Does functional homogenization accompany taxonomic homogenization of British birds and how do biotic factors and climate affect these processes?

Environmental change has reshuffled communities often causing taxonomic homogenization rather than differentiation. Some studies suggest that this increasing similarity of species composition between communities is accompanied by an increase in similarity of trait composition—functional homogenization—although different methodologies have failed to come to any consistent conclusions. Functional homogenization could have a large effect on ecosystem functioning and stability. Here, we use the general definition of homogenization as “reduced spatial turnover over time” to compare changes in Simpson's beta diversity (taxonomic turnover) with changes in Rao's quadratic entropy beta diversity (functional turnover) in British breeding birds at three spatial scales. Using biotic and climatic variables, we identify which factors may predispose a site to homogenization. The change in turnover measures between two time periods, 20 years apart, was calculated. A null model approach was taken to identify occurrences of functional homogenization and differentiation independent of changes in taxonomic turnover. We used conditional autoregressive models fitted using integrated nested Laplace approximations to determine how environmental drivers and factors relating to species distributions affect changes in spatial turnover of species and functional diversity. The measurement of functional homogenization affects the chance of rejection of the null models, with many sites showing taxonomic homogenization unaccompanied by functional homogenization, although occurrence varies with spatial scale. At the smallest scale, while temperature‐related variables drive changes in taxonomic turnover, changes in functional turnover are associated with variation in growing degree days; however, changes in functional turnover become more difficult to predict at larger spatial scales. Our results highlight the multifactorial processes underlying taxonomic and functional homogenization and that redundancy in species traits may allow ecosystem functioning to be maintained in some areas despite changes in species composition.     

Does canopy position affect wood specific gravity in temperate forest trees?

The radial increases in wood specific gravity known in many tree species have been interpreted as providing mechanical support in response to the stresses associated with wind loading. This interpretation leads to the hypothesis that individuals reaching the canopy should (1) be more likely to have radial increases in specific gravity and (2) exhibit greater increases than individuals in the subcanopy. Wood specific gravity was determined for three species of forest trees (Acer rubrum, Fagus grandifolia and Tsuga canadensis) growing in central Massachusetts, USA. Acer rubrum shows radial increases in specific gravity, but these increases are not more pronounced in canopy trees; the other two species show a pattern of radial decreases. The degree of radial increase or decrease is influenced by tree height and diameter. Of the dominant tree species for which we have data, A. rubrum, Betula papyrifera and Pinus strobus show radial increases in specific gravity, whereas F. grandifolia, T. canadensis and Quercus rubra show decreases. The occurrence of radial increases in B. papyrifera and P. strobus, which are often canopy emergents, suggests that it is overall adaptive strategy that is important rather than position (canopy vs. subcanopy) of any individual tree. It is suggested that radial increases in specific gravity are associated with early-successional status or characteristics and decreases with late-successional status or persistence in mature forest.     

Climate change – Bad news for montane forest herb layer species?

Global warming presents a threat to plant species distributed at montane or alpine altitudes if the topography does not allow upward shifts in distribution ranges. Nevertheless, the species might also benefit from increasing temperatures and secondary effects on dominant species (e.g. bark beetle outbreaks or summer drought affecting the canopy species). As a consequence, disturbance frequency in montane forests might increase and light availability for herb layer species will increase. We addressed these interactions in a common garden experiment in Central Germany at different altitudes, representing cold and moist vs. warm and dry conditions. We investigated three montane species with different life forms, including a herb (Trientalis europaea), a grass (Calamagrostis villosa) and a dwarf shrub (Vaccinium myrtillus) under three shading treatments (3%, 28% and 86% of full sunlight). We hypothesized that montane species are at a disadvantage in the lowland, with the dwarf shrub suffering more than the grass. Furthermore, we hypothesized an antagonistic interaction of increased temperature and increased light conditions. While T. europaea and V. myrtillus showed only slightly responses to low altitude conditions, C. villosa displayed a nearly fifteen fold increase in biomass production, despite higher observed herbivory levels in the lowland. We failed to show an antagonistic effect of increased temperature and increased light availability, as all study species suffered from deep shade conditions and grew best under full light conditions at both sites. In conclusion, both improved temperature and light conditions might be principally beneficial for the investigated boreal species, in particular for the grass species C. villosa.     

Seasonal dynamics of shoot biomass of dominant clonal herb species in an oak–hornbeam forest herb layer

Seasonal fluctuations of light availability, nutrient concentrations, and moisture affect plant population traits like density, standing biomass, and flowering. We analyzed seasonal changes of density and shoot biomass of the four most frequent herb species growing in an oak–hornbeam forest community, i.e., Anemone nemorosa, Ficaria verna, Galeobdolon luteum, and Galium odoratum. In 2010 and 2011 plant biomass was harvested from 7 to 10 randomly situated square sample plots (0.36 m²) in the homogenous oak–hornbeam forest community every week in the spring and every two weeks in the summer and autumn. The highest abundance of Anemone nemorosa reached over 1000 shoots per m², of Ficaria verna 459.5 shoots per m², of Galium odoratum 83.6 shoots per m², and of Galeobdolon luteum 98.4 shoots per m² (means for 2010 and 2011, based on all sample plots). We did not observe negative correlation between density and shoot biomass. Growth rates of vegetative shoot biomass amounted to 0.857 mg day^?1 for Anemone nemorosa, 0.467 mg day^?1 for Ficaria verna, 0.722 mg day^?1 for Galium odoratum, and 0.448 mg day^?1 for Galeobdolon luteum (means for 2010 and 2011). Spring ephemerals had much higher densities of shoots than summer-greens. Summer-greens reached higher biomass of individual shoots than spring ephemerals. Flowering shoots constituted only 4, 2, and 11% of all shoots for A. nemorosa, F. verna, and G. odoratum, respectively. More resource availability resulting in high shoot biomass did not translate to higher share of flowering shoots.

     

Changing taxonomic and functional β-diversity of cladoceran communities in Northeastern and South Brazil

Hydrobiologia - Disturbances caused by both natural and anthropogenic forces can drive changes in alpha and beta taxonomic diversity and be accompanied by losses or gains in freshwater ecosystem...     

Does forest fragmentation cause an increase in forest temperature?

Forest fragmentation is considered by many to be a primary cause of the current biodiversity crisis. The underlying mechanisms are poorly known, but a potentially important one is associated with altered thermal conditions within the remaining forest patches, especially at forest edges. Yet, large uncertainty remains about the effect of fragmentation on forest temperature, as it is unclear whether temperature decreases from forest edge to forest interior, and whether this local gradient scales up to an effect of fragmentation (landscape attribute) on temperature. We calculated the effect size (correlation coefficient) of distance from forest edge on air temperature, and tested for differences among forest types surrounded by different matrices using meta-analysis techniques. We found a negative edge-interior temperature gradient, but correlation coefficients were highly variable, and significant only for temperate and tropical forests surrounded by a highly contrasting open matrix. Nevertheless, it is unclear if these local-scale changes in temperature can be scaled up to an effect of fragmentation on temperature. Although it may be valid when considering “fragmentation” as forest loss only, the landscape-scale inference is not so clear when we consider the second aspect of fragmentation, where a given amount of forest is divided into a large number of small patches (fragmentation per se). Therefore, care is needed when assuming that fragmentation changes forest temperature, as thermal changes at forest edges depend on forest type and matrix composition, and it is still uncertain if this local gradient can be scaled up to the landscape.     

Does disturbance regime change community assembly of angiosperm plant communities in the boreal forest?

Aims To examine if and how species and phylogenetic diversity change in relation to disturbance, we conducted a review of ecological literature by testing the consistency of the relationship between phylogenetic diversity and disturbance and compared taxonomic groups, type of disturbance and ecosystem/habitat context. We provide a case study of the phylogenetic diversity–disturbance relationship in angiosperm plant communities of a boreal forest region, compared with types of natural and anthropogenic disturbances and plant growth forms.Methods Using a large-scale sampling plot network along a complete (0–100%) anthropogenic disturbance gradient in the boreal biome, we compared the changes of angiosperm plant community structure and composition across plots. We estimated natural disturbance with historical records of major fires. We then calculated phylogenetic diversity indexes and determined species richness in order to compare linear and polynomial trends along disturbance gradients. We also compared the changes of community structure for different types of anthropogenic disturbances and examined how the relationships between species and phylogenetic diversity and disturbance regimes vary among three different life forms (i.e. forbs, graminoids and woody plants).Important findings Phylogenetic diversity was inconsistently related to disturbance in previous studies, regardless of taxon, disturbance type or ecosystem context. In the understudied boreal ecosystem, angiosperm plant communities varied greatly in species richness and phylogenetic diversity along anthropogenic disturbance gradients and among different disturbance types. In general, a quadratic curve described the relationship between species richness and anthropogenic disturbance, with the highest richness at intermediate anthropogenic disturbance levels. However, phylogenetic diversity was not related to disturbance in any consistent manner and species richness was not correlated with phylogenetic diversity. Phylogenetic relatedness was also inconsistent across plant growth forms and different anthropogenic disturbance types. Unlike the inconsistent patterns observed for anthropogenic disturbance, community assembly among localities varying in time since natural disturbance exhibited a distinct signature of phylogenetic relatedness, although those trends varied among plant growth forms.     

Ecological filters and variability in stream macroinvertebrate communities: do taxonomic and functional structure follow the same path?

We examined the community-environment relationships of lotic macroinvertebrates in near-pristine headwater streams, and the correlation between patterns in taxonomic and functional structure at two regional extents. The across-ecoregion scale comprised five ecoregions spanning all of Finland, while the within-ecoregion scale comprised of north boreal and middle boreal ecoregions. We expected that taxonomic structure should exhibit stronger relationships than functional structure to spatial gradients, while the reverse should be true for local environmental factors. We found some support for this notion, because spatial variables were marginally more important for taxonomic than functional structure. Furthermore, within the two ecoregions, local environmental variables were slightly more important for functional than taxonomic structure. Geographical location (i.e. spatial variables) was more influential at the across-ecoregions extent than within the two ecoregions. Largely the same local environmental variables accounted for variation in both taxonomic and functional structure, including water pH, nutrients, colour, and stream size. Similar responses to local environmental features likely contributed to the significant correlation between patterns in taxonomic and functional structure. It has been suggested that functional traits may be rather insensitive to natural variation, yet our functional categorization showed distinct variation along local environmental, catchment, and spatial gradients across near-pristine streams. Thus, natural ecological filters should be accounted for prior to the examination of the effects of anthropogenic filters on stream macroinvertebrate community structure.     

Does morphological convergence imply functional similarity? A test using the evolution of quadrupedalism in ornithischian dinosaurs

Convergent morphologies are thought to indicate functional similarity, arising because of a limited number of evolutionary or developmental pathways. Extant taxa displaying convergent morphologies are used as analogues to assess function in extinct taxa with similar characteristics. However, functional studies of extant taxa have shown that functional similarity can arise from differing morphologies, calling into question the paradigm that form and function are closely related. We test the hypothesis that convergent skeletal morphology indicates functional similarity in the fossil record using ornithischian dinosaurs. The rare transition from bipedality to quadrupedality occurred at least three times independently in this clade, resulting in a suite of convergent osteological characteristics. We use homology rather than analogy to provide an independent line of evidence about function, reconstructing soft tissues using the extant phylogenetic bracket and applying biomechanical concepts to produce qualitative assessments of muscle leverage. We also optimize character changes to investigate the sequence of character acquisition. Different lineages of quadrupedal ornithischian dinosaur stood and walked differently from each other, falsifying the hypothesis that osteological convergence indicates functional similarity. The acquisition of features correlated with quadrupedalism generally occurs in the same order in each clade, suggesting underlying developmental mechanisms that act as evolutionary constraints.     

Does prescribed burning result in biotic homogenization of coastal heathlands?

Biotic homogenization due to replacement of native biodiversity by widespread generalist species has been demonstrated in a number of ecosystems and taxonomic groups worldwide, causing growing conservation concern. Human disturbance is a key driver of biotic homogenization, suggesting potential conservation challenges in seminatural ecosystems, where anthropogenic disturbances such as grazing and burning are necessary for maintaining ecological dynamics and functioning. We test whether prescribed burning results in biotic homogenization in the coastal heathlands of north‐western Europe, a seminatural landscape where extensive grazing and burning has constituted the traditional land‐use practice over the past 6000 years. We compare the beta‐diversity before and after fire at three ecological scales: within local vegetation patches, between wet and dry heathland patches within landscapes, and along a 470 km bioclimatic gradient. Within local patches, we found no evidence of homogenization after fire; species richness increased, and the species that entered the burnt Calluna stands were not widespread specialists but native grasses and herbs characteristic of the heathland system. At the landscapes scale, we saw a weak homogenization as wet and dry heathland patches become more compositionally similar after fire. This was because of a decrease in habitat‐specific species unique to either wet or dry habitats and postfire colonization by a set of heathland specialists that established in both habitat types. Along the bioclimatic gradient, species that increased after fire generally had more specific environmental requirements and narrower geographical distributions than the prefire flora, resulting in a biotic ‘heterogenisation’ after fire. Our study demonstrates that human disturbance does not necessarily cause biotic homogenization, but that continuation of traditional land‐use practices can instead be crucial for the maintenance of the diversity and ecological function of a seminatural ecosystem. The species that established after prescribed burning were heathland specialists with relatively narrow geographical ranges.     

How do sheep affect plant communities and arthropod populations in temperate grasslands?

Grasslands being used in sheep farming systems are managed under a variety of agricultural production, recreational and conservational objectives. Although sheep grazing is rarely considered the best method for delivering conservation objectives in seminatural temperate grasslands, the literature does not provide unequivocal evidence on the impact of sheep grazing on pasture biodiversity. Our aim was therefore to review evidence of the impacts of stocking rate, grazing period and soil fertility on plant communities and arthropod populations in both mesotrophic grasslands typical of agriculturally improved areas and in native plant communities. We therefore conducted a literature search of articles published up to the end of the year 2010 using ‘sheep’ and ‘grazing’ as keywords, together with variables describing grassland management, plant community structure or arthropod taxa. The filtering process led to the selection of 48 articles, with 42 included in the stocking rate dataset, 9 in the grazing period dataset and 10 in the soil fertility dataset. The meta-analysis did not reveal any significant trends for plant species richness or plant community evenness along a wide stocking rate gradient. However, we found frequent shifts in functional groups or plant species abundance that could be explained by the functional properties of the plants in the community. The meta-analysis confirmed that increasing soil fertility decreased plant species richness. Despite the very limited dataset, plant species richness was significantly greater in autumn-grazed pastures than in ungrazed areas, which suggests that choosing an appropriate grazing period would be a promising option for preserving biodiversity in sheep farming systems. Qualitative review indicated that low grazing intensity had positive effects on Orthoptera, Hemiptera (especially phytophagous Auchenorrhyncha) and, despite a diverse range of feeding strategies, for the species richness of Coleoptera. Lepidoptera, which were favoured by more abundant flowering plants, also benefited from low grazing intensities. Spider abundance and species richness were higher in ungrazed than in grazed pastures. In contrast, there are insufficient published studies to draw any firm conclusions on the benefits of late grazing or stopping fertilization on insect diversity, and no grounds for including any of this information in decision support tools at this stage.     

Low functional β-diversity despite high taxonomic β-diversity among tropical estuarine fish communities

The concept of β-diversity, defined as dissimilarity among communities, has been widely used to investigate biodiversity patterns and community assembly rules. However, in ecosystems with high taxonomic β-diversity, due to marked environmental gradients, the level of functional β-diversity among communities is largely overlooked while it may reveal processes shaping community structure. Here, decomposing biodiversity indices into α (local) and γ (regional) components, we estimated taxonomic and functional β-diversity among tropical estuarine fish communities, through space and time. We found extremely low functional β-diversity values among fish communities (<1.5%) despite high dissimilarity in species composition and species dominance. Additionally, in contrast to the high α and γ taxonomic diversities, α and γ functional diversities were very close to the minimal value. These patterns were caused by two dominant functional groups which maintained a similar functional structure over space and time, despite the strong dissimilarity in taxonomic structure along environmental gradients. Our findings suggest that taxonomic and functional β-diversity deserve to be quantified simultaneously since these two facets can show contrasting patterns and the differences can in turn shed light on community assembly rules.     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

How pervasive is biotic homogenization in human‐modified tropical forest landscapes?

Land‐cover change and ecosystem degradation may lead to biotic homogenization, yet our understanding of this phenomenon over large spatial scales and different biotic groups remains weak. We used a multi‐taxa dataset from 335 sites and 36 heterogeneous landscapes in the Brazilian Amazon to examine the potential for landscape‐scale processes to modulate the cumulative effects of local disturbances. Biotic homogenization was high in production areas but much less in disturbed and regenerating forests, where high levels of among‐site and among‐landscape β‐diversity appeared to attenuate species loss at larger scales. We found consistently high levels of β‐diversity among landscapes for all land cover classes, providing support for landscape‐scale divergence in species composition. Our findings support concerns that β‐diversity has been underestimated as a driver of biodiversity change and underscore the importance of maintaining a distributed network of reserves, including remaining areas of undisturbed primary forest, but also disturbed and regenerating forests, to conserve regional biota.     

Does functional redundancy exist?

Functional redundancy has often been assumed as an intuitive null hypothesis in biodiversity experiments, but theory based on the classical Lotka-Volterra competition model shows that functional redundancy sensu stricto is incompatible with stable coexistence. Stable coexistence requires differences between species which lead to functional complementarity and differences between the yields of mixtures and monocultures. Only a weaker version of functional redundancy, i.e. that mixture yields lie within the range of variation of monoculture yields, is compatible with stable coexistence in Lotka-Volterra systems. Spatial and temporal environmental variability may provide room for functional redundancy at small spatial and temporal scales, but is not expected to do so at the larger scales at which environmental variations help maintain coexistence. Neutral coexistence of equivalent competitors, non-linear per capita growth rates, and lack of correlation between functional impact and biomass may provide the basis for the existence of functional redundancy in natural ecosystems. Overall, there is a striking parallel between the conditions that allow stable coexistence and those that allow overyielding.     

Does niche overlap control relative abundance in French lacustrine fish communities? A new method incorporating functional traits

1. The mechanisms that structure biological communities hold the key to understanding ecosystem functioning and the maintenance of biodiversity. Patterns of species abundances have been proposed as a means of differentiation between niche-based and neutral processes, but abundance information alone cannot provide unequivocal discrimination. 2. We combined species niche information and species' relative abundances to test the effects of two opposing structuring mechanisms (environmental filtering and niche complementarity) on species' relative abundances in French lacustrine fish communities. The test involved a novel method comparing the abundance-weighted niche overlap within communities against that expected when relative abundances were randomized among species within the community. 3. Observed overlap was consistently significantly lower than expected at random for two (swimming ability and trophic status) of four primary niche axes across lakes of differing physical environments. Thus, for these niche axes, pairs of abundant species tended to have relatively low niche overlap, while rare species tended to have relatively high niche overlap with abundant species. 4. This suggests that niche complementarity may have acted to enhance ecosystem function and that it is important for species coexistence in these fish communities. The method used may be easily applied to any sort of biological community and thus may have considerable potential for determining the generality of niche complementarity effects on community structure.     

Neural signalling: Does colocalization imply cotransmission?

Recent results suggest that neurons that contain multiple neurotransmitters may make synaptic connections with different target neurons that are mediated by only a subset of their transmitter complement.