Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Analyzing mechanisms regulating diversity in rangelands through comparative studies: a case in the southwestern Pyrennees

The preservation of the floristic diversity of natural ecosystems and the maintenance of extensive livestock grazing are two of the major priorities of European Union agricultural and environmental policies. Temperate natural and seminatural grasslands are among the most threatened ecosystems in Europe. In order to establish the most appropriate management practices that prevent degradation and maintain the richness of these ecosystems, detailed analyses of the mechanisms that regulate biodiversity of the communities are particularly necessary. In this study we make a comparative analysis of the species richness and the diversity of the main rangeland communities in a widespread, heterogeneous area located in the Western Spanish Pyrennees. The nature and the contribution of perennials and annuals to the diversity patterns encountered are studied and related to the mechanisms suggested to regulate diversity in each case. Higher diversity values were reported in more environmentally stressed, xeric and moist communities. High spatial and temporal heterogeneity and low rates of competitive displacement related to the scarcity of basic resources are among the factors suggested to regulate the diversity of these communities. In areas where environmental factors are more favorable, such as perennial-dominated mesic rangelands, the enhancement of the diversity seemed more dependent on biotic disturbances, such as herbivory, that decreased the cover of perennials and favored the recruitment of short-lived species. Dispersal processes related to traditional nomadic grazing allow a flow of species from one rangeland to another but also enhance the establishment of species coming from lowland habitats. Most of the annuals enhancing the diversity of the mesic community were ruderals. The consideration of ruderals as exotics or as species belonging to this grazing ecosystem poses an interesting, controversial topic related to the ecological quality of the diversity estimated for these communities.     

The vertical distribution of below-ground biomass in grassland communities in relation to grazing regime and habitat characteristics

Abstract. 40 sites, representing different pasture types in Northwest Spain, were sampled in respect of their floristic composition, distribution of above and below-ground biomass and environmental and physical variables. Five plant community types were identified by classification techniques of plant species composition. These communities were then characterized in terms of the percentage of ground covered by herbaceous and shrub vegetation, stones, rocks and gaps as well as their topographic location and characteristics of the shallow soil (pH, organic matter, nitrogen and calcium content). Bio-mass was assessed in terms of above-ground structures, surface crowns and three below-ground layers to a depth of 10 cm. Three types of grazing regime were distinguished: Concentrated Intense Grazing in early spring (CIG), Extended Intense Grazing throughout the spring (EIG), and Non-Intense Grazing (NIG). Grazing regime showed the highest association with plant community type and three broad categories were identified: xeric stressed pastures, which nevertheless received CIG, mesic pastures with EIG, and three kinds of NIG mesic pastures. The xeric communities had the highest proportion of aboveground biomass, as a consequence of their greater proportion of woody perennials. These xeric communities displayed a more gradual reduction in below-ground biomass with depth than mesic pastures, a likely consequence of the low water content in the upper soil layers. The mesic communities had a high concentration of below-ground biomass in the upper layers when they were intensely grazed. However, when grazing was low (i.e. NIG situations), these communities had greater variability in biomass profiles than any of the other pasture types. Possible causes of the patterns in biomass distribution of the intensely grazed pastures are discussed.     

Impacts of the invasive exotic <Emphasis Type="Italic">Prosopis juliflora</Emphasis> (Sw.) D.C. on the native flora and soils of the UAE

The effects of the invasive exotic Prosopis juliflora shrubs on the natural plant communities and soil chemical characters were assessed in two regions of the United Arab Emirates (UAE). Five sites were selected subjectively: three in Sharja with 73 stands and two in Ras Al-Khima with 37 stands. Stands were located randomly within each site to cover density variation in Sharja and size variation in Ras Al-Khima. Density, frequency, richness and evenness of the associated annual and perennial species were studied in nine quadrats distributed under, at the margin and outside the canopy of a P. juliflora shrub located in the center of each stand. The results indicated that the effect of P. juliflora on the associated flora depends significantly on the density and size of the canopy. Larger individuals and greater densities have significantly greater negative impacts on the associated plants. All the studied community attributes were significantly lower under P. juliflora canopies than outside. Annuals were inhibited more than perennials. The number of annuals with significant reductions in density and/or frequency under P. juliflora canopies was significantly greater than the number of perennials. Density of more than 50% of the associated annuals was significantly inhibited under P. juliflora canopies. Density of P. juliflora seedlings was greater underneath the canopy of the same species than away from them, indicating little or no self allelopathic effect (auto-inhibition) under field conditions. P. juliflora ameliorated some soil characters, through significant pH reduction and increase in K, N and P and organic matters. Hence, plant diversity might be enhanced following eradication of P. juliflora.     

Epifaunal community structure in Acropora spp. (Scleractinia) on the Great Barrier Reef: implications of coral morphology and habitat complexity

The role of microhabitat in structuring epifaunal communities on four corals of varying morphology in the genus Acropora (A. millepora, A. hyacinthus, A. pulchra, A. formosa) was determined on two fringing reefs in the central Great Barrier Reef. Greater abundance and species richness of epifauna on tightly branched coral species in comparison to their rarity or absence on open-branched species suggests that protection afforded by complex habitats is important in structuring coral epifaunal communities. Within species, neither total colony space nor live surface area of corals was correlated with either the abundance or species richness of associated epifauna. However, space between branches significantly affected the size of Tetralia crabs associated with different coral species. Patterns in the size distribution of Tetralia on two species of Acropora suggest that crabs select coral hosts according to branch spacing, changing host species as they grow larger.     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.     

Variation in community structure of gall‐inducing insects associated with a tropical plant supports the hypothesis of competition in stressful habitats

Environmental factors act as drivers of species coexistence or competition. Mesic environments favor the action of parasites and predators on gall communities, while the factors that determine the structure of gall communities in xeric environments remain unknown. We evaluated the structure of gall communities along an environmental gradient defined by intrinsic plant characteristics, soil fertility, and aridity, and investigated the role of competition as a structuring force of gall communities in xeric environments. We created null models to compare observed and simulated patterns of co‐occurrence of galls and used the C‐score index to assess community aggregation or segregation. We used the NES C‐score (standardized C‐score) to compare patterns of co‐occurrence with parameters of environmental quality. Xeric environments had poorer and more arid soils and more sclerophyllous plants than mesic environments, which was reflected in the distribution patterns of gall communities. Values of the C‐score index revealed a segregated distribution of gall morphospecies in xeric environments, but a random distribution in mesic environments. The low availability of resources for oviposition and the high density of gallers in xeric environments reinforce interspecific competition as an important structuring force for gall communities in these environments.     

Galling arthropod diversity in adjacent swamp forests and restinga vegetation in Rio Grande do Sul, Brazil

Galling arthropods create plant structures inside which they find shelter. Factors acting on galler diversity are still being discussed, with this fauna considered more diverse in xeric than mesic environments (higrothermic stress hypothesis, HSH), and also in more plant diverse sites. Here we compare galler abundance (N), equitability (E), species richness (S) and composition between adjacent restinga (xeric) and swamp forests (mesic) in Parque Estadual de Itapeva (29°21' S, 49°45' W), Rio Grande do Sul, southern Brazil. Five trails, two in swamp forest and three in restingas, were sampled four times each (January/December 2005). After an effort of 60h/person, 621 galled plant individuals belonging to 104 gall morphotypes were recorded. This suggests a high galler diversity for the Park, comparable to the richest places known. No differences were found for N, E or S between restingas and swamp forests. However, faunal composition differs significantly between the vegetation types. The dominant (most abundant) species are different in either vegetation type, and are rare or absent on the other vegetation type. Such species composition analysis is still largely ignored for gallers, and stresses the fact that the HSH cannot explain this pattern, since the latter is based on preferences by the ovipositing galler for xeric sites instead of mesic ones. The two habitats differ in microclimate, but species richness, as would be predicted by the HSH, does not differ. This small scale pattern can perhaps be attributed to biogeographic processes on larger scales, as suggested by the resource synchronisation hypothesis.     

Stress‐Driven Changes in the Strength of Facilitation on Tree Seedling Establishment in West African Woodlands

The strength of competitive and facilitative interactions in plant communities is expected to change along resource gradients. Contrasting theoretical models predict that with increasing abiotic stress, facilitative effects are higher, lower, or similar than those found under more productive conditions. While these predictions have been tested in stressful environments such as arid and alpine ecosystems, they have hardly been tested for more productive African woodlands. We experimentally assessed the strength of tree seedling facilitation by nurse trees in mesic and dry woodlands in Benin, West Africa. We planted seedlings of the drought‐sensitive Afzelia africana and the drought‐tolerant Khaya senegalensis under three microsite conditions (closed woodland, woodland gap, and open fields). Seedling survival was greater within woodlands compared with open fields in both the mesic and dry woodlands. The relative benefits in seedling survival were larger at the dry site, especially for the drought‐sensitive species. Nevertheless, plant interactions became neutral or negative during the dry season in the drier woodland, indicating that the net positive effects may be lost under very stressful abiotic conditions. We conclude that facilitation also occurs in the relatively more productive conditions of African woodlands. Our results underscore the role of environmental variation in space and time, and the stress tolerance of species, in explaining competitive and facilitative interactions within plant communities. Abstract in French is available at http://www.blackwell‐synergy.com/loi/btp .     

Role of topography and soils in grassland structuring at the landscape and community scales

The relative importance of abiotic factors in community assembly is debated and thought to be dependent on the scale. I investigated the relative role of topography and soils as structuring agents at the landscape and the community scales in 126 subalpine calcareous grasslands in the Pyrenees, in terms of species composition and abundance. I wished to know: (1) the role of abiotic factors in the organization of plant communities across the landscape; (2) how much of the variation in community distribution was accounted for by abiotic factors; and (3) how well their role applied to the distribution of dominant species at the landscape and the community scales. The hypothesis was: abiotic factors play an important role in community distribution in the landscape, but species interactions are more important within communities. Multivariate methods generated four communities, organized in two contrasting groups along the main vegetation axis, which explained 13% of the variation: mesic grasslands (Nardus stricta and Festuca nigrescens communities) and xeric grasslands (Carex humilis and Festuca gautieri communities). Mesic communities were more acidic and fertile than xeric communities. Changes in the abiotic environment, accounting for up to 80% of the variation in the vegetation, were smooth, while the transition between xeric and mesic grasslands was sharp in terms of species composition. The distribution in the landscape of the first main species from each community was closely related to abiotic factors, which modeled poorly the abundance of the main species at smaller scales. At the within-community scale, the explanatory power of biotic relationships was community dependent, producing the most significant models for plants highly dominant within their communities, such as N. stricta and F. gautieri. Contrary to current hypothesis, there was a shift from mainly positive relationships among dominant species in fertile mesic communities to mainly negative in infertile xeric ones.     

Plant diversity,herbivory and resistance of a plant community to invasion in Mediterranean annual communities

Several components of the diversity of plant communities, such as species richness, species composition, number of functional groups and functional composition, have been shown to directly affect the performance of exotic species. Exotics can also be affected by herbivores of the native plant community. However, these two possible mechanisms limiting invasion have never been investigated together. The aim of this study was to investigate the relationships between plant diversity, herbivory and performance of two annual exotics, Conyza bonariensis and C. canadensis, in Mediterranean annual communities. We wanted to test whether herbivory of these exotics was influenced either by species richness, functional-group richness or functional-group composition. We also studied the relationship between herbivory on the exotic species and their performance. Herbivory increased with increasing species and functional-group richness for both Conyza species. These patterns are interpreted as reflecting a greater number of available herbivore niches in a richer, more complex, plant community. The identities of functional groups also affected Conyza herbivory, which decreased in the presence of Asteraceae or Fabaceae and increased in the presence of Poaceae. Increasing herbivory had consequences for vegetative and demographic parameters of both invasive species: survival, final biomass and net fecundity decreased with increasing herbivory, leading to a loss of reproductive capacity. We conclude that communities characterised by a high number of grass species instead of Asteraceae or Fabaceae may be more resistant to invasion by the two Conyza species, in part due to predation by native herbivores.     

Vegetation recovery and plant facilitation in a human-disturbed lava field in a megacity: searching tools for ecosystem restoration

Unplanned urban development threatens natural ecosystems. Assessing ecosystem recovery after anthropogenic disturbances and identifying plant species that may facilitate vegetation regeneration are critical for the conservation of biodiversity and ecosystem services in urban areas. At the periphery of Mexico City, illegal human settlements produced different levels of disturbance on natural plant communities developed on a lava field near the Ajusco mountain range. We assessed natural regeneration of plant communities 20 years after the abandonment of the settlements, in sites that received low (manual harvesting of non-timber forest products), medium (removal of aboveground vegetation), and high (removal of substrate and whole vegetation) disturbance levels. We also tested the potential facilitative role played by dominant tree and shrub species. Plant diversity and vegetation biomass decreased as disturbance level increased. Sites with high disturbance level showed poor regeneration and the lowest species similarity compared to the least disturbed sites. Six dominant species (i.e., those with the highest abundance, frequency, and/or basal area) were common to all sites. Among them, three species (the tree Buddleja cordata, and two shrubs, Ageratina glabrata and Sedum oxypetalum) were identified as potential facilitators of community regeneration, because plant density and species richness were significantly higher under their canopies than at open sites. We propose that analyzing community structural traits of the successional vegetation (such as species diversity and biomass) and identifying potential facilitator species are useful steps in assessing the recovery ability of plant communities to anthropogenic disturbances, and in designing restoration strategies.     

Responses of different herb life-history groups to a dominant shrub species along a dune stabilization gradient

Shrubs play a pivotal role in reversing desertification and in promoting ecological rehabilitation in severe environments. However, how the interactions between shrubs and their understorey species change during restoration remains unclear. Here, we report the results of an observational study conducted in the Mu Us Desert of northern China. This study explored how dune stabilization and the size of individual shrubs affect shrub–herb interactions. In particular, we aimed to determine how different life-history groups and performance indicators (e.g., biomass and richness) of subordinate species respond to shrub–herb interactions during dune stabilization. The shrub Artemisia ordosica had positive effects on understorey species in this dune system. The ability of the shrub to promote the growth of understorey species increased with dune stabilization, but decreased from small to large shrubs. This effect was due to an increase in the relative abundance and biomass of perennials and their higher sensitivity to the positive effects of the shrubs. In contrast, the ability of shrubs to improve the richness of understorey species decreased with dune stabilization, but increased from small to large shrubs. This effect occurred because perennials suffered high strain during recruitment in disturbed open patches of the semi-fixed dunes, particularly below small shrubs. Our results support the theory claiming that communities are sets of hidden interaction groups that have contrasting responses (neutral for annuals, but facilitative for perennials) to dominant neighbors in a single community, depending on their functional strategies. Furthermore, our study highlights the high efficiency of A. ordosica in facilitating ecological restoration of dry and disturbed sandy communities.     

Fungal community composition in neotropical rain forests: the influence of tree diversity and precipitation

Plant diversity is considered one factor structuring soil fungal communities because the diversity of compounds in leaf litter might determine the extent of resource heterogeneity for decomposer communities. Lowland tropical rain forests have the highest plant diversity per area of any biome. Since fungi are responsible for much of the decomposition occurring in forest soils, understanding the factors that structure fungi in tropical forests may provide valuable insight for predicting changes in global carbon and nitrogen fluxes. To test the role of plant diversity in shaping fungal community structure and function, soil (0-20?cm) and leaf litter (O horizons) were collected from six established 1-ha forest census plots across a natural plant diversity gradient on the Isthmus of Panama. We used 454 pyrosequencing and phospholipid fatty acid analysis to evaluate correlations between microbial community composition, precipitation, soil nutrients, and plant richness. In soil, the number of fungal taxa increased significantly with increasing mean annual precipitation, but not with plant richness. There were no correlations between fungal communities in leaf litter and plant diversity or precipitation, and fungal communities were found to be compositionally distinct between soil and leaf litter. To directly test for effects of plant species richness on fungal diversity and function, we experimentally re-created litter diversity gradients in litter bags with 1, 25, and 50 species of litter. After 6?months, we found a significant effect of litter diversity on decomposition rate between one and 25 species of leaf litter. However, fungal richness did not track plant species richness. Although studies in a broader range of sites is required, these results suggest that precipitation may be a more important factor than plant diversity or soil nutrient status in structuring tropical forest soil fungal communities.     

Effects of plant species richness and evenness on soil microbial community diversity and function

Understanding the links between plant diversity and soil communities is critical to disentangling the mechanisms by which plant communities modulate ecosystem function. Experimental plant communities varying in species richness, evenness, and density were established using a response surface design and soil community properties including bacterial and archaeal abundance, richness, and evenness were measured. The potential to perform a representative soil ecosystem function, oxidation of ammonium to nitrite, was measured via archaeal and bacterial amoA genes. Structural equation modeling was used to explore the direct and indirect effects of the plant community on soil diversity and potential function. Plant communities influenced archaea and bacteria via different pathways. Species richness and evenness had significant direct effects on soil microbial community structure, but the mechanisms driving these effects did not include either root biomass or the pools of carbon and nitrogen available to the soil microbial community. Species richness had direct positive effects on archaeal amoA prevalence, but only indirect impacts on bacterial communities through modulation of plant evenness. Increased plant evenness increased bacterial abundance which in turn increased bacterial amoA abundance. These results suggest that plant community evenness may have a strong impact on some aspects of soil ecosystem function. We show that a more even plant community increased bacterial abundance, which then increased the potential for bacterial nitrification. A more even plant community also increased total dissolved nitrogen in the soil, which decreased the potential for archaeal nitrification. The role of plant evenness in structuring the soil community suggests mechanisms including complementarity in root exudate profiles or root foraging patterns.     

The Impact of Seeding Method on Diversity and Plant Distribution in Two Restored Grasslands

Previous studies have compared grassland restoration techniques based on resulting species richness and composition. However, none have determined if different techniques generate different plant distributions in space, which may further impact restoration success. This study tests if there are quadrat‐scale (1 m²) differences between paired drilled and broadcast plantings in diversity, composition, and plant distributions. Higher competition intensity in and more contiguous spaces between rows in drill‐seeded restorations were hypothesized to result in larger patches of native grasses and exotic species. Two paired drill‐ and broadcast‐seeded plantings were sampled in June 2007 in Iowa, U.S.A. Within 10 quadrats in each planting, we measured species abundance with point intercept sampling and plant distributions by dividing the quadrat into 64 cells and recording the most abundant species in each cell. Drilled and broadcast plantings at both sites had similar Simpson’s diversity and evenness. However, the effect of planting type on species richness, composition, and plant distribution was site dependent. Native warm‐season grasses in one site, and exotic species in the second, occupied more space and were distributed in larger patches in drilled plantings. Furthermore, drilled canopies consistently captured more light than broadcast canopies. This suggests that initial differences in seed placement can affect resulting plant distributions, resource use, and potentially long‐term species turnover. Mechanisms structuring vegetation in these communities need to be further investigated to determine if this approach can provide more information on long‐term diversity maintenance in restorations than traditional measures.     

Patterns of functional clonal traits and clonal growth modes in contrasting grasslands in the central Apennines,Italy

Abstract. Aim: Patterns of plant functional traits related to clonality (clonal growth modes; CGM) in plant communities were studied and hypotheses on the importance of the selected traits in plant communities supported by soils differing in moisture and nutrient status were tested. Material and Methods: Selected plant functional traits, such as the position of the mother‐daughter plants connections, length of spacers, frequency of multiplication, persistency of ramets connections, presence of storage organs and bud protection were studied in two contrasting plant communities (xeric and mesic abandoned pastures) typical of central Apennines, Italy. Results and Discussion: Clonality was shown to be of great importance in both mesic and xeric grasslands. The major differences between the two communities were due to the dominant CGMs: turf graminoids (having effective protection of growth meristems in dense tussocks) dominated xeric grasslands, while rhizomatous graminoids (typical of competitive resource‐rich environments) dominated mesic grasslands. Below‐ground CGOs (clonal growth organs), shorter spacers, higher multiplication potential, permanent ramet connection, large bud bank and increased importance of bud protection were found to be of importance in water stressed xeric grassland. Contrary to our expectations, the mesic (less stressed) grasslands have the higher number of clonal plants possessing storage organs.     

Impacts of ecosystem engineers on community attributes: effects of cushion plants at different elevations of the Chilean Andes

Ecosystem engineers are organisms able to modulate environmental forces and, hence, may change the habitat conditions for other species. In so doing, ecosystem engineers may affect both species richness and evenness of communities and, in consequence, change species diversity. If these changes in community attributes are related to the magnitude of the habitat changes induced by the engineers, it seems likely that engineer species will have greater effects on diversity in sites where they cause larger habitat changes. We addressed this issue by evaluating the effects of three alpine cushion plants on species richness, evenness, and diversity of high-Andean plant communities. Given that the difference in microclimatic conditions between cushions and the external environment increases with elevation, we proposed that these organisms should have greater effects on community attributes at higher than at lower elevation sites. Results showed that the three cushion species had positive effects on species richness, diversity, and evenness of plant communities. It was also observed that the magnitude of these effects changed with elevation: positive effects on species richness and diversity increased towards upper sites for the three cushions species, whereas positive effects on evenness increased with elevation for one cushion species but decreased with elevation for other two cushion species. These results suggest that the presence of cushions is important to maintain plant diversity in high-Andean communities, but this positive effect on diversity seems to increase as the difference in environmental conditions between cushions and the external environment increases with elevation.     

Variation in serotiny of three Banksia species along a climatic gradient

The degree of serotiny (i.e. the proportion of follicles remaining closed in each year's crop of cones since the last fire) was measured in Bank-sia attenuata, B. menziesii and B. prionotes at five sites along a climatic gradient extending 500 km north of Perth, Western Australia. The decrease in annual rainfall and increase in average temperature along the gradient paralleled a decrease in plant height and an increase in the degree of serotiny of all species. Extreme serotiny was recorded in the scrub-heath at the xeric end of the gradient whereas two species were essentially non-serotinous in the low woodland at the most mesic site. It is concluded that degree of serotiny is related to the fire characteristics of the site which depend on plant height. In xeric scrub-heath, the entire canopies of the Banksia spp. are consumed by fire which promotes massive release of seed. This facilitates recruitment in an otherwise unpredictable and unreliable seedbed. In mesic woodland, where cones rarely come into contact With flames, seeds are released spontaneously and site conditions are more conducive to recruitment in the inter-fire period.     

Rapid Increases in Forest Understory Diversity and Productivity following a Mountain Pine Beetle (Dendroctonus ponderosae) Outbreak in Pine Forests

The current unprecedented outbreak of mountain pine beetle (Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests of western Canada has resulted in a landscape consisting of a mosaic of forest stands at different stages of mortality. Within forest stands, understory communities are the reservoir of the majority of plant species diversity and influence the composition of future forests in response to disturbance. Although changes to stand composition following beetle outbreaks are well documented, information on immediate responses of forest understory plant communities is limited. The objective of this study was to examine the effects of D. ponderosae-induced tree mortality on initial changes in diversity and productivity of understory plant communities. We established a total of 110 1-m² plots across eleven mature lodgepole pine forests to measure changes in understory diversity and productivity as a function of tree mortality and below ground resource availability across multiple years. Overall, understory community diversity and productivity increased across the gradient of increased tree mortality. Richness of herbaceous perennials increased with tree mortality as well as soil moisture and nutrient levels. In contrast, the diversity of woody perennials did not change across the gradient of tree mortality. Understory vegetation, namely herbaceous perennials, showed an immediate response to improved growing conditions caused by increases in tree mortality. How this increased pulse in understory richness and productivity affects future forest trajectories in a novel system is unknown.