Evaluating different harvest intensities over understory plant diversity and pine seedlings,in a <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait. natural stand of Spain

Although modern forestry takes into consideration the analysis of the effects of forest management on plant structure, diversity and seedlings, little is known about how those parameters respond to harvest techniques in the Mediterranean region. We investigated the effect of three different harvest intensities, respect to uncut controls, on understory plant species functional groups, richness, diversity and pine seedlings in a natural Maritime pine stand in Spain, three years after harvesting. The harvest treatments produced a reduction of the number of Pinus pinaster seedlings and woody species cover, and an increase of species richness (total and of annual species) and plant cover of annual species respect to control plots (CO). The Shannon diversity values showed no differences between treatments. These results emphasize that the tree harvest treatments analyzed are not suitable for the management of this P. pinaster stand. Otherwise, the reduction of pine seedling density by harvest treatments and the changes in richness and cover of functional groups would not induce the natural regeneration of this stand maintaining the understory plant layer.     

Forest plant community as a driver of soil biodiversity: experimental evidence from collembolan assemblages through large‐scale and long‐term removal of oak canopy trees Quercus petraea

Plant–soil interactions are increasingly recognized to play a major role in terrestrial ecosystems functioning. However, few studies to date have focused on slow dynamic ecosystems such as forests. As they are vertically stratified by multiple vegetation strata, canopy tree removal by thinning operations could alter forest plant community through tree canopy opening. Very little is known about cascading effects on soil biodiversity. We conducted a large‐scale, multi‐site assessment of collembolan assemblage response to long‐term canopy tree removal in sessile oak Quercus petraea temperate forests. A total of 33 experimental plots were studied covering a large gradient of canopy tree basal area, stand age and local abiotic contexts. Collembolan abundance strongly declined with canopy tree removal in early forest successional stage and this was mediated by negative effect of understory plant community composition changes, i.e. shift from moss and forb to tree seedling, fern, shrub and grass species. Negative effect of this composition shift on collembolan species richness was largely offset by positive effect of the increase in understory plant species richness. This gives support to both the plant mass‐ratio and functional diversity hypotheses. Collembolan functional groups had contrasting response patterns, which were mediated by different ecological factors. Epedaphic (r‐strategist) abundance and species richness increased with canopy tree removal in relation with the increase in understory plant species richness. In contrast, euedaphic (K‐strategist) abundance and species richness declined with canopy tree removal in early forest successional stage in relation with changes in understory plant community composition and species richness, as well as microclimatic conditions. Overall, our study provides experimental evidence that forest plant community can be a strong driver of collembolan assemblages. It also emphasizes the role of trees as foundation species of forest ecosystems that can shape soil biodiversity through their regulation of understory plant community and ecosystem abiotic conditions.     

Establishment and Control of Hay-scented Fern: A Native Invasive Species

Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m² and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.     

不同间伐强度对南亚热带杉木人工林林下植物功能群的影响

间伐作为我国森林经营的主要措施之一,对提高森林生态系统内部生物多样性起着不可忽视的作用。然而,不同间伐强度对森林林下植物功能群的影响程度尚不明确。以广西凭祥热林中心经过4种不同强度抚育间伐8年后的杉木人工林为研究对象(轻度间伐ST,34%;中度间伐MT,50%;重度间伐HT,74%;不进行间伐的对照,CK),对林下植被及其环境因子进行调查和测定。研究结果表明:不同间伐强度对禾草植物功能群的物种丰富度均无显著影响(P0.05),MT显著增加杂草植物功能群的物种丰富度(P0.05),HT显著提高蕨类植物功能群的物种丰富度(P0.05),ST显著增加木本植物功能群的物种丰富度(P0.05),与CK相比,ST、MT和HT均能显著增加藤本植物功能群的物种丰富度(P0.05);不同间伐强度对禾草和木本植物功能群的重要值均无显著影响(P0.05),但HT显著降低了杂草植物功能群的重要值(P0.05),ST显著降低了蕨类植物功能群的重要值(P0.05),ST显著增加藤本植物功能群的重要值(P0.05)。主成分分析(PCA)表明,ST和HT的林下植物功能群组成和分布与对照组相比均发生了显著的变异。方差分解分析(Variation partitioning analyses)结果显示,林分和土壤因子对林下植物功能群变异的解释率相似,二者在决定杉木人工林林下植物功能群变异中都起着重要作用。     

Effects of large herbivore browsing on the functional groups of woody plants in a southern African savanna

This study examined the effect of different large herbivore species and stocking rates in savanna ecosystems of Zimbabwe on the richness and abundances of woody plant functional groups and woody plant functional attributes. Seven fence-lines with different herbivore species and stocking rates on either side of the fence were sampled. Plots were placed on both sides of each fence at each of 18 randomly selected positions. The size and species of each woody plant was recorded for each plot. It was found that the number of species with different functional attributes of spinescence, leaf longevity, fruit type and dispersal mechanism and in the functional groups of palatability were not different on the different sides of the fence. However, there were differences in plant abundances for 26 out of the 35 tests carried out on plant abundances with different functional attributes and functional groups. It was hypothesised that the time needed to change woody plant species richness is hundreds of years in these systems, whereas the time needed to change woody plant abundances is decades.     

除草剂对桉树人工林下植物及土壤微生物群落的影响

除草剂在桉树人工林中的应用越来越普遍,但关于除草剂对桉树人工林林下植物和土壤微生物群落的影响知之甚少。通过桉树人工林低剂量高频率(LHF)、中剂量中频率(MMF)、高剂量低频率(HLF)除草剂喷施试验,并与人工除草(MT)为对照,比较分析不同剂量、不同频率除草剂施用对林下植物和土壤微生物群落的影响。结果表明,施用除草剂导致桉树人工林林下植物种类和功能群组成发生显著变化,但并未显著降低林下植物群落物种丰富度和多样性,随除草剂施用频率的降低及恢复时间的增加,物种丰富度及多样性指数呈恢复趋势。除草剂施用也导致土壤养分含量降低。除草剂通过对林下植物群落和土壤养分的负面影响间接影响土壤微生物群落。LHF显著降低藤本植物而显著提高蕨类植物功能群的重要值,从而显著降低了微生物群落、真菌和放线菌的磷脂脂肪酸(PLFA)含量。MMF显著降低木本和藤本植物而显著提高禾草植物功能群的重要值,导致土壤微生物群落和放线菌的PLFA含量显著降低。HLF未显著影响林下植物及土壤微生物群落,但土壤全磷含量显著降低,速效磷含量也大幅下降。施用除草剂显著降低了土壤微生物生物量碳、氮的含量。因此,生产上应减少除草剂的施用,...     

Woody Encroachment Removal from Midwestern Oak Savannas Alters Understory Diversity across Space and Time

Recovering biodiversity is a common goal during restoration; however, for many ecosystems, it is not well understood how restoration influences species diversity across space and time. I examined understory species diversity and composition after woody encroachment removal in a large-scale savanna restoration experiment in central Iowa, United States. Over a 4-year time series, restoration had profound effects across space and time, increasing richness at local and site-level scales. Restoration sites had increased α (within sample) Simpson's diversity and α and γ (site level) species richness relative to control sites, although γ and β (among sample) Simpson's diversity, β richness, and α species evenness were not affected. Changes in richness were driven by graminoids at the α and γ scales and woody species (and some evidence for forbs) at the α scale. Interestingly, indicator species analysis revealed that at least some species from all functional groups were promoted by restoration, although no species were significant indicators of pre-treatment or control sites. Both savanna and nonsavanna species were indicators of restored sites. Restoration promoted exotic species at both scales, although species with spring phenologies were unaffected. Woody encroachment removal may be a means to promote species establishment in savannas; however, in this study, it resulted in establishment and proliferation of native and exotic and savanna and nonsavanna species. Future work might consider reintroduction of key savanna species to supplement those that have established. Work like this demonstrates the utility of restoration experiments for conducting research on large- and multiscale processes, such as species diversity.     

Passive Restoration of Atlantic Forest Following Pinus taeda Harvesting in Southern Brazil

The understory of exotic tree plantations can have non‐negligible native species richness. Ecological restoration of these sites may include the harvest of trees, depending on the tradeoff between timber income and harvest impacts on biodiversity. This study aimed to investigate how a site can recover from harvest disturbance, by comparing the regeneration of woody species in the understory of two types of 37‐year‐old Pinus taeda plantation (P1 and P2, high and low relative density of pine seedlings in the understory, respectively), with stands that were similar to P2 but subjected to harvest and then abandoned for 15 years (R sites). Secondary forests (SF) were used as references. We sampled three different sites for each stand condition; soil chemical properties, estimations of litter mass, and canopy cover were measured. P1 had low species diversity, and P2 and R had 50 and 46% of SF richness, respectively. The R site contained few pine saplings and was floristically similar to P2; this indicated that 15 years was sufficient for the recovery of plant diversity to near pre‐harvesting levels. Soil fertility was highest in SF and lowest in P1. Thus old plantations of P. taeda with low relative density of pine juveniles can be cost‐effective starting points for restoration. Despite the destructive effects of pine harvest, recovery of native species can occur rapidly. In situations in which clearcutting of pine stands is not planned or possible, modest thinning of P. taeda adults and/or intensive thinning of juveniles could expedite restoration.     

Alien Plant Invasions,Introduced Ungulates,and Alternative States in a Mesic Forest in Hawaii

Restoration by natural successional processes after removal of perturbations may not be feasible for many degraded ecosystems. Controlling major ecological threats such as non‐native ungulates is often a critical first step toward restoring native communities but past degradation, interactions with alien species and abiotic features may create conditions requiring additional intervention to ensure effective conservation. We monitored a series of fenced plots within diverse mesic forest on western Kauai in the Hawaiian Islands from 1998 to 2005 to determine the effects of ungulate removal on native and alien plant species. Relative to unfenced control plots, germination of seedlings and frequency of understory species of both native and alien species increased in the fenced plots. Density of both native and alien canopy and understory species declined more in unfenced than fenced plots, but density of native species declined more than alien species density in both fenced and unfenced plots. In fenced plots, the frequency of larger alien woody species and cover of an alien, mat‐forming fern species increased over time, indicating that fencing may encourage alien species that could interfere with regeneration of native species. Our study suggests that effective conservation of this and other remnant native Hawaiian forests will require both ungulate exclusion, removal of alien plant species with especially detrimental effects on native species, and proactive restoration programs for native species without natural sources of propagules. As the effects of invasive species continue to escalate, continental ecosystems lacking high endemism may also require similar interventions to preserve their biodiversity.     

Exploring the functional significance of forest diversity: A new long-term experiment with temperate tree species (BIOTREE)

Effects of biodiversity on ecosystem functioning have been mainly studied in experiments that artificially create gradients in grassland plant diversity. Woody species were largely excluded from these early experiments, despite the ecological and socioeconomic importance of forest ecosystems. We discuss conceptual aspects of mechanistically driven research on the biodiversity–ecosystem functioning relationship in forests, including the comparison of scientific approaches like ‘observational studies’, ‘removal experiments’, and ‘synthetic-assemblage experiments’. We give a short overview on the differences between herbaceous and forest ecosystems, focusing on canopy characteristics, and the possibilities for individual versus population-based investigations.We present detailed information about the first large-scale, multisite and long-term biodiversity–ecosystem functioning experiment with tree species of temperate forests (BIOTREE – BIOdiversity and ecosystem processes in experimental TREE stands). At three sites of differing geology and local climate, we planted 200,000 saplings on a total area of 70 ha. At two sites, diversity gradients were established by varying the number of tree species (BIOTREE-SPECIES). At a third site, only functional diversity at a constant level of tree species richness was manipulated by selecting mixtures that differ in the functional trait values of the corresponding species (BIOTREE-FD). Additional experimental treatments at the subplot level include silvicultural management options, the addition of subdominant species, and the reduction of genetic diversity. Response variables focus on productivity, biogeochemical cycles and carbon sequestration, and resource use complementarity.We explore the use of different measures of functional diversity for a posteriori classifications of functional richness and their use in the analysis of our tree diversity experiment. The experiment is thought to provide a long-term research platform for a variety of scientific questions related to forest biodiversity and ecosystem processes.     

Convergence in Arthropod Assemblages with Various Restoration Approaches for Canadian Deciduous Forests

Silvicultural practices are traditionally aimed at increasing forest profits; however, recent approaches to forest conservation have broadened to include nature-based silviculture for regenerating forests. In southern Ontario (Canada), originally dominated by deciduous forests, conifer plantations were established on abandoned agricultural sites. Currently, there is an increasing interest to convert these conifer stands to a state that mimics the original deciduous forest. We investigated arthropod abundance, species richness of carabid beetles, and abundance of arthropod assemblages (trophic and prey groups) under five silvicultural treatments conducted to regenerate deciduous forests (the natural forest type) from the old conifer plantations. The treatments included: (1) uniform canopy removal; (2) uniform canopy removal and understory removal; (3) group canopy removal; (4) group canopy removal and understory removal; and (5) untreated control plots (relatively pure red pine). Insects were sampled annually using sweepnets and pitfall traps. Results revealed treatment effects on the abundance of Coleoptera, Heteroptera, herbivores, and small arthropods (<3 mm) caught in sweepnet samples, where plots subjected to group shelterwood removal and understory removal supported higher abundances than the control plots. There was no treatment effect on the abundance of other arthropod groups or on the species richness and abundance of carabid beetles. The silvicultural treatments used to encourage natural regeneration did not seem to affect arthropod food availability for insectivorous vertebrates. Thus, the type of silvicultural strategy used to convert pine plantations to a stage that mimics the natural deciduous forests had little overall impact on arthropods.     

氮添加对红松人工林草本层植物多样性的影响

氮沉降是驱动生物多样性变化的重要因素之一。为了探索氮添加对红松（Pinus koraiensis）人工林草本层植物多样性的影响及其驱动机制,以黑龙江凉水国家级自然保护区红松人工林为研究对象,设置N0（对照处理,0 kg hm^-2 a^-1）、N20（低氮处理,20 kg hm^-2 a^-1）、N40（中氮处理,40 kg hm^-2 a^-1）和N80（高氮处理,80 kg hm^-2 a^-1）4个施氮水平,进行6年的氮添加实验。结果表明：（1）氮添加显著降低草本层3个功能群的密度和盖度,而对高度无显著影响;（2）6年氮添加使对照与施氮处理间群落相似度随施氮水平的增加而减小;（3）氮添加显著降低草本植物的丰富度和Shannon-Wiener多样性指数,而未对蕨类和木本植物的丰富度和Shannon-Wiener多样性指数产生显著影响,对草本层3个功能群的Pielou均匀度指数均无显著影响;（4）氮添加对草本植物的C、N、P含量、N：P、C：P产生显著影响,对木本植物的P含量、N：P、C：P产生显著影响,对蕨类植物的C：N：P生态化学计量均无显著影响;（5）草本植物多样性与土壤化学性质无显著的相关关系,草本植物丰富度、Shannon-Wiener多样性指数与植物盖度、密度呈显著的正相关关系,丰富度与植物N含量呈显著的负相关关系,Shannon-Wiener多样性指数与植物N：P呈显著的负相关关系。研究表明6年氮添加改变植物草本层中物种组成和群落结构,3个功能群密度和盖度显著降低,高度未产生显著变化,仅降低草本植物的丰富度和多样性。造成该现象的原因可能是,不同物种对于氮的利用特性和耐受程度存在差异,氮添加引起草本植物养分失衡,改变物种组成和群落结构,从而影响草本植物多样性。研究结果可为我国温带森林生态系统持续性管理提供数据和理论基础。     

Long-term consequences of mechanical fuel management for the conservation of Mediterranean forest herb communities

Mechanical clearing of understory vegetation is increasingly used in Euro-Mediterranean forests to reduce fire hazard, yet its long-term consequences for biodiversity remain poorly understood. This study analysed the influence of time since understory management and management frequency, on herbaceous species richness, cover and composition, functional richness and composition, and richness and cover within functional groups (life and growth forms, dispersal strategy, clonality, and plant height), using a chronosequence of cork oak (Quercus suber) stands spanning about 70 years. Overall species richness was virtually constant over time, but the richness of species with annual life form and plasticity in height was much higher in recently and recurrently treated stands; the opposite was found for perennial (mainly hemicryptophytes and chamaephytes), tussock-forming and clonal species richness, and functional richness. Overall herbaceous cover and that of annual, semi-basal, non-clonal and plastic species (in height) were favoured by recent and recurrent fuel treatments; cover by perennial (hemicryptophytes and chamaephytes), short basal, tussock-forming, and clonal species tended to increase for >10–20 years after management, and declined with management frequency. There was a marked shift in species and functional composition associated with time since understory management and management frequency. These findings suggest that widespread fuel management at <10 year intervals may shift understory herb communities to early-successional stages, impairing the persistence of species and functional groups recovering slowly after disturbance. Fuel management needs to balance the dual goals of fire hazard reduction and biodiversity conservation, retaining undisturbed patches in landscapes otherwise managed to reduce fuel accumulation.     

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.     

桉树与红锥混交对土壤养分及林下植物功能群的影响

桉树人工林对生态环境的影响一直是全球性争议的热点问题,桉树与珍贵乡土树种混交的生态环境效应备受关注。为探究桉树混交营林措施对林地土壤养分及林下植物功能群的影响,该研究以桉树纯林(PE)、桉树×红锥混交林(MEC)和红锥纯林(PCH)为对象,开展了林下植物群落及环境因子的调查测定。研究结果显示：不同林分的土壤理化性质具有显著差异,混交林的土壤pH、有效氮(AN)和有效磷(AP)含量显著高于纯林;而土壤含水量(SMC)、有机碳(SOC)、总氮(TN)含量及C∶N和C∶P在混交林中没有显著优势,并呈现PEMEC>PCH的趋势。混交林显著增加林下木本植物功能群(WFG)的物种丰富度,而PCH显著增加蕨类植物功能群(FeFG)的物种丰富度。混交林的WFG和禾草植物功能群(GFG)的重要值均显著高于红锥纯林,而红锥纯林FeFG的重要值显著高于混交林。主坐标分析结果表明,MEC与PE的林下植物功能群组成差异不显著,但与PCH的差异显著;冗余分析结果揭示了AN和AP是WFG占优势的主要影响因子,SMC、TN...     

Does Restoration Enhance Regeneration of Seasonal Deciduous Forests in Pastures in Central Brazil?

The goal of restoration is to accelerate ecosystem recovery, but in ecosystems that naturally regenerate rapidly restoration techniques need to be selected carefully to facilitate rather than impede natural recovery. We compared the effects of five restoration techniques, such as plowing the soil, removing grasses, adding forest litter, seeding, and planting nursery‐growing seedlings, on the regeneration of seasonal deciduous forest trees in four abandoned pastures in central Brazil. We monitored all woody stems immediately prior to treatments and again 14 months after the treatments. We recorded an average of 16,663 tree stems per hectare and a total of 83 species before implementing treatments. Planting strongly increased species richness; adding litter and seeding had weaker positive effects on richness; and plowing and grass removal had no effect. Plowing substantially reduced the density of naturally established stems. Despite the high survival of planted seedlings, stem density in planting treatments did not change because the tractor and digging holes to plant seedlings caused mortality of naturally regenerating seedlings. Tree stems grew more in the grass release plots than in the control plots. Our results suggest that early succession of seasonal deciduous forest in pastures in the region studied does not need to be stimulated once the perturbation is stopped and that intensive restoration efforts may actually slow recovery. We recommend only enrichment planting of seedlings that are not able to resprout.     

Woody species as landscape modulators: their effect on the herbaceous plants in a Mediterranean maquis

“Landscape modulators” are ecosystem engineers that have an impact on community structure by creating patches in the landscape mosaic. Our aim was to study the effect of evergreen-trees, as landscape modulators, on herbaceous plants in a Mediterranean maquis system in northern Israel. We examined the effects of canopy removal and cattle grazing on species richness, plant functional types, and rare plant species in two patch-types: (1) woody—under tree canopy (or the location of a removed canopy); (2) herbaceous—in open areas with no tree canopy. Patch-type and tree removal affected species richness and plant functional types. The extreme negative effect of the woody patch-type on species richness disappeared soon after the removal of the landscape modulator canopy. We conclude that the dominant effect of the evergreen woody landscape modulators can be regulated by canopy removal and grazing for maintaining patch-type and landscape diversities, and consequent high species richness in Mediterranean ecosystems, which is a main goal of global nature conservation policy.     

Ecological consequences of dead wood extraction in an arid ecosystem

Coarse woody debris (CWD) plays a key role in ecosystems, reducing erosion and affecting soil development, storing nutrients and water, providing a major source of energy and nutrients, serving as a seedbed for plants and as habitat for decomposers and heterotrophs. We asked whether removal of CWD affected the structure and functioning of an arid woodland ecosystem in mid-western Argentina. These woodlands are protected by national laws and inhabited by indigenous local pastoralists who have land rights to use natural resources, including wood for fuel and construction material. We hypothesized that removal of CWD affected negatively the populations of wood-nesting pollinators, the reproductive performance of Prosopis flexuosa (the dominant tree species), plant cover, richness and composition, and nutrient cycling in the soil. We conducted a manipulative experiment consisting of four pairs of 70 m radius circular plots, each pair with an experimental (CWD removal) and a control (no removal) plots. Experimental CWD extraction affected negatively flower visitor abundance, although the magnitude of this effect decreased over time. In contrast, extraction had a significant, positive effect on seed production of P. flexuosa. No significant effects of extraction were found on cover, richness and composition of understory plants and soil properties. Thus, CWD did not have the generally negative effects expected under our hypotheses.     

Species versus genotypic diversity of a nitrogen-fixing plant functional group in a metacommunity

Exploring species and genetic diversity interactions provides new opportunities for furthering our understanding of the ecology and evolution of population and community dynamics, and for predicting responses of ecosystems to environmental change. Theory predicts that species diversity within communities and genetic diversity within populations will covary positively, because either species and genetic diversity interact synergistically or they respond in parallel fashion to common habitat conditions. We tested the hypothesis of positive covariation between species and genotypic diversity in a metacommunity of the species-rich southwest Australian flora. We hypothesised that the connection between genotypic diversity and species diversity is strong within functional groups, but weak or non-existent if the species considered extend beyond the functional group. We show that allelic richness of Daviesia triflora, an ant-dispersed pea, covaries positively with the species richness of six co-occurring nitrogen-fixing legume species. No pattern can be detected between allelic richness of D. triflora and species richness of ant-dispersed species when four non-legumes are added. We also show that genetic diversity of D. triflora is not governed by the same environmental factors that determine the presence of a group of large-shrub/small-tree species in the same metacommunity. This study shows that species and genetic diversity covariation are more likely to be confined to within, rather than between, plant functional groups.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.