Demography of the long‐lived conifer Agathis ovata in maquis and rainforest,New Caledonia

Abstract. The endemic New Caledonian conifer Agathis ovata occurs as an emergent tree in fire‐prone shrublands (maquis), and fire‐sensitive rainforest. Growth, survivorship and recruitment over 5 yr were compared for populations from forest and maquis on ultramafic substrates in New Caledonia to investigate whether demographic behaviour varied in response to the strongly contrasting forest and shrubland environments. Growth of seedlings and of small (30–100 cm height) and large (100 cm height; 5 cm DBH) saplings was slow, but varied significantly among stages, site types and years. The greatest difference in growth rates was among stages, seedlings growing 0.34 cm.yr^?1, small saplings 1.06 cm.yr^?1 and large saplings 2.13 cm.yr^?1. Tree DBH increased by only 0.05 cm.yr^?1 and, based on these rates, individuals with DBH of 30 cm are estimated to be more than 700 yr old. Few trees (3.5%) produced cones in any year and seedling recruitment was low, but some recruitment was recorded each year in both maquis and forest. Rates of recruitment per parent were highest in forest (1.28.yr^?1, cf 0.78.yr^?1), but the higher density of trees in maquis meant that overall recruitment was greater there (92 ha^?1.yr^?1, cf 56 ha^?1.yr^?1). Seedling mortality ranged from 0.9 to 2.9% among years with no significant difference between maquis and forest. No sapling mortality was recorded, but annual tree mortality ranged from 0 to 1.4%. Evidence from a recently burned site indicated that while trees may survive fire, seedlings and saplings do not. Post‐fire seedling recruitment per ha from surviving trees was four times lower than in unburned sites, but growth rates were four times higher. Similar demographic attributes, including high survivorship, low growth rate and low rates of recruitment over a long reproductive life, characterize Agathis ovata populations in both maquis and rainforest in New Caledonia and are indicative of a broad tolerance of light environments that is unusual among tree species. These demographic attributes help to explain the long‐term persistence of the species in these strongly contrasting habitats.     

Long‐term forest structure and regeneration after wildfire in Russian Karelia

We examined forest structure and regeneration in a 350‐ha forest dominated by Pinus sylvestris 31 yr after a wildfire in the Vienansalo wilderness, Russian Karelia. In most parts of the area, the 1969 fire was not stand replacing but had left larger trees alive so that the area generally remained forest covered. In some localities, however, all trees apparently died and distinct gaps were formed, suggesting that the fire severity varied considerably, contributing to increased variation in stand structure. Living and dead wood volumes were similar, 112 and 96 m³.ha^‐1, respectively. The tree species proportions of dead vs living wood indicated that prior to fire disturbance Picea was more common in the area. Regeneration was abundant (saplings, ca. 14 000 ind.ha^‐1, height 20 ‐200 cm) and tree seedling recruitment had occurred over a long period of time. Regeneration density was highest on the mesic Vaccinium‐Myrtillus forest site type, decreasing towards nutrient‐poor site types. The most common regeneration microsites were level ground (56% of saplings), immediate surroundings of decayed wood (23%) and depressions (11%). The high proportion of saplings on level ground suggests that after the fire regeneration conditions have been favourable across the whole forest floor. Nevertheless, the areas in the vicinity of decayed wood have been particularly important microsites for seedling establishment. The results provide an example of the effects of wildfire on forest structure in a natural Pinus sylvestris dominated forest, demonstrating the non stand replacing character of fire, high variability in stand structure and the abundance of post‐fire regeneration.     

Patch to landscape patterns in post fire recruitment of a serotinous conifer

Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m² but older patches had thinned to one tree every 3–15 m². Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.     

Ectomycorrhiza succession patterns in Pinus sylvestris forests after stand-replacing fire in the Central Alps

Fires shape fundamental properties of many forest ecosystems and climate change will increase their relevance in regions where fires occur infrequently today. In ecosystems that are not adapted to fire, post-fire tree recruitment is often sparse, a fact that might be attributed to a transient lack of mycorrhizae. Ectomycorrhizal (EcM) fungi play an important role for recruitment by enhancing nutrient and water uptake of their hosts. The questions arise whether and for how long the EcM community is transformed by fire. We investigated the resistance and resilience of EcM fungal communities on a chronosequence of 12 Pinus sylvestris stands in Valais (Switzerland) and Val d’Aosta (Italy) affected by fire between 1990 and 2006. Soil samples from burnt and non-burnt forests were analyzed with respect to EcM fungi by means of a bioassay. The number of EcM species was significantly lower in samples from recently (2–5 years) burnt sites than non-burnt forest, and increased with time since fire reaching levels of adjacent forests after 15–18 years. Community composition changed after fire but did not converge to that of non-burnt sites over the 18 year period. Only Rhizopogon roseolus and Cenococcum geophilum were abundant in both burnt sites and adjacent forest. Our data indicate fire resistance of some EcM fungal species as well as rapid resilience in terms of species number, but not in species composition. As long as the function of different EcM species for seedling establishment is unknown, the consequences of long-term shifts in EcM community composition for tree recruitment remain unclear.     

Responses to Fire in Selected Tropical Dry Forest Trees1

Fire is a frequent disturbance in the tropical dry forests of Central America, yet very little is known about how native species respond to such events. We conducted an experimental burn in a tropical dry forest of western Nicaragua to evaluate plant responses to fire with respect to survivorship and recruitment. Measurements of woody vegetation of all size classes were carried out prior to the prescribed burn and three successive years post fire. We selected the 15 most abundant species <10 cm DBH to assess percent survivorship and sprouting responses post fire. Changes in seedling densities for these 15 most abundant species and the 15 least abundant species were analyzed using a repeated measure ANOVA. We also assessed changes in seedling densities for three species of international conservation concern. We found three major fire‐coping strategies among common dry forests plants: resisters (low fire‐induced mortality), resprouters (vigorous sprouting), and recruiters (increased seeding post‐fire). While survivorship was generally high relative to tropical moist forest species, those species with lower survivorship used either seeding or sprouting as an alternative strategy for persisting in the forest community. Seed dispersal mechanisms, particularly wind dispersal, appear to be an important factor in recruitment success post‐fire. Burn treatment led to a significant increase in the density of seedlings for two species of conservation concern: Guaiacum sanctum and Swietenia humilis. Results of this study suggest that common dry forest species in western Nicaragua are fire tolerant. Further study of individual species and their fire responses is merited.     

Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

Limited Edge Effects Along a Burned‐Unburned Bornean Forest Boundary Seven Years after Disturbance

Large parts of the everwet tropics have been burned, leaving many unburned–burned forest edges. Here we studied a Bornean forest edge to determine: (1) how unburned and burned forest differ in vegetation structure, diversity, composition and plant functional traits 7 yr after fire, and (2) if these variables showed significant edge effects. Environmental and inventory data from 120 plots (0.01 ha each), covering both sides of a ～1.3 km forest boundary were sampled. Differences in vegetation structure, diversity, composition and plant functional traits were analyzed in relation to disturbance type (Mann–Whitney tests) and edge distance (partial correlation analysis that controlled for confounding effects of elevation, slope and fire intensity). Seven years after fire, burned forest differed significantly from unburned forest in most measured variables while few significant edge effects were detected, i.e., there existed a sharp delimitation between the two forest types. The regeneration of the burned forest depended almost entirely on in situ recruitment with little input of late successional species from the neighboring old growth forest. On the other hand, old growth forest showed few signs of edge degradation. A possible explanation for these results might be related to the absence of a mast fruiting event during these first 7 yr of forest recovery, resulting in low levels of late successional species seed input into the burned forest, combined with the quick development of a closed canopy in the burned forest by early successional species that shielded the unburned forest from adverse edge effects.     

Effects of fire and drought in a tropical eucalypt savanna colonized by rain forest

Aim This study documents the effects of multiple fires and drought on the woody structure of a north Australian savanna never grazed by domestic stock. Location The study was conducted in a 500 ha pocket of Eucalyptus‐dominated savanna surrounded by a late Quaternary lava flow. The flow is known as the Great Basalt Wall, located c. 50 km northeast of Charters Towers in semi‐arid north‐eastern Australia. This region was exposed to the largest 5‐year rainfall deficit on record between 1992 and 1996. Methods All individual woody plants were tagged within a 1.56 ha plot. Species were segregated into their habitat affinities (rain forest, ecotone, savanna) and regeneration strategy (resprouter, seeder). The survivorship of plants within these categories was analysed in relation to fire intensity from the first fire, and to each of four fires lit between 1996 and 2001. Results Before the first fire, the plot contained thirty‐one tree species including twenty‐one typical of the surrounding dry rain forest. These rain forest species were represented by small individuals and constituted <1% of the total basal area of woody plants. The basal area of savanna trees was 7.5 m² ha^?1 at the commencement of monitoring, although 31% had recently died and others had major crown damage. Further death of the drought debilitated savanna trees was substantial during the first year of monitoring and the basal area of live savanna trees declined to 1.1 m² ha^?1 after 5 years. Most species from both rain forest and savanna were classified as resprouters and are capable of regenerating from underground organs after fire. Species without this ability (rain forest seeders and ecotone seeders) were mostly eliminated after the first two consecutive fires. Among resprouters, survivorship declined as fire intensity increased and this was more pronounced for rain forest than for savanna species. Repeated burning produced a cumulative effect of decreasing survivorship for rain forest resprouters relative to savanna resprouters. Main conclusions The study provides evidence that savanna and rain forest trees differ in fire susceptibility and that recurrent fire can explain the restricted distribution of rain forest in the seasonally arid Australian tropics. The time of death of the savanna trees is consistent with the regional pattern after severe drought, and highlights the importance of medium term climate cycles for the population dynamics of savanna tree species and structure of Australian savannas.     

Stand structure of the emergent conifer Agathis ovata in forest and maquis, Province Sud, New Caledonia

Abstract. The size structure of the endemic New Caledonian conifer Agathis ovata is reported for sample stands in forest and maquis from three areas on ultramafic substrates in the south of the main island (Grand Terre). In closed forest Agathis ovata is typically represented by a low density of emergent adult trees with only limited evidence of seedling recruitment. In maquis, Agathis ovata is represented by individuals of all sizes, with seedlings and saplings abundant in most sample stands. Preliminary evidence from tree-ring studies indicates that rings may be annual. Estimated diameter growth rate is about 2 mm y^?1 for trees ≥ 10 cm d.b.h., and ring counts suggest tree ages of up to 400 years in maquis and 500 years in forest. Agathis ovata, and three other members of the Araucariaceae found in New Caledonia (Araucaria laubenfelsii, A. montana and A. rulei), are the only tree species which regularly occur scattered in maquis in this way, creating an unusual structural assemblage. No angiosperm tree species show this behaviour. The circumstances under which the Agathis ovata– maquis stands arise and are maintained are the subject of further investigation. Preliminary evidence for tree ages indicates that these stands predate European arrival in New Caledonia and so are not the result of recent increases in the frequency and intensity of human disturbances. The presence of fire scars on many individuals, and location of most stands on slopes and spurs with outcropping laterite (cuirasse), suggests that this assemblage may owe its existence to the interplay of fire regime, topography and rockiness, and a resistance to fire in Agathis ovata which increases with plant size and age.     

Mesophication in temperate Europe: A dendrochronological reconstruction of tree succession and fires in a mixed deciduous stand in Białowieża Forest

1. The shift from shade‐intolerant species to shade‐tolerant mesophytic species in deciduous and mixed forests of the temperate zone is well described in studies from North America. This process has been termed mesophication and it has been linked to changes in fire regime. Fire suppression results in the cessation of establishment of heliophytic, fire‐dependent tree species such as oak (Quercus) and pine (Pinus). Due to the scarcity of old‐growth forests in Europe, data on long‐term compositional changes in mixed forests are very limited, as is the number of studies exploring whether fire played a role in shaping the dynamics.
2. The aim of this study was to reconstruct tree succession in a 43‐ha natural mixed deciduous forest stand in Bia?owie?a Forest (BF), Poland using dendrochronological methods. In addition, the presence of aboveground fire legacies (charred and fire‐scarred deadwood) enabled the fire history reconstruction.
3. Dendrochronological data revealed tree establishment (Quercus) back to the end of the 1500s and fires back to 1659. Under a regime of frequent fires until the end of the 18th century, only oak and pine regenerated, sporadically. A shift in the fire regime in the first half of the 19th century triggered oak and pine cohort regeneration, then gradually spruce (Picea) encroached. Under an increasingly dense canopy and less flammable conditions, regeneration of shade‐tolerant Carpinus, Tilia, and Acer began simultaneously with the cessation of oak and pine recruitment.
4. Synthesis. The study reports the first evidence of mesophication in temperate Europe and proves that fire was involved in shaping the long‐term dynamics of mixed deciduous forest ecosystems. Our data suggest that fire exclusion promoted a gradual recruitment of fire‐sensitive, shade‐tolerant species that inhibited the regeneration of oak and pine in BF.

     

The relationship between fire history and an exotic fungal disease in a deciduous forest

Exotic diseases have fundamentally altered the structure and function of forest ecosystems. Controlling exotic diseases across large expanses of forest has proven difficult, but fire may reduce the levels of diseases that are sensitive to environmental conditions. We examined Cornus florida populations in burned and unburned Quercus–Carya stands to determine if burning prior to anthracnose infection has reduced the impacts of an exotic fungal disease, dogwood anthracnose, caused by Discula destructiva. We hypothesized that fire has altered stand structure and created open conditions less conducive to dogwood anthracnose. We compared C. florida density, C. florida health, and species composition and density among four sampling categories: unburned stands, and stands that had burned once, twice, and 3 times over a 20-year period (late 1960s to late 1980s). Double burn stands contained the greatest density of C. florida stems (770 stems ha⁻¹) followed by triple burn stands (233 stems ha⁻¹), single burn stands (225 stems ha⁻¹) and unburned stands (70 stems ha⁻¹; P < 0.01). We observed less crown dieback in small C. florida trees (<5 cm diameter at breast height) in burned stands than in unburned stands (P < 0.05). Indicator species analysis showed that burning favored species historically associated with Quercus–Carya forests and excluded species associated with secondary succession following nearly a century of fire suppression. Our results suggest that fire may mitigate the decline of C. florida populations under attack by an exotic pathogen by altering forest structure and composition. Further, our results suggest that the burns we sampled have had an overall restorative effect on forest communities and were within the fire return interval of the historic fire regime. Consequently, prescribed fire may offer a management tool to reduce the impacts of fungal disease in forest ecosystems that developed under historic burning regimes.     

Population age structure of Pinus banksiana at the southern edge of the Canadian boreal forest

Abstract. To assess the effects of site type, forest initiation periods and fire regimes on the dynamics of Pinus banksiana (Jack pine), the age structure of 69 populations of the species was analyzed. Two landscapes with different fire regimes were selected in the southern part of the Canadian boreal forest in Québec: the ‘mainland landscape’ is characterized by a fire regime of large lethal fires, the ‘island landscape’ is affected by a complex fire regime including lethal and non-lethal fires. Age structure was compared between forest initiation periods and site types (mesic mainland, xeric mainland and xeric island) using the Shannon regularity index. An even-aged population structure was found within the first 100 yr following a lethal fire, while after that period the population structure becomes more uneven-aged. Under mesic conditions, populations tend to have an even-aged structure, under xeric conditions an uneven-aged structure. Natural openings present in xeric sites allow for recruitment in the absence of fire. This permits the self-maintenance of Pinus banksiana. Xeric island populations show more uneven-aged structures than xeric mainland populations. The occurrence of non-lethal fires on the islands creates uneven-aged structures. Further, the results suggest that the selection pressure of the island fire regime, favouring non-serotinous and mixed P. banksiana individuals, is one of the factors responsible for a higher recruitment in the absence of fire on islands than on the mainland.     

Contrasting Population Dynamics of the Endemic New Caledonian Conifer Araucaria laubenfelsii in Maquis and Rain Forest

This study compares demographic parameters and population dynamics for high disturbance (maquis) and low disturbance (rain forest) environments of the montane conifer, Araucaria laubenfelsii, in New Caledonia. The establishment, growth, survival and reproduction of ca 2500 individuals were followed in permanent plots over 10 yr. Growth and survival rates for A. laubenfelsii show that it is a long-lived, slow growing tree, with evidence of suppression in the sapling size classes in mature rain forest. Growth rates for all size classes are generally faster in maquis than rain forest. Transition matrix analyses estimated positive rates of population increase (λ values>1), with populations expanding in maquis, and stable in mature forest. Araucaria laubenfelsii is able to regenerate continuously in maquis and early successional rain forest, but recruitment is limited in older stands. Life table response experiment analyses showed that reproduction, and transitions from sapling to mature tree stage, contributed positively to λ in maquis, but negatively in forest. Araucaria laubenfelsii on Mont Do can be considered a long-lived pioneer, with early maquis colonizers helping to drive succession from maquis to forest. While opportunities for recruitment decline with time as rain forest sites develop a closed canopy, occasional gap phase recruitment, combined with disturbance by cyclones, landslides and fire, provide opportunities to ensure species persistence. Understanding contrasting population dynamics of A. laubenfelsii in maquis and rain forest will better facilitate conservation management of this species, particularly given current high rates of land conversion and degradation in New Caledonia. Abstract in French is available at http://www.blackwell-synergy.com/loi/btp     

Predicting the Recovery of <Emphasis Type="Italic">Pinus halepensis</Emphasis> and <Emphasis Type="Italic">Quercus ilex</Emphasis> Forests after a Large Wildfire in Northeastern Spain

Quercus ilex and Pinus halepensis are two of the most common tree species of the western Mediterranean basin. Both species regenerate reliably after fire: P. halepensis colonizes recently disturbed areas by effective seedling recruitment, while Q. ilex resprouts vigorously after disturbances. For this reason, the natural regeneration of these species after fire should ensure the re-establishment of a forest similar to that which existed before the fire. This study analyzes with a simple simulation model whether or not the relative abundance of monospecific and mixed forests of these species in the landscape is altered by fire. We also analyze the topographic factors and the forest structure before the fire that determine the changes in forest composition after fire. This study has been carried out in a large fire that occurred in NE Spain. Overall, 33% of plots changed to another community type, but this probability of change varied considerably among community types before the fire. Monospecific forests of P. halepensis or Q. ilex had a high probability of remaining in their original composition after the fire, whereas the resilience of mixed forests of these two species was quite low. Mixed forests changed for the most part to monospecific P. halepensis or Q. ilex forests. Analysis of several factors determining these changes indicated that only elevation as a significant topographical variable. The effect of fire was to increase the altitudinal differentiation between the two species. P. halepensis forests that changed to mixed or Q. ilex forests were those of highest elevation, while the mixed and Q. ilex plots that changed to P. halepensis forests were those located at the lowest elevations. Concerning structural variables before fire, density of Q. ilex trees before the fire showed a much greater effect than P. halepensis density in determining the post-fire community. Finally, burn severity also influenced the changes observed. For both P. halepensis and Q. ilex forests, plots that changed to another forest type were mainly those that burned more severely. In the case of mixed forests, even low fire severities involved high probabilities of change to monospecific forests.     

Long‐term post‐fire changes in the northeastern boreal forest of Quebec

Abstract. Natural dynamics in the boreal forest is influenced by disturbances. Fire recurrence affects community development and landscape diversity. Forest development was studied in the northeastern boreal forest of Quebec. The objective was to describe succession following fire and to assess the factors related to the changes in forest composition and structure. The study area is located in northeastern Quebec, 50 km north of Baie‐Comeau. We used the forest inventory data gathered by the Ministère des Ressources naturelles du Québec (MRNQ). In circular plots of 400 m², the diameter at breast height (DBH) of all stems of tree species greater than 10 cm was recorded and in 40 m² subplots, stems smaller than 10 cm were measured. A total of 380 plots were sampled in an area of 6000 km². The fire history reconstruction was done based on historical maps, old aerial photographs and field sampling. A time‐since‐fire class, a deposit type, slope, slope aspect and altitude were attributed to each plot. Each plot was also described according to species richness and size structure characteristics. Traces of recent disturbance were also recorded in each plot. Changes in forest composition were described using ordination analyses (NMDS and CCA) and correlated with the explanatory variables. Two successional pathways were observed in the area and characterized by the early dominance of intolerant hardwood species or Picea mariana. With time elapsed since the last fire, composition converged towards either Picea mariana, Abies balsamea or a mixture of both species and the size structure of the coniferous dominated stands got more irregular. The environmental conditions varied between stands and explained part of the variability in composition. Their effect tended to decrease with increasing time elapsed since fire, as canopy composition was getting more similar. Gaps may be important to control forest dynamics in old successional communities.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Impact of the invasive alien grass Melinis minutiflora at the savanna-forest ecotone in the Brazilian Cerrado

Exotic grasses are a serious threat to biodiversity in the cerrado savannas of central Brazil. Of particular concern is the possible role they may have in impeding tree regeneration at gallery (riverine) forest edges and increasing fire intensity, thereby driving gallery forest retreat. Here we quantify the effect of roads and distance from gallery forests on the abundance of the African grass Melinis minutiflora Beauv. and test for an effect of this species on woody plant regeneration and leaf area index. Melinis was present at approximately 70% of the sites near gallery forest edges, with its frequency declining sharply at greater distances from the edge. Melinis frequency was 2.8 times greater where roads were present nearby. Leaf area index (LAI) of the ground layer was 38% higher where Melinis was present than where it was absent. LAI was strongly correlated to fine fuel mass (r² = 0.80), indicating higher fuel loads where Melinis was present. The abundance of tree and shrub species in the ground layer was negatively related to LAI and to the presence of Melinis. The greater fuel accumulation and reduced tree regeneration caused by Melinis may cause a net reduction in forest area by increasing fire intensity at the gallery forest edge and slowing the rate of forest expansion.     

Effect of fire on recruitment of two dominant perennial grasses with different palatability from semi-arid grasslands of NW Patagonia (Argentina)

Fire is a non-selective disturbance that impacts equally plant species that could be selected differentially by livestock. Post-fire recruitment dynamics is an important ecological process that has been barely studied in Patagonian grass species. This work analyzes the effect of fire on seed germination, seedling growth, and survival of Pappostipa speciosa (ex Stipa speciosa) and Festuca pallescens, two dominant perennial grasses from NW Patagonia that differ in palatability. We hypothesized that physical and chemical factors derived from fire differentially affect recruitment of these species. We performed experiments in the field and under laboratory and greenhouse conditions to study the integral effect of fire and of related abiotic factors (i.e., smoke, heat, charcoal, and ash) on different phases of recruitment of both species. Experimental burning promoted P. speciosa emergence over time, but they did not affect F. pallescens total emergence. Experimental burning decreased P. speciosa seedling growth (i.e., few leaves and small size), but they did not affect seedling survival. Smoke from laboratory experiments stimulated P. speciosa germination. Exposing F. pallescens seeds to 120°C decreased germination and seedling growth. Fire might act as a selective force on recruitment of both species, as well as changing competitive interactions during postfire regeneration. The effect of fire on the recruitment dynamics of the studied species depended strongly on both intrinsic species characteristics and meteorological conditions.     

Fire effects on ecosystem nitrogen cycling in a Californian bishop pine forest

Fire can cause severe nitrogen (N) losses from grassland, chaparral, and temperate and boreal forest ecosystems. Paradoxically, soil ammonium levels are markedly increased by fire, resulting in high rates of primary production in re-establishing plant communities. In a manipulative experiment, we examined the influence of wild-fire ash residues on soil, microbial and plant N pools in a recently burned Californian bishop pine (Pinus muricata D. Don) forest. Ash stimulated post-fire primary production and ecosystem N retention through direct N inputs from ash to soils, as well as indirect ash effects on soil N availability to plants. These results suggest that redistribution of surface ash after fire by wind or water may cause substantial heterogeneity in soil N availability to plants, and could be an important mechanism contributing to vegetation patchiness in fire-prone ecosystems. In addition, we investigated the impact of fire on ecosystem N cycling by comparing ¹⁵N natural abundance values from recently burned and nearby unburned P. muricata forest communities. At the burned site, ¹⁵N natural abundance in recolonising species was similar to that in bulk soil organic matter. By contrast, there was a marked ¹⁵N depletion in the same species relative to the total soil N pool at the unburned site. These results suggest that plant uptake of nitrate (which tends to be strongly depleted in ¹⁵N because of fractionation during nitrification) is low in recently burned forest communities but could be an important component of eco- system N cycling in mature conifer stands. Received: 29 June 1999 / Accepted: 24 October 1999     

A long‐term record of Quercus decline,logging and fires in a southern Swedish Fagus‐Picea forest

Abstract. We reconstructed forest development and disturbance events (fire and logging) during the last 1000 yr with tree‐ring data, pollen and charcoal analysis from a semi‐natural Fagus sylvatica‐Picea abies forest (ca. 1 km²) in the hemiboreal zone. According to pollen analysis, Quercus robur together with Pinus sylvestris was abundant in the forest until the turn of the 18th/19th centuries when these species disappeared completely (Quercus) or nearly completely (Pinus) and were replaced by Fagus and Picea. The disappearance of Quercus was corroborated by the remarkable discovery of a single Quercus stump that had been cut in the 18th century and had become overgrown and preserved by a very old Picea. In total 11 fires were dated from 1555 to 1748 from fire scars in several Pinus stumps cut 100 ‐ 200 yr ago. Since the last fire in 1748, no Quercus or Pinus have regenerated in the core of the reserve apart from single pines in neighbouring managed forest (80 yr ago). During the period of documented fires Fagus was protected from fires in a refuge made up of large boulders. Picea colonized the region at the time when the fires ceased 250 yr ago. We hypothesize that most of the fires were probably of human origin because of their patchiness and high frequency compared to the natural background levels of lightning ignitions in the region. On a 300‐yr time scale, logging and fire suppression seem to strongly overshadow the effect of climate change on forest composition and dynamics.