基于树轮火疤塔河蒙克山樟子松林火灾的频度分析

大兴安岭地区是我国重要林区,又是林火的多发区,林火是森林生态系统中的重要干扰因子,对整个森林生态系统结构、功能和动态都有重要影响。樟子松为欧洲赤松的一个变种,在我国主要分布于大兴安岭地区。近年来,频繁的森林火灾造成樟子松林大面积减少。因此,重建大兴安岭樟子松林火历史,掌握樟子松林火灾规律已显得十分迫切。在大兴安岭北部塔河县蒙克山林场采集了11棵樟子松火疤圆盘,利用树木年轮年代学方法重建了大兴安岭北部塔河县蒙克山樟子松林的火灾历史,获得1个以树轮年代学为基础的樟子松火疤年表。利用火历史分析软件得到蒙克山樟子松林火灾间隔期和轮回期分别为24.8a和33a。由火疤年表得到的蒙克山樟子松林火灾历史大致可分为3个历史时期:满清中期(1723-1859年)、清末民国时期(1860-1949年)和建国后(1950年至今)。41个火疤记录中早早材火(E)所占比例最大,占全部火疤数目的61%,晚材火(A)、未确定火(U)和休眠季节火(D)则相对较少。大区域性火灾事件平均间隔期为32.5a,最大火灾间隔期为61a。本研究为进一步探究大兴安岭地区寒温带针叶林火灾历史的时间和空间格局及其之间的联系提供了基础数据。     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.     

Replacement patterns and species coexistence in an Andean Araucaria‐Nothofagus forest

Questions: Fire appears to affect both replacement patterns and coexistence of Araucaria araucana‐Nothofagus pumilio forests in the Andean Araucarian region of south‐central Chile. A quantitative assessment of coexistence in the absence of recent fires, however, is lacking. In this study, we considered the life‐history attributes, time of recruitment and spatial pattern of individuals of both tree species to address the following questions. How regular has recruitment of both species been in time? Is there any temporal niche differentiation? Are the two species positively or negatively associated in space and, if so, at what scale and for what age and size classes? Is there any spatial niche differentiation? Location: Andean Araucarian region of Chile, Villarrica National Park (39°35′S, 71°31′W; 1300 m a.s.l.). Methods: We stem‐mapped and cored a total of 1073 trees in a 1‐ha plot in a late‐successional post‐fire stand to examine spatiotemporal patterns of establishment. We used semivariogram modelling and the pair‐correlation function to distinguish between regeneration modes and describe species interactions. Results: The two species differ in their regeneration mode: whereas A. araucana appeared to recruit more continuously in time and space, episodic pulses of establishment were dominant for N. pumilio. At small scales, younger age‐class stems of A. araucana were randomly distributed, while older age‐class stems were aggregated. This was in contrast to common patterns for temperate tree species, including N. pumilio, following processes of self‐thinning. Younger age classes of A. araucana were distributed independently of older trees of both species, but younger age classes of N. pumilio had a negative association with older conspecifics at scales similar to crown diameter. Conclusions: In the absence of recent fires, it is likely that A. araucana would dominate the stand alone, given its greater shade tolerance, greater longevity and continuous recruitment. However, while canopy closure is still incomplete, the shade‐intolerant N. pumilio will be able to recruit in those open areas after seed masting and will coexist with A. araucana.     

Regeneration dynamics and life history differences in southern Chilean Nothofagus forests: a synthesis

Large, infrequent natural disturbances have been proposed as a key component in determining the distribution and abundance of Nothofagus in southern Andean temperate forests. In this study, a comparison of influences of small-to-intermediate-sized natural disturbances and effects of selective logging on the establishment and growth of Nothofagus forests in south-central Chile are synthesized with the results of other forest studies to develop general hypotheses on the regeneration dynamics of southern Chilean Nothofagus alpina forests. A synthesis is given for regeneration pattern in the abundance of 18 temperate rain forests in the Andean Range, Chile. The study aimed to determine the role of life history differences in promoting coexistence of the five main tree species (N. alpina, N. dombeyi, N. pumilio, N. macrocarpa and Laurelia philippiana) in N. alpina-dominated forests. Age data reported for N. alpina, a shade mid-tolerant emergent tree in the temperate rain forests of southern Chile, indicate maximum lifespans > 650 years, figures unprecedented for N. alpina. In low elevation stands, N. alpina coexists with broad-leaved evergreen tree species, such as L. philippiana, L. sempervirens, Persea lingue, and in these stands an intermittent establishment of Nothofagus occurred and appeared to be most dependent on small-to-intermediate disturbances. In high-elevation stands, in contrast, Nothofagus establishment was less dependent on disturbance, regeneration being much more continuous even in the absence of canopy openings. The forests studied provide another example of the general pattern of increasing dependence of Nothofagus on disturbance towards the more productive end of the environmental gradients. As a long-lived pioneer and despite its dependence on disturbance in lowland sites, N. alpina has been subject to selection for complementary growth, adult survivorship, and mid-tolerance to shade. Thus, interspecific differences in juvenile and adult life history characteristics of N. alpina and its competitors may be sufficient to maintain its persistence in the landscape. In conclusion, this study of population structures and replacement patterns provides a comprehensive picture on our understanding of regeneration dynamics and trait differentiation in southern temperate forests by recognizing both the influences of environmental gradients (i.e., altitude) on competing species and the disturbance regimes. This revised version was published online in June 2006 with corrections to the Cover Date.     

树木年轮在干扰历史重建中的应用

干扰是影响森林生态系统结构和功能的重要因子,重建森林群落干扰历史可以掌握干扰发生的机制和规律,对森林经营有重要意义。年轮记载了树木逐年的生长历史,利用年轮可以重建不同尺度上干扰的时空格局,具有重建历史长、定年准确和材料容易获得的优点,是森林动态历史研究不可替代的资料。干扰对树木个体的影响可以分为伤害干扰和生长干扰两种类型。不同类型的干扰的特点和对树木的生长影响不同所以重建方法也不同,对树干造成直接伤害的干扰可以利用树干的疤痕重建,如火灾和泥石流等;生长干扰对树木的生长势造成影响,可以通过识别年轮中的生长抑制和释放发生和持续的时间来确定干扰的时间和强度。但是对破坏性的干扰事件要通过群落中个体的建成时间高峰来判断。欧洲和北美地区利用树木年轮重建森林干扰历史的研究已经很广泛,主要包括火灾、虫灾、地质灾害和气象灾害等干扰类型,在中国利用年轮重建群落干扰的研究还处于起步阶段。本文探讨了应用树木年轮重建不同森林干扰事件的方法,总结了不同的重建方法和干扰重建取得的进展,并指出了以后研究的方向,为以后的干扰重建提供了参考。     

Alsodes Australis,A New Species of Leptodactylid Frog from the Temperate Nothofagus Forest of Southern Chile and Argentina

A new species of frog, Alsodes australis, is described from the temperate Nothofagus forest of southern Chile and Argentina, based on adults and tadpoles. Up to now, no sympatric occurrence of another congeneric species is known from the type locality. The genus comprises three species groups. From a karyological point of view this species is included in the monticola group, whose species have 26 chromosomes.     

Fire legacies in eastern ponderosa pine forests

Disturbance legacies structure communities and ecological memory, but due to increasing changes in disturbance regimes, it is becoming more difficult to characterize disturbance legacies or determine how long they persist. We sought to quantify the characteristics and persistence of material legacies (e.g., biotic residuals of disturbance) that arise from variation in fire severity in an eastern ponderosa pine forest in North America. We compared forest stand structure and understory woody plant and bird community composition and species richness across unburned, low‐, moderate‐, and high‐severity burn patches in a 27‐year‐old mixed‐severity wildfire that had received minimal post‐fire management. We identified distinct tree densities (high: 14.3 ± 7.4 trees per ha, moderate: 22.3 ± 12.6, low: 135.3 ± 57.1, unburned: 907.9 ± 246.2) and coarse woody debris cover (high: 8.5 ± 1.6% cover per 30 m transect, moderate: 4.3 ± 0.7, low: 2.3 ± 0.6, unburned: 1.0 ± 0.4) among burn severities. Understory woody plant communities differed between high‐severity patches, moderate‐ and low‐severity patches, and unburned patches (all p < 0.05). Bird communities differed between high‐ and moderate‐severity patches, low‐severity patches, and unburned patches (all p < 0.05). Bird species richness varied across burn severities: low‐severity patches had the highest (5.29 ± 1.44) and high‐severity patches had the lowest (2.87 ± 0.72). Understory woody plant richness was highest in unburned (5.93 ± 1.10) and high‐severity (5.07 ± 1.17) patches, and it was lower in moderate‐ (3.43 ± 1.17) and low‐severity (3.43 ± 1.06) patches. We show material fire legacies persisted decades after the mixed‐severity wildfire in eastern ponderosa forest, fostering distinct structures, communities, and species in burned versus unburned patches and across fire severities. At a patch scale, eastern and western ponderosa system responses to mixed‐severity fires were consistent.     

Structure and tree-fall gap dynamics of old-growth Nothofagus forests in Tierra del Fuego,Argentina

Abstract. Tree size and age structure, tree-fall and gap characteristics, and regeneration in gaps were studied in Nothofagus-dominated old-growth forests in Tierra del Fuego, Argentina. Gap-phase regeneration has resulted in all-aged populations for N. pumilio, N. betuloides, and Drimys winteri, and regeneration in gaps appears to be maintaining coexistence between species in mixed stands. N. betuloides fills many gaps via advance regeneration and some individuals persist for > 150 yr in the understory. Multiple periods of release and suppression indicate that N. betuloides may take advantage of several gap events to reach the main canopy. Likewise, Drimys grows well under closed canopy and can rapidly respond to gap formation, sometimes impeding the regeneration of N. betuloides. In contrast, N. pumilio regenerates in gaps mainly from seed or from advance regeneration of small, ephemeral seedlings. Gap turnover times in Fuegian forests were estimated at 300 - 500 yr, although gap formation was highly episodic and possibly associated with regionally extensive windstorms, earthquakes, and stand-level dieback. 92 % of gaps involved multiple tree-falls, and at least 53 % involved secondary expansion. Gap and tree-fall characteristics in Tierra del Fuego were similar to results from northern Patagonia, Chile, and New Zealand; however, we emphasize that regeneration of Nothofagus spp. and Drimys winteri in gaps depends on associated vegetation and varies along both local and regional environmental gradients.     

Damage caused by mistletoe Misodendrum punctulatum Banks Ex Dc. on architecture and radial growth of Nothofagus pumilio (Poepp. et Endl.) Krasser forests of southern Chile

Parasitic plants rely completely or partially on host plant, namely holoparasitic or hemiparasitic, respectively. The effect of parasitism on host plants is not fully known. Misodendrum punctulatum (Misodendraceae) is the most common hemiparasite species in the southernmost forests of Chile and Argentina, and it has been identified as an important sanitary problem in forestry. In this study we evaluated the M. punctulatum effect on growth rates and architecture of pure Nothofagus pumilio (Nothofagaceae) forests in southern South America (52°10′S; 71°55′W). We established three plots of 30 × 30 m, and 10 cores were extracted from diameter at breast height (d.b.h. = 1.3 m) from living trees in each plot, using increment borers. Tree rings were measured, and anova was used to compare the annual growth rate of both infected and uninfected N. pumilio. The results showed that N. pumilio trees severely infected by M. punctulatum reduce growth rates in contrast with uninfected individuals (P < 0.05). Besides, trees with high‐infection levels evidenced deterioration in its architecture, showing asymmetric and suppressed canopies, according to ad hoc numerical indexes. This study provides a starting point to understand the parasite–host relationship in the Patagonian forests of southern Chile.     

The significance of life history strategies in the developmental history of mixed beech (Nothofagus) forests,New Zealand

Size and age structure analysis, dated past disturbances, treefall replacement patterns, and spatial pattern analysis were used to reconstruct the developmental history of two old-growth Nothofagus fusca/N. menziesii stands, South Island, New Zealand. Diameter and height class distributions suggested that N. menziesii was replacing N. fusca, however, stand history reconstruction analysis showed that both species had regenerated intermittently after small-scale disturbances. Although large-scale disturbances such as blowdowns may occasionally generate even-aged stands, gap-phase regeneration maintains the forests in compositional equilibrium. In the absence of other competing tree species and understorey plants the two species appear to coexist by way of different life history strategies, where one species (N. menziesii) has low juvenile mortality and the other (N. fusca) has faster height growth rates and greater longevity and adult survivorship.     

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.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

Tree population dynamics in Kibale National Park, Uganda 1975–1998

Changes in species composition, stem abundance, and basal area of trees taller than or equal to 10 m in a medium altitude tropical rain forest at the Ngogo study area, Kibale National Park, Uganda are described for the period between 1975 and 1998 ( n  = 23 years). Two enumeration episodes were conducted in 263 plots of 5 m by 50 m during 1975–80 and 1997–98. During this period, species richness decreased by 3% (from 92 to 89). Species diversity (H') also declined slightly from 2.97 to 2.86. The number of trees recorded in the sample plots decreased by 8% (from 2545 to 2329), while basal area decreased from 49.48 m² ha⁻¹ to 48.68 m² ha^−1. However, stem abundance and basal area increased for some species.     

Natural and anthropogenic influences on coarse woody debris stocks in Nothofagus–Araucaria forests of northern Patagonia,Argentina

Coarse woody debris (CWD) is an important element driving ecological processes, strengthening ecosystem resilience and for biodiversity within forest ecosystems. However, the abundance and distribution of CWD and their relation to natural and human factors are poorly known in southern South America. In this work we studied the density and volume of CWD types in Nothofagus–Araucaria stands in northern Patagonia (Neuquén – Argentina) and relationships with forest composition and structure. We also studied their relationships with fire history, topography and human‐related variables. Twenty‐three stands with Nothofagus pumilio, Nothofagus antarctica and/or Araucaria araucana were sampled to estimate quantities of logs, snags and dead branches using the planar‐intersect method. CWD density and volume in these forests were moderate and varied across the landscape with a spatial pattern determined by biotic, abiotic and human use–related variables. Mean CWD volume was 52.9 m³ ha^?1 (range: 1.6–143.7) and significantly varied among forest types and watersheds. CWD was positively related to dbh, tree height and slope, but negatively related to tree density. CWD was clearly influenced by composition and structural characteristics of stands, where the tree species traits had an important role. As well, the observed amount and type of CWD, whereby most of the stands showed low levels of old (pre‐disturbance) logs/snags and poor new inputs of deadwood, may be explained by fire frequency. Firewood gathering and livestock grazing negatively affected deadwood stocks and topography counteracts this effect by limiting human access. Fire disturbance history, windthrow and dieback pulses produced by insect outbreaks and human access seemed to be the main causes that best explained CWD spatial distribution and abundance patterns in north‐western Patagonian forests.