Facilitation and Inhibition of Seedlings of an Invasive Tree (Acer platanoides) by Different Tree Species in a Mountain Ecosystem

Facilitation is known to be an important process structuring natural plant communities. However, much less is known about its role in facilitating the invasion of ecosystems by non-native plant species. In this study we evaluated the effects of invasive (Acer platanoides) and native (Pseudotsuga menziesii) forest types on the performance of A. platanoides seedlings, and related these effects to structural and functional properties associated with the two forest types, in a native P. menziesii forest that is being invaded by A. platanoides. Acer platanoidesseedlings had higher densities, recruitment, and survival, and experienced less photoinhibition and water stress when beneath conspecific canopies than in the adjacent P. menziesii forest. Soil moisture and canopy cover were greater in the invaded patch than the native forest. There was no difference in soil fertility or understory light levels between locations. These demographic (i.e. seedling survival), physiological, and environmental differences appeared to be due to the effects of A. platanoides and P. menziesii trees. Thus, Acer trees appear to produce a more mesic environment by modifying the structure and phenology of the forest canopy and by altering the timing of transpirational water loss relative to P. menziesii. Environmental modification by invaders that lead to positive effects on conspecifics may help us to understand the dramatic success and lag periods of some invasive species     

Impact of invasion of Acer platanoides on canopy structure and understory seedling growth in a hardwood forest in North America

Invasion by exotic plant species is known to affect native communities and ecosystems, but the mechanisms of the impacts are much less understood. In a field study, we examined the effects of a tree invader, Acer platanoides (Norway maple, NM), on canopy structure and seedling growth in the understory of a North American deciduous forest. The experimental site contains a monospecific patch of A. platanoides and a mixed patch of A. platanoides with its native congener, A. rubrum (red maple, RM). In the study, we examined canopy characteristics of three types of trees in the forests, i.e., RM trees in the mixed forest, NM trees in the mixed forest, and NM trees in its monospecific patch. Height growth and biomass production of RM and NM seedlings under intact canopies and newly created gaps of the three types of trees were followed for two growing seasons. We found that removal of half of the canopy from focal trees increased canopy openness and light transmission to the forest floor, but to a greater extent under NM trees than under RM trees. Seedlings of these two Acer species varied greatly in biomass production under canopies of the same type of trees and in their responses to canopy opening. For example, seedlings of the exotic NM grown under the native RM trees in the mixed forests increased biomass production by 102.4% compared to NM seedlings grown under conspecific trees. The native RM seedlings grown under NM trees, however, reduced biomass production by 23.5% compared to those grown under conspecific trees. It was also observed that RM was much more responsive in biomass production to canopy opening than NM. For instance, total seedling biomass increased by 632.2% in RM, but by only 134.6% in NM in response to the newly created gaps. In addition, we found that NM seedlings allocated a greater portion of biomass below-ground as canopy openness increased, whereas the same trend was not observed in RM seedlings. Our results thus demonstrated that invasion of NM significantly altered canopy structure and community dynamics in the hardwood forest. Because the exotic NM seedlings are able to grow well under the native RM trees, but not vice versa, NM will likely expand its distribution in the forests and make it an ever increasingly serious tree invader in its non-native habitats, including North America.     

Reduced risk for positive soil-feedback on seedling regeneration by invasive trees on a very nutrient-poor soil in Seychelles

Invasive plants sometimes alter habitat conditions so as to promote further invasion, either by the same or by other non-native species. Such positive feedbacks often occur because the non-native species increases soil fertility, thereby favouring recruitment of non-native seedlings. This has been demonstrated in nitrogen-poor habitats invaded by nitrogen-fixing species, but it is unclear whether similar processes operate in habitats limited by phosphorus and other nutrients. I compared the growth of seedlings of Cinnamomum verum, an abundant invasive tree on phosphorus-poor soils in the Seychelles, in soils taken from beneath different tree species. I expected that soil phosphorus availability would be higher beneath stands of C. verum than beneath stands of either the native Northea hornei or the non-native nitrogen-fixing species, Falcataria moluccana. I therefore predicted that C. verum seedlings would grow faster in soil taken from beneath C. verum trees than in soil taken from beneath either of the other two species. To test this hypothesis, I performed a bioassay experiment with seedlings of C. verum grown in soils from stands of C. verum, F. moluccana and N. hornei. Different nutrient treatments (control, plus phosphorus (P), plus nitrogen (N), plus N and P, and plus complete fertilizer) were applied to investigate how nutrient availabilities modulate the effects of the trees. In the control treatment without added nutrients, there was a weak tendency for seedlings to perform better in the soils from beneath invasive than native trees. However, seedling growth in soils from beneath invasive species was markedly higher following the addition of phosphorus in the case of the F. moluccana soil, and complete fertilizer in the case of the C. verum soil. These results indicate that on very nutrient-poor soils, a low supply of nutrients other than N may reduce the risk of a soil-feedback by invasive trees on seedling regeneration.     

Canopy manipulations of exotic Bitter Willow (Salix elaeagnos) forest for indigenous seedling recruitment: A pilot study

The invasive exotic tree species Bitter Willow (Salix elaeagnos; Salicaceae) has colonised areas of rank exotic grassland and has been found to contain indigenous seed, dispersed by frugivorous birds into the monospecific stands. This small pilot study examined whether indigenous seedlings that have germinated in the understorey of exotic Bitter Willow stands could be stimulated to establish through the creation of small‐scale canopy gaps. In Bitter Willow forest, four single Bitter Willow trees were poisoned to create canopy gaps. Light transmission and seedling regeneration of tree and shrub species were assessed beneath both the four manipulated and three comparable intact Bitter Willow canopies. Over 3 years, seedling height and density increased more beneath opened compared to intact Bitter Willow canopies. These results suggest that Bitter Willow can fill the roles of both a facilitative nurse and a perch tree. Larger‐scale canopy manipulation experiments of both Bitter Willow and other Salix species are needed to determine the full potential of canopy manipulations for forest restoration.     

Performance of seedlings of the invasive alien tree Schinus molle L. under indigenous and alien host trees in semi-arid savanna

We assessed the importance of host trees in influencing invasion patterns of the alien tree Schinus molle L. ( Anacardiaceae ) in semi-arid savanna in South Africa. Recruitment of S. molle is dependent on trees in its invaded habitat, particularly Acacia tortilis Hayne. Another leguminous tree, the invasive alien mesquite ( Prosopis sp.), has become common in the area recently, but S. molle rarely recruits under canopies of this species. Understanding of the association between these species is needed to predict invasion dynamics in the region. We conducted experiments to test whether: (i) seedling survival of S. molle is better beneath A. tortilis than beneath mesquite canopies; (ii) growth rates of S. molle seedlings are higher beneath A. tortilis than beneath mesquite. Results showed that growth and survival of S. molle did not differ significantly beneath the native A. tortilis and the alien Prosopis species. This suggests that microsites provided by canopies of mesquite are as good for S. molle establishment as those provided by the native acacia. Other factors, such as the failure of propagules to arrive beneath mesquite trees, must be sought to explain the lack of recruitment beneath mesquite.     

<Emphasis Type="Italic">Acer rubrum</Emphasis> (red maple) growth is negatively affected by soil from forest stands dominated by its invasive congener (<Emphasis Type="Italic">Acer platanoides</Emphasis>, Norway maple)

Invasive species continue to alter the plant communities of the eastern United States. To better understand the mechanisms and characteristics associated with invasive success, we studied competition between two Acer species. In a greenhouse, we tested (1) the effect of forest soil type (beneath an invasive and native stand) on seedling growth of the invasive Acer platanoides (Norway maple) and native A. rubrum (red maple), and the (2) effects of full (above- and below-ground) and partial inter-specific competition on species growth. We found A. rubrum growth was negatively affected by soil from the invaded stand, as it had lower above-ground (32%) and below-ground (26%) biomass, and number of leaves (20%) than in the native soil. The root:shoot resource allocations of A. platanoides depended on soil type, as it had 14% greater root:shoot mass allocation in the native soil; this ability to change root:shoot allocation may be contributing to the ecological success of the species. Widely published as having a large ecological amplitude, A. rubrum may be a useful species for ecological restoration where A. platanoides has been present, but the impacts of A. platanoides on soil functioning and subsequent plant interactions must be addressed before protocols for native reintroductions are improved and implemented.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

Dispersal and recruitment limitation in native versus exotic tree species: life‐history strategies and Janzen‐Connell effects

Life‐history traits of invasive exotic plants are typically considered to be exceptional vis‐à‐vis native species. In particular, hyper‐fecundity and long range dispersal are regarded as invasive traits, but direct comparisons with native species are needed to identify the life‐history stages behind invasiveness. Until recently, this task was particularly problematic in forests as tree fecundity and dispersal were difficult to characterize in closed stands. We used inverse modelling to parameterize fecundity, seed dispersal and seedling dispersion functions for two exotic and eight native tree species in closed‐canopy forests in Connecticut, USA. Interannual variation in seed production was dramatic for all species, with complete seed crop failures in at least one year for six native species. However, the average per capita seed production of the exotic Ailanthus altissima was extraordinary: > 40 times higher than the next highest species. Seed production of the shade tolerant exotic Acer platanoides was average, but much higher than the native shade tolerant species, and the density of its established seedlings (≥ 3 years) was higher than any other species. Overall, the data supported a model in which adults of native and exotic species must reach a minimum size before seed production occurred. Once reached, the relationship between tree diameter and seed production was fairly flat for seven species, including both exotics. Seed dispersal was highly localized and usually showed a steep decline with increasing distance from parent trees: only Ailanthus altissima and Fraxinus americana had mean dispersal distances > 10 m. Janzen‐Connell patterns were clearly evident for both native and exotic species, as the mode and mean dispersion distance of seedlings were further from potential parent trees than seeds. The comparable intensity of Janzen‐Connell effects between native and exotic species suggests that the enemy escape hypothesis alone cannot explain the invasiveness of these exotics. Our study confirms the general importance of colonization processes in invasions, yet demonstrates how invasiveness can occur via divergent colonization strategies. Dispersal limitation of Acer platanoides and recruitment limitation of Ailanthus altissima will likely constitute some limit on their invasiveness in closed‐canopy forests.     

Biogeographical contrasts to assess local and regional patterns of invasion: a case study with two reciprocally introduced exotic maple trees

Quantitative comparisons of distribution and abundance of exotic species in their native and non‐native ranges represent a first step when studying invaders. However, this approach is rarely applied 2 particularly to tree species. Using biogeographical contrasts coupled with regional dispersal surveys, we assessed whether two exotic maple tree species, Acer negundo and Acer platanoides, can be classified as invasive in the non‐native regions surveyed. We also examined the importance of biogeography in determining the degree of invasion by exotic species using this reciprocal approach. Local‐scale surveys were conducted in a total of 34 forests to compare density, relative abundance, age structure of native and introduced populations, and whether the two introduced maple species negatively affected native tree species density. Regional‐scale surveys of a total of 136 forests were then conducted to assess distribution in the introduced regions. Introduced populations of A. negundo were denser than populations measured in their native range and negatively related to native tree species density. Age structure did not differ between regions for this species. At the regional scale, this species has invaded most of the riparian corridors sampled in France. Conversely, the density of A. platanoides introduced populations was similar to that of native populations and was not related to native tree species density. Although seedling recruitment was higher away than at home, this species has invaded only 9% of the forests sampled in southern Ontario, Canada. Although reported invasive, these two exotic maple species differed in their relative demographic parameters and regional spread. Acer negundo is currently invasive in southern France while A. platanoides is not aggressively invasive in southern Ontario. Importantly, this study effectively demonstrates that biogeography through structured contrasts provide a direct means to infer invasion of exotic species.     

Role of nurse plants in Araucaria Forest expansion over grassland in south Brazil

Abstract Araucaria Forest expansion over grassland takes place under wet climate conditions and low disturbance and it is hypothesized that isolated trees established on grassland facilitate the establishment of forest woody species beneath their canopies. Forest with Araucaria angustifolia is a particular type of Brazilian Atlantic Forest and the main forest type on the highland plateau in south Brazil, often forming mosaics with natural Campos grassland. The objectives of this paper were to evaluate the role of isolated shrubs and trees as colonization sites for seedlings of Araucaria Forest woody species on grassland, to determine which species function as preferential nurse plants in the process and the importance of vertebrate diaspore dispersal on the structure of seedling communities beneath nurse plants. The study was conducted in São Francisco de Paula, Rio Grande do Sul State, where we sampled isolated shrubs and trees in natural grassland near Araucaria Forest edges. Seedlings were counted and identified, and seedling diaspore dispersal syndromes, size and colour were registered. We detected 11 woody species with a potential role in nucleating grassland colonization by forest species. Beneath the canopies of nurse plants more forest species seedlings were found compared with open field grassland and the seedlings had diaspores mostly dispersed by vertebrates. Also, more seedlings were found under the canopy of A. angustifolia than beneath other nurse plant species. We conclude that A. angustifolia trees established on grassland act as nurse plants, by attracting disperser birds that promote colonization of the site by other forest species seedlings, and that under low level of grassland disturbance, conservation of frugivorous vertebrate assemblages may increase forest expansion over natural grassland and also facilitate the regeneration of degraded forest areas.     

Facilitation of tree seedling establishment in a sand dune succession

Abstract. A nucleated pattern of establishment by pine seedlings of Pinus strobus and P. resinosa around pre-established oak trees of Quercus rubra in a sand dune succession in Ontario, Canada was examined using field observations, seed planting and habitat manipulation. Densities of young pines beneath oak canopies were approximately six times greater than in treeless areas, and densities on the north sides of the trees were significantly greater than on the south sides. However, oaks younger than 35 yr showed no preferential establishment beneath them, while the pine population structures beneath older oaks indicated single periods of successful recruitment. Pine seed planted beneath and beyond oak canopies of three sizes germinated primarily beneath the canopies of medium and large-sized oaks, but subsequent survivorship of seedlings over two growing seasons was poor. Several micro-environmental conditions were changed by oak canopies, but only shade showed a pattern closely corresponding to that of seedling establishment. A habitat manipulation experiment confirmed the primary role of shade in facilitating pine seedling establishment beneath oaks. Failure of pine to continue recruiting successfully beneath facilitating trees is tentatively attributed to intraspecific competition among pine individuals.     

Interactions between annuals and woody perennials in a Western Australian nature reserve

Abstract. We studied the interactions between woody perennial species and native and non-native annual species in a number of vegetation types within a nature reserve in the Western Australian wheatbelt. In particular, we examined the responses of annuals to perennial canopy removal, fire, soil disturbance and nutrient additions, and the effects of removal of annuals on perennial seedling regeneration. Experimental shrub removal significantly increased the abundance of annuals in a dense shrubland dominated by Allocasuarina campestris, but had no effect in a more open species-rich sandplain heath. Soil disturbance and nutrient addition in the heath area had no significant influence on annual abundance until three years after treatment. Fire had no clear effect on annual abundance in the heath within the reserve, but promoted a large increase in non-native species within an adjacent roadverge. A pattern of increased soil nutrient levels was accompanied by greatly increased non-native annual abundance beneath individual trees of Santalum spicatum. Exploratory laboratory bioassay experiments indicated that several woody perennials produced leachates that were capable of reducing the germination or growth of the introduced grass Avena fatua, indicating that allelopathy may be an important component of the interaction between the annual and perennial components. Within a woodland community, fire temporarily reduced the abundance of annual species and increased the establishment of perennial seedlings. Field experiments showed that annuals significantly reduced the survival of seedlings of the shrub Allocasuarina campestris. Our results indicate that intact native vegetation canopies effectively prevent invasion by non-native annuals, and that regeneration by native perennials is likely to be inhibited by the presence of an abundant annual cover.     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.     

Nutrition ofPinus caribaea in its native savanna habitat

Summary Fertility levels in soils beneathPinus caribaea trees were examined in the Mountain Pine Ridge savannas, Belize, where fire control has precipitated the development of pine woodland. Slight surface soil enrichment was recorded beneath pine canopies, but to levels well below those found beneath associated hardwoods. Estimates of total nutrient pools beneath trees showed modest cation accumulation beneath a 73 year old tree but some defecits in Ca and Mg beneath a 24 year old tree. A tap root cutting experiment on trees of the same species revealed no significant declines in foliar nutrient levels after 19 months. It is concluded that no pronounced long-term deterioration in soil fertility levels is developing beneath stands ofP. caribaea in the savanna, although some temporary nutrient declines may exist beneath young pine stands. Atmospheric inputs are the most likely source of nutrient accretion and it is suggested that the establishment of hardwood associates with pine may enhance the rates of nutrient capture from this source.     

Transpirational Water Loss in Invaded and Restored Semiarid Riparian Forests

The invasive tree, Tamarix sp., was introduced to the United States in the 1800s to stabilize stream banks. The riparian ecosystem adjacent to the middle Rio Grande River in central New Mexico consists of mature cottonwood (Populus fremontii) gallery forests with a dense Tamarix understory. We hypothesized that Populus would compensate for reduced competition by increasing its water consumption in restored riparian plots following selective Tamarix removal, resulting in similar transpiration (T) among stands. The northern study site included a Populus stand invaded by Tamarix (INV_N) and a restored Populus‐only stand (RES_N), as did a southern site (INV_S and RES_S) approximately 80 miles south. At each site, 20 × 20–m plots were established where up to 16 stems were monitored throughout the 2004 growing season using thermal dissipation sapflow sensors. Populus sapflux rates were greater in restored stands, suggesting those trees compensated for understory removal by using more water. Sapflow was scaled to estimate stand‐level T based on a quantitative assessment of sapwood basal area (A_sw) by species. Although exotic species represented 85 and 91% of the total stems in the invaded stands, it amounted to only 3% (INV_S) and 4% (INV_N) of the total A_sw, contributing proportionately less to T compared to Populus. Our results indicate that removing Tamarix from the Populus understory in this riparian forest had a minimal impact on stand water balance. Riparian restoration of the type discussed herein should focus primarily on enhancing riparian health rather than generating water.     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

Acacia trees as keystone species in Negev desert ecosystems

Abstract. The only trees in most of the Negev desert are 3 native Acacia species. We tested the hypothesis that they act as keystone species as a result of the improved soil conditions under their canopies. Furthermore, because many Acacia populations suffer high levels of mortality due to water stress, we tested whether trees in high mortality populations had diminished effects on plant species and soil quality under their canopies. We show that plant species diversity beneath the tree canopies is higher than in the surrounding areas. There was also a clearly identifiable suite of species with higher occurrence under the trees. Plant species composition differed significantly between high and low mortality sites. However, there was higher species diversity in high mortality sites and under trees with higher water stress. Soil nutrient content was higher under the trees than in the open areas, especially under larger trees and trees with higher water status. The results indicate that there is a combination of positive and negative effects of Acacia trees on the under‐canopy environment, which may include positive effects of higher soil nutrients and a negative influence of higher soil salinity.     

Influence of Invasive Tree Kill Rates on Native and Invasive Plant Establishment in a Hawaiian Forest

Fire tree (Morella faya) has invaded extensive areas of wet and mesic forest on the Island of Hawai’i, forming nearly monospecific stands. Our objective was to identify a method of controlling M. faya, which would allow native plants to establish while minimizing establishment by invasive plants. Treatments (logging all trees, trees left standing but girdled, and incremental girdling over 20 months) were selected to kill M. faya stands at different rates, thereby creating different conditions for species establishment. Leaf litter was either removed or left in place; seeds and seedlings of three native pioneer species, three native forest species, and three alien invasive species were then added to determine their ability to establish. Native pioneer species established best in the log and girdle treatments, whereas seedling emergence of native forest species was higher in the girdle and incremental girdle treatments. Seedlings of invasive species emerged faster than the natives, but each of them responded differently to the stand treatments. Leaf litter reduced seedling emergence for all species, with small‐seeded species (<1 mg/seed) affected most under low light conditions. No single method eliminated all invaders, but girdling of M. faya provided suitable conditions for most native species. If combined with selective removal of the most disruptive alien species and native seed additions, girdling could be an effective general strategy for restoring native forests that have been overwhelmed by woody invaders.     

Distinctive vegetation communities are associated with the long‐lived conifer Agathis australis (New Zealand kauri,Araucariaceae) in New Zealand rainforests

The conifer Agathis australis (New Zealand kauri; Araucariaceae) has a significant influence on soil processes beneath its canopies, reducing soil pH, stalling nitrogen cycling processes, and sometimes forming podzols. Distinctive plant species assemblages have been anecdotally observed to occur in association with A. australis stands; however, the authenticity of these proposed associations has not been formally assessed. Owing to the effects of A. australis on its soil environment and the recorded vegetation patterns, we hypothesized that this species may act as a foundation species, playing a significant role in structuring plant community composition in its vicinity. To test this, we investigated the influence of proximity to A. australis on plant community composition at stand and individual tree scales. We also investigated compositional variation with distance from the conifer Dacrydium cupressinum (rimu, Podocarpaceae) within the same forests to directly compare A. australis effects to those of another large conifer. We examined changes in stand composition relative to the abundance of each of these conifers at two forests, and measured changes in environmental conditions and plant composition with increasing distance from mature individuals at one of the same and one other study site. The organic soil formed beneath A. australis individuals was highly acidic, with high levels of NH₄‐N, carbon and total nitrogen, but low levels of NO₃‐N. We recorded a difference in species composition in the vicinity of A. australis compared to forest without this species in the same environment, describing three groups of species: stress‐tolerant species dependent on the presence of A. australis within mature forest; those dependent on areas with A. australis absent; and those with distributions unaffected by A. australis presence. Such effects on the abiotic and biotic environments were not recorded in the vicinity of individuals of D. cupressinum. These results highlight the substantial effect that A. australis has in enhancing landscape‐scale habitat heterogeneity and influencing overall forest diversity.     

Canopy and age structures of some old sub-boreal Picea stands in British Columbia

Abstract. 14 old, unlogged, Picea-dominated stands in the moist cool Sub-Boreal Spruce biogeoclimatic subzone of central British Columbia, Canada, were sampled to describe canopy heterogeneity, regeneration patterns and tree population age structures. These stands are composed of Picea engelmannii × glauca hybrids, Abies lasiocarpa and lesser amounts of Pinus contorta and Populus tremuloides, and had survived 124–343 yr since the last stand-destroying wildfire. Canopy cover was patchy and highly variable (ranging from 30.5 % to 86.4 %) but was not significantly related to stand age. Vertical canopy structure was less variable, reflecting the shade-tolerance and live crown ratios (length of live canopy expressed relative to tree height) of component species: 18.8 % for Populus, 20.2 % for Pinus, 46.7 % for Picea and 51.4 % for Abies. Individual stands varied considerably in their population structures and in their stand development trajectories, yet some patterns are evident. Survivors of the initial post-disturbance cohort of trees took 51 to 118 yr (mean = 80, s.d. = 20) to establish. Some stands had all tree species present during stand initiation, while other stands indicated early successional roles for Populus and Pinus, or a late successional role for Abies. Abies recruitment, while often slow in the beginning, occurs uniformly throughout the history of most stands, reflecting the high shade-tolerance of this species. Picea is often recruited in high densities early in stand development, and then (after long periods of exclusion) may be displaced by Abies in some stands but maintains itself in others. Minor, single-tree disturbances (due to bark beetles, root rot, and windthrow) were important in accelerating the reinitiation of Picea in the understory. Results thus suggest that stands from this region can be self-perpetuating in the absence of fire. Yet, post-fire tree populations still clearly dominate these spruce-fir forests, for only the oldest stand had greater basal area in the replacement cohort than in the initial cohort.