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.     

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     

Linking dominant Hawaiian tree species to understory development in recovering pastures via impacts on soils and litter

Large areas of tropical forest have been cleared and planted with exotic grass species for use as cattle pasture. These often remain persistent grasslands after grazer removal, which is problematic for restoring native forest communities. It is often hoped that remnant and/or planted trees can jump‐start forest succession; however, there is little mechanistic information on how different canopy species affect community trajectories. To investigate this, I surveyed understory communities, exotic grass biomass, standing litter pools, and soil properties under two dominant canopy trees—Metrosideros polymorpha (‘ōhi‘a) and Acacia koa (koa)—in recovering Hawaiian forests. I then used structural equation models (SEMs) to elucidate direct and indirect effects of trees on native understory. Native understory communities developed under ‘ōhi‘a, which had larger standing litter pools, lower soil nitrogen, and lower exotic grass biomass than koa. This pattern was variable, potentially due to historical site differences and/or distance to intact forest. Koa, in contrast, showed little understory development. Instead, data suggest that increased soil nitrogen under koa leads to high grass biomass that stalls native recruitment. SEMs suggested that indirect effects of trees via litter and soils were as or more important than direct effects for determining native cover. It is suggested that diverse plantings which incorporate species that have high carbon to nitrogen ratios may help ameliorate the negative indirect effects of koa on natural understory regeneration.     

Invasive plant suppresses the growth of native tree seedlings by disrupting belowground mutualisms

The impact of exotic species on native organisms is widely acknowledged, but poorly understood. Very few studies have empirically investigated how invading plants may alter delicate ecological interactions among resident species in the invaded range. We present novel evidence that antifungal phytochemistry of the invasive plant, Alliaria petiolata, a European invader of North American forests, suppresses native plant growth by disrupting mutualistic associations between native canopy tree seedlings and belowground arbuscular mycorrhizal fungi. Our results elucidate an indirect mechanism by which invasive plants can impact native flora, and may help explain how this plant successfully invades relatively undisturbed forest habitat.     

Foliar production and decomposition rates in urban forests invaded by the exotic invasive shrub, <Emphasis Type="Italic">Lonicera maackii</Emphasis>

Exotic invasive shrubs can form dense monocultures in forest understories, which can have cascading effects on ecosystem structure and function. Amur honeysuckle, an exotic shrub that forms dense canopies in eastern forests, has the potential to alter plant community structure and ecosystem functions, such as primary production and decomposition. The goal of this study was to examine foliar productivity and leaf litter decomposition in forests invaded by Amur honeysuckle (Lonicera maackii) and to determine the extent to which the presence of this dominant exotic species may alter ecosystem function in these forests. We found that forests invaded by Amur honeysuckle had 16 times greater honeysuckle foliar biomass and 1.5 times lower total foliar biomass than forests of equivalent tree basal area, but having few honeysuckle shrubs. This suggests that productivity of native tree and shrub species may be reduced where honeysuckle density is high. Additionally, honeysuckle litter decayed four times faster and released nitrogen more rapidly than sugar maple litter, and sugar maple litter decayed 19% faster in forests invaded by Amur honeysuckle. These findings suggest that forests invaded by Amur honeysuckle may exhibit lower rates of organic matter accrual and less nitrogen retention in the forest floor. Since honeysuckle leaves develop in early spring before those of other shrubs or trees in the area, the rapid release of nitrogen from honeysuckle litter that we measured in early spring is timed to benefit this invasive species. The temporally coincident phenologies of nitrogen release during decomposition with the foliar growth needs of this shrub indicates that a potential positive feedback loop may exist between these processes that promotes continued growth and dominance of honeysuckle shrubs in these forested systems.     

Water flux and canopy conductance of natural versus planted forests in Patagonia, South America

In NW Patagonia, South America, natural shrublands and mixed forests of short Nothofagus antarctica (G. Forst.) Oerst. trees are currently being replaced by plantations with Pseudotsuga menziesii (Mirb) Franco. This land use change is controversial because the region is prone to drought, and replacement of native vegetation by planted forests may increase vegetation water use. The goal of this study was to examine the physiological differences, especially the response of water flux and canopy conductance to microclimate, that lead to greater water use by exotic trees compared to native trees. Meteorological variables and sapflow density of P. menziesii and four native woody species were measured in the growing season 2005–2006. Canopy conductance (gc) was estimated for both the exotic (monoculture) and native (multi-species) systems, including the individual contributions of each species of the native forest. Sapflow density, stand-level transpiration and gc were related to leaf-to-air vapor pressure difference (VPD). All native species had different magnitudes and diurnal patterns of sapflow density compared to P. menziesii, which could be explained by the different gc responses to VPD. Stomatal sensitivity to VPD suggested that all native species have a stronger stomatal control of leaf water potential and transpiration due to hydraulic limitations compared to P. menziesii. In conclusion, differences in water use between a P. menziesii plantation and a contiguous native mixed forest of similar basal area could be explained by different gc responses to VPD between species (higher sensitivity in the native species), in addition to particular characteristics of the native forest structure.     

Apparent competition: an impact of exotic shrub invasion on tree regeneration

Invasion of habitats by exotic shrubs is often associated with a decrease in the abundance of native species, particularly trees. This is typically interpreted as evidence for direct resource competition between the invader and native species. However, this may also reflect indirect impacts of the exotic shrubs through harboring high densities of seed predators––known as apparent competition. Here I present data from separate seed predation experiments conducted with two shrub species exotic to North America; Rosa multiflora, an invader of abandoned agricultural land, and Lonicera maackii, an invader of disturbed or secondary forest habitats. Both experiments showed significantly greater risks of seed predation for tree seeds located under shrub canopies when compared to open microhabitats within the same site. These results indicate the potential importance of indirect impacts of exotic species invasions on native biota in addition to the direct impacts that are typically the focus of research.     

Brown's Woods: An Early Gravel Pit Forest Restoration Project, Ontario, Canada

Few early examples of forest restoration projects are extant in the formerly forested parts of eastern North America. In this paper I present the history and status of an early forest restoration project in a denuded gravel pit in Ontario, Canada. The site was part of a deciduous forest until 1840, at which time forest clearing occurred. From 1874 to 1886 the site was exploited as a gravel pit. In 1887, under the direction of William Brown, it was planted with 14 species of coniferous and deciduous trees, of which 10 are still present. No soil preparation was carried out. The trees were pruned for 7 years, but in 1892 intensive maintenance ceased. In the 107 years since planting, the site has acquired some structural characteristics similar to the surrounding native deciduous forest, but it retains characteristics of an artificial community. Canopy cover has increased from 85% to over 95% since 1930 and is primarily Juglans nigra and Acer platan aides, Survivorship and current growth rates of native and exotic taxa have been similar. Recruitment patterns suggest that J. nigra and A. platanoides will dominate the canopy over the next century. The site is a useful example of the progress and problems created by attempts at forest restoration today.     

Alternative pathways of liana communities in the forests of northwestern Argentina

Liana dynamics in secondary and mature forests are well known in tropical areas dominated by native tree species. Outside the tropics and in secondary forests invaded by exotic species, knowledge is scarce. In this study, we compare liana communities between secondary and mature forests dominated by native species in a subtropical montane area of Sierra de San Javier, Tucuman, Argentina. Additionally, we evaluate changes of liana communities in secondary forests with increasing densities of Ligustrum lucidum and Morus alba, two of the most invasive exotic trees of the area. We surveyed liana species richness and density in three 30-year secondary patches, four 60-year secondary patches, and four mature patches dominated by native tree species, to analyze changes in liana communities with forest age. Within each patch, we sampled 10–25 20 × 20 m quadrats. Additionally, we surveyed liana density and species richness in secondary forest patches with different densities of L. lucidum and M. alba. In native-dominated forests, liana species richness increased and showed a tendency of increasing basal area from 30-year secondary forests to mature forests. Liana density was highly variable, and most of the species were shared between native-dominated secondary and mature forests. Liana density and species richness decreased with L. lucidum density, whereas in secondary forests highly dominated by M. alba, lianas increased in density. Overall, lianas followed different pathways influenced by native forest succession and exotic tree invasions.     

长白山阔叶红松林不同演替阶段林下红松幼苗能量与养分季节动态

为了解林下红松幼苗生长和养分存储季节动态,以长白山原始阔叶红松林(原始林)和次生杨桦林(次生林)林下2年生红松幼苗为对象,研究林下光合有效辐射(PAR)、幼苗生物量、非结构性碳水化合物(NSC)、全氮(N)和全磷(P)等指标的季节变化,分析两林分林下光照的季节动态及其差异对红松幼苗生长和养分积累的影响。结果表明: 原始林和次生林林下月PAR累积量季节变化都呈“双峰”型,夏季为郁闭期,两林分林下光线弱。春季和秋季为阔叶树无叶期,林下光照条件变好,且次生林林下光照明显好于原始林;原始林和次生林红松幼苗的生物量、NSC、全N和全P浓度的季节动态与林下光照的季节变化基本一致,在春季和秋季表现为显著增加,在夏季呈下降趋势。春季幼苗的淀粉浓度增加,夏季淀粉和可溶性糖浓度均逐渐降低,到8月达到最低值,秋季可溶性糖浓度显著升高。春季和秋季次生林林下幼苗的生物量和NSC浓度整体上均显著高于原始林,而夏季两林分差异不显著。因此,春季和秋季的林下光照条件差异是影响原始林和次生林中红松幼苗养分积累和生长更新差异的主要原因。     

Does forest restoration using taungya foster tree species diversity? The case of Afram Headwaters Forest Reserve in Ghana

The taungya agro‐forestry system is an under‐researched means of forest restoration that may result in high tree diversity. Within a forest reserve in Ghana, the forest core and its surrounding Teak‐ and Cedrela‐taungya on logged, cropped and burned land were mapped with ALOS satellite imagery. Native trees, seedlings and saplings were enumerated in 70 random, nested plots, equally divided between forest and taungya. The native tree regeneration was assessed by species richness (SR), Shannon‐Wiener Index (SWI), Shannon Evenness Index (SEI) and species density (SeD) for seedlings, saplings and trees separately and combined and subsequently correlated with canopy covers (CC) in taungya. As anticipated, the taungya diversity was lower than the forest diversity but higher than reported from nontaungya exotic plantations. In the forest, the diversity of native trees increased from seedlings through saplings to trees. The reverse was found in the taungya. Taungya seedling diversity was not significantly different from the forest, while the sapling and tree diversity were significantly lower. Weak correlations of CC with SR, SWI, SEI and SeD were found. Our results suggest the need for treatment to maintain the tree diversity beyond the seedling stage in the taungya.     

Breakdown of the species-area relationship in exotic but not in native forest patches

We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothesized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns.     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

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.     

Native,insect herbivore communities derive a significant proportion of their carbon from a widespread invader of forest understories

Research on natural enemies demonstrates the potential for exotic plants to be integrated into foodwebs through the activities of native herbivores. The quantitative importance of exotics as a food resource to herbivores is more difficult to ascertain. In addition, some widespread invaders appear to have minimal herbivore loads. Microstegium vimineum is one example. It is an annual, C₄ grass that invades forest understories and is widespread across the eastern US. Its invasion alters the structure and composition of forests. We sampled invertebrates in a tree-canopy gap and under canopy area, and used the unique carbon isotope value of M. vimineum to estimate the quantitative importance of the invader as a food resource relative to native plants. Seven of the eight invertebrate species derived on average >35% of their biomass carbon from M. vimineum, and some individuals representing both ‘chewing’ and ‘sucking’ feeding guilds derived their biomass carbon exclusively from M. vimineum. Our results show that M. vimineum can be a significant food resource for a multi-species, multi-guild, assemblage of native, invertebrate herbivores. Future work is required to assess whether M. vimineum is acquiring herbivores in other parts of its introduced range, and if so what might be the ecological consequences.     

Bird predation enhances tree seedling resistance to insect herbivores in contrasting forest habitats

According to the associational resistance hypothesis, neighbouring plants are expected to influence both the insect herbivore communities and their natural enemies. However, this has rarely been tested for the effects of canopy trees on herbivory of seedlings. One possible mechanism responsible for associational resistance is the indirect impact of natural enemies on insect herbivory, such as insectivorous birds. But it remains unclear to what extent such trophic cascades are influenced by the composition of plant associations (i.e. identity of ‘associated’ plants). Here, we compared the effect of bird exclusion on insect leaf damage for seedlings of three broadleaved tree species in three different forest habitats. Exclusion of insectivorous birds affected insect herbivory in a species-specific manner: leaf damage increased on Betula pendula seedlings whereas bird exclusion had no effect for two oaks (Quercus robur and Q. ilex). Forest habitat influenced both the extent of insect herbivory and the effect of bird exclusion. Broadleaved seedlings had lower overall leaf damage within pine plantations than within broadleaved stands, consistent with the resource concentration hypothesis. The indirect effect of bird exclusion on leaf damage was only significant in pine plantations, but not in exotic and native broadleaved woodlands. Our results support the enemies hypothesis, which predicts that the effects of insectivorous birds on insect herbivory on seedlings are greater beneath non-congeneric canopy trees. Although bird species richness and abundance were greater in broadleaved woodlands, birds were unable to regulate insect herbivory on seedlings in forests of more closely related tree species.     

<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.     

Crown structure and leaf area of the understorey species Prunus serotina

A detailed biometrical study of the exotic understorey invader Prunus serotina (Ehrh.) was carried out in a mixed coniferous forest stand in northern Belgium. Based on detailed destructive measurements of eight selected model trees, allometric relations of tree height, crown projected area, woody and leaf dry mass and leaf area on tree diameter at breast height (DBH) were derived. The scaling-up procedure from the tree to the stand level was done using the frequency distribution of DBH obtained at the selected experimental plot. The vertical and radial distributions of the tree foliage were estimated by the “cloud” technique. The vertical profile of leaf area showed a bimodal distribution pattern with maxima at heights of 4 and 6 m above the ground. The leaf area index (LAI) of the understorey Prunus serotina as estimated by the described up-scaling procedure (5.1) was significantly higher than the LAI (2.6) as measured by a plant canopy analyser and was also significantly higher than the LAI of the overstorey species Scots pine (1.5–3.0). The LAI of a neighbouring Rhododendron understorey reached only 1.25. This study emphasises the importance of an exotic understorey species in the total leaf area of mixed coniferous forests which might have important implications for the energy and mass exchanges of the entire forest.     

Flowering phenology and reproductive success of native shrub Berberis darwinii Hook. along different light environments

Studies evaluating flowering phenology and reproductive success are necessary when we want to direct a domestication project in a species with a potential productive value. We studied flowering phenology and reproductive success of Berberis darwinii growing under different light conditions in its native distribution area in the Andean Patagonian forests of Argentina. We test the hypothesis that plants grown under conditions of high-light availability exhibit advanced phenology and higher reproductive success than those grown under conditions of lower light availability. Phenology and reproductive success were determined in three contrasting light conditions at two forest sites, which were, canopy, gap and forest edge. Plants did not bloom under the forest canopy. Flowering and fruiting period lengths were similar in both sites and light conditions of gap and forest edge during spring and summer. Although gap plants had more racemes per shoot, racemes of edge plants had more flowers, fruits and a higher proportion of flowers producing ripe fruit. We show that B. darwinii reproduction studied in the Andean Patagonian forests is conditioned by the canopy openness. Regarding reproductive success, edge plants invest less resources in flower production than gap plants to have similar fruit production.