Seed Bank Dynamics of French Broom in Coastal California Grasslands: Effects of Stand Age and Prescribed Burning on Control and Restoration

Abstract Successful control of invasive exotic plants depends to a large degree on the regeneration potential of the target exotic species and other species that might also be influenced by the removal effort. In coastal grasslands of California, exotic French broom (Genista monspessulana) forms both dense stands aboveground and abundant seed banks belowground. Land managers attempt to reduce broom cover and the seed bank through prescribed burning, but before this study, no investigators had examined the effect of repeated burning on French broom and associated grassland species. We found that the soil seed bank of stands that were burned had fewer broom seeds than unburned areas but that repeated burning did not reduce the seed bank beyond what was observed after one fire. Fire also did not have any consistent effect on the seed banks of other grassland species. We also examined the relationship between broom stand age and seed bank size but did not find a strong relationship between them. Our data suggested that the broom seed bank stays constant or declines slightly with stand age. We did, however, find that nonbroom seed numbers decreased as broom stands aged. Our results suggest that fire does reduce the size of the broom seed bank and that control of broom need not be limited to only the youngest stands.     

Shrubby today but not tomorrow? Structure,composition and regeneration dynamics of direct seeded revegetation

Summary The establishment of direct seeded revegetation is well researched. However, there is little understanding of whether revegetation simplifies with age and loses many of the short‐lived understorey shrub species that provide critical resources for birds and other fauna, or regenerates sufficiently to be self‐sustaining. We sought to address this by investigating the change in structure and composition of 33 direct seeded sites established by Greening Australia between 1990 and 1996 in the Southern Tablelands of NSW. Transects were used to collect data describing the abundance and richness of woody plants in 1998 and 2008, and the abundance of woody plant regeneration in 2008. Our analysis showed the predicted number of live stems per metre declined exponentially from 5.8 stems/m (～17 000 stems/ha) at age 1.5 years to 1.5 stems/m (～4500 stems/ha) at age 17.5 years. Predicted woody plant species richness also declined with age, with a linear relationship. The number of species in the seed mix affected predicted woody plant species richness. However, large increases in seed mix richness produced relatively small increases in predicted species richness. Regeneration (new stems) was present at high levels (>100 stems/ha); however, 82% of regeneration appeared to originate from delayed germination of sown seed rather than seed from established plants (recruitment). The predicted abundance of new stems (delayed germination and recruitment) declined with age, and for a given age increased with row width. Young stands (11.5 years), seeded with wide rows (4 m), had approximately six times the new stems of similar aged stands seeded with narrow rows (2 m). Our results indicate direct seeded stands simplify with age, becoming less dense and containing fewer species. Maintaining a diversity of shrub species in direct seeded sites may therefore require ongoing disturbances (scarifying, scalping, fire, thinning) of established sites and changes to establishment techniques for new sites. We suggest further research to establish the status of soil seed banks in direct seeded sites, testing different forms of disturbances to trigger regeneration in older revegetation sites and establishing new sites using wide rather than narrow rows and seed mixes enriched with species from genera other than Eucalyptus or Acacia.     

Response of Callitris intratropica R.T. Baker & H.G. Smith to fire protection,Murgenella, Northern Australia

Pattern analysis of stems > 10 cm d.b.h. on 514 one-tenth ha quadrats showed that Callitris intratropica has a clumped distribution throughout the more uniformly distributed Eucalyptus forests at Murgenella in Northwest Arnhem Land. Callitris clumps are typically located on sites with sandy soils. Eucalypts occur within Callitris clumps. Fire suppression over the past 18 years has allowed some C. intratropica to establish on Eucalyptus forest sites with fine textured soils It is argued that distribution of Callitris preceding fire management was controlled by the interactive effects of fire, soils and understorey vegetation. High grass fuel loads (and thus intense fires), and competition are thought to have previously limited establishment of C intratropica on fine textured soils. Early dry season burning by Aboriginals may also have been important in limiting fire intensities and accumulation of fuel in Callitris stands. Dry electrical storms commonly cause fires prior to the summer rains. Such wildfires did not completely kill stands of Callitris saplings or trees at Murgenella. Survivors were found to be significantly bigger and to have thicker bark than the dead stems. Sapling size is related to age and density, thus fire may be an important mechanism in thinning heavily stocked stands. The continuous regeneration of Callitris and Eucalyptus is in marked contrast to the inhibition of Pinus caribaea recruitment following fire protection in the monsoonal neotropics. The lack of seral tree species in northern Australia suggests an equilibrium between forest distribution and the physical environment.     

Natural regeneration patterns of the introduced larch, Larix kaempferi (Pinaceae), on the volcano Mount Koma, northern Japan

The 1929 eruption of Mount Koma (1140 m in altitude) completely deforested the vegetation on the summit area of the volcano. An alien species of larch, Larix kaempferi (=L. leptolepis) was planted on the lower slopes of the volcano between 1953 and 1963, but since then has become abundant on the summit. To determine the regeneration patterns of the larch, we measured stem densities on aerial photographs taken in 1963, 1973 and 1994, and in the field during 1996. Larix kaempferi stems with crown diameters > 2 m were mapped on aerial photographs divided into 1083 100 m×100 m grids. Stem densities on the summit increased from 0.8/ha in 1963, to 14.1/ha in 1994. Willows and birches also established there but did not grow as large as the larch. All these species are wind dispersed, and larch densities were higher in areas closer to the plantations, suggesting that wind intensity and direction determined seed migration. Four environmental factors—slope gradient, direction, elevation, and distance from the plantations in each grid—were correlated with larch stem densities. Multiple regression showed that establishment patterns of L. kaempferi in the early stages were mainly related to distance from seed source (the plantations). Later, geographical disturbances and/or physiological stresses became more important. Density effects on tree invasion have been weak until now. We concluded that revegetation primarily depends on the chance of seed immigration, and that the larches may be an earlier successional species than any other native tree species.     

Non weaving spiders on native woodlands and <Emphasis Type="Italic">Eucalyptus</Emphasis> plantations in Western Mexico: diversity and distribution patterns

Eucalyptus spp. are commonly planted, forming non-native plantations, including the tropics and their wildlife conservation value is relatively unknown. Recent studies have concluded that secondary forests and tree plantations are less diverse than well-developed tropical rain forests. However, introduced Eucalyptus stands harbored similar species richness to surrounding native woodland in temperate woodlands in North America though the identity of the species present may differ. Species composition, as well as dominance curves and differences in community structure add additional insight to understanding faunistic responses to replacement of native woodland by Eucalyptus plantations. Here, we compared species richness, diversity patterns, and the distribution of non-weaving spiders between native woodlands and Eucalyptus plantations in a temperate region of Mexico. We found more Lycosidae species in all plantation stands. Other community attributes were not consistently different between plantations and native woodlands. This is explained by similarities between, and differences within, the understory of the two main vegetation types. Multivariate analyses identified three spider groups and five spider species could be identified as indicators of these groups. A comparison of the number of species of the wandering spiders between the two vegetation types suggests a compensation pattern that is reported here for the first time.     

Site quality effects on post‐fire regeneration of Pinus brutia forest on a Greek island

Abstract. Natural stands of Pinus brutia were compared to burned areas after a great fire in 1985 on the island of Thasos, North Greece, in relation to site quality effects on the success of natural regeneration and understorey vegetation. The number of seedlings in the natural stands of P. brutia was 1124/ha against 3188/ha in the burned area. In both cases natural regeneration was successful. Under the canopy of P. brutia forests the predominant shrubs were Quercus coccifera and vPhillyrea latifolia. A high percentage (up to 40%) of the total area was covered by these species. In the burned areas, 12 yr after the wildfire of 1985, Cistus incanus and C. salvifolius played a significant role in the survival of P. brutia seedlings. These species had covered the burned area long before the pine seed germination.     

Diversity of understory birds in old stands of native and Eucalyptus plantations

We compared the vegetation structure between old (>70 year) stands of planted diversified native forests and stands of Eucalyptus tereticornis embedded in a mosaic of Eucalyptus stands. We then tested for differences in the abundance, species richness, species composition, and ecological traits (forest dependence, sensitivity to forest fragmentation, and diet) of the understory bird assemblages inhabiting both kinds of stands. We expected differences in the structure of the bird assemblages because of the different origins and management strategies (contrary to native stands, Eucalyptus stands were selectively logged in the past). Three stands of each habitat (native and Eucalyptus) were sampled with mist nets during 11 months. Eucalyptus stands had a denser understory, whereas native plantations had a more developed vertical structure and a greater density of native trees. The abundance distribution of bird species was more homogeneous in Eucalyptus than in native stands. Eucalyptus had slightly higher species richness (36 species) than native stands (32 species). The composition of species and the occurrence of the diet, forest dependence, and sensitivity to forest fragmentation categories were similar between habitats. Some bird species (e.g. Turdus leucomelas), however, were more abundant in one habitat over the other. Old stands of Eucalyptus and planted native forest can harbor a diverse bird community similar in structure but not exactly equivalent for individual bird species. Planting native diversified forests and keeping set‐aside stands of the exotic tree should be viewed as complementary rather than alternative strategies for maintaining bird diversity within plantations.     

The influence of dominant shrubs,fire, and time since fire on soil seed banks in mixed chaparral

The composition and density of soil seed banks beneath co-occurring Adenostoma fasciculatum and Ceanothus greggii shrubs from three chaparral stands last burned 9, 35 and 85 years before 1986 were investigated. The overall density of seeds in the soil, as estimated by germinations under greenhouse conditions, increased with time since fire (ca. 8000 to 25000/m²). However, this increase was due entirely to the accumulation of A. fasciculatum seed in the soil (ca. 2000 to 21000/m²). In contrast, the density of C. greggii seed was different in each of the three stands, but was not correlated with time since fire: maximum densities were recorded from the 35 year old stand (ca. 2000/m²).A total of 31 taxa germinated and 17 occurred in sufficient numbers to be analyzed statistically. Germinable seed densities of three herb species were not influenced by soil source (beneath A. fasciculatum or C. greggii), time since fire, or the direct effects of a controlled fire treatment. Germinable seed densities of a further nine species were significantly influenced by the elapsed time since stands last burned. The densities of four decreased and five increased. Four of the species that increased in seed density over the three stands were annuals, suggesting that the chaparral sub-canopy habitat is not as unfavorable for annuals as is often assumed. The fire treatment decreased germinable seed densities of four annual species by 40–70%, but increased the germinable seed densities of the shrubs A. fasciculatum and C. greggii, and the annual Phacelia brachyloba. Our results indicate that seeds of A. fasciculatum will increase in the soil bank for at least 85 years after fire in chaparral where it is dominant. In contrast, seed reserves of C. greggii appear to be influenced primarily by site-specific patterns of seed production and by the intensity of post-dispersal seed predation.     

The potential of small Eucalyptus plantations in farmscapes to foster native woody plant diversity: local and landscape constraints

The major focus of ecological restorations has been on understanding local factors. However, landscape factors such as dispersal limitation of individuals or propagules across the surrounding matrix can also constrain the restoration progress. We investigated to what extent native woody species colonize and thrive in plantations, focusing on both the role of local factors such as grazing and canopy cover as well as on landscape factors. We recorded all native tree and shrub species in 60 small Eucalyptus plantations embedded in an open agricultural landscape at 0.1–12 km from a remnant continuous forest in central Ethiopia. We found a total of 1,571 individuals of native woody plants belonging to 55 species. Number of such species in a plantation increased significantly with the height of the grass sword indicating their sensitivity to grazing. Moreover, the number of woody species in the patches decreased significantly with distance to the forest. Our results illustrate the need for regulating the grazing pressure for a successful regeneration of native species in Eucalyptus plantations. In addition, sowing or planting native trees will be necessary in most plantations, as only few remnant natural forests that could act as seed sources occur across the Ethiopian highlands. Another main obstacle might be the prohibition of selling timber of native trees, which indirectly discourage farmers from letting native trees regenerate. Thus, the increasing cover of Eucalyptus seen across the country will not automatically foster a recovery of native woody plant biodiversity, even if managed to optimize local environmental conditions.     

Allelopathic Effects of Tree Species on Some Soil Microbial Populations and Herbaceous Plants

The allelopathic potential of four tree species on soil microbial populations and some herbaceous plants (two understory species and one general biotest species) was investigated. Effects of three nonindigenous tree species, Eucalyptus globulus Labill, Pinus radiata D.Don and Acacia melanoxylon R.Br., on microorganisms participating in the cycle of nitrogen were evaluated, comparing them with those produced by the autochthonous Quercus robur L. Influence of the trees on Lactuca sativa L., Dactylis glomerata L. and Trifolium repens L. was also checked in bioassays. Cell numbers of Nitrosomonas sp. were negatively affected by Acacia and Eucalyptus stands, mainly during spring, when flowers are especially abundant on the ground. Proteolytic microorganisms were also negatively affected by Eucalyptus and Pinus stands, whilst Quercus stand did not show any toxicity. Soil bioassays showed clear inhibitory effects on germination and growth of understory plants, particularly soils from Eucalyptus and Acacia stands. The greatest effects had the soil from Acacia stand, which was phytotoxic during the whole period of germination and growth of understory plants. Allelopathic phenomena could be, at least partially, responsible of the low species diversity in the understory of the nonindigenous tree stands.     

Dispersal and recruitment dynamics in the fleshy‐fruited Persoonia lanceolata (Proteaceae)

Question: What is the role of dispersal, persistent soil seed banks and seedling recruitment in population persistence of fleshy‐fruited obligate seeding plant species in fire‐prone habitats? Location: Southeastern Australia. Methods: We used a long‐term study of a shrubby, fleshy‐fruited Persoonia species (Proteaceae) to examine (1) seed removal from beneath the canopy of adult plants; (2) seedling recruitment after fire; (3) the magnitude and location of the residual soil seed bank; and (4) the implications for fire management of obligate seeding species. We used demographic sampling techniques combined with Generalised Linear Modelling and regression to quantify population changes over time. Results: Most of the mature fruits (90%) on the ground below the canopy of plants were removed by Wallabia bicolor (Swamp wallaby) with 88% of seeds extracted from W. bicolor scats viable and dormant. Wallabies play an important role in moving seeds away from parent plants. Their role in occasional long distance dispersal events remains unknown. We detected almost no seed predation in situ under canopies (< 1%). Seedling recruitment was cued to fire, with post‐fire seedling densities 6‐7 times pre‐fire adult densities. After fire, a residual soil seed bank was present, as many seeds (77‐100%) remained dormant and viable at a soil depth where successful future seedling emergence is possible (0‐5 cm). Seedling survival was high (> 80%) with most mortality within 2 years of emergence. Plant growth averaged 17 cm per year. The primary juvenile period of plants was 7–8 years, within the period of likely return fire intervals in the study area. We predicted that the study population increased some five‐fold after the wildfire at the site. Conclusions: Residual soil seed banks are important, especially in species with long primary juvenile periods, to buffer the populations against the impact of a second fire occurring before the seed bank is replenished.     

Effects of European colonization on indigenous ecosystems: post-settlement changes in tree stand structures in Eucalyptus–Callitris woodlands in central New South Wales, Australia

Aim There has been considerable debate about pre‐settlement stand structures in temperate woodlands in south‐eastern Australia. Traditional histories assumed massive tree losses across the region, whereas a number of recent histories propose that woodlands were originally open and trees regenerated densely after settlement. To reconcile these conflicting models, we gathered quantitative data on pre‐settlement stand structures in Eucalyptus–Callitris woodlands in central New South Wales Australia, including: (1) tree density, composition, basal area and canopy cover at the time of European settlement; and (2) post‐settlement changes in these attributes. Location Woodlands dominated by Eucalyptus species and Callitris glaucophylla, which originally occupied approximately 100,000 km² in central New South Wales, Australia. Methods We recorded all evidence of pre‐settlement trees, including stumps, stags and veteran trees, from 39 relatively undisturbed 1‐ha stands within 16 State Forests evenly distributed across the region. Current trees were recorded in a nested 900 m² quadrat at each site. Allometric relationships were used to estimate girth over bark at breast height, tree basal area, and crown diameter from the girth of cut stumps. A post‐settlement disturbance index was developed to assess correlations between post‐settlement disturbance and attributes of pre‐settlement stands. Results The densities of all large trees (> 60 cm girth over bark at breast height) were significantly greater in current stands than at the time of European settlement (198 vs. 39 trees ha^?1). Pre‐settlement and current stands did not differ in basal area. However, the proportional representation of Eucalyptus and Callitris changed completely. At the time of settlement, stands were dominated by Eucalyptus (78% of basal area), whereas current stands are dominated by Callitris (74%). On average, Eucalyptus afforded 83% of crown cover at the time of settlement. Moreover, the estimated density, basal area and crown cover of Eucalyptus at the time of settlement were significantly negatively correlated with post‐settlement disturbance, which suggests that these results underestimate pre‐settlement Eucalyptus representation in the most disturbed stands. Main conclusions These results incorporate elements of traditional and recent vegetation histories. Since European settlement, State Forests have been transformed from Eucalyptus to Callitris dominance as a result of the widespread clearance of pre‐settlement Eucalyptus and dense post‐settlement recruitment of Callitris. Tree densities did increase greatly after European settlement, but most stands were much denser at the time of settlement than recent histories suggest. The original degree of dominance by Eucalyptus was unexpected, and has been consistently underestimated in the past. This study has greatly refined our understanding of post‐settlement changes in woodland stand structures, and will strengthen the foundation for management policies that incorporate historical benchmarks of landscape vegetation changes.     

Organisation of the soil seed bank in mixed chaparral

Seed densities, estimated from germinations under glasshouse conditions, were determined for fire-treated and control soils sampled from north and south facing stands of mixedAdenostoma fasciculatum-Ceanothus greggii chaparral that last burned 10, 17, 36, 62 and 86 years before 1987. A total of 53 species was recorded. Total germinable seed density was not influenced by aspect nor by an experimental fire treatment, however, densities varied significantly, but not predictably, over the time since fire sequence. Species diversity was significantly lower in the 86 year old stand, but neither diversity or richness was influenced by the fire treatment. Soils from north facing slopes supported a higher species diversity overall, and seed densities of six species were significantly higher on north than on south slopes.We identify two functional groups in the seed bank on the basis of the direct effects of fire on seed densities: a fire-dependent group, comprising 6 species and about 21% of the total seed bank, in which germinable densities increased after the fire treatment, and a fire-independent group, made up of 11 species and almost two-thirds of the seed bank. The fire-independent group is further separated into two: a shrub-centred sub-group (Crassula erecta, Filago californica, Pterostegia drymarioides, Streptanthus heterophyllus andCryptantha intermedia) maintained larger seed banks in older stands and was associated with shrub cover; and an opportunistic sub-group (Camissonia hirtella, Gnaphalium chilense, Mimulus pilosus andSenecio vulgaris) tended to have larger seed banks in younger stands and was associated with canopy gaps.Overall, the soil seed assemblage from this mixed chaparral does not appear to be tightly coupled to fire-generated opportunities for recruitment. The reason for this is that the fire-dependent group represents only 6 species. In addition, among the fire-independent species, the shrub-centred group of 5 annuals made up almost 40% of total soil seed density. This group shows ecological and taxonomic affinities with desert annuals.     

Introduced populations of <Emphasis Type="Italic">Genista monspessulana</Emphasis> (French broom) are more dense and produce a greater seed rain in California,USA, than native populations in the Mediterranean Basin of Europe

Some invasive plants perform better in their area of introduction than in their native region, and this is often attributed either to phenotypic responses and/or to adaptive evolution following exposure to new environmental conditions. Genista monspessulana (French broom) is native to Europe, but highly invasive and abundant along the Pacific Coast of the USA. In this study, the population density and age structure, plant growth and reproductive traits, and seed bank characteristics of 13 native (Mediterranean Basin) and 15 introduced (California, USA) field populations of G. monspessulana were compared. Mean population density, plant height and stem diameter were greater in introduced populations, with the latter two traits explained by a greater mean plant age. Age structure also showed a greater percentage of seedling plants in introduced populations. Fecundity was higher in introduced populations when measured in terms of mature seeds per pod, but lower when comparing seed production per plant (number of pods and mature seeds). Thus, seed rain and seed bank size was considerably higher in introduced populations. Results from this study indicate that G. monspessulana performs better in its introduced region. We hypothesize that release from natural enemies and competitors together with more favorable environmental conditions in the introduced region may explain the invasion success of G. monspessulana. As a result, an integrated management approach using introduced seed predators to suppress seed production and selected management practices to reduce seed banks may be needed for effective long-term control in California.     

Seed bank dynamics of two obligate seeders, <Emphasis Type="Italic">Cistus monspeliensis</Emphasis> and <Emphasis Type="Italic">Rosmarinus officinalis</Emphasis>, in relation to time since fire

Many species in Mediterranean-type ecosystems regenerate after fire by seed germination from soil seed banks. Seed bank dynamics of two of those obligate seeders, Cistus monspeliensis and Rosmarinus officinalis, were investigated in relation to stand age since fire in southwestern Portugal. Soil seed density, annual seed input, annual seed losses through germination and seed persistence were compared between species at stands differing in age since fire (5, 10 and 35 years). Soil seed density and seed input increased over the first decade after fire and were lowest at 35-year-old stands for C. monspeliensis. In R. officinalis, few seeds were produced and found in the soil at early stages, and maximum seed input and soil seed density were attained at 35-year-old stands. Soil seed density was mostly driven by seed production in both species, which is largely dependent on plant traits and population dynamics related to fire. Overall, stand age since fire had a negligible effect on seed germination, seed persistence and viability. Ten to 39% of buried seeds were not recovered after 1 year, and viability of seeds recovered was 97–100% for C. monspeliensis and only 0–3% for R. officinalis. Variation in plant traits within the seeder syndrome was evidenced by this study. R. officinalis evidenced lower seed persistence, lower proportion of viable seed produced and lower density of viable soil seed than C. monspeliensis at any stage after fire. R. officinalis is expected to depend largely on previous year seed production for population replacement after fire.     

Icacina oliviformis (Icacinaceae): A close look at an underexploited food plant. III. Ecology and production

Data are presented on the ecology and production ofIcacina oliviformis (Poiret) Raynal in natural stands in northern Central African Republic. The species grows on a variety of soils and in numerous plant communities. It is fire adapted, and thrives when burned in the early dry season. The plant is primarily bee pollinated and has a number of pests that infest the stems and young fruits. Plant densities are high, with a maximum of ca. 5345 plants/ha in the study sites. Standing tuberous root biomass can be considerable, with a maximum of ca. 19904 kg/ha for the study sites. Seed yield in the sample sites was relatively low with a maximum of ca. 214 kg/ha. With proper management, yields of both seed and tuberous roots could be substantially increased.     

Influence of overstory composition on understory colonization by native species in plantations on a degraded tropical site

Abstract. Patterns of understory colonization by native and naturalized trees and shrubs were evaluated in 4.5-year-old plantations of three exotic tree species, Casuarina equisetifolia, Eucalyptus robusta, and Leucaena leucocephala, on a degraded coastal grassland site with reference to overstory composition and understory environmental conditions. 19 secondary forest species were established in the plantation understories (with a total area of 0.52 ha), while no natural regeneration occurred in unplanted, though protected, control areas. The majority of these species (90 %) and the total seedling population (97 %) were zoochorous, indicating the importance of frugivorous bats and particularly birds as facilitators of secondary forest species colonization. Understory species richness and seedling densities were affected significantly by overstory composition, the most abundant regeneration occurring beneath Leucaena and least under Casuarina. Understory colonization rates within mixed-species stands were intermediate between those of single-species stands of the trees comprising their overstories. Significant negative correlations were found between understory species richness and seedling density, and forest floor depth and dry mass, especially for small-seeded ornithochorous species. Higher colonization rates near the peripheries of plantation plots relative to plot interiors were due in part to roosting site preferences by frugivores, particularly bats. The study results indicate that overstory species selection can exert a significant influence on subsequent patterns of colonization by secondary forest species and is an important consideration in the design of plantations for ‘catalyzing’ succession on deforested, degraded sites.     

Canopy cover and tree regeneration in old-growth cove forests of the Appalachian Mountains

Relationships between canopy cover and tree regeneration were determined for various species in cove forests of the Great Smoky Mountains. Old-growth stands were sampled with six plots covering a total area of 4.8 ha. Each plot was subdivided into contiguous 10×10 m quadrats. Canopy cover overlying each of the 480 quadrats was characterized with three different indices based on visual estimates of cover. Influences of: (1) overlying cover, (2) proximate openings, and (3) total area of proximate openings on quadrat regeneration densities were determined. Most species reproducing by seed and some species reproducing by vegetative means had higher densities in quadrats with openings, but only the intolerants were highly dependent on gaps. Tsuga canadensis, a very shade-tolerant species, was one of the few species with abundant regeneration beneath dense canopy cover. In general, understory areas near gaps had somewhat higher regeneration densities than other areas with overlying cover. Several shade-tolerant species showed a positive regeneration density response to canopy openings and an ability to regenerate in gaps 0.01–0.03 ha in area. These openings were too small for intolerant species. Many species exhibited a positive response to total size of the proximate opening(s). A sharp increase in regeneration density with area of the opening(s) was evident at approximately 0.04 ha for the shade-intolerant species.     

Influence of rocky landscape features and fire regime on vegetation dynamics in Appalachian Quercus forests

Question : How do interactions between rocky landscape features and fire regime influence vegetation dynamics? Location : Continental Eastern USA. Methods : We measured vegetation, disturbance and site characteristics in 40 pairs of rocky and non‐rocky plots: 20 in recently burned stands, and 20 in stands with no evidence of recent fire (‘unburned’ stands). Two‐way analysis of variance (ANOVA) was used to assess the main and interaction effects of fire and rock cover on plant community composition. Results : In burned stands, rock cover had a strong influence on vegetation. Non‐rocky ‘matrix’ forests were dominated by Quercus, and had abundant ground cover and advance regeneration of early and mid‐successional tree species. Burned rocky patches supported greater density of fire‐sensitive species such as Acer rubrum, Sassafras albidum and Nyssa sylvatica and had little advance regeneration or ground cover. Quercus had fewer fire scars and catfaces (open, basal wounds) on rocky patches, suggesting that rocky features mitigate fire severity. In unburned stands, differences between rocky and non‐rocky patches were less distinct, with both patch types having sparse ground cover, little tree regeneration, and high understorey densities of relatively shade tolerant A. rubrum, N. sylvatica and Betula lenta. Conclusion : Under a sustained fire regime, heterogeneity in rock cover created a mosaic where fire‐adapted species such as Quercus dominate the landscape, but where fire‐sensitive species persisted in isolated pockets of lower fire severity. Without fire, species and landscape richness may decline as early‐mid successional species are replaced by more shade tolerant competitors.