Genotypic selection shapes patterns of within-species diversity in experimental plant populations

1.  An increasing body of evidence suggests that within-species diversity plays an important role for community and ecosystem functioning, alters complex trophic interactions and affects patterns of species diversity and coexistence. Nonetheless, we lack a good understanding of how genotypic trait variation translates into shifts in the relative abundance of genotypes within populations.
2.  In this study, we show that genotypic selection strongly alters dominance relationships among genotypes over a period of 5 years. This resulted in remarkably consistent changes in the proportional representation of genotypes, and in a concomitant decline of diversity and evenness in our experimental populations.
3.  High growth rates and the production of large offspring were positively associated with genotypic performance. Vegetative abundances of genotypes translated monotonically into flowering frequencies.
4.   Synthesis . We conclude that genotypic selection markedly affects patterns of diversity and consistently alters genotypic abundance and mean trait distributions in plant populations over a relatively short period of time.     

Effects of hydrological variation and planktivorous competition on macroinvertebrate community structure in experimental billabongs

1. To study two factors which are predicted as causing changes to community structure in cut-off meanders (colloquially known in Australia as billabongs, a term of aboriginal origin), 16 experimental billabongs were constructed. These were designed to test two hypotheses: (a) that the structure of macrophyte and invertebrate communities within billabongs is altered by changing the pattern of flooding; and (b) that the presence of small planktivorous fish alters invertebrate community structure and diversity within billabongs.
2. An increase in the duration of flooding seems to favour animals better adapted to a greater availability of macrophyte habitat. Changing the seasonality of flooding resulted in prolonging of the time water was available over the summer months.
3. The presence of a planktivorous fish appears to affect macroinvertebrate communities through competition with other planktivores. Variable top-down pressure may create differing successional patterns and ultimately different communities at lower trophic levels.     

Phyloecology of urban alien floras

1.  Understanding the mechanisms that affect invasion success of alien species is a major issue in current ecological research. Although many studies have searched for either functional or habitat attributes that drive invasion mechanisms, few researchers have addressed the role of phylogenetic diversity of alien species.
2.  Here, using data from 21 urban floras located in Europe and eight in the USA, we show that the phylogenetic diversity of alien species is significantly lower than that of native species, both at the continental scale and at the scale of single cities.
3.  Second, we show that if archaeophytes and neophytes (non-native species introduced into Europe before and after AD 1500, respectively) are analysed separately, archaeophytes show lower phylogenetic diversity than neophytes, while the phylogenetic structure of neophytes is indistinguishable from a random sample of species from the entire species pool.
4.  Our results suggest that urban aliens are subject to environmental filters that constrain their phylogenetic diversity, although these filters act more strongly upon archaeophytes than neophytes.
5.   Synthesis. Despite the huge taxonomic diversity of plants imported into European and American cities, the strong environmental filters imposed by cities constrain the functional diversity of urban floras, which is reflected in their generally low phylogenetic diversity. Urban alien floras are mainly composed of phylogenetically related species that are well adapted to anthropogenic habitats, although these filters are stronger for species groups with longer residence times.     

How does habitat diversity affect the species–area relationship?

Aim   To examine the way in which 'area' and 'habitat diversity' interact in shaping species richness and to find a simple and valid way to express this interaction.
Location   The Natura 2000 network of terrestrial protected areas in Greece, covering approximately 16% of the national territory.
Methods   We used the Natura 2000 framework, which provides a classification scheme for natural habitat types, to quantify habitat heterogeneity. We analysed data for the plant species composition in 16,143 quadrats in which 5044 species and subspecies of higher plants were recorded. We built a simple mathematical model that incorporates the effect of habitat diversity on the species–area relationship (SAR).
Results   Our analysis showed that habitat diversity was correlated with area. However, keeping habitat diversity constant, species richness was related to area; while keeping area constant, species richness was related to habitat diversity. Comparing the SAR of the 237 sites we found that the slope of the species–area curve was related to habitat diversity.
Main conclusions   Discussion of the causes of the SAR has often focused on the primacy of area per se versus habitat heterogeneity, even though the two mechanisms are not mutually exclusive and should be considered jointly. We find that increasing habitat diversity affects the SAR in different ways, but the dominant effect is to increase the slope of the SAR. While a full model fit typically includes a variety of terms involving both area and habitat richness, we find that the effect of habitat diversity can be reduced to a linear perturbation of the slope of the species accumulation curve.     

Consequences of removing a keystone herbivore for the abundance and diversity of arthropods associated with a cruciferous shrub

Abstract. 1. The effect of the removal of Timarcha lugens (Chrysomelidae), one of the main herbivores of Hormathophylla spinosa (Cruciferae), on the abundance of co-occurring phytophagous insects, the abundance of non-phytophagous arthropods (detritivores, predators, and parasitoids), and the structure and diversity of the entire arthropod community, was studied for 3 years (1999–2001).
2. There was competition between T. lugens and co-occurring herbivores; the removal of T. lugens was correlated with an increase in the abundance of sap-suckers, flower-feeders, and, above all, folivores.
3. Timarcha lugens also had an indirect effect on arthropods belonging to other trophic levels; the abundance of predators increased significantly after the removal of T. lugens .
4. Community composition was affected by the experimental removal. In addition, the diversity of the overall community increased after removal of T. lugens .
5. The study demonstrated experimentally that T. lugens has a significant effect not only on other species belonging to the same trophic level, but also on the abundance of species belonging to higher trophic levels, and, consequently, on the entire structure and diversity of the complex community in which it is immersed.     

Microhabitat specialization in a species-rich palm community in Amazonian Ecuador

1 Relationships between microhabitat variables (altitude, inclination, topographic position, drainage, canopy height) and the distribution and abundance of palms and palm-like plants in 50 ha of old-growth terra firme rain forest in the Yasuní National Park, lowland Amazonian Ecuador, were examined using 118 20 × 20 m plots laid out in a stratified random design.
2 If microhabitat niche differentiation is important for maintaining the species richness of the community, then (i) the distribution of the palms will be strongly influenced by microhabitat heterogeneity and (ii) palms of similar growth form will show antagonistic microhabitat relationships.
3 Mantel and cluster analyses showed that palm species distributions were strongly structured by topography. The main difference in species composition was between plots in the bottomland and plots on the upper slopes and hill tops.
4 Logistic and logit analyses showed that 20 of the 31 palm and palm-like taxa analysed had distributions that were significantly related to the microhabitat variables measured, mainly to topography but also to drainage and canopy height.
5 Spatial autocorrelation in the overall community structure was not explained by the microhabitat variables. Analyses of distributions or abundances of single species showed neighbourhood effects for seven taxa.
6 Antagonistic patterns of microhabitat preferences were recognizable among some species pairs of small palms, medium-sized palms and palm-like plants, but not among canopy palms.
7 It is concluded that microhabitat specialization is an important factor in maintaining the diversity of this palm community, while mass effects might also be important.     

Community structure of arboreal caterpillars within and among four tree species of the eastern deciduous forest

Abstract.  1. A seasonally replicated experimental design was used to address the question of how differences within and among host tree species affect arboreal caterpillar communities.
2. Seasonal variation influenced caterpillar community composition most significantly, and the similarity among caterpillar assemblages did not necessarily follow the pattern of phylogenetic relatedness among host trees.
3. Species richness and abundance of caterpillars were higher on oaks and maples than on American beech. Diversity partitioning models revealed that β diversity was only occasionally greater or less than expected by chance alone.
4. When β diversity was significant, values tended to be greater than expected by chance among replicate trees within each species and lower than expected by chance among the four tree species.
5. Differences among trees appeared important for determining patterns of species presence/absence for rare species and influencing patterns of species dominance within caterpillar assemblages. Differences among tree species had a significant effect on overall lepidopteran community composition and mean species diversity (i.e. α diversity).
6. Because β diversity of caterpillars among host trees was lower than expected by chance, host specificity within the Lepidoptera may be less prevalent than thought previously.     

Odonates as biological indicators of grazing effects on Canadian prairie wetlands

Abstract.  1. Aquatic macro-invertebrates have frequently been used as biological indicators in lotic environments but much less commonly so in lentic habitats. Dragonflies and damselflies (Order Odonata) satisfy most selection criteria for lentic bioindicators of grazing impacts.
2. Intensive cattle grazing affects most of the Canadian prairie pothole region but the effects of grazing on wetlands are poorly understood.
3. Here the vegetation structure and invertebrate community composition of 27 prairie potholes in Alberta, Canada were studied and compared. Wetlands were evenly divided into three treatments of different grazing regimes.
4. Removal of emergent vegetation by cattle grazing decreased odonate abundance and reproductive effort. Shorter Scirpus acutus stems resulted in significantly fewer damselflies (Suborder Zygoptera) and lower reproductive efforts.
5. Overall odonate diversity was affected by the height of key plant species, highlighting the importance of the vegetation structure of both emergent vegetation for breeding and adjacent upland vegetation for nocturnal roosts. Wetland vegetation structure was more important than vegetation composition to the life history of odonates.
6. Wetland water quality parameters of nitrogen, phosphorus, total dissolved solids (TDS), and chlorophyll-a concentration did not change due to the presence of grazing cattle at wetlands so water quality influences were rejected as mechanisms of change.
7. Larval odonate diversity and abundance was positively correlated with overall aquatic macro-invertebrate diversity and abundance, hence it was concluded that the larval odonate community can be an accurate bioindicator of intactness and diversity of overall aquatic macro-invertebrate communities in Canadian prairie wetlands.     

Comparative demographic impacts of `info-chemicals' and exploitative competition: an empirical test using Daphnia magna

1. Crowding `info-chemicals', metabolites released into the environment that act as signals to conspecifics as well as other organisms, have often been shown to cause phenotypic plasticity in cladoceran life-history traits including morphology, reproductive strategy and sex induction. Effects on population demography and diversity, however, have not been examined directly.
2. Replicate laboratory populations of the zooplankter, Daphnia magna, were started with 250 juveniles and allowed to develop for 8 weeks in 10 L flow-through aquaria with food levels of 1 mg CL^–1. Two crowded water treatments from different clonal sources were compared with untreated water as a control.
3. The presence of crowding chemicals affected initial population structure, through reduction in parthenogenetic female body lengths, brood sizes and juvenile densities. However, the timing of population density maxima and subsequent population structure, clonal composition and diversity were similar among all treatments.
4. Clonal richness remained slightly higher in untreated populations because of increased prevalence of rare clones.
5. Exploitation competition had greater impact on population structure than the relative concentration of crowding chemicals, indicating that density dependent effects were mediated primarily by competition for common resources.
6. Crowding info-chemicals may play a greater role in community structure than in single species population structure because of taxonomic diversity in species' response that may be unrelated to the source species density.     

Additive apportioning of lepidopteran and coleopteran species diversity across spatial and temporal scales in a cool–temperate deciduous forest in Japan

Abstract.  1. Forest entomofauna retain high diversity, and examining beta diversity, or species turnover, among assemblages in a forest community is vital to elucidate the source of this diversity.
2. Under the DIVERSITAS in Western Pacific and Asia–International Biodiversity Observation Year (DIWPA–IBOY) project for simultaneously documenting biodiversity throughout the Western Pacific and Asian Region, 892 lepidopteran species (51 742 specimens) and 355 coleopteran species (11 633 specimens) were collected in 2001 by light traps in a cool–temperate forest in northern Japan.
3. This study evaluated the beta diversity of lepidopteran and coleopteran communities by ecological categories (i.e. trap location, forest strata, sampling days, and months), and assessed the habitat preferences of lepidopteran and coleopteran species.
4.  anova -like additive apportioning models were used to quantify the beta diversity among the categories. The models simultaneously provide assessments of whether species distributions are biased in favour of particular habitats.
5. Significantly high beta diversity was observed among months for both Lepidoptera and Coleoptera. The category of months corresponded fairly well to the number of specialist species detected in the category, although a remarkably large number of significant specialists in Coleoptera were observed on strata.
6. The high beta diversity and number of specialist species among strata in both communities indicate that stratification between canopy and ground, and seasonal variation, played major roles in species composition and the rich entomofauna in the forest. Highly mobile adults were influenced by the vertical spatial scale, as previously suggested for larvae.     

Ecosystem engineering across ecosystems: do engineer species sharing common features have generalized or idiosyncratic effects on species diversity?

Aim  To integrate the effects of ecosystem engineers (organisms that create, maintain or destroy habitat for other species) sharing the same archetype on species diversity, and assess whether different engineer species have generalized or idiosyncratic effects across environmentally similar ecosystems.
Location  High-Andean habitats of Chile and Argentina, from 23° S to 41° S.
Methods  We measured and compared the effects of eight alpine plants with cushion growth-form on species richness, species diversity (measured as the Shannon–Wiener index) and evenness of vascular plant assemblages across four high-Andean ecosystems of Chile and Argentina.
Results  The presence of cushion plants always increased the species richness, diversity (measured as the Shannon–Wiener index) and evenness of high-Andean plant assemblages. However, while the presence of different cushion species within the same ecosystem controlled species diversity in the same way, these effects varied between cushion species from different ecosystems.
Main conclusions  Results consistently supported the idea that increases in habitat complexity due to the presence of ecosystem engineers, in this case cushion plants, would lead to higher community diversity. Results also indicate that effects of the presence of different cushion species within the same ecosystem could be generalized, while the effects of cushion species from different ecosystems should be considered idiosyncratic.     

Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses

Questions: What is the observed relationship between plant species diversity and spatial environmental heterogeneity? Does the relationship scale predictably with sample plot size? What are the relative contributions to diversity patterns of variables linked to productivity or available energy compared to those corresponding to spatial heterogeneity? Methods: Observational and experimental studies that quantified relationships between plant species richness and within‐sample spatial environmental heterogeneity were reviewed. Effect size in experimental studies was quantified as the standardized mean difference between control (homogeneous) and heterogeneous treatments. For observational studies, effect sizes in individual studies were examined graphically across a gradient of plot size (focal scale). Relative contributions of variables representing spatial heterogeneity were compared to those representing available energy using a response ratio. Results: Forty‐one observational and 11 experimental studies quantified plant species diversity and spatial environmental heterogeneity. Observational studies reported positive species diversity‐spatial heterogeneity correlations at all points across a plot size gradient from ～1.0 × 10^?1 to ～1.0 × 10¹¹ m², although many studies reported spatial heterogeneity variables with no significant relationships to species diversity. The cross‐study effect size in experimental studies was not significantly different from zero. Available energy variables explained consistently more of the variance in species richness than spatial heterogeneity variables, especially at the smallest and largest plot sizes. Main conclusions: Species diversity was not related to spatial heterogeneity in a way predictable by plot size. Positive heterogeneity‐diversity relationships were common, confirming the importance of niche differentiation in species diversity patterns, but future studies examining a range of spatial scales in the same system are required to determine the role of dispersal and available energy in these patterns.     

The non-indigenous diatom Didymosphenia geminata alters benthic communities in New Zealand rivers

1. Blooms of the benthic, stalked diatom Didymosphenia geminata were first observed in New Zealand in 2004. Since then, D. geminata has spread to numerous catchments in the South Island and is also spreading in its native range. The species is a rare example of an invasive alga in lotic systems.
2. Ecosystem effects may be expected as D. geminata attains unusually high biomass in rivers. We examined data from three independent studies in three South Island, New Zealand, rivers for evidence of effects on periphyton biomass and benthic invertebrate communities.
3. The combined results confirmed that the presence of D. geminata was associated with greatly increased periphyton biomass and, in most cases, increased invertebrate densities. We also recorded shifts in community composition, dominated by increased densities of Oligochaeta. Chironomidae, Cladocera and Nematoda also generally increased in density with D. geminata . Significant increases or declines in other invertebrate taxa were inconsistent among rivers.
4. In all three studies, increased spatial invertebrate community homogeneity was associated with high D. geminata biomass at the within-river scale. However, no declines in taxon richness or diversity were detected.
5. Although ecosystem effects of D. geminata on existing periphyton biomass and invertebrate communities are measurable, no inferences can be made from the present data about effects on higher trophic levels (fish).     

Temporal patterns in shore fly (Diptera, Ephydridae) community structure in a salt marsh habitat

Abstract.  1. An ecological investigation of the shore fly community was carried out in a salt marsh in Novosedly in the Czech Republic.
2. Sampling was carried out in 1997 and 1999 at approximately 14-day intervals during the seasonally active period, using yellow water traps.
3. Altogether 14 374 specimens of the family Ephydridae were collected (6040 in 1997 and 8334 in 1999), belonging to 48 shore fly species (37 in 1997 and 41 in 1999).
4. Community structure, species richness, species diversity, evenness, and seasonal and inter-annual changes were studied. The species richness, species diversity, and evenness of the shore fly community in 1999 were higher than in 1997 but inter-annual difference in species diversity was not statistically significant. Paracoenia fumosa , Scatella tenuicosta , and Hydrellia albilabris were found to be the most abundant species in both study years.
5. Seasonal patterns and phenology of the most abundant species are discussed. Hydrellia albilabris showed similar seasonal distribution of abundance in 1997 and 1999, whereas seasonal distribution of P. fumosa and S. tenuicosta varied between the study years. Paracoenia fumosa , Hydrellia griseola , H. albilabris , Scatella paludum , S. tenuicosta , and Philotelma defecta were found to occur throughout the whole observation period from March until November while Notiphila cinerea , Hydrellia ischiaca , and Psilopa nigritella appeared in a limited period only.     

The effects of disturbance events on phytoplankton community structure in a small temperate reservoir

1 The effects of disturbances, in the form of storm events, on phytoplankton community structure were examined over the course of four years in Eau Galle Reservoir, Wisconsin, USA.
2 Disturbances consistently brought about significant, but highly transient, increases in apparent phytoplankton species richness. It is likely that these resulted from temporary increases in the biomass of previously undetected rare species.
3 Substantial shifts in community dominance were confined to large, early season events, and were seldom long-lived. Later 'climax' communities were highly resistant to any changes in dominance, even when increases in species richness occurred.
4 Regardless of when they occurred, disturbances tended to favour species from a narrow range of the successional sequence.     

Experimental evidence that soil heterogeneity enhances plant diversity during community assembly

Aims Environmental heterogeneity is a primary mechanism explaining species coexistence and extant patterns of diversity. Despite strong theoretical support and ample observational evidence, few experimental studies in plant communities have been able to demonstrate a causal link between environmental heterogeneity and plant diversity. This lack of experimental evidence suggests that either fine-scale heterogeneity has weak effects on plant diversity or previous experiments have been unable to effectively manipulate heterogeneity. Here, we utilize a unique soil manipulation to test whether fine-scale soil heterogeneity will increase plant richness through species sorting among experimental patch types.Methods This experiment was conducted in the tallgrass prairie region of south-central Kansas, USA. We utilized the inherent variation found in the vertical soil profile, which varied in both biotic and abiotic characteristics, and redistributed these strata into either homogeneous or heterogeneous spatial arrangements in 2.4×2.4 m plots. After the soil manipulation, 34 native prairie species were sown into all plots. We conducted annual censuses at peak biomass to quantify species composition and plant density by species within the experimental communities.Important findings After 2 years, species richness was significantly higher in heterogeneous relative to homogeneous plots and this pattern was independent of total plant density. In the heterogeneous plots, 13 species had higher establishment in a specific patch type representing one of the three soil strata. Conversely, no species had greater establishment in the mixed stratum, which comprised the homogeneous plots, relative to the heterogeneous strata. These species sorting patterns suggest that fine-scale heterogeneity creates opportunities for plant establishment due to niche differences, which translates into increased plant diversity at the plot scale. Species richness was more strongly related to plant density among patches comprising homogenous plots—where fine-scale heterogeneity was minimized, but weak in heterogeneous plots. This pattern is consistent with the idea that richness–density relationships dominate when neutral processes are important but are weak when niche processes operate. Unlike many previous attempts, our results provide clear, experimental evidence that fine-scale soil heterogeneity increases species richness through species sorting during community assembly.     

Functional benefits of predator species diversity depend on prey identity

Abstract.  1. Determining the functional significance of species diversity in natural enemy assemblages is a key step towards prediction of the likely impact of biodiversity loss on natural pest control processes. While the biological control literature contains examples in which increased natural enemy diversity hinders pest control, other studies have highlighted mechanisms where pest suppression is promoted by increased enemy diversity.
2. This study aimed to test whether increased predator species diversity results in higher rates of predation on two key, but contrasting, insect pest species commonly found in the rice ecosystems of south-east Asia.
3. Glasshouse experiments were undertaken in which four life stages of a planthopper ( Nilaparvata lugens ) and a moth ( Marasmia patnalis ) were caged with single or three-species combinations of generalist predators.
4. Generally, predation rates of the three-species assemblages exceeded expectation when attacking M. patnalis , but not when attacking N. lugens. In addition, a positive effect of increased predator species richness on overall predation rate was found with M. patnalis but not with N. lugens .
5. The results are consistent with theoretical predictions that morphological and behavioural differentiation among prey life stages promotes functional complementarity among predator species. This indicates that emergent species diversity effects in natural enemy assemblages are context dependent; they depend not only on the characteristics of the predators species, but on the identity of the species on which they prey.     

Linking species diversity to the functioning of ecosystems: on the importance of environmental context

There is currently much interest in understanding how loss of biodiversity might alter ecological processes vital to the functioning of ecosystems. Unfortunately, ecologists have reached little consensus regarding the importance of species diversity to ecosystem functioning because empirical studies have not demonstrated any consistent relationship between the number of species in a system and the rates of ecological processes. We present the results of a simple model that suggests there may be no single, generalizable relationship between species diversity and the productivity of an ecosystem because the relative contributions of species to productivity change with environmental context. The model determined productivity for landscapes varying in species diversity (the number of species in the colonist pool), spatial heterogeneity (the number of habitat types composing the landscape), and disturbance regimes (+/? a non‐selective mortality). Linear regressions were used to relate species diversity and productivity for each of the environmental contexts. Disturbance changed the form of the diversity/productivity relationship by reducing the slope (i.e. the change in productivity per species added to the colonist pool), but spatial heterogeneity increased or decreased this slope depending on the particular habitat types composing the landscape. The cause of the diversity/productivity relationship also changed with environmental context. The amount of variation in productivity explained by species diversity always increased with spatial heterogeneity, while the amount of variation explained by species composition (i.e. the particular species composing the colonist pool) tended to increase with disturbance. These results lead us to conclude that the form and cause of the relationship between species diversity and productivity may be highly dynamic‐changing over both time and space. Because the trends resulted from well‐known mechanisms by which environmental variation alters the absolute and relative abundances of taxa, we suspect this conclusion may be applicable to many different systems.     

Species and genotype diversity drive community and ecosystem properties in experimental microcosms

Species diversity is important to ecosystems because of the increased probability of including species that are strong interactors and/or because multiple-species communities are more efficient at using resources due to synergisms and resource partitioning. Genetic diversity also contributes to ecosystem function through effects on primary productivity, community structure and resilience, and modulating energy and nutrient fluxes. Lacking are studies investigating the relationship between ecosystem function and diversity where hierarchical levels of biological diversity are systematically varied during experimentation. In this experiment, we manipulated both species and genotypic diversity of two Daphnia species in microcosms initially seeded with Chlamydomonas and measured community- and ecosystem-level properties to determine which level of diversity was most important for explaining variation in the property. Our results show that species diversity alters bacterial community composition while high genotypic diversity reduces bacterial richness and primary productivity. In addition, the highest levels of genotypic and species richness appear to increase community and ecosystem stability. These findings reveal that species and genotypic diversity are significant drivers of community and ecosystem properties and stability.