Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

Functional rarity of plants in German hay meadows — Patterns on the species level and mismatches with community species richness

Functional rarity (FR) — a feature combining a species'' rarity with the distinctiveness of its traits — is a promising tool to better understand the ecological importance of rare species and consequently to protect functional diversity more efficiently. However, we lack a systematic understanding of FR on both the species level (which species are functionally rare and why) and the community level (how is FR associated with biodiversity and environmental conditions). Here, we quantify FR for 218 plant species from German hay meadows on a local, regional, and national scale by combining data from 6500 vegetation relevés and 15 ecologically relevant traits. We investigate the association between rarity and trait distinctiveness on different spatial scales via correlation measures and show which traits lead to low or high trait distinctiveness via distance‐based redundancy analysis. We test how species richness and FR are correlated, and use boosted regression trees to determine environmental conditions that are driving species richness and FR. On the local scale, only rare species showed high trait distinctiveness while on larger spatial scales rare and common species showed high trait distinctiveness. As infrequent trait attributes (e.g., legumes, low clonality) led to higher trait distinctiveness, we argue that functionally rare species are either specialists or transients. While specialists occupy a particular niche in hay meadows leading to lower rarity on larger spatial scales, transients display distinct but maladaptive traits resulting in high rarity across all spatial scales. More functionally rare species than expected by chance occurred in species‐poor communities indicating that they prefer environmental conditions differing from characteristic conditions of species‐rich hay meadows. Finally, we argue that functionally rare species are not necessarily relevant for nature conservation because many were transients from surrounding habitats. However, FR can facilitate our understanding of why species are rare in a habitat and under which conditions these species occur.     

Biocontrol potential varies with changes in biofuel–crop plant communities and landscape perenniality

We examined the potential local‐ and landscape‐level impacts of different biofuel production systems on biocontrol, an important service provided by arthropod natural enemies. Specifically, we sampled natural enemies with sweep nets and measured predation of sentinel pest eggs in stands of corn, switchgrass and mixed prairie in Michigan and Wisconsin (total n=40 for natural enemy sampling, n=60 for egg predation), relating them to crop type, forb cover and diversity, and the composition and heterogeneity of the surrounding landscape. Grasslands with intermediate levels of forb cover and flower diversity supported two‐orders of magnitude more natural enemy biomass, fourfold more natural enemy families, and threefold greater rates of egg predation than corn. Data suggest this was in part due to a general increase in biomass, richness and predation in perennial grasslands compared with corn, combined with a positive effect of intermediate levels of forb cover and flower diversity. Specifically, natural enemy biomass and family richness showed hump‐shaped relationships to forb cover that peaked in sites with 5–25% forbs, while egg predation increased with floral diversity. At the landscape scale, both natural‐enemy biomass and egg predation increased with the area of forest in the landscape, and egg predation almost doubled as the area of herbaceous, perennial habitats within 1.5 km of study sites increased. Our results suggest that floristically diverse, perennial grasslands support diverse and abundant predator communities that contribute to natural pest suppression. In addition, large‐scale production of biofuel crops could positively or negatively affect biocontrol services in agricultural landscapes through associated changes in the area of perennial habitats. Biofuel landscapes that incorporate perennial grasslands could support a variety of beneficial organisms and ecosystem services in addition to producing biomass.     

Rapid redistribution of agricultural land alters avian richness,abundance, and functional diversity

The conversion of natural, or seminatural, habitats to agricultural land and changes in agricultural land use are significant drivers of biodiversity loss. Within the context of land‐sharing versus land‐sparing debates, large‐scale commercial agriculture is known to be detrimental to biodiversity, but the effects of small‐scale subsistence farming on biodiversity are disputed. This poses a problem for sustainable land‐use management in the Global South, where approximately 30% of farmland is small‐scale. Following a rapid land redistribution program in Zimbabwe, we evaluated changes in avian biodiversity by examining richness, abundance, and functional diversity. Rapid land redistribution has, in the near term, resulted in increased avian abundance in newly farmed areas containing miombo woodland and open habitat. Conversion of seminatural ranched land to small‐scale farms had a negative impact on larger‐bodied birds, but species richness increased, and birds in some feeding guilds maintained or increased abundance. We found evidence that land‐use change caused a shift in the functional traits of the communities present. However, functional analyses may not have adequately reflected the trait filtering effect of land redistribution on large species. Whether newly farmed landscapes in Zimbabwe can deliver multiple benefits in terms of food production and habitat for biodiversity in the longer term is an open question. When managing agricultural land transitions, relying on taxonomic measures of diversity, or abundance‐weighted measures of function diversity, may obscure important information. If the value of smallholder‐farmed land for birds is to be maintained or improved, it will be essential to ensure that a wide array of habitat types is retained alongside efforts to reduce hunting and persecution of large bird species.     

Environmental gradients shift the direction of the relationship between native and alien plant species richness

Aim

To assess how environmental, biotic and anthropogenic factors shape native–alien plant species richness relationships across a heterogeneous landscape.

Location

Banks Peninsula, New Zealand.

Methods

We integrated a comprehensive floristic survey of over 1200 systematically located 6 × 6 m plots, with corresponding climate, environmental and anthropogenic data. General linear models examined variation in native and alien plant species richness across the entire landscape, between native‐ and alien‐dominated plots, and within separate elevational bands.

Results

Across all plots, there was a significant negative correlation between native and alien species richness, but this relationship differed within subsets of the data: the correlation was positive in alien‐dominated plots but negative in native‐dominated plots. Within separate elevational bands, native and alien species richness were positively correlated at lower elevations, but negatively correlated at higher elevations. Alien species richness tended to be high across the elevation gradient but peaked in warmer, mid‐ to low‐elevation sites, while native species richness increased linearly with elevation. The negative relationship between native and alien species richness in native‐dominated communities reflected a land‐use gradient with low native and high alien richness in more heavily modified native‐dominated vegetation. In contrast, native and alien richness were positively correlated in very heavily modified alien‐dominated plots, most likely due to covariation along a gradient of management intensity.

Main conclusions

Both positive and negative native–alien richness relationships can occur across the same landscape, depending on the plant community and the underlying human and environmental gradients examined. Human habitat modification, which is often confounded with environmental variation, can result in high alien and low native species richness in areas still dominated by native species. In the most heavily human modified areas, dominated by alien species, both native and alien species may be responding to similar underlying gradients.

     

Conventional land‐use intensification reduces species richness and increases production: A global meta‐analysis

Most current research on land‐use intensification addresses its potential to either threaten biodiversity or to boost agricultural production. However, little is known about the simultaneous effects of intensification on biodiversity and yield. To determine the responses of species richness and yield to conventional intensification, we conducted a global meta‐analysis synthesizing 115 studies which collected data for both variables at the same locations. We extracted 449 cases that cover a variety of areas used for agricultural (crops, fodder) and silvicultural (wood) production. We found that, across all production systems and species groups, conventional intensification is successful in increasing yield (grand mean + 20.3%), but it also results in a loss of species richness (?8.9%). However, analysis of sub‐groups revealed inconsistent results. For example, small intensification steps within low intensity systems did not affect yield or species richness. Within high‐intensity systems species losses were non‐significant but yield gains were substantial (+15.2%). Conventional intensification within medium intensity systems revealed the highest yield increase (+84.9%) and showed the largest loss in species richness (?22.9%). Production systems differed in their magnitude of richness response, with insignificant changes in silvicultural systems and substantial losses in crop systems (?21.2%). In addition, this meta‐analysis identifies a lack of studies that collect robust biodiversity (i.e. beyond species richness) and yield data at the same sites and that provide quantitative information on land‐use intensity. Our findings suggest that, in many cases, conventional land‐use intensification drives a trade‐off between species richness and production. However, species richness losses were often not significantly different from zero, suggesting even conventional intensification can result in yield increases without coming at the expense of biodiversity loss. These results should guide future research to close existing research gaps and to understand the circumstances required to achieve such win‐win or win‐no‐harm situations in conventional agriculture.     

Long‐term phosphite application maintains species assemblages,richness and structure of plant communities invaded by Phytophthora cinnamomi

The impact of the plant pathogen Phytophthora cinnamomi and the fungicide phosphite on species assemblages, richness, abundance and vegetation structure was quantified at three sites in Kwongkan communities in the Southwest Australian Floristic Region. Healthy and diseased vegetation treated with phosphite over 7–16 years was compared with non‐treated healthy and diseased vegetation. After site differences, disease had the greatest effect on species assemblages, species richness and richness within families. Disease significantly reduced cover in the upper and lower shrub layers and increased sedge and bare ground cover. Seventeen of 21 species assessed from the families Ericaceae, Fabaceae, Myrtaceae and Proteaceae were significantly less abundant in non‐treated diseased vegetation. In diseased habitats, phosphite treatment significantly reduced the loss of shrub cover and reduced bare ground and sedge cover. In multivariate analysis of species assemblages, phosphite‐treated diseased plots grouped more closely with healthy plots. Seven of 17 susceptible species were significantly more abundant in phosphite‐treated diseased plots compared with diseased non‐treated plots. The abundance of seven of 10 Phytophthora‐susceptible species was significantly higher along transects in phosphite‐treated vegetation. Comparison of the floristics of healthy non‐treated with healthy‐treated plots showed no significant differences in species assemblages. Of 21 species assessed, three increased in abundance and only one decreased significantly in phosphite‐treated healthy plots. In three Kwongkan communities of the SWAFR, P. cinnamomi had a profound impact on species assemblages, richness, abundance and vegetation structure. There was no evidence of adverse effects of phosphite treatment on phosphorus‐sensitive species, even after fire. Treatment with phosphite enhanced the survival of key susceptible species and mitigated disease‐mediated changes in vegetation structure. In the absence of alternative methods of control in native communities, phosphite will continue to play an important role in the protection of high priority species and communities at risk of extinction due to P. cinnamomi.     

Lichen species richness is highest in non-intensively used grasslands promoting suitable microhabitats and low vascular plant competition

Lichens are very sensitive to habitat changes and their species richness is likely to decline under intensive land use. Currently, a comprehensive study analyzing lichen species richness in relation to land-use types, extending over different regions and including information on habitat variables, is missing for temperate grasslands. In three German regions we studied lichen species richness in 490 plots of 16 m² representing different land-use types, livestock types, and habitat variables. Due to the absence of low-intensity pastures and substrates such as woody plants, deadwood and stones, there were no lichens in the 78 plots in Schorfheide-Chorin. In the two other regions, the richness of lichen species was 45 % higher in pastures than in meadows, and 77 % higher than in mown pastures, respectively. Among the pastures, the richness of all lichen species was on average 10 times higher in sheep-grazed pastures than in the ones grazed by cattle or horses. On average, the richness of all lichen species increased by 3.3 species per additional microhabitat. Furthermore, the richness of corticolous lichens increased by 1.2 species with 10 % higher cover of woody plants, lignicolous lichen species richness increased by 4.8 species with 1 % higher cover of deadwood, and saxicolous lichen species richness increased by 1.0 species with 1 % higher cover of stones. Our findings highlight the importance of low-intensity land use for lichen conservation. In particular, the degradation of grasslands rich in microhabitats and the destruction of lichen substrates by intensification, and conversion of unfertilized pastures formerly grazed at low intensity to meadows should be avoided to maintain lichen diversity.     

The phylogenetic properties of native‐ and exotic‐dominated plant communities

Phylogenetic properties of communities (phylogenetic diversity and phylogenetic structure) allow for the characterisation of phylogenetic patterns and provide the information necessary to infer mechanisms of species assembly. Because humans have introduced exotic species and modified the physical conditions of landscapes, the phylogenetic properties of communities should change according to the proportion of natives to exotics hosted by sites and to the strength of the conditions that act as habitat filters in human‐disturbed habitats. To assess the effects of the introduction of exotic plant species, we characterized the phylogenetic properties of 67 plant communities with different degrees of exotic species dominance in a region of central Chile with a Mediterranean climate. Five indices were used to estimate the phylogenetic properties. The Faith index (FPD), the mean pairwise distance (MPD) and the mean nearest neighbour distance (MNND) were used to estimate phylogenetic diversity, and the nearest relative index (NRI) and the nearest taxon index (NTI) were used as estimators of the phylogenetic structure (the phylogenetic distribution of taxa in a community) of species assemblages. We observed greater phylogenetic diversity of natives versus exotic plants despite the fact that natives accounted for a fewer number of taxa among the studied communities. Second, assemblages exhibited a phylogenetically clustered structure, which is attributable to an over‐representation of some families of exotic flora (Asteraceae, Brassicaceae, Fabaceae, Papaveraceae, Poaceae) and suggests habitat filtering processes that could have acted by selecting species with traits that permit adaptation to the harsh conditions of human‐disturbed sites.     

Coexistence through spatio-temporal heterogeneity and species sorting in grassland plant communities

The effect of spatial heterogeneity on species coexistence relies on the degree of niche heterogeneity in the habitat and the ability of species to exploit the available niche opportunities. We studied species coexistence in a perennial grassland, and tested whether small-scale disturbances create environmental heterogeneity that affects coexistence and whether the functional diversity of species in the species pool affects the ability of community composition to reflect heterogeneity through species sorting. We manipulated the spatio-temporal heterogeneity of disturbance and the functional diversity of species added as seed and measured their impact on the spatial turnover of species composition. Disturbance increased environmental heterogeneity and spatial turnover, and the effect of heterogeneity on turnover was greatest in the presence of a functionally diverse species pool, showing the importance of trait variation among species for exploiting environmental heterogeneity, and suggesting that coexistence occurred due to species sorting among heterogeneous niches.     

Grain-dependent relationships between plant productivity and invertebrate species richness and biomass in calcareous grasslands

The relationships among productivity, species richness and consumer biomass are of fundamental importance for understanding determinants of biodiversity. These relationships may depend on grain size. We examined the relationships between productivity (above-ground phytomass) and plant species richness and between productivity and species richness and biomass of gastropods and grasshoppers using sampling units of different sizes (0.5, 2.75 and 23 m²) in nutrient-poor, calcareous grasslands in north-western Switzerland in two successive years. Species richness of forbs had a unimodal relationship with productivity in sampling units of 0.5 m² and was negatively correlated with productivity at the other two plot sizes in one year. In the other year, forb species richness tended to decrease with productivity in sampling units of 23 m². No similar relationship was found for grasses. Gastropod biomass had a unimodal relationship with productivity at 0.5 m² in the first year. Grasshopper species richness was correlated with forb species richness at plot sizes of 2.75 and 23 m². This study demonstrates that patterns detected between productivity and diversity and between productivity and biomass of consumers depend on the grain size used in the investigation and vary among years.Die Zusammenhänge zwischen Produktivität, Artenreichtum und Biomasse von Konsumenten sind wichtig, um zu verstehen, was Biodiversität beeinflußt. Diese Zusammenhänge können von der Größe der Untersuchungsfläche abhängig sein. Wir untersuchten während zwei aufeinanderfolgenden Jahren die Zusammenhänge zwischen Produktivität (oberirdische Pflanzenbiomasse) und Artenreichtum von Gefäßpflanzen, sowie zwischen Produktivität und Artenreichtum und Biomasse von Schnecken und Heuschrecken bezüglich dreier räumlicher Skalen (0,5, 2,75 und 23 m²) in Kalkmagerrasen in der Nordwestschweiz. Der Zusammenhang zwischen dem Artenreichtum von Kräutern und der Produktivität war unimodal in Flächeneinheiten von 0,5 m² und negativ in Flächeneinheiten von 2,75 und 23 m² im ersten Jahr und war tendenziell negativ in Flächeneinheiten von 23 m² im zweiten Jahr, während kein solcher Zusammenhang bei Gräsern gefunden wurde. Der Zusammenhang zwischen Produktivität und Biomasse von Schnecken war unimodal in Flächeneinheiten von 0,5 m² im ersten Jahr. Außerdem bestand ein Zusammenhang zwischen dem Artenreichtum von Kräutern und Heuschrecken in Flächeneinheiten von 2,75 und 23 m². Diese Arbeit zeigt, daß Zusammenhänge zwischen Produktivität und Diversität sowie zwischen Produktivität und Biomasse von Konsumenten von der Größe der Untersuchungsfläche abhängen und zwischen Jahren variieren.     

Woody plant encroachment reduces species richness of herb‐rich woodlands in southern Australia

Abstract Woody plants have been increasing in many woodland and savanna ecosystems owing to land use changes in recent decades. We examined the effects of encroachment by the indigenous shrub Leptospermum scoparium (Myrtaceae) on herb‐rich Eucalyptus camaldulensis woodlands in southern Australia. Species richness and compositional patterns were examined under the canopy of L. scoparium and in surrounding open areas to determine the species most susceptible to structural changes. Richness was significantly lower in areas of moderate to high L. scoparium cover (>15%), suggesting that a threshold shrub cover caused major change in this ecosystem. Shrubs were associated with a significant reduction in above‐ground biomass of the ground‐layer flora and a significant shift in community composition. The few species that were positively associated with high L. scoparium cover were also common in the woodland flora; no new species were recorded under the shrub canopy. Important environmental changes associated with L. scoparium cover were decreased light availability and increased litter cover, which were likely a consequence of encroachment. Leptospermum scoparium cover was also associated with greater surface soil moisture, which may be a consequence of increased shading under the shrub canopy or indicate favourable soil conditions for L. scoparium establishment. Reductions in species richness and abundance of the germinable seed bank were found in soil samples taken from under L. scoparium. With ongoing recruitment of L. scoparium and consequent increases in shrub cover, ground‐layer diversity in these species‐rich woodlands should continue to decline over time.     

Biophysical and human influences on plant species richness in grasslands: Comparing variegated landscapes in subtropical and temperate regions

A survey of grassy woodlands in the Queensland subtropics was conducted, recording herbaceous species richness at 212 sites on three properties (2756 ha). A range of habitats typical of cattle grazing enterprises was sampled and site variables included lithology, slope position, tree density, soil disturbance, soil enrichment and grazing. Results were compared with a previously published survey of temperate grasslands. Lithology, slope position and tree density had relatively minor effects on plant species richness, although in both surveys there was some evidence of lower species richness on the more fertile substrates. Soil disturbance and soil enrichment significantly reduced the richness of native species in both surveys, while exotic species were insensitive (subtropics) or increased (temperate) with disturbance. Rare native species were highly sensitive to disturbances, including grazing, in the temperate study. Although some trends were similar for rare species in the subtropics, the results were not significant and there were complex interactions between grazing, lithology and slope position. Grazing did not have a negative effect on native species richness, except in the closely grazed patches within pastures, and then only on the most intensively developed property. At the scale recorded (30 m²), the native pastures, roadsides and stock routes sampled in the subtropics appear to be among the most species‐rich grasslands ever reported, both nationally and globally. Native species richness was approximately 50% higher than the temperate survey figures across all the comparable habitats. While there are no clear reasons for this result, potential explanations are proposed.