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

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

The Role of Cloud Combing and Shading by Isolated Trees in the Succession from Maquis to Rain Forest in New Caledonia1

This study examined the role of shading and cloud combing of moisture by scattered trees of the emergent conifer Araucaria laubenfelsii (Corbass.) in montane shrubland‐maquis at Mont Do, New Caledonia, in facilitating the succession from shrubland to rain forest. Water collection experiments showed that these trees combed significant amounts of water from low clouds on days when no rainfall was recorded and deposited this moisture on the ground beneath the tree canopy. Analysis of photosystem II function in A. laubenfelsii and five other plant species using fluorometry revealed much lower photosystem stress in plants beneath scattered A. laubenfelsii than for individuals exposed to full sunlight in the open maquis. Transition matrix analyses of vegetation change based on “the most likely recruit to succeed” indicated that the transition from maquis to forest was markedly faster when emergent trees of A. laubenfelsii acted as nuclei for forest species invasion of die maquis. On the basis of these lines of evidence, it is argued that increased moisture and shading supplied to the area directly below the crown of isolated A. laubenfelsii trees in the maquis facilitates the establishment of both conifer seedlings and other rain forest tree and shrub species. In the absence of fire, rain forest can reestablish through spread in two ways: first, by expansion from remnant patches, and second, from coalescence of small rain forest patches formed around individual trees of A. laubenfelsii.     

Dynamics of forests with Araucariaceae in the western Pacific

Abstract. Several species of Araucaria and Agathis (Araucariaceae) occur as canopy emergents in rain forests of the western pacific region, often representing major components of total stand biomass. New data from permanent forest plots (and other published work) for three species (Araucaria hunsteinii from New Guinea, A. laubenfelsii from New Caledonia, and Agathis australis from New Zealand) are used to test the validity of the temporal stand replacement model proposed by Ogden (1985) and Ogden & Stewart (1995) to explain the structural and compositional properties of New Zealand rain forests containing the conifer Agathis australis. Here we propose the model as a general one which explains the stand dynamics of rain forests with Araucariaceae across a range of sites and species in the western Pacific. Forest stands representing putative stages in the model were examined for changes through time in species recruitment, growth and survivorship, and stand richness, density and basal area. Support for the model was found on the basis of: 1. Evidence for a phase of massive conifer recruitment following landscape-scale disturbances (e.g. by fire at the Huapai site, New Zealand for Agathis australis); 2. Increasing species richness of angiosperm trees in the pole stage of forest stand development (i.e. as the initial cohort of conifers reach tree size; >10 cm DBH); 3. A high turnover rate for angiosperms (<100 yr), and low turnover for conifers (≥ 100 yr) in the pole stage, but similar turnover rates for both components (50–100 yr) as forests enter the mature to senescent phase for the initial conifer cohort; 4. Very low rates of recruitment for conifers within mature stands, and projected forest compositions which show increasing dominance by angiosperm tree species; 5. A low probability of conifer recruitment in large canopy gaps created by conifer tree falls during the initial cohort senescent phase, which could produce a second generation low density stand in the absence of landscape scale disturbance; 6. Evidence that each of the three species examined required open canopy conditions (canopy openness > 10 %) for successful recruitment. The evidence presented here supports the temporal stand replacement model, but more long-term supporting data are needed, especially for the phase immediately following landscape level disturbance.     

Disturbance processes and spatial patterns of two emergent conifers in New Caledonia

Abstract Araucaria laubenfelsii Corbasson and Araucaria montana Brongn et Gris are emergent conifers in maquis and forest communities which are subjected to a combination of fire and cyclonic disturbances. Both species are able to survive fire once stems reach a critical size, but most seedlings and saplings are killed. Both species were found to be clumped at most spatial scales for both saplings and trees in maquis, probably resulting from a combination of patchy fire and a limited ability to disperse seeds. Comparisons of burned and unburned A. montana sites suggest that clumping increases only slightly after fire. The degree of clumping in the A. laubenfelsii sites, suggested that fire disturbances may occur relatively frequently in the region. Cyclonic disturbances are infrequent but may result in blow‐down of large individuals within both maquis and forest. All of the individuals blown‐down during the study had been previously fire‐scarred. Around tree blow‐downs, seedling and sapling densities can be high. This likely reflects both the low dispersability of Araucaria seeds and enhanced moisture from the shading of the adult, when it was alive. Disturbance by fire and wind play an important role in the regeneration dynamics and spatial pattern of these species in maquis in New Caledonia.     

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

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

Humans,fire and landscape pattern: understanding a maquis‐forest complex,Mont Do,New Caledonia,using a spatial ‘state‐and‐transition’ model

Abstract Aim The research explores how changes in disturbance regime resulting from human settlement may affect landscape structure. A spatially explicit grid‐based simulation model is used to explore the interplay between humans, fire regime and landscape composition. Location The study site for this research is the botanical reserve at Mont Do, New Caledonia. The endemic conifer Araucaria laubenfelsii (Araucariaceae) forms a key component of the landscape at Mont Do. This species is unusual in that it is found scattered as an emergent in maquis and as a canopy species in adjacent rain forest patches. Although now dominated by a low maquis, prior to human settlement of New Caledonia, montane landscapes such as Mont Do are likely to have been heavily forested. Methods A spatially explicit simulation model, based on field data and palaeoecological information, was used to explore interactions between disturbance regime and the landscape. The model is described briefly here and more fully in Perry & Enright (2002) Ecological Modelling, 152 , 279. Results The model suggests that human‐influenced changes to the fire regime at Mont Do have been important in generating the current landscape structure. The origin and maintenance of forest landscapes and maquis‐forest mosaic landscapes are considered in the context of alternative stable states. Strong feedback loops between fire size and landscape composition, mediated at the smaller scale by other similar mechanisms, are capable of driving landscape change. The utility of a spatial state and transition modelling approach is demonstrated. Main conclusions The current landscape pattern on Mont Do is likely the result of changes to the fire regime occurring since human settlement. The specific mechanisms for this change outlined here may occur in a number of other similar systems. Understanding the origin and persistence of these ‘fire landscapes’ in New Caledonia and in the south‐west Pacific in general is crucial for their effective management.     

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

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

Threats to the conifer species found on New Caledonia’s ultramafic massifs and proposals for urgently needed measures to improve their protection

Forty-one endemic conifer species occur on New Caledonia’s ultramafic substrates (known locally as “terrains miniers”), the source of nickel ore deposits being exploited at a rapidly increasing rate. Impacts of the removal of native vegetation and its destruction by the deposition of mine wastes are compounded by fire, which has dramatically reduced and fragmented the original cover. A new threat evaluation of these conifers, now being incorporated into the IUCN Red List, is presented. A conservation action plan is proposed to ensure their long term survival. Four species are classified as Critically Endangered (CR), 13 Endangered (EN), 6 Vulnerable (VU), 7 Nearly Threatened (NT) and 11 Least Concern. Wetland habitats contain two threatened species (i.e., CR, EN or VU), all inadequately protected in a single reserve. High altitude forest and maquis (a characteristic scrub-like vegetation) have four threatened and three NT conifers only partially encompassed in protected areas, some open to mining. Thirteen threatened species are restricted to low- and mid-elevation forests, and another three that can also occur on non-ultramafic substrates have isolated, unprotected populations in small forest remnants. Fire and land clearing for mining threaten two conifer species in low- to mid-elevation maquis along with subpopulations of five primarily forest species. Conserving the threatened conifers on terrains miniers will require coordinated measures including: comprehensive protection of forest remnants by forbidding mine waste stockpiling; establishment of new reserves, especially in key unprotected massifs; effective fire prevention; restoration of forest corridors between forest remnants; and multiplication/transplanting of selected species.     

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.     

Fire and succession in the ultramafic maquis of New Caledonia

Aim This study investigates the role of fire and post fire succession in determining the structure and composition of vegetation on ultramafic iron crust soils. Location The study was conducted in the Plaines des Lacs region of southern New Caledonia. Methods A survey was made of eighty-eight sites, recording floristic composition, trunk size-class distributions, regeneration after fire, growth ring counts of Dacrydium araucarioides (Podocarpaceae) and historical information on past fires. Floristic data was ordinated using multidimensional scaling and an index of succession based on structural and historical information. A transition matrix model was developed to predict the effect of fire frequency on vegetation composition. Results The vegetation is undergoing postfire succession from maquis to forest, after about 75 years, and eventually to rainforest. Gymnostoma deplancheanum has a key role as an early colonist that produces shade, the bulk of the litter, and forms nitrogen fixing nodules with Frankia sp. However, the open canopy of Gymnostoma and slow litter decay creates flammable conditions. Though many species resprout from rootstocks, only thirty-nine persist through fires while 114 others colonize at later successional stages, as the litter layer and shade increase. Some early successional species are later excluded but these can persist locally in swamps and on rocky hill tops. Forest and rainforest are less flammable and the matrix model suggests that ignition frequency has a critical role in determining the abundance of maquis or forest. Main conclusions The vegetation mosaic represents a post fire succession from open maquis to forest. Palynological and charcoal records from late Pleistocene sediments suggest that fire has been a major factor determining the development of maquis vegetation since before the arrival of humans. Recently, frequent fires have converted much of the vegetation to maquis, posing a threat to some forest species and largely eliminating rainforest from iron crust soils.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Influence of bioclimatic variables on tree-line conifer distribution in the Greater Yellowstone Ecosystem: implications for species of conservation concern

Aim Tree‐line conifers are believed to be limited by temperature worldwide, and thus may serve as important indicators of climate change. The purpose of this study was to examine the potential shifts in spatial distribution of three tree‐line conifer species in the Greater Yellowstone Ecosystem under three future climate‐change scenarios and to assess their potential sensitivity to changes in both temperature and precipitation. Location This study was performed using data from 275 sites within the boundaries of Yellowstone and Grand Teton national parks, primarily located in Wyoming, USA. Methods We used data on tree‐line conifer presence from the US Forest Service Forest Inventory and Analysis Program. Climatic and edaphic variables were derived from spatially interpolated maps and approximated for each of the sites. We used the random‐forest prediction method to build a model of predicted current and future distributions of each of the species under various climate‐change scenarios. Results We had good success in predicting the distribution of tree‐line conifer species currently and under future climate scenarios. Temperature and temperature‐related variables appeared to be most influential in the distribution of whitebark pine (Pinus albicaulis), whereas precipitation and soil variables dominated the models for subalpine fir (Abies lasiocarpa) and Engelmann spruce (Picea engelmannii). The model for whitebark pine substantially overpredicted absences (as compared with the other models), which is probably a result of the importance of biological factors in the distribution of this species. Main conclusions These models demonstrate the complex response of conifer distributions to changing climate scenarios. Whitebark pine is considered a ‘keystone’ species in the subalpine forests of western North America; however, it is believed to be nearly extinct throughout a substantial portion of its range owing to the combined effects of an introduced pathogen, outbreaks of the native mountain pine beetle (Dendroctonus ponderosae), and changing fire regimes. Given predicted changes in climate, it is reasonable to predict an overall decrease in pine‐dominated subalpine forests in the Greater Yellowstone Ecosystem. In order to manage these forests effectively with respect to future climate, it may be important to focus attention on monitoring dry mid‐ and high‐elevation forests as harbingers of long‐term change.     

Modeling above‐ground litterfall in eastern Mediterranean conifer forests using fractional tree cover,and remotely sensed and ground data

Question: How can we model above‐ground litterfall in Mediterranean conifer forests using remotely sensed and ground data, and geographic information systems (GIS)? Location: Eastern Mediterranean conifer forest of Turkey. Methods: Above‐ground litterfall from Mediterranean forest stands of Pinus nigra, Cedrus libani, Pinus brutia and Juniperus excelsa and mixed Abies cilicica, C. libani and P. nigra was modeled as a function of fractional tree cover using a regression tree algorithm, based on IKONOS and Landsat TM/ETM+data. Landsat TM/ETM+images for the study area were used to map actual stand patterns, based on a land‐cover map of species stands using a supervised classification. Results: Total amount of annual above‐ground litterfall for the entire study area (12 260 km²) was estimated at 417.2 Mg ha^?1 for P. brutia, 291.1 Mg ha^?1 for the mixed stand, 115.5 Mg ha^?1 for P. nigra, 54.6 Mg ha^?1 for J. excelsa and 45.9 Mg ha^?1 for C. libani. The maps generated indicate the distribution of the seasonal amount of total above‐ground litterfall for different species and the distribution of species stands in the study area. There was an increase in the amount of above‐ground litterfall for P. brutia stand in summer, for J. excelsa in autumn and for C. libani, P. nigra and the mixed stand of A. cilicica, P. nigra and C. libani in winter. Conclusion: Application of this model helps to improve the accuracy of estimated litterfall input to soil organic carbon pools in the Mediterranean conifer forests.     

Fire Interval Effects on Successional Trajectory in Boreal Forests of Northwest Canada

Although succession may follow multiple pathways in a given environment, the causes of such variation are often elusive. This paper describes how changes in fire interval mediate successional trajectory in conifer-dominated boreal forests of northwestern Canada. Tree densities were measured 5 and 19 years after fire in permanent plots and related to pre-fire vegetation, site and fire characteristics. In stands that were greater than 75 years of age when they burned, recruitment density of conifers was significantly correlated with pre-fire species basal area, supporting the expectation of stand self-replacement as the most common successional pathway in these forests. In contrast, stands that were under 25 years of age at the time of burning had significantly reduced conifer recruitment, but showed no change in recruitment of trembling aspen (Populus tremuloides). As a result, young-burned stands had a much higher probability of regenerating to deciduous dominance than mature-burned stands, despite the dominance of both groups by spruce (Picea mariana and Picea glauca) and pine (Pinus contorta) before the fire. Once initiated, deciduous-dominated stands may be maintained across subsequent fire cycles through mechanisms such as low on-site availability of conifer seed, competition with the aspen canopy, and rapid asexual regeneration of aspen after fire. We suggest that climate-related increases in fire frequency could trigger more frequent shifts from conifer to deciduous-dominated successional trajectories in the future, with consequent effects on multiple ecosystem processes.     

黑龙江省大兴安岭林区火烧迹地森林更新及其影响因子

林火干扰是大兴安岭森林更新的影响因子之一,研究火烧迹地森林更新的影响因子(立地条件、火前植被、火干扰特征)对理解生态系统的结构、功能和火后演替轨迹具有重要意义。选取呼中及新林林业局55个代表性火烧样地,利用增强回归树分析法分析了火烧迹地森林更新的影响因素。结果表明:(1)立地条件是影响针、阔叶树更新苗密度的主要因素;海拔对针叶树更新苗密度的影响最大;坡度对阔叶树更新苗密度影响最大;(2)距上次火烧时间对针叶树更新苗比重影响最大,其次是林型;(3)中度林火干扰后森林更新状况好于轻度和重度火烧迹地。根据火烧迹地森林更新调查分析可知:林型影响火后演替模式,火前为针叶树或阔叶树纯林,火后易发生自我更新(火后树种更新组成与火前林型相同),而针阔混交林在火干扰影响下易于发生序列演替(火后初期以早期演替树种更新为主)。     

The abundance,distribution and structural characteristics of tree‐holes in Nothofagus forest,New Zealand

Tree‐holes provide an important microhabitat that is used for feeding, roosting and breeding by numerous species around the world. Yet despite their ecological importance for many of New Zealand's endangered species, few studies have investigated the abundance or distribution of tree‐holes in native forests. We used complementary ground and climbed tree surveys to determine the abundance, distribution and characteristics of tree‐holes in undisturbed Nothofagus forest in the Lewis Pass, New Zealand. We found that hole‐bearing trees were surprisingly abundant compared with many other studies, including Australian Eucalyptus species and American beech. In fact, we estimated as many as 3906 tree‐holes per hectare, of which 963 holes per hectare were potentially large enough to provide roost sites for hole‐nesting bats in New Zealand, while only eight holes per hectare were potentially suitable for specialist hole‐nesting birds. This was of great interest as primary cavity‐excavating animals are absent from New Zealand forests, compared with North America and Australia. Moreover, tree‐hole formation in New Zealand is likely to be dominated by abiotic processes, such as branch breakage from windstorms and snow damage. As has been found in many other studies, tree‐holes were not uniformly distributed throughout the forest. Tree‐holes were significantly more abundant on the least abundant tree species, Nothofagus fusca, than on either N. menziesii or N. solandri. In addition to tree species, tree size was also an important factor influencing the structural characteristics of tree‐holes and their abundance in this forest. Moreover, these trends were not fully evident without climbed tree surveys. Our results revealed that ground‐based surveys consistently underestimated the number of tree‐holes present on Nothofagus trees, and illustrate the importance of using climbed inspections where possible in tree‐hole surveys. We compare our results with other studies overseas and discuss how these are linked to the biotic and abiotic processes involved in tree‐hole formation. We consider the potential implications of our findings for New Zealand's hole‐dwelling fauna and how stand dynamics and past and future forest management practices will influence the structural characteristics of tree‐holes and their abundance in remnant forest throughout New Zealand.     

Post-Glacial Climate–Fire Interactions Control Tree Composition of Mesic Temperate Forests in Eastern North America

Stand-scale gap-phase dynamics is generally viewed as the main driver of development in mesic deciduous forests of the temperate biome. Soil charcoal of temperate forests in eastern North America are unnoticed in most surveys, thus explaining why fire is undervalued as a driver of forest succession. The extent to which gap-phase, fire, or other processes are responsible for the regeneration and maintenance of mesic deciduous forests is unknown because paleoecological evidence is lacking. We tested the fire-driven succession hypothesis on the development of this major forest type. Based on charcoal ¹⁴C dates of two sites, 44 and 55 fires occurred since early Holocene, with a mean interval of 170 to 215 years. The vegetation of both sites followed comparable post-glacial trajectories consisting of three distinct periods. Conifers dominated the two first periods during 5200–6000 years and were replaced by hardwoods–conifers over the last 3500 years. The first period was represented by boreal conifers, whereas the second period, dominated by white pine (Pinus strobus) forests, persisted during 3000–4300 years. The third period marked the development of hardwood (sugar maple, Acer saccharum) forests. Fires occurred continuously on the sites since early Holocene likely under dry conditions during the conifer periods and cooler and moister conditions during the hardwood–conifer period. Recurrent fires appear with climate as key drivers of the long-term dynamics of several temperate forests in eastern North America. Similar studies on other temperate forests should be pursued to test the hypothesis of climate–fire interactions influencing tree composition change.

     

Population collapse and retreat to fire refugia of the Tasmanian endemic conifer Athrotaxis selaginoides following the transition from Aboriginal to European fire management

Untangling the nuanced relationships between landscape, fire disturbance, human agency, and climate is key to understanding rapid population declines of fire‐sensitive plant species. Using multiple lines of evidence across temporal and spatial scales (vegetation survey, stand structure analysis, dendrochronology, and fire history reconstruction), we document landscape‐scale population collapse of the long‐lived, endemic Tasmanian conifer Athrotaxis selaginoides in remote montane catchments in southern Tasmania. We contextualized the findings of this field‐based study with a Tasmanian‐wide geospatial analysis of fire‐killed and unburned populations of the species. Population declines followed European colonization commencing in 1802 ad that disrupted Aboriginal landscape burning. Prior to European colonization, fire events were infrequent but frequency sharply increased afterwards. Dendrochronological analysis revealed that reconstructed fire years were associated with abnormally warm/dry conditions, with below‐average streamflow, and were strongly teleconnected to the Southern Annular Mode. The multiple fires that followed European colonization caused near total mortality of A. selaginoides and resulted in pronounced floristic, structural vegetation, and fuel load changes. Burned stands have very few regenerating A. selaginoides juveniles yet tree‐establishment reconstruction of fire‐killed adults exhibited persistent recruitment in the period prior to European colonization. Collectively, our findings indicate that this fire‐sensitive Gondwanan conifer was able to persist with burning by Aboriginal Tasmanians, despite episodic widespread forest fires. By contrast, European burning led to the restriction of A. selaginoides to prime topographic fire refugia. Increasingly, frequent fires caused by regional dry and warming trends and increased ignitions by humans and lightning are breaching fire refugia; hence, the survival Tasmanian Gondwanan species demands sustained and targeted fire management.