Landscape-scale connectivity and fragment size determine species composition of grassland fragments

As a consequence of agricultural intensification and habitat fragmentation since the mid-20th century, biological diversity has declined considerably throughout the world, particularly in Europe. We assessed how habitat and landscape-scale heterogeneity, such as variation in fragment size (small vs. large) and landscape configuration (measured as connectivity index), affect plant and arthropod diversity. We focused on arthropods with different feeding behaviour and mobility, spiders (predators, moderate dispersal), true bugs (mainly herbivores and omnivores with moderate dispersal), wild bees (pollinators with good dispersal abilities), and wasps (pollinators, omnivores with good dispersal abilities). We studied 60 dry grassland fragments in the same region (Hungarian Great Plain); 30 fragments were represented by the grassland component of forest-steppe stands, and 30 were situated on burial mounds (kurgans). Forest-steppes are mosaics of dry grasslands with small forests in a matrix of plantation forests. Kurgans are ancient burial mounds with moderately disturbed grasslands surrounded by agricultural fields. The size of fragments ranged between 0.16–6.88 ha (small: 0.16–0.48 ha, large: 0.93–6.88 ha) for forest-steppes and 0.01–0.44 ha (small: 0.01–0.10 ha and large: 0.20–0.44 ha) for kurgans. Fragments also represented an isolation gradient from almost cleared and homogenous landscapes, to landscapes with relatively high compositional heterogeneity. Fragment size, connectivity, and their interaction affected specialist and generalist species abundances of forest-steppes and kurgans. Large fragments had higher species richness of ground-dwelling spiders, and the effect of connectivity was more strongly positive for specialist arthropods and more strongly negative for generalists in large than in small fragments. However, we also found a strong positive impact of connectivity for generalist plants in small kurgans in contrast to larger ones. We conclude that besides the well-known effect of enhancing habitat quality, increasing connectivity between fragments by restoring natural and semi-natural habitat patches would help to maintain grassland biodiversity.     

Plasticity and overcompensation in grass responses to herbivory

Several hypotheses predict defoliation-induced increases in individual plant fitness. In this paper we examine three such hypotheses: the Herbivore Optimization Hypothesis (HOH); the Continuum of Responses Hypothesis (CRH); and the Growth Rate Model (GRM). All three have in common predictions based on responses of defoliated individuals with the objective of explaining community and higher level phenomena. The latter two extend theory by specifying conditions for overcompensatory responses. They differ in whether overcompensation is sensitive to conditions external (CRH) or internal (GRM) to the plant. We tested these hypotheses with field experiments in a grassland system in which two native, perennial grass species replace each other along a short topographic/resource gradient. We detected positive, neutral, and negative changes in plant mass in response to partial defoliation. Patterns of responses to the edaphic and competitive environment combinations were unique to each species and neither the CRH nor the GRM were able to consistently predict responses in these grasses. Predictions of the HOH were fully supported only by the species naturally limited to lower-resource environments: overcompensation occurred in natural environments and it occurred at herbivory levels these plants experience naturally. Thus, the overcompensatory response can be important for the maintenance of local plant population distributions. However, new mechanistic theory is needed to account for the trend common to both species: overcompensatory responses to herbivory were greater in the edaphic environment in which each species was naturally most abundant.     

A simple method to determine leaf angles of grass species

There are several very accurate methods to determine leaf angles in closed canopies. However, these are generally very time-consuming or require special equipment. Average canopy leaf angles were derived from simple height and blade length measurements. An exponential relationship between the height/length ratio and the average blade leaf angle was used. The method was tested for two grass species, Dactylis glomerata and Festuca arundinacea, grown under different UV-B levels. The results clearly show that the method is reasonably accurate and able to identify UV-B induced changes in leaf angle. To get these results only 50 measurements of leaf blade height and length were necessary to calculate the allometric relationship, after which 10 length and height measurements from a canopy were used to calculate the average canopy leaf angle.     

Interactive effects of plant-available soil silicon and herbivory on competition between two grass species

Background and Aims

The herbivore defence system of true grasses (Poaceae) is predominantly based on silicon that is taken up from the soil and deposited in the leaves in the form of abrasive phytoliths. Silicon uptake mechanisms can be both passive and active, with the latter suggesting that there is an energetic cost to silicon uptake. This study assessed the effects of plant-available soil silicon and herbivory on the competitive interactions between the grasses Poa annua, a species that has previously been reported to accumulate only small amounts of silicon, and Lolium perenne, a high silicon accumulator.

Methods

Plants were grown in mono- and mixed cultures under greenhouse conditions. Plant-available soil silicon levels were manipulated by adding silicon to the soil in the form of sodium silicate. Subsets of mixed culture pots were exposed to above-ground herbivory by desert locusts (Schistocerca gregaria).

Key Results

In the absence of herbivory, silicon addition increased biomass of P. annua but decreased biomass of L. perenne. Silicon addition increased foliar silicon concentrations of both grass species >4-fold. Under low soil-silicon availability the herbivores removed more leaf biomass from L. perenne than from P. annua, whereas under high silicon availability the reverse was true. Consequently, herbivory shifted the competitive balance between the two grass species, with the outcome depending on the availability of soil silicon.

Conclusions

It is concluded that a complex interplay between herbivore abundance, growth–defence trade-offs and the availability of soil silicon in the grasses'' local environment affects the outcome of inter-specific competition, and so has the potential to impact on plant community structure.     

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change – fertilisation and herbivore loss – are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.     

Body size and tree species composition determine variation in prey consumption in a forest‐inhabiting generalist predator

Trophic interactions may strongly depend on body size and environmental variation, but this prediction has been seldom tested in nature. Many spiders are generalist predators that use webs to intercept flying prey. The size and mesh of orb webs increases with spider size, allowing a more efficient predation on larger prey. We studied to this extent the orb‐weaving spider Araneus diadematus inhabiting forest fragments differing in edge distance, tree diversity, and tree species. These environmental variables are known to correlate with insect composition, richness, and abundance. We anticipated these forest characteristics to be a principle driver of prey consumption. We additionally hypothesized them to impact spider size at maturity and expect shifts toward larger prey size distributions in larger individuals independently from the environmental context. We quantified spider diet by means of metabarcoding of nearly 1,000 A. diadematus from a total of 53 forest plots. This approach allowed a massive screening of consumption dynamics in nature, though at the cost of identifying the exact prey identity, as well as their abundance and putative intraspecific variation. Our study confirmed A. diadematus as a generalist predator, with more than 300 prey ZOTUs detected in total. At the individual level, we found large spiders to consume fewer different species, but adding larger species to their diet. Tree species composition affected both prey species richness and size in the spider''s diet, although tree diversity per se had no influence on the consumed prey. Edges had an indirect effect on the spider diet as spiders closer to the forest edge were larger and therefore consumed larger prey. We conclude that both intraspecific size variation and tree species composition shape the consumed prey of this generalist predator.     

Tree hollows can affect epiphyte species composition

Tree hollows often harbor animals and microorganisms, thereby storing nutritive resources derived from their biological activities. The outflows from tree hollows can create unique microenvironments, which may affect communities of epiphytic organisms on trunk surfaces below the hollows. In this study, we tested whether the species richness and composition of epiphytic bryophytes (liverworts and mosses) and lichens differ above and below tree hollows of Aria japonica and Cercidiphyllum japonicum in a Japanese temperate forest. The species richness of epiphytic bryophytes and lichens did not differ above and below hollows; however, the species composition of bryophytes differed significantly above and below hollows. Indicator species analyses showed that the moss species Anomodon tristis and the liverwort species Porella vernicosa were significantly more common below than above hollows, while the liverwort species Radula japonica and four lichen species, including Leptogium cyanescens, occurred more frequently above than below hollows. Our results highlight that tree hollows can produce unique microenvironments on trunk surfaces that potentially contribute to the maintenance of epiphytic diversity on a local scale.     

Siltation and hydrologic regime determine species composition in herbaceous floodplain communities

Interest in the restoration of riparian habitat is increasing. However, little is known about factors responsible for riparian communities, especially grasslands. In order to construct plant communities for a restoration project in the floodplain of a large river in the Midwestern United States, we sampled four floodplains with various disturbance regimes located in Illinois and Missouri. They were chosen to be representative of different plant communities of floodplains, with a focus on herbaceous communities. The areas included backwater lakes, alluvial fans, groundwater seep marshes, oxbow marshes, seasonally inundated grassland, and non-inundated grassland. Vegetation, soils and groundwater or standing water depth were measured at various intervals along transects. Communities were produced using TWINSPAN and tested for differences in environmental factors. The soil morphology, taxonomic classification, and fertility parameters were similar among sites. Environmental factors influencing community composition were the presence of permanent water and silt deposition. We conclude that water depth determines species composition in permanently wet areas. Silt deposition determines composition in seasonally inundated grassland. Where silt deposition is high enough to inhibit seedling emergence, dominance is attained by plants able to reproduce vegetatively by rhizomes. Such a reproductive process leads to nearly monotypic stands produced by large clones. Results are discussed in relation to models of riparian processes and succession.     

Local filters limit species diversity,but species pools determine composition

In a series of three experiments, we tested for effects of species pools, resource stress, and species interactions on four aspects of community structure: species richness, evenness, species composition, and functional group composition. We also examined whether the impacts of species interactions on the community varied with resource availability or species pool. Communities of sand dune annuals grew from seed bank samples collected from two sites in three different years, so that the species pool differed at two levels: the source site and the year of seed bank collection. Communities experienced one of three irrigation treatments and a range of sowing densities, which varied resource supply (stress) and the potential for species interactions, respectively.Species richness and evenness were most affected by local factors: higher densities and lower water availability decreased species diversity. In contrast, species composition was influenced most by the species pool. Functional group composition had an intermediate response, and was affected by both species pools and local filters.Resource stress and species interactions strongly filtered species from the community, but the identity of species remaining was variable. Furthermore, the magnitude of species interaction effects on richness and evenness varied with species pools. Thus, the outcome of species sorting among biotic and abiotic environments was dependent on the pool of available species. Contrary to predictions from theory, the effects of species interactions on the community did not vary consistently with resource levels.     

Tree species identity and interactions with neighbors determine nutrient leaching in model tropical forests

An ecosystem containing a mixture of species that differ in phenology, morphology, and physiology might be expected to resist leaching of soil nutrients to a greater extent than one composed of a single species. We tested the effects of species identity and plant-life-form richness on nutrient leaching at a lowland tropical site where deep infiltration averages >2 m year(-1). Three indigenous tree species with contrasting leafing phenologies (evergreen, dry-season deciduous, and wet-season deciduous) were grown in monoculture and together with two other life-forms with which they commonly occur in tropical forests: a palm and a giant, perennial herb. To calculate nutrient leaching over an 11-year period, concentrations of nutrients in soil water were multiplied by drainage rates estimated from a water balance. The effect of plant-life-form richness on retention differed according to tree species identity and nutrient. Nitrate retention was greater in polycultures of the dry-season deciduous tree species (mean of 7.4 kg ha(-1) year(-1) of NO(3)-N lost compared to 12.7 in monoculture), and calcium and magnesium retention were greater in polycultures of the evergreen and wet-season deciduous tree species. Complementary use of light led to intensification of soil exploitation by roots, the main agent responsible for enhanced nutrient retention in some polycultures. Other mechanisms included differences in nutrient demand among species, and avoidance of catastrophic failure due to episodic weather events or pest outbreaks. Even unrealistically simple multi-life-form mimics of tropical forest can safeguard a site's nutrient capital if careful attention is paid to species' characteristics and temporal changes in interspecific interactions.     

Pine species determine fungal microbiome composition in a common garden experiment

The factors shaping the composition of microbial communities in trees remain poorly understood. We evaluated whether the core and satellite fungal communities in five pine species (Pinus radiata, Pinus pinaster, Pinus sylvestris, Pinus nigra, and Pinus uncinata) were shaped by the host species identity. Because the trees had earlier been inoculated with a fungal pathogen (Fusarium circinatum), we also explored the possibilities to detect its presence and potential co-occurrence networks. We found interspecific variation in the fungal community composition and abundance among the different tree species and the existence of a core microbiome that was independent of the host species. The presence of F. circinatum was confirmed in some samples through qPCR but the pathogen did not co-occur with a specific fungal community. The results highlight the importance of host species as a determinant of microbiome assembly in common environments.     

Ploidy level interacts with population size and habitat conditions to determine the degree of herbivory damage in plant populations

Recently it has been suggested that ploidy level of a plant population may have important effects on plant‐animal interactions. Plant‐animal interactions can also be strongly altered by factors such as plant population size and habitat conditions. It is, however, not known how these factors interact to shape the overall pattern of plant‐animal interactions. I studied the interaction between a perennial plant, Aster amellus, and a monophagous herbivorous moth, Coleophora obscenella, and investigated the effect of ploidy level of the plant population, plant population size, isolation and habitat conditions on density of the insect, damage by the insect, and plant performance. Ploidy level, plant population size and habitat conditions, but not isolation, strongly influence plant‐herbivore interactions. Furthermore, there are significant interactions between effects of ploidy level and plant population size and between ploidy level and isolation. Hexaploid plants suffer higher seed damage by the herbivore, but their seed production is still higher than that of diploids. Herbivores thus partly limit the evolutionary success of the hexaploid plants. Plant‐animal interactions are also strongly determined by plant population size. Small populations of A. amellus (below forty flowering ramets) host no C. obscenella larvae, indicating a minimum A. amellus population size that can sustain a viable C. obscenella population. Negative and positive effects of plant population size balance and result in no relationship between plant population size and number of developed seeds per flower head. The results also show a significant interaction between ploidy level and plant population size, indicating that the increase in density of C. obscenella larvae with plant population size is greater in hexaploid than in diploid populations. The results also indicate that the effect of ploidy level on plant‐herbivore interactions can be altered by plant population size, which suggests that plant‐herbivore interactions are driven by a complex of interactions among different factors. Studying each factor separately could thus lead to biased conclusions about patterns of interactions in such systems.     

Influence of soil and microclimate on species composition and grass encroachment in heath succession

Aims and Methods Mostly due to land use changes, European heathlands have become increasingly rare. In addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses and a loss in species diversity. Despite many investigations, information about the precise environmental parameters that determine the development and maintenance of heathland vegetation is still insufficient. In order to determine the environmental factors that control heath succession and grass encroachment, and to develop appropriate management schemes, we studied the influence of several soil and microclimate parameters on species composition and vegetation characteristics in five successional stages in a coastal heathland on the island of Hiddensee, north-east Germany, where the encroachment of Carex arenaria has become a major problem.Important findings We recorded the highest plant species richness in grey dune and birch forest plots, while the encroachment of C. arenaria let to a significant decline in plant species richness. The most important environmental factors influencing species richness and distribution of single species were microclimate, soil moisture, soil pH and the C/N ratio. While many studies reported the importance of differences in nutrient availability, we found no significant correlations between soil nutrient availability and vegetation pattern. Environmental conditions in dense C. arenaria stands, especially soil properties (e.g. soil pH), showed great differences in comparison to the other successional stages. However, no correlations between the encroachment of C. arenaria and single environmental factors were found. Our results show that not only soil nutrients are important abiotic factors in heaths but that also microclimate and soil moisture play an important role and that many factors are involved in heath succession and in the promotion of grass encroachment. Management plans for the conservation and restoration of heathlands should therefore focus on the specific site conditions and should take several abiotic and biotic factors into account.     

Tree seedling richness,but not neighborhood composition,influences insect herbivory in a temperate deciduous forest community

Insect herbivores can serve as important regulators of plant dynamics, but their impacts in temperate forest understories have received minimal attention at local scales. Here, we test several related hypotheses about the influence of plant neighborhood composition on insect leaf damage in southwestern Pennsylvania, USA. Using data on seedlings and adult trees sampled at 36 sites over an approximately 900 ha area, we tested for the effects of total plant density, rarefied species richness (i.e., resource concentration and dietary‐mixing hypotheses), conspecific density (i.e., Janzen–Connell hypothesis), and heterospecific density (i.e., herd‐immunity hypothesis), on the proportion of leaf tissue removed from 290 seedlings of 20 species. We also tested for the effects of generic‐ and familial‐level neighborhoods. Our results showed that the proportion of leaf tissue removed ranged from zero to just under 50% across individuals, but was generally quite low (<2%). Using linear mixed models, we found a significant negative relationship between insect damage and rarefied species richness, but no relationship with neighborhood density or composition. In addition, leaf damage had no significant effect on subsequent seedling growth or survival, likely due to the low levels of damage experienced by most individuals. Our results provide some support for the resource concentration hypothesis, but suggest a limited role for insect herbivores in driving local‐scale seedling dynamics in temperate forest understories.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.     

Tree species composition and diversity in natural temperate forests of the North-Western Himalayas

The present study was undertaken in the natural temperate Himalayan forests of Himachal Pradesh, India, to assess the tree species composition and diversity. For this purpose, six major forest types (FT) viz., FT1- Upper Himalayan Pinus roxburghii forest, FT2- Quercus leucotrichophora forest, FT3- Low-level P. wallichiana forest, FT4- Moist C. deodara forest, FT5- Western Mixed Coniferous Forest, FT6- Pinus gerardiana forest were selected. Detailed sampling was carried out in these forest types, and the sample plots in each forest type were laid out using the stratified random approach. Tree stem density varied from 191.11 N ha^?1 (FT6) to 441.11 N ha^?1 (FT2), whereas the tree total basal cover varied from 20.01 m² ha^?1 (FT6) to 47.59 m² ha^?1 (FT5). The diversity indices reflected that a total of 21 tree species (16 genera, 11 families) were identified, with tree species richness ranging from 3 (FT6) to 9 (FT1 & FT5). The forest type FT5 recorded the maximum Shannon index of diversity (2.36), Simpson Index of diversity (0.75), Margalef's Index of richness (1.37), pielou equitability (0.74), menheink index of species richness (0.49), whereas the highest Shannon Index of diversity (0.73) and species heterogeneity (0.85) in FT6 forest type. Furthermore, dominance-diversity (d-d) curves drawn that all the six forest types showed geometric curves reveals that one or two tree species are dominant in a particular forest type. Simultaneously, the research area's species diversity, tree stem density, and tree total basal cover were equivalent to those seen in other sections of Western Himalayas.     

Tree species composition and diversity gradients in white-water forests across the Amazon Basin

Aim Attention has increasingly been focused on the floristic variation within forests of the Amazon Basin. Variations in species composition and diversity are poorly understood, especially in Amazonian floodplain forests. We investigated tree species composition, richness and α diversity in the Amazonian white‐water (várzea) forest, looking particularly at: (1) the flood‐level gradient, (2) the successional stage (stand age), and (3) the geographical location of the forests. Location Eastern Amazonia, central Amazonia, equatorial western Amazonia and the southern part of western Amazonia. Methods The data originate from 16 permanent várzea forest plots in the central and western Brazilian Amazon and in the northern Bolivian Amazon. In addition, revised species lists of 28 várzea forest inventories from across the Amazon Basin were used. Most important families and species were determined using importance values. Floristic similarity between plots was calculated to detect similarity variations between forest types and over geographical distances. To check for spatial diversity gradients, α diversity (Fisher) of the plots was correlated with stand age, longitudinal and latitudinal plot location, and flood‐level gradient. Results More than 900 flood‐tolerant tree species were recorded, which indicates that Amazonian várzea forests are the most species‐rich floodplain forests worldwide. The most important plant families recorded also dominate most Neotropical upland forests, and c. 31% of the tree species listed also occur in the uplands. Species distribution and diversity varied: (1) on the flood‐level gradient, with a distinct separation between low‐várzea forests and high‐várzea forests, (2) in relation to natural forest succession, with species‐poor forests in early stages of succession and species‐rich forests in later stages, and (3) as a function of geographical distance between sites, indicating an increasing α diversity from eastern to western Amazonia, and simultaneously from the southern part of western Amazonia to equatorial western Amazonia. Main conclusions The east‐to‐west gradient of increasing species diversity in várzea forests reflects the diversity patterns also described for Amazonian terra firme. Despite the fine‐scale geomorphological heterogeneity of the floodplains, and despite high disturbance of the different forest types by sedimentation and erosion, várzea forests are dominated by a high proportion of generalistic, widely distributed tree species. In contrast to high‐várzea forests, where floristic dissimilarity increases significantly with increasing distance between the sites, low‐várzea forests can exhibit high floristic similarity over large geographical distances. The high várzea may be an important transitional zone for lateral immigration of terra firme species to the floodplains, thus contributing to comparatively high species richness. However, long‐distance dispersal of many low‐várzea trees contributes to comparatively low species richness in highly flooded low várzea.     

Direct and indirect interactions co‐determine species composition in nurse plant systems

Facilitation by nurse plants plays an important role in determining community composition in severe environments. Although the unidirectional effect of nurses on beneficiary species has received considerable research interest, nurse‐mediated interactions among beneficiary species (so‐called indirect interactions) are less known. Consequently, community composition in nurse plant systems is generally considered as a simple consequence of the facilitative effect of the nurse even though beneficiary species may significantly contribute to community assembly and modulate the direct nurse effects on the community. In an observational study we assessed nurse effects and nurse‐mediated beneficiary interactions in two contrasting nurse plant systems in dry environments using a newly developed framework. We quantified plant–plant interaction intensity using the relative interaction index (RII) at the community and species level for three Retama sphaerocarpa shrub size‐classes in a semiarid shrubland and four Arenaria tetraquetra cushion plant communities differing in aspect and elevation in dry alpine gravel habitats. The observed RII was split into nurse and beneficiary effects, and related to individual mass, species frequency and abundance using generalized linear mixed models. Results showed predominantly positive nurse effects and negative beneficiary interactions. The effect size of nurse plants, however, was significantly higher than the effect size of beneficiary species in both systems. Individual plant mass and abundance of species was dependent on the combined effects of nurse and beneficiary species whereas species occurrence was related to nurse effects only. Despite evident differences, the semiarid and alpine nurse plant systems showed strong functional parallelisms. We found interdependence between the effects of nurse and beneficiary species on beneficiary plant assemblages emphasizing their combined role on community assembly in both systems. Our results highlight the need to consider indirect interactions to understand fully plant community dynamics.