Simply the best: the transition of savanna saplings to trees

Tree cover in savannas is determined as much by disturbances from fire and herbivory as by rainfall and soil resources. Fire especially acts to limit tree cover via a demographic bottleneck, limiting the recruitment of tree saplings to adults. Because sapling growth rates determine rates of sapling to tree recruitment, predicting changes in tree cover requires data on sapling growth rates, commonly expressed as population means. Here, we discuss the variability in sapling growth rates in Acacia populations in a savanna in Hluhluwe iMfolozi Park in South Africa. Saplings growing at mean rates under typical fire regimes in African savannas would likely never escape the fire‐trap to become adults. Only the fastest growing saplings could grow above the flame zone between fires. We suggest that maximum growth rates are more ecologically relevant than mean growth rates in natural populations and experiments. Maximum growth rates are better than mean growth rates as predictors of sapling release within species, as shown here, and probably of which species are likely ‘winners’ in savanna tree communities.     

Fire and nutrient gradient effects on the sapling ecology of four Acacia species in the presence of grass competition

The variation among tree species in their sapling responses to disturbances such as fire, herbivory and grass competition results in different probabilities of recruitment into savanna communities. A sapling??s ability to persist under these disturbances depends on its investment in growth, defences and stored reserves, based on soil available nutrients. We examined the survival and investments in growth and defence of the saplings of four Acacia species, in the presence of competing grasses, under varying soil nutrient levels. We also have evaluated the effects of fire on these saplings. We performed the study in a greenhouse at the UKZN Botanical Garden, Pietermaritzburg, South Africa. Nutrients had no significant effect on the survival of tree saplings regardless of fire treatment. Only 22.5?% saplings survived after fire treatment compared with 84?% of control saplings. Acacia sieberiana showed the highest resprout survival, while A. karroo did not survive the fire. Growth in A. nigrescens increased only at intermediate levels (0.5N, 1N) of the fertiliser provided. Condensed tannins in A. karroo also decreased at an intermediate level (1N) of fertiliser. Resprouts grew faster in stem height than in diameter and did not invest in chemical defences. No treatment had an effect on the proportion of root and shoot dry biomass. In conclusion, African acacias are not dependent on soil nutrients for their survival and differ in their ability to resprout after fire. Intermediate levels of nutrient availability are beneficial to sapling development as opposed to zero and high nutrient levels, due to nutrient-grass dynamics. The benefits of post-fire grass removal and an increase in nutrients can be effectively utilised by only a few species of acacias.     

Demographic implications of life-history stage characteristics in two African acacias at a Makeni savanna plot in Zambia

Aims In spite of the importance of African acacias in vegetation succession and provision of goods and services, little is known about life-history variations within and among species. Much of the work done on African acacias has focused on seed predation and germination and seedling establishment, especially of Acacia tortilis, Acacia nilotica and Acacia karroo. The primary aim of the present work is to investigate differences in the demography of Acacia polyacantha and Acacia sieberiana and the relationship between life-history characteristics and population size. A secondary objective is to assess how fire, an important ecological factor in savanna vegetation, might modify the growth and demographic dynamics of the two acacias.Methods The study was conducted at the Makeni savanna plot in central Zambia, southern Africa. Seedling emergence from both non-scarified and scarified seeds sown at different times in the wet season and the fate and growth of seedlings and saplings were monitored over a period of 4 years. Annual growth of permanently marked sample trees in annually burnt and fire-protected blocks was recorded over a 6-year period (2002–08) in order to assess inter-specific differences and how fire modifies tree growth patterns. Censuses of natural saplings and trees were conducted periodically in sample blocks to determine recruitment into these life-history stages.Important findings Seedling emergence and sapling survival rates were much higher in A. sieberiana than in A. polyacantha. However, both seedling and sapling growth rates were higher in A. polyacantha than in A. sieberiana but tree growth rates were similar in the two species. Under fire protection tree growth was significantly influenced by tree size and year while under annual burning only tree size significantly affected tree growth. The dominance of A. sieberiana over A. polyacantha at the study site was attributed to higher seedling emergence rate, higher sapling survival rate and a large sapling bank that forms a reliable source of tree recruitment. The life-history stage characteristics of A. polyacantha suggest that this is an early successional species.     

Defence against vertebrate herbivores trades off into architectural and low nutrient strategies amongst savanna Fabaceae species

Herbivory contributes substantially to plant functional diversity and in ways that move far beyond direct defence trait patterns, as effective growth strategies under herbivory require modification of multiple functional traits that are indirectly related to defence. In order to understand how herbivory has shaped plant functional diversity, we need to consider the physiology and architecture of the herbivores and how this constrains effective defence strategies. Here we consider herbivory by mammals in savanna communities that range from semi‐arid to humid conditions. We posited that the saplings of savanna trees can be grouped into two contrasting defence strategies against mammals, namely architectural defence versus low nutrient defence. We provide a mechanistic explanation for these different strategies based on the fact that plants are under competing selection pressures to limit herbivore damage and outcompete neighbouring plants. Plant competitiveness depends on growth rate, itself a function of leaf mass fraction (LMF) and leaf nitrogen per unit mass (N_m). Architectural defence against vertebrates (which includes spinescence) limits herbivore access to plant leaf materials, and partly depends on leaf‐size reduction, thereby compromising LMF. Low nutrient defence requires that leaf material is of insufficient nutrient value to support vertebrate metabolic requirements, which depends on low N_m. Thus there is an enforced tradeoff between LMF and N_m, leading to distinct trait suites for each defence strategy. We demonstrate this tradeoff by showing that numerous traits can be distinguished between 28 spinescent (architectural defenders) and non‐spinescent (low nutrient defenders) Fabaceae tree species from savannas, where mammalian herbivory is an important constraint on plant growth. Distributions of the strategies along an LMF‐N_m tradeoff further provides a predictive and parsimonious explanation for the uneven distribution of spinescent and non‐spinescent species across water and nutrient gradients.     

Defoliation depletes the carbohydrate reserves of resprouting <Emphasis Type="Italic">Acacia</Emphasis> saplings in an African savanna

Over the past century there has been a global trend towards tree expansion and densification in rangelands and savannas. This phenomenon is commonly referred to as bush encroachment. In South Africa Acacia karroo is one of the key species responsible for bush encroachment. It has been suggested that the combination of fire and browsing might limit bush encroachment by A. karroo more effectively than either browsing or fire alone. We hypothesized that these repeated disturbances progressively deplete root carbohydrates and compromise resprouting ability. This was tested by burning and then manually defoliating A. karroo once a month for 1 year. Manual defoliation did not inhibit the rapid shoot elongation after topkill of A. karroo saplings. During this initial phase, the growth of the new shoots of A. karroo was dependent more on mobilised root reserves than on photoassimilates from the new shoots. Frequent manual defoliation of resprouting A. karroo saplings prevented the replenishment of starch reserves. We suggest a mechanism for how the interaction of browsing and fire can suppress and perhaps reverse bush encroachment in African savannas. Saplings that have reduced starch reserves at the end of dry season due to browsing will struggle to resprout if they are burnt. Even if they do not die, they will be less able to escape fire damage in the next fire than if they had been able to resprout unimpeded.     

Fire-tolerance mechanisms of common woody plant species in a semiarid savanna in south-western Zimbabwe

The frequency of fire has increased in savannas yet few studies have assessed how plants persist when subjected to long‐term disturbance by fire. We investigated the contributions of bark thickness and resprouting to the persistence of woody plants in two fire trials that were started in 1948 and 1949. The number of resprouts per individual, bark thickness, basal diameter and height of woody plants were measured in unburnt plots and those burnt annually, triennially and quinquennially during the late dry season. Changes in tree density, number of resprouts and individuals in different height classes between 1963 and 2002 were assessed. Bark thickness varied among species and also increased with increases in basal diameter. Generally, plants with thick bark survived fire more than those with thin bark. Resprouting was the major fire survival strategy for most species. The number of resprouts produced per plant ranged from 4 ± 3 (Acacia rehmanniana) to 14 ± 9 (Pseudolachnostylis maprouneifolia). Fire reduced species richness in plots burnt annually and triennially by 47% and 6% respectively. Species richness increased in unburnt plots (5%) and those burnt quinquennially (16%). Most woody species survived fire through a combination of traits.     

Investigating the vulnerability of an African savanna tree (Sclerocarya birrea ssp. caffra) to fire and herbivory

Sclerocarya birrea ssp. caffra (marula), a typical savanna tree, is vulnerable to the effects of fire, herbivory and their combination. This paper investigated the relative importance of these agents of disturbance, at the level of the individual stem, by specifically focusing on the following questions: (i) What is the greatest cause of mortality in adult marula stems in conservation areas with both elephants and fire? (ii) Does fire interact with bark stripping to cause adult stem mortality and if so what is the dominant mechanism? (iii) At what stem diameter are marulas resistant to fire? Field surveys quantified the extent of damage in marula individuals in the Kruger National Park, South Africa, highlighting the high levels of extreme herbivory such as toppling (7%) and pollarding (8%), relative to bark stripping (only 6% with more than 50% of the circumference stripped). In addition to extreme herbivory, the progression from bark stripping through to invasion of the soft, exposed heartwood by wood borers, often facilitated by fire, through to toppling of the weakened stem after successive fires, appears to be the dominant mechanism by which fire interacts with herbivory to cause adult stem death. Bark stripping and fire manipulation experiments indicated that bark stripping failed to increase the vulnerability of stems to fire directly through transport tissue damage. However, the combination of bark stripping and fire reduced the ability of the stem to regrow bark, increasing the vulnerability of the exposed stem to boring insects and future fires. Fire manipulation experiments were used to identify the minimum stem diameter of resistance to fire. Marula resisted stem death when greater than 3.4 cm in basal diameter. This paper emphasizes the importance of both fire and herbivory in the development of woody plant population structure and by extension, the relative proportion of trees and grasses in savanna landscapes.     

Density-dependent shoot-borer herbivory increases the age of first reproduction and mortality of neotropical tree saplings

Shoot herbivory by the sapling specialist shoot-borer Cromarcha stroudagnesia (Lepidoptera, Pyralidae, Chrysauginae) is shown to have large direct and indirect effects on the rates of height increment and mortality of saplings of its host tree, Tabebuia ochracea (Bignoniaceae), in the secondary successional tropical dry forests of the Area de Conservación Guanacaste, Costa Rica. Experiments and field observations over 3-4 years show a substantial reduction in sapling height increments due to C. stroudagnesia herbivory, of equivalent magnitude to the difference in height increments between undamaged saplings in canopy gaps and full understorey shade. Extrapolating this data at average amounts of C. stroudagnesia herbivory increases the duration of the pre-reproductive sapling life stage by about 40% relative to undamaged plants. This is an underestimate, as top shoot herbivory by C. stroudagnesia also increased the probability of canopy gap saplings being overtopped and shaded by surrounding vegetation. Sapling mortality was increased by C. stroudagnesia herbivory, with 11.8% of the most heavily damaged young saplings dying in 3 years while no undamaged saplings died. Cromarcha stroudagnesia herbivory strongly increases with the number of conspecific T. ochracea saplings and the number of conspecific shoots within 50 m of focal saplings. It is therefore likely to disproportionately decrease the number of saplings and rate of recruitment to reproductive age in areas of high conspecific sapling density. These results suggest that sapling herbivory, especially herbivory of terminal meristems, has an important but largely unexplored influence on the population dynamics of tropical tree species. They further demonstrate that sapling herbivory by insects, in addition to the well-studied insect predation and herbivory of seedlings, is likely to influence tree species coexistence in tropical forests.     

Differences in insect species richness and faunal composition of birch seedlings, saplings and trees: the importance of plant architecture

Abstract. 1. Data are presented on the species richness and faunal composition of herbivorous insects on birch seedlings, saplings and trees at one site in Northern England.
2. Species richness of insect herbivores in equal-sized samples from birch seedlings and trees was similar through most of the season.
3. Effects of plant architecture were confined to the first sampling date, when seedling faunas were species poor compared with trees – possibly due to safe overwintering sites on the extensive bark, twigs and buds of trees.
4. The faunal composition of birch seedlings, saplings and trees was also similar. Out of a total of 112 recorded species of herbivores, only one aphid species was confined to seedlings.
5. Similarly, no evidence for clear-cut vertical stratification of insects within trees was found.
6. Species turnover as host plants mature ('horizontal' stratification) and vertical stratification within trees add little to the high overall species richness of birch-feeding insects in Britain, contrary to the predictions of Lawton (1983).     

Impact of fire on leaf nutrients,arthropod fauna and herbivory of native and exotic eucalypts in Kings Park,Perth, Western Australia

The vegetation of Kings Park, near the centre of Perth, Western Australia, once had an overstorey of Eucalyptus marginata (jarrah) or Eucalyptus gomphocephala (tuart), and many trees still remain in the bushland parts of the Park. Avenues and roadsides have been planted with eastern Australian species, including Eucalyptus cladocalyx (sugar gum) and Eucalyptus botryoides (southern mahogany), both of which have become invasive. The present study examined the effect of a recent burn on the level of herbivory on these native and exotic eucalypts. Leaf damage, shoot extension and number of new leaves were measured on tagged shoots of saplings of each tree species in unburnt and burnt areas over an 8‐month period. Leaf macronutrient levels were quantified and the number of arthropods on saplings was measured at the end of the recording period by chemical knockdown. Leaf macronutrients were mostly higher in all four species in the burnt area, and this was associated with generally higher numbers of canopy arthropods and greater levels of leaf damage. It is suggested that the pulse of soil nutrients after the fire resulted in more nutrient‐rich foliage, which in turn was more palatable to arthropods. The resulting high levels of herbivory possibly led to reduced shoot extension of E. gomphocephala, E. botryoides and, to a lesser extent, E. cladocalyx. This acts as a negative feedback mechanism that lessens the tendency for lush, post‐fire regrowth to outcompete other species of plants. There was no consistent difference in the levels of the various types of leaf damage or of arthropods on the native and the exotic eucalypts, suggesting that freedom from herbivory is not contributing to the invasiveness of the two exotic species.     

Synergistic interactions between leaf beetle herbivory and fire enhance tamarisk (Tamarix spp.) mortality

The combined effects of herbivory and fire on plant mortality were investigated using prescribed burns of tamarisk (Tamarix ramosissima Lebed) exposed to herbivory by the saltcedar leaf beetle (Chrysomelidae: Diorhabda carinulata Desbrocher). Tamarix stands in the Humboldt Sink (NV, USA) were divided into three treatments: summer burn (August 2006), fall burn (October 2006) and control (unburned), and litter depth was manipulated to vary fire intensity within burn seasons. A gradient of existing herbivory impact was described with three plant condition metrics prior to fire: reduced proportions of green canopy, percent root crown starch sampled at the height of the growing season (August 2006), and percent root crown starch measured during dormancy (December 2006). August root crown starch concentration and proportion green canopy were strongly correlated, although the proportion green canopy predicted mortality better than August root crown starch. December root crown starch concentration was more depleted in unburned trees and in trees burned during the summer than in fall burn trees. Mortality in summer burned trees was higher than fall burned trees due to higher fire intensity, but December root crown starch available for resprouting in the spring was also lower in summer burned trees. The greatest mortality was observed in trees with the lowest December root crown starch concentration which were exposed to high fire intensity. Disproportionate changes in the slope and curvature of prediction traces as fire intensity and December starch reach reciprocal maximum and minimum levels indicate that beetle herbivory and fire intensity are synergistic.     

The impact of elephants, Loxodonta africana, on woody vegetation through selective debarking in Samburu and Buffalo Springs National Reserves, Kenya

The impact of elephants on the woody plant community through debarking was investigated in Samburu and Buffalo Springs National Reserves, Kenya. Acacia elatior Brenan , the most abundant tree species in the riverine zone, accounted for 68% (n = 1375) of woody plants. A. tortilis (Forsskal) Hayne dominated plots away from the river. Debarking incidences were significantly higher for A. elatior than for other species indicating selective utilization. The riverine zone by virtue of having more trees of the preferred species, A. elatior, had the highest debarking incidences. Presence of very few saplings along the river is attributed to both elephant trampling and herbivory by other species. An estimated 38.5% and 22.5% of the riverine A. elatior and A. tortilis trees respectively, were bound to die within the next 4–5 years because of severe debarking, ≥75% of bark circumference. Debarking was positively correlated with stem circumference; the medium-sized trees being the worst affected by the elephants' selective debarking behaviour. Intense debarking incidences were recorded during the dry season. Through the elephants' selective debarking, the riverine habitat is bound to open up gradually, leading to considerable habitat change in the near future. Elephant impact on vegetation is less away from the river and increases with their densities.     

Impacts of a mineral lick‐centred land use system on woody vegetation cover in an East African Savannah

Evaluation of woody vegetation changes with distance from a salt crater was conducted in the semi‐arid rangelands of southern Ethiopia. Data on live woody plants were collected over three seasons at 0, 1, 4, 6, 9 and 12 km from the salt crater. The density and diversity of woody plants differed significantly (p < .01) along the distance gradient. Six woody plant families were identified of which Fabaceae and Burseraceae were the dominant families. Acacia drepanolobium, Acacia nilotica, Commiphora africana and Acacia mellifera were among the severely encroaching woody species. There were high proportions of seedlings and saplings recorded closer to the salt crater showing a vigorous recruitment by woody plants. Woody plant encroachment along the 12‐km transect ranged from a low to severe encroachment, which could be translated into poor rangeland condition. Changes in soil characteristics increased grazing pressure and sedentary settlement around the salt crater, and the breakdown of traditional institutions seems to be major contributing factors to these vegetation changes. We suggest that severely encroached areas could be improved through a combination of methods such as bush clearing, prescribed fire, browsing animals and proper grazing management.     

Bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition

The mosaic of trees, shrubs and open grassland in mesic African savannas is highly dynamic and strongly influenced by mammal herbivory and fire. We investigated the bird fauna in four different savanna habitats to help assess the impacts of vegetation change on this component of faunal diversity. Birds were censused, plant species were identified and vegetation structure was measured in four different vegetation types (Acacia nilotica woodland, Acacia nigrescens woodland, broadleaf thicket and open grassland) in the Hluhluwe-Umfolozi Park in northern KwaZulu Natal, South Africa. Multivariate ordination analyses were used to determine the relative importance of vegetation structure and floristic composition in defining bird assemblages. The bird communities of the grasslands, the acacia woodlands, and the broadleaf woodlands were clearly separated on the first axis of the detrended canonical correspondence analysis (DCCA). Canopy cover and foliage height diversity (FHD) were strongly correlated with the first axis of DCCA, possibly reflecting a secondary successional series from grassland to woodland, known as bush encroachment. Floristic composition (based on presence–absence data only) seemed to be less important for bird community composition than vegetation structure. The results indicate that changes in vegetation structure, caused by bush encroachment, could cause concomitant changes in bird community composition.     

Herbivory effects on saplings are influenced by nutrients and grass competition in a humid South African savanna

Woody plant encroachment is a common consequence of disturbance in savannas. Grazers and browsers interfere with sapling establishment dynamics by direct consumption of plant tissue, changing soil nutrient status (through fertilization and trampling) and grass competition. Studies evaluating the effects of herbivory on sapling establishment have mostly been extrapolated from single species. In a controlled field experiment, we studied the effects of clipping (simulating grazing and browsing), nutrients, grass competition, and their interactive effects on sapling survival and growth of four dominant humid and four dominant mesic savanna species. We conducted this experiment in a humid South African savanna. We found no effects on sapling survival by the treatments provided. However, clipped saplings of all species increased their investment in relative growth rate of stem length (RGR_L). Clipping had a greater negative impact on relative growth rate of more humid than mesic species in terms of stem diameter (RGR_D), total dry biomass and proportion of leaf biomass. Nutrients had a positive effect on the RGR_L and sapling biomass of three mesic species. Positive effects of nutrients on RGR_L of one humid and two mesic species were observed in their clipped saplings only. Grass competition had a strong negative impact on all growth parameters measured. Clipped saplings of one humid and two mesic species had lower RGR_L with grass competition whereas intact saplings showed no significant response. After clipping, humid savanna species were more vulnerable to grass competition than mesic species, with reduced ability to use nutrients. In conclusion, herbivory increases sapling vulnerability to grass competition, with humid species being more susceptible than mesic species, indicating that woody-plant control strategies are more likely to be effective in humid savannas.     

Plant traits affecting herbivory on tree recruits in highly diverse subtropical forests

Differences in herbivory among woody species can greatly affect the functioning of forest ecosystems, particularly in species-rich (sub)tropical regions. However, the relative importance of the different plant traits which determine herbivore damage remains unclear. Defence traits can have strong effects on herbivory, but rarely studied geographical range characteristics could complement these effects through evolutionary associations with herbivores. Herein, we use a large number of morphological, chemical, phylogenetic and biogeographical characteristics to analyse interspecific differences in herbivory on tree saplings in subtropical China. Unexpectedly, we found no significant effects of chemical defence traits. Rather, herbivory was related to the plants' leaf morphology, local abundance and climatic niche characteristics, which together explained 70% of the interspecific variation in herbivory in phylogenetic regression. Our study indicates that besides defence traits and apparency to herbivores, previously neglected measures of large-scale geographical host distribution are important factors influencing local herbivory patterns among plant species.     

Nutrient status not secondary metabolites drives herbivory and pathogen infestation across differently mycorrhized tree monocultures and mixtures

Research aimed at understanding the mechanisms underlying the relationship between tree diversity and antagonist infestation is often neglecting resource-use complementarity among plant species. We investigated the effects of tree species identity, species richness, and mycorrhizal type on leaf herbivory and pathogen infestation. We used a tree sapling experiment manipulating the two most common mycorrhizal types, arbuscular mycorrhiza and ectomycorrhiza, via respective tree species in monocultures and two-species mixtures. We visually assessed leaf herbivory and pathogen infestation rates, and measured concentrations of a suite of plant metabolites (amino acids, sugars, and phenolics), leaf elemental concentrations (carbon, nitrogen, and phosphorus), and tree biomass. Tree species and mycorrhizal richness had no significant effect on herbivory and pathogen infestation, whereas species identity and mycorrhizal type had. Damage rates were higher in arbuscular mycorrhizal (AM) than in ectomycorrhizal (EM) trees. Our structural equation model (SEM) indicated that elemental, but not metabolite concentrations, determined herbivory and pathogen infestation, suggesting that the investigated chemical defence strategies may not have been involved in the effects found in our study with tree saplings. Other chemical and physical defence strategies as well as species identity as its determinant may have played a more crucial role in the studied saplings. Furthermore, the SEM indicated a direct positive effect of AM trees on herbivory rates, suggesting that other dominant mechanisms, not considered here, were involved as well. We found differences in the attribution of elemental concentrations between the two rates. This points to the fact that herbivory and pathogen infestation are driven by distinct mechanisms. Our study highlights the importance of biotic contexts for understanding the mechanisms underlying the effects of biodiversity on tree-antagonist interactions.     

Effects of different herbivores on an actinorhizal species in Northwest Patagonia

Above-ground herbivory has a direct impact on plant life cycles, particularly at more sensitive stages, due to reduction of vegetative biomass. However, this effect may not be negative if it results in net biomass compensation. As sapling stage could be the best stage for native species to be outplanted, understanding the impact of aboveground herbivory on tree saplings is necessary for restoration purposes. We studied the effect of herbivory on saplings of Ochetophila trinervis (Rhamnaceae), a native woody species from North-west Patagonia, which forms an actinorhizal symbiosis with the N₂-fixing actinobacteria Frankia. This tree species has the potential to be used for recovering degraded lands. Nevertheless, there is a perplexing contradiction between the high seed output of O. trinervis and the scarcity of saplings in the field. For 4 months, 1-year-old O. trinervis saplings were exposed to aboveground herbivory by generating different protection degrees (unprotected, protected against some kind of walking herbivores—protected saplings; and protected against all kind of walking herbivores—excluded saplings). The impact of herbivores over sapling survival was minimal (92?±?3%, mean?±?SE) and it was similar among saplings exposed to different protection degrees. The highest frequency of foliar damage in excluded saplings suggests the attack of flying herbivores. The increased emergence of new sprouts and root length growth in saplings highly damaged by herbivores (about three fold and two fold higher than in excluded saplings, respectively), evidenced the capacity of O. trinervis to develop a compensatory growth. The results contradict the assumption that herbivory explains the low density of saplings despite high seed production. Given the high-sapling survival and biomass compensation of O. trinervis after herbivory, we suggest that this species might be appropriate for restoration of degraded areas in the region.     

Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.