Ecological Restoration of Native Forest at Aratiatia, North Island, New Zealand

Successional pathways in native forest, planted 15–33 years ago on reconstructed surfaces to restore aesthetic values destroyed by hydro‐electric dam construction at Aratiatia, central North Island, New Zealand, were compared with those on similar surfaces left unplanted. Only native species were planted. Classification identified three canopy communities and several ground layer communities with significant inter‐stratum relationships: Pittosporum tenuifolium‐Sophora tetraptera short forest with ground layers dominated by litter; P. tenuifolium‐Kunzea ericoides short forest over adventive grasses on planted sites; and adventive Cytisus scoparius shrubland over grasses on unplanted sites. Planted communities mirror young secondary forests on intact substrates in the district, but have lower density and similar or higher basal area than such forests elsewhere. Established seedlings of seven planted canopy trees, mostly early successional bird‐dispersed species, are reasonably widespread in floristically rich Pittosporum‐Sophora forest. Seedlings of only two species are widespread in floristically poor Pittosporum‐Kunzea forest, and none on unplanted sites. This first large‐scale attempt at ecological restoration in New Zealand, by mass planting of new surfaces with early successional native woody species, has created aesthetically‐pleasing stands of indigenous forest on sites which would otherwise remain in relatively stable adventive shrubland communities for the foreseeable future. Only continued monitoring will show whether further management is necessary and whether natural processes are operating at a level sufficient to ensure that artificially initiated successions will continue along more or less natural pathways.     

Floristic changes over 30 years in a Canterbury Plains kānuka forest remnant, and comparison with adjacent vegetation types

The Canterbury Plains have lost most of their pre-Polynesian indigenous vegetation, primarily forest and shrubland. One of the few remaining areas is the 2.3 ha Eyrewell Scientific Reserve which consists mostly of low kānuka (Kunzea ericoides) forest and a small area of grassland. We assessed the Reserve vegetation using a combination of plots and transect surveys at different times of the year between 2001–2003. For comparison with the Reserve vegetation we also assessed plots in an adjacent grazed kānuka remnant, adjacent cultivated pasture and Eyrewell Forest, a pine plantation. Our study of the Eyrewell Reserve in 2001–2003 found that since an assessment of the Reserve in 1972, 28 indigenous species were no longer present but 14 indigenous species and 48 adventive species were newly recorded. The dramatic invasion of the Reserve is illustrated by the fact that 60% of the 118 species recorded in 2003 were adventives compared to 34% in 1972. Despite this invasion and the loss of indigenous species, Reserve plots still have more than twice as many species as plots in the adjacent pine plantation. The Reserve also included several species of high conservation value such as the “Chronically Threatened” Leptinella serrulata, and the “At Risk” Aciphylla subflabellata, Coprosma intertexta and Pterostylis tristis. Plots in the Reserve grassland and adjacent pasture had the lowest percentage of indigenous species of all habitat types, with the pasture plots having no indigenous vascular plant species. In contrast the understorey of the old pine stands had the highest percentage of indigenous species of any of the habitats and in places was dominated by kānuka up to 4 m tall, indicating that these plantations also have conservation value. Eyrewell Reserve and the few other remaining kānuka remnants in the Canterbury Plains represent an important pool of indigenous species for conservation. Options for the future management of the Reserve are discussed.     

Birds and small mammals in kanuka (

Native kanuka (Kunzea ericoides) and adventive gorse (Ulex europaeus) stands aged 10-14 years, and not grazed by domestic stock, were studied near Nelson, New Zealand. The aim was to determine their use by introduced small mammals, and native and adventive birds, and the effects of these animals on seed rain and seedling dynamics as factors influencing vegetation succession. Seed traps were established where they could catch only bird-dispersed or wind-blown seed, and seedling emergence and growth were monitored. Bird abundance was estimated by five-minute bird counts, and small mammal abundance by trapping. The summed frequencies of all birds, and those likely to disperse seeds, were similar in kanuka and gorse. The endemic native bird species, bellbirds (Anthornis melanura) which are omnivorous, brown creepers (Mohoua novaeseelandiae) and grey warblers (Gerygone igata) which are insectivorous, were more frequent in kanuka than in gorse, while fantails (Rhipidura fuliginosa) were equally frequent in both stands. Non-endemic silvereyes (Zosterops lateralis) which are omnivorous were the most abundant seed-dispersing species, and they were significantly more frequent in gorse, as were adventive California quail (Callipepla californica) which are granivorous. Other small adventive granivores and omnivores were either more frequent in kanuka or gorse, or equally common in both stands. Ship rats (Rattus rattus) and possums (Trichosurus vulpecula) were in low numbers throughout. Mice (Mus domesticus) were more frequent in the gorse, probably because of the shelter offered by the dry gorse litter, and food supply, e.g. gorse seed. More seeds of native, fleshy-fruited shrubs fell in the kanuka, largely those of Coprosma spp. and Cyathodes juniperina, which grow in the kanuka. Seed species richness was similar in kanuka and gorse. In both cases, the seed rain appeared more influenced by the local seed source than by the different bird communities. In both kanuka and gorse, the relationship between seed rain and seedling numbers was close only for the most common fleshy-fruited species. Seedling emergence and survival was greater in gorse because of openings in the canopy, and the lower density of the introduced hares and rabbits. Overall, the different morphology and structure of the adventive gorse probably have the greatest influence in seedling dynamics, and ultimately on vegetation succession.     

Bone-seed (

Bone-seed, Chrysanthemoides monilifera ssp. monilifera (L.), is an environmental weed of coastal vegetation communities scattered throughout New Zealand. To assess the long-term implications for native forest regeneration in sites where bone-seed is present, we selected four study sites around Wellington, New Zealand, where bone-seed was abundant. We compared seed bank composition in bone-seed-invaded sites with nearby native forest patches, and monitored bone-seed and native seedling recruitment with and without control of mature bone-seed plants. We also tested the potential effects of fire on bone-seed recruitment in these communities by heating seeds prior to germination. Bone-seed, gorse (Ulex europaeus), and native species emerged from seed bank samples taken from bone-seed-invaded sites, but only native species and (less) gorse emerged from seed bank samples taken from native forest patches. Gorse germination was strongly promoted by heat but bone-seed germination was less affected by heat. Bone-seed seedling abundance increased dramatically following canopy removal, whereas native seedling abundance decreased dramatically. This suggests that disturbance of any form is likely to favour recruitment of bone-seed (and gorse) over native species, although in the long term, native seedlings can establish beneath the canopy of mature bone-seed plants. It is not yet known if, in the absence of further disturbance, regenerating native vegetation will eventually replace bone-seed in New Zealand.     

The role of blackbirds (

The naturalised European blackbird (Turdus merula) is the most widely distributed avian seed disperser in New Zealand. Together with the native silvereye (Zosterops lateralis) they are the major seed dispersers over large areas of New Zealand. I review the international literature on aspects of the ecology and behaviour of blackbirds relevant to their potential for dispersing weeds in New Zealand. Blackbirds eat a wide range of native and exotic fruit including many naturalised species. Their habitat preferences and behaviour result in germinable seeds being deposited in a range of sites, particularly in shrubby habitats, where seedling establishment is likely. Most seeds will be deposited within 50 m but some may be carried a kilometre or more to develop new invasive loci. Blackbirds therefore probably make a major contribution to the development of novel plant communities of naturalised woody weeds. These communities provide fruit more suited to non- endemic native birds and naturalised birds, than to endemic birds. The relative contribution of blackbirds and silvereyes to seed dispersal of native and exotic species requires investigation. The outcome may suggest potential for managing blackbirds as a vector of weed invasions.     

Succession of Exotic and Native Species Assemblages within Restored Floodplain Forests: A Test of the Parallel Dynamics Hypothesis

Exotic plants pose a threat to restoration success in post‐agricultural bottomlands, but little information exists on their dynamics during succession of actively restored sites. We hypothesized that exotic assemblages would establish during succession and that their compositional trends during succession time would mirror those published for native species in other systems, with an early peak in herbaceous richness followed by a decline as woody species establish. In the summer of 2008, we sampled 16 sites across an 18‐year chronosequence of restored forests, with an additional four mature forest stands for comparison, within the Cypress Creek NWR, Illinois, U.S.A. We identified all vascular plant species and quantified canopy openness at three canopy strata, and soil texture and chemistry. Trends in exotic assemblages were significantly correlated with canopy openness at all strata. Richness of exotic and native herbaceous species was related to stand age and consistent with a Weibull regression model. Native and exotic herbaceous cover followed an exponential decay model. Woody native richness over time conformed to a logistic model; woody exotics exhibited no relationship with stand age and were present in sites of all ages. Our results indicate that although their rates of decline differ, herbaceous exotics and natives exhibit similar successional dynamics; therefore, herbaceous exotics may not pose a lasting threat to restoration success in reforested floodplains. Woody exotics can establish across a range of successional stages and persist under closed canopy conditions. Bottomland restorations are vulnerable to the invasion and expansion of exotic plant species even after canopy closure.     

Removal of cattle accelerates tropical dry forest succession in Northwestern Mexico

Domestic livestock influence patterns of secondary succession across forest ecosystems. However, the effects of cattle on the regeneration of tropical dry forests (TDF) in Mexico are poorly understood, largely because it is difficult to locate forests that are not grazed by cattle or other livestock. We describe changes in forest composition and structure along a successional chronosequence of TDF stands with and without cattle (chronic grazing or exclusion from grazing for ~ 8 year). Forest stands were grouped into five successional stages, ranging from recently abandoned to mature forest, for a total of 2.7 ha of the sampled area. The absence of cattle increased woody plant (tree and shrub) density and species richness, particularly in mid-successional and mature forest stands. Species diversity and evenness were generally greater in sites where cattle were removed and cattle grazing in early successional stands reduced establishment and/or recruitment of new individuals and species. Removal of cattle from forest stands undergoing succession appears to facilitate a progressive and non-linear change of forest structure and compositional attributes associated with rapid recovery, while cattle browsing acts as a chronic disturbance factor that compromises the resilience and structural and functional integrity of the TDF in northwestern Mexico. These results are important for the conservation, management, and restoration of Neotropical dry forests.     

Is patience a virtue? Succession, light, and the death of invasive glossy buckthorn (Frangula alnus)

Non-native, invasive glossy buckthorn (Frangula alnus P. Mill.) threatens North American forests by inhibiting tree regeneration. While glossy buckthorn commonly invades younger, developing forests, we hypothesized that this species is competitively excluded as secondary succession proceeds. Specifically, we tested the hypothesis that glossy buckthorn mortality in New Hampshire forests is associated with low levels of photosynthetically active radiation (PAR), low levels of nutrients in the soil, and greater abundance of shade tolerant tree species. Twenty-six living and 26 dead glossy buckthorn individuals were randomly selected on three, mid-successional study sites in southeastern New Hampshire, USA. Shrub ages, indicated by wood ring counts, showed that both living and dead shrubs were members of the same cohort, roughly 16–18 years of age. Living and dead shrubs were compared as to (1) tree basal area in the surrounding forest (estimated by plot sampling), (2) light intensity in the vicinity of the buckthorn crowns (measured by Sunfleck Ceptometer), (3) canopy openness and % total transmitted PAR (estimated by hemispheric photography), and (4) concentrations of available Ca, P, Mg, and K in the B horizon. Relative to live buckthorn shrubs, dead individuals were associated with greater basal area of shade tolerant species, decreased PAR in the vicinity of shrub crown, and decreased soil concentrations of Ca, P, and Mg. The results are consistent with the hypothesis that competition for light and perhaps nutrients limits the ability of glossy buckthorn to persist in late successional stands. If these relationships are causal, active removal of glossy buckthorn might not be required to reduce its abundance in late successional stands.     

Insect assemblages in a native (kanuka –

In New Zealand, the European shrub gorse (Ulex europaeus) is becoming the initial post-disturbance shrub, replacing the native myrtaceous manuka (Leptospermum scoparium) and kanuka (Kunzea ericoides) scrub in this role. Change in the dominant vegetation is likely to affect the native invertebrate community. To quantify these changes, we compared the assemblages of four selected insect taxa (Coleoptera, Lepidoptera, and two groups of Diptera, the Tachinidae and the fungus gnats, represented in New Zealand by the families Ditomyiidae, Keroplatidae and Mycetophilidae) in neighboring stands of kanuka and gorse using Malaise and pitfall traps set during December. We sorted 34 387 specimens into 564 recognisable taxonomic units. Ordinations showed that insect assemblages associated with each habitat were distinct for all four insect groups. The gorse habitat was species rich compared with kanuka for tachinids, fungus gnats and Malaise-trapped beetles, and both habitats contained few adventive species. Many species were unique to each habitat. Some species loss might occur if the kanuka-dominated community continues to be replaced by gorse, but gorse is nevertheless considered to be valuable as a habitat for native invertebrates.     

Spontaneous Establishment of Woody Plants in Central European Derelict Sites and their Potential for Reclamation

The performance of woody plants was analyzed in 15 successional seres starting at bare ground in central European manmade habitats. The total cover of woody species after 10 years of succession was significantly related neither to initial soil moisture nor to nitrogen (expressed using Ellenberg indicator values). But the comparison of seres indicates that establishment of woody plants was easier under moderate environmental conditions and retarded in extreme habitats (dry, nutrient-poor, or acid). The arrival of the first woody plants was delayed in dry sites. No significant differences were found between primary and secondary seres, either with respect to the total cover of woody plants reached after 10 years of succession or considering the time of their arrival. In total, 24 woody species (10 shrubs and 14 trees) appeared in the series investigated. Their successional performance (in terms of the number of seres in which the species occurred and maximum cover reached in any sere) was not related to species traits (life strategy, type of mycorrhizae, mode of dispersal, diaspore weight), except for the regeneration strategy, species with seasonal regeneration by seeds were capable of creating higher cover. Betula pendula (European birch) was the most successful species in spontaneous succession, especially on moist sites. Practical suggestions for the management of particular habitats (sites disturbed by mining, sites reclaimed after acid rain deforestation, urban sites, abandoned fields) are provided regarding the establishment of woody plants.     

Leaf palatability and decomposability increase during a 200‐year‐old post‐cultural woody succession in New Zealand

Question: How do aggregate trait values and functional diversity of leaf traits linked to palatability and decomposability change during a woody post‐cultural succession spanning 200 years? Location: Coastal Marlborough, South Island, New Zealand. Methods: The biomass of all woody species was determined in 32 20‐m × 20‐m plots ranging from 10 to 200 years in time since last disturbance. Species abundances were combined with data on leaf nutrient, secondary metabolite and structural carbohydrate content to calculate biomass‐weighted trait means (i.e. aggregate trait values) and functional diversity index values for each plot. Aggregate trait values and functional diversity were regressed on successional age and total live above‐ground carbon content to examine functional shifts with succession and one consequence of succession – increasing above‐ground carbon. Results: Almost all significant regressions between aggregate trait values and both successional age and above‐ground carbon indicated a shift toward increased leaf palatability and decomposability during succession. The relationships were all non‐linear, with aggregate trait value shifts occurring relatively early in the successional sequence. There was weak evidence for an increase in functional richness with succession, but this was a secondary effect relative to the shifts in aggregate trait values. Conclusions: These results are in direct contrast with studies of the early stages of herbaceous post‐cultural successions from grasslands to shrublands, which have found a shift towards communities of decreasing palatability and decomposability, suggesting that functional shifts in woody succession may be fundamentally different.     

Population structure and succession in temperate forests of southwestern Japan

In the temperate forests of southwestern Japan, the population density of woody plants in the community increases in the early stage of secondary succession, reaches a peak in the old oak-chestnut forest, and decreases towards the climax beech forest. The species richness and diversity of woody plants also show a trend similar to that of the population density. The canopy-tree population decreases in the course of the succession while the basal area increases, showing a self-thinning process. The species richness, diversity and population density of herbaceous plants are much influenced by the dominance of the bamboo, Sasa palmata. The life-history traits of trees, lower trees and shrubs are discussed in relation to their shoot system, reproductive pattern and successional processes.     

Restoration of Tropical Moist Forests on Bauxite-Mined Lands in the Brazilian Amazon

We evaluated forest structure and composition in 9- to 13-year-old stands established on a bauxite-mined site at Trombetas (Pará), Brazil, using four different reforestation techniques following initial site preparation and topsoil replacement. These techniques included reliance on natural forest regeneration, mixed commercial species plantings of mostly exotic timber trees, direct seeding with mostly native early successional tree species, and mixed native species plantings of more than 70 tree species (the current operational restoration treatment at this site). Replicated fixed-radius plots in each treatment and in undisturbed primary forest were used to quantify the canopy and understory structure and the abundance and diversity of all vascular plant species. Treatment comparisons considered regeneration density, species richness and diversity for all floristic categories, and, for trees and shrubs, the relative contribution of initial planting and subsequent regeneration from soil seed banks and seed inputs from nearby primary forests. With the possible exception of the stands of mixed commercial species, which were superior to all others in terms of tree basal-area development but relatively poor in species richness, all treatments were structurally and floristically diverse, with a high probability of long-term restoration success. Of these, the mixed native species plantings appeared to be at least risk of arrested succession due to the dominance of a broader range of tree species of different successional stages or expected life spans. In all treatments, several locally important families of primary forest trees (Annonaceae, Chrysobalanaceae, Lauraceae, Palmae and Sapotaceae) were markedly underrepresented due to a combination of poor survival of initial plantings and limitations on seed dispersal from the surrounding primary forest.     

Distribution and abundance of endemic coelostomidiid scale insects (Hemiptera: Coelostomidiidae) in Auckland forests, New Zealand

Sap-feeding insects can excrete considerable quantities of sugar-rich honeydew. In New Zealand, South Island beech (Nothofagus spp.) forests are shaped by the extensive honeydew resource produced by two endemic coelostomidiid species (Ultracoelostoma assimile and U.?brittini) and geckos on northern islands are known to feed on the honeydew of a third endemic coelostomidiid, Coelostomidia zealandica. There are six other endemic coelostomidiid species in New Zealand that utilise a range of plant hosts but the ecological role of these species is poorly understood. A survey of mainland forests in the Auckland Ecological Region was conducted in February?April 2006 to investigate the distribution and abundance of coelostomidiids in this area. Three coelostomidiid species were detected in the survey (C.?zealandica, C.?pilosa and C.?wairoensis) and five new host?scale insect associations were identified. C.?zealandica was uncommon, C.?pilosa was widespread in broadleaved?podocarp forest but only formed light infestations, and C.?wairoensis was present in all teatree stands examined, often forming heavy infestations on k?nuka (Kunzea ericoides). Infested k?nuka trees had sooty moulds growing on them and exotic wasps were regularly seen feeding on C.?wairoensis honeydew. The extent and intensity of C. wairoensis infestation on k?nuka suggests it will have community-level impacts.     

Underplanting degraded exotic Pinus with indigenous conifers assists forest restoration

We propose that nonharvest plantations could provide important opportunities for restoration of indigenous forest cover and related ecosystem services. We assessed the relative performance of three Podocarpaceae (podocarps) species planted into a degraded Ponderosa Pine (Pinus ponderosa) plantation, central North Island, New Zealand. We hypothesised that the degraded pine plantation overstorey could provide suitable conditions for the development of a podocarp‐dominated forest structure within ca. 50 years of underplanting and that podocarp growth would differ depending on the species suitability to the site. Rimu (Dacrydium cupressinum) significantly outperformed both Totara (Podocarpus totara) and Kahikatea (Dacrycarpus dacrydioides) in height and diameter growth. Rimu was now the structurally dominant tree where it occurred rather than pine. Per annum scaled carbon storage within Rimu stands was significantly greater than the Totara, Kahikatea or Pine stands. All podocarp species had attained a greater stand density compared to the pine overstorey. Possible reasons for the differing podocarp growth performance include different light requirements, response to soil nutrients, elevational distributions and frost susceptibility. There were significant differences in understorey species richness among the different stands of podocarp species. Underplanting accelerated successional development by incorporating late‐successional indigenous canopy dominants within the forest succession and overcame limitations imposed on forest succession at the site from its isolation from indigenous forest tree seed sources.     

Plant species richness under

Exotic pine plantations constitute a significant landscape feature in the North Island of New Zealand but their conservation value for native plant species is not often documented. Pine stem density, height and basal area of nine plantations of Pinus radiata ranging in age from 6 to 67 years in Kinleith Forest was determined. Pines reached heights of 60 m, and stand basal areas up to 183 ± 14 m(2)ha(-1). The abundance of woody shrubs, tree ferns and ground ferns was assessed in each stand. Understorey composition of shrubs and ferns was reflected on the first two axes of DCA ordinations and correlated with the age of the pines. Adventive shrubs predominated in stands < 20 years old. Light-demanding native shrubs with bird dispersed fruits predominated in older stands, with more shade-tolerant species in the oldest site. Species richness increased rapidly in the first 11 years, but thereafter more slowly. Twelve native shrub species and 22 ferns were recorded from the most diverse stands. Richness and species composition were related to stand age, and probably also to topographical heterogeneity and aspect. Tree ferns reached densities of 2000—2500 ha(-1) and basal areas of 20—30 m(2)ha(-1) in the older stands. Initially the tree fern population was strongly dominated by Dicksonia squarrosa, which comprised 84% of individuals overall. Five species were present by 29 years. The faster growing Cynthea medullaris and C. smithii achieved greater heights than the Dicksonia spp., and their relative biomass was greatest in the oldest stands.     

Interactions between vegetation development and island formation in the Alpine river Tagliamento

Abstract. Early-successional stages of woody vegetation on gravel bars were studied in an island-braided section of the River Tagliamento in northeastern Italy. We mapped landscape-level changes in the study area (125 ha) by GIS-based analysis of aerial photographs for two time periods (1984–1986, 1986–1991); we surveyed island vegetation, and estimated island age by tree ring analysis. The study area experienced considerable changes between 1984 and 1991 due to at least two major floods in 1987 and 1990. The development of woody vegetation on bars follows three distinct phases: (1) gravel bars plus large woody debris (LWD), (2) pioneer islands, and (3) established islands. Established islands have sections dominated by shrubs of Salix elaeagnos, S. purpurea, S. daphnoides and S. triandra, and tree-dominated sections with Populus nigra, Salix alba and Alnus incana. Large woody debris seems to play a key role for plant colonization on gravel bars. The succession from bars to established islands took about 10 - 20 yr, and the probability of an island being washed away decreased with island age. Erosion produced new LWD which again initiated successional processes in the active zone of the river. Most species were already present in the early-successional stages, although the number of species increased with island development. Established islands were characterized by a distinctive species composition, including an assemblage of species less tolerant of inundation. The results are discussed within the framework of island dynamics and its significance for restoration of early-successional habitats in more regulated rivers.     

Evaluating nurse plants for restoring native woody species to degraded subtropical woodlands

Harsh habitats dominated by invasive species are difficult to restore. Invasive grasses in arid environments slow succession toward more desired composition, yet grass removal exacerbates high light and temperature, making the use of “nurse plants” an appealing strategy. In this study of degraded subtropical woodlands dominated by alien grasses in Hawai'i, we evaluated whether individuals of two native (Dodonaea viscosa, Leptocophylla tameiameia) and one non‐native (Morella faya) woody species (1) act as natural nodes of recruitment for native woody species and (2) can be used to enhance survivorship of outplanted native woody species. To address these questions, we quantified the presence and persistence of seedlings naturally recruiting beneath adult nurse shrubs and compared survival and growth of experimentally outplanted seedlings of seven native woody species under the nurse species compared to intact and cleared alien‐grass plots. We found that the two native nurse shrubs recruit their own offspring, but do not act as establishment nodes for other species. Morella faya recruited even fewer seedlings than native shrubs. Thus, outplanting will be necessary to increase abundance and diversity of native woody species. Outplant survival was the highest under shrubs compared to away from them with few differences between nurse species. The worst habitat for native seedling survival and growth was within the unmanaged invasive grass matrix. Although the two native nurse species did not differentially affect outplant survival, D. viscosa is the most widespread and easily propagated and is thus more likely to be useful as an initial nurse species. The outplanted species showed variable responses to nurse habitats that we attribute to resource requirements resulting from their typical successional stage and nitrogen fixation capability.     

Canopy manipulation as a tool for restoring mature forest conifers under an early‐successional angiosperm canopy

Low‐light environments in early‐successional forests that have established after abandonment of farming often restrict the establishment of later successional species resulting in an arrested succession. This 6‐year study tested the potential of different canopy manipulations to facilitate the establishment of a light‐demanding canopy tree species, tōtara (Podocarpus totara), within a regenerating kānuka (Kunzea robusta) stand. Results highlighted the effectiveness of artificial gaps over other methods (ring‐barking and edge‐planting) in accelerating the growth of planted tōtara. Seedlings under gaps grew consistently taller and faster over time indicative of an improved understorey light environment. Ring‐barking did not have a significant effect on tōtara growth because only a portion of the treated trees died, and after 6 years dead trees remained standing with intact branches resulting in insignificant increases in light transmission. At the forest edge sites, tōtara growth was highly variable. Although some seedlings grew as tall as in the gaps, others did not. Survival was also lower in the edge sites than in other treatments, which was likely due to enhanced herbivory from ungulates which impacted some plants at these sites. Gap creation is likely to be an important tool for restoring late‐successional canopy species in regenerating stands both through providing ideal sites for the growth of light‐demanding species such as tōtara and through natural establishment of other future canopy trees into the gaps.     

Multiple successional pathways of boreal forest stands in central Canada

Predicting forest composition change through time is a key challenge in forest management. While multiple successional pathways are theorized for boreal forests, empirical evidence is lacking, largely because succession has been inferred from chronosequence and dendrochronological methods. We tested the hypotheses that stands of compositionally similar overstory may follow multiple successional pathways depending on time since last stand‐replacing fire (TSF), edaphic conditions, and presence of intermediate disturbances. We used repeated measurements from combining sequential aerial photography and ground surveys for 361 boreal stands in central Canada. Stands were measured in 8–15 yr intervals over a ~ 60 yr period, covering a wide range of initial stand conditions. Multinomial logistic regression was used to analyze stand type transitions. With increasing TSF, stands dominated by shade‐intolerant Pinus banksiana, Populus sp., and Betula papyrifera demonstrated multiple pathways to stands dominated by shade‐tolerant Picea sp., Abies balsamea, and Thuja occidentalis. Their pathways seemed largely explained by neighborhood effects. Succession of stands dominated by shade‐tolerant species, with an exception of stands dominated by Picea sp., was not related to TSF, but rather dependent on edaphic conditions and presence of intermediate disturbances. Varying edaphic conditions caused divergent pathways with resource limited sites being dominated by nutrient‐poor tolerant species, and richer sites permitting invasion of early successional species and promoting species mixtures during succession. Intermediate disturbances promoted deciduous persistence and species diversity in A. balsamea and mixed‐conifer stands, but no evidence was detected to support “disturbance accelerated succession”. Our results demonstrate that in the prolonged absence of stand‐replacing disturbance boreal forest stands undergo multiple succession pathways. These pathways are regulated by neighborhood effects, resource availability, and presence of intermediate disturbance, but the relative importance of these regulators depends on initial stand type. The observed divergence of successional pathways supports the resource‐ratio hypothesis of plant succession.