trajectory of the relationship between species and functional trait diversity in communities undergoing land‐use change. The maintenance of ecosystem function and of species diversity in the face of increasing land‐use change are complementary goals. The use of a more ecologically realistic model of responses of species and functional traits will improve our ability to make wise management decisions to achieve both aims in specific at‐risk systems.     

Does richness of emergent plants affect CO2 and CH4 emissions in experimental wetlands?

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Do protected areas in urban and rural landscapes differ in species diversity?

Previous studies from Central Europe and North America showed that species richness is higher in urban than in rural landscapes. Do protected areas, which can be found in both city and countryside, reflect this species richness pattern? The impact of urban land-use might reduce conservation success and necessitate special management strategies. We compared species richness and species spatial turnover of selected animal and plant taxa (carabids, butterflies, snails, birds, lichens, mosses, vascular plants) in 30 protected areas in the city of Halle and 56 protected areas in the adjacent rural district of Saalkreis (Central Germany). Species were mapped by experienced biologists within a systematic species inventory. We corrected species numbers for the effects of landscape structure (e.g. size, shape and distance of habitats) which might influence species diversity beyond urbanisation effects. Butterflies, birds and lichens had significantly higher species numbers in the rural protected areas. Species spatial turnover was higher among urban areas than among rural areas or pairs of urban and rural areas for most taxa. Diversity in all taxa depended on the size of a protected area. We discussed these patterns in the context of the general urban-rural species diversity patterns. Our results indicate an increasing isolation of species assemblages with urbanisation and highlight that space for protected areas is even more limited in urban than rural areas. An effective conservation of urban species diversity should include both typical urban and semi-natural habitats to cover the full range of species living in cities.     

Is there a trade-off between species diversity and genetic diversity in forest tree communities?

The two most important components of biodiversity, species diversity and genetic diversity, have generally been treated as separate topics, although a coordination between both components is believed to be critical for ecosystem stability and resilience. Based on a new trait concept that allows for the assessment of genetic diversity across species, the relationship between species diversity and genetic diversity was examined in eight forest tree communities composed of different tree genera including both climax and pioneer species. It was intended to check whether a trade-off exists between the two diversity components as was found in a few studies on animal species.Using several isozyme-gene systems as genetic markers, the genetic diversity across species within each of the tree communities was determined by two measures, the commonly used intraspecific genetic diversity averaged over species and the recently developed transspecific genetic diversity per species. Both data sets were compared with the corresponding community-specific species diversity resulting in a positive relationship between the two diversity components. A statistically significant positive correlation was established between the transspecific genetic diversity per species and the species diversity for three isozyme-gene systems. Beyond that, consistent results were obtained using different parameters of the diversity measure which characterize the total, the effective and the number of prevalent variants. The number of prevalent variants reflected most significantly the non-randomness of the observed diversity patterns.These findings can be explained by the observation that the pioneer tree species reveal a by far higher genetic diversity than the climax tree species, which means that an increase in species diversity, due to the addition of several pioneer species at the expense of one or two climax species, goes along with an increase in the level of genetic diversity. Forest tree communities with the highest degree of species diversity exhibit therefore the highest transspecific genetic diversity per species. This result was discussed with regard to the particular composition and stability of forest tree communities.