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.     

Study on canopy disturbance regime and mechanism of tree species diversity maintenance in the lower subtropical evergreen broad-leaved forest,South China

ABSTRACT

The canopy disturbance, the gap environment, gap regeneration and maintenance of tree species diversity in the lower subtropical evergreen broad-leaved forest (LSEBF) of South China were studied in this paper. The most common manner of gap formation in the forest was by stem breakage. Most gaps were formed by two gap makers. The sizes of most expanded gaps (EG) and canopy gaps (CG) were in the range of 100 – 300 m² and 50 – 100 m², respectively. The ecological factors in gaps were analysed on the basis of contrasting measurement of the microclimatic regimes in gaps of different sizes and in non-gap stands. Tree species in the LSEBF were classified into 5 ecological species groups on the bases of their changes in order of importance values in gaps and in non-gap stands. Most of the species reached their peak of regeneration density around the gap sizes of 100 m² and 500 m². The curves of regeneration density vs. gap age for major species revealed two types. Regeneration densities of most species, and species diversity indices in gaps were greater than those in non-gap stands.     

Growth and posture control strategies in Fagus sylvatica and Acer pseudoplatanus saplings in response to canopy disturbance

Background and Aims

Forest tree saplings that grow in the understorey undergo frequent changes in their light environment to which they must adapt to ensure their survival and growth. Crown architecture, which plays a critical role in light capture and mechanical stability, is a major component of sapling adaptation to canopy disturbance. Shade-adapted saplings typically have plagiotropic stems and branches. After canopy opening, they need to develop more erect shoots in order to exploit the new light conditions. The objective of this study was to test whether changes in sapling stem inclination occur after canopy opening, and to analyse the morphological changes associated with stem reorientation.

Methods

A 4-year canopy-opening field experiment with naturally regenerated Fagus sylvatica and Acer pseudoplatanus saplings was conducted. The appearance of new stem axes, stem basal diameter and inclination along the stem were recorded every year after canopy opening.

Key Results

Both species showed considerable stem reorientation resulting primarily from uprighting (more erect) shoot movements in Fagus, and from uprighting movements, shoot elongation and formation of relay shoots in Acer. In both species, the magnitude of shoot uprighting movements was primarily related to initial stem inclination. Both the basal part and the apical part of the stem contributed to uprighting movements. Stem movements did not appear to be limited by stem size or by stem growth.

Conclusions

Stem uprighting movements in shade-adapted Fagus and Acer saplings following canopy disturbance were considerable and rapid, suggesting that stem reorientation processes play a significant role in the growth strategy of the species.     

Heterogeneous genetic structure in a Fagus crenata population in an old-growth beech forest revealed by microsatellite markers

The within-population genetic structure of Fagus crenata in a 4-ha plot (200 x 200 m) of an old-growth beech forest was analysed using microsatellite markers. To assess the genetic structure, Moran's I spatial autocorrelation coefficient was calculated. Correlograms of Moran's I showed significant positive values less than 0.100 for short-distance classes, indicating weak genetic structure. The genetic structure within the population is created by limited seed dispersal, and is probably weakened by overlapping seed shadow, secondary seed dispersal, extensive pollen flow and the thinning process. Genetic structure was detected in a western subplot of 50 x 200 m with immature soils and almost no dwarf bamboos (Sasa spp.), where small and intermediate-sized individuals were distributed in aggregations with high density because of successful regeneration. By contrast, genetic structure was not found in an eastern subplot of the same size with mature soils and Sasa cover, where successful regeneration was prevented, and the density of the small and intermediate-sized individuals was low. Moreover, genetic structure of individuals in a small-size class (diameter at breast height < 12 cm) was more obvious than in a large-size class (diameter at breast height >/= 12 cm). The apparent genetic structure detected in the 4-ha plot was therefore probably the result of the structure in the western portion of the plot and in small and intermediate-sized individuals that successfully regenerated under the favourable environment. The heterogeneity in genetic structure presumably reflects variation in the density that should be affected by differences in regeneration dynamics associated with heterogeneity in environmental conditions.     

Natural disturbance and tree species coexistence in an old-growth beech - dwarf bamboo forest,southwestern Japan

Abstract. The structure and composition of a cool-temperate old-growth beech (Fagus crenata) - dwarf bamboo (Sasa spp.) forest, partially affected by landslide disturbance, in the Daisen Forest Reserve of southwestern Japan, were investigated in relation to forest floor and canopy conditions. All stems ≥ 4 cm DBH were mapped on a 4-ha plot and analyses were made of population structure, spatial distribution and spatial association of major tree species. The dominant species, F. crenata, which had the maximum DBH among the species present, had the highest stem density. However, for other species, larger-sized species had lower stem density with few smaller stems or saplings, while smaller-sized species had higher stem density with many smaller stems or saplings. Canopy trees of F. crenata were distributed randomly in the plot, while its stems in the other layers and all other species were distributed patchily. Small patches represent gap-phase regeneration. Larger patches correlate with landslide disturbance, difference in soil age, or the presence/absence of Sasa. Cluster analysis for spatial associations among species and stems in the different layers revealed that the forest community consists of several groups. One main group was formed on sites not covered with Sasa. This group contained a successional subgroup (from Betula grossa to Acer mono and/or F. crenata) initiated by landslide disturbance and a subgroup of tree species that avoid Sasa. Another group was formed on sites with mature soils covered largely with Sasa. This contained associations of canopy trees of F. crenata and smaller-sized tree species such as Acanthopanax sciadophylloides and Acer japonicum. It is found that the community of this old-growth beech forest is largely organized by natural disturbance and heterogeneous conditions of the forest floor (difference in soil age and presence/absence of Sasa). The existence of these different factors and the different responses of species to them largely contribute to the maintenance of tree species diversity in this forest.; Keywords: Cluster analysis; Fagus crenata; Forest dynamics; Gap; Landslide; Spatial pattern.     

Effects of CO2 enrichment on whole-plant carbon budget of seedlings of Fagus grandifolia and Acer saccharum in low irradiance

Carbon exchange rates (CER) and whole-plant carbon balances of beech (Fagus grandifolia) and sugar maple (Acer saccharum) were compared for seedlings grown under low irradiance to determine the effects of atmospheric CO₂ enrichment on shade-tolerant seedlings of co-dominant species. Under contemporary atmospheric CO₂, photosynthetic rate per unit mass of beech was lower than for sugar maple, and atmospheric CO₂ enrich ment enhanced photosynthesis for beech only. Aboveground respiration per unit mass decreased with CO₂ enrichment for both species while root respiration per unitmass decreased for sugar maple only. Under contemporary atmoapheric CO₂, beech had lower C uptake per plant than sugar maple, while C losses per plant to nocturnal aboveground and root respiration were similar for both species. Under elevated CO₂, C uptake per plant was similar for both species, indicating a significant relative increase in whole-seedling CER with CO₂ enrich ment for beech but not for sugar maple. Total C loss per plant to aboveground respiration was decreased for beech only because increase in sugar maple leaf mass counterbalanced a reduction in respiration rates. Carbon loss to root respiration per plant was not changed by CO₂ enrichment for either species. However, changes in maintenance respiration cost and nitrogen level suggest changes in tissue composition with elevated CO₂. Beech had a greater net daily C gain with CO₂ enrichment than did sugar maple in contrast to a lower one under contemporary CO₂. Elevated CO₂ preferentially enhances the net C balance of beech by increasing photosynthesis and reducing respiration cost. In all cases, the greatest C lost was by roots, indicating the importance of belowground biomass in net C gain. Relative growth rate estimated from biomass accumulation was not affected by CO₂ enrichment for either species possibly because of slow growth under low light. This study indicates the importance of direct effects of CO₂ enrichment when predicting potential change in species distribution with global climate change.     

Structure and dynamics of a <Emphasis Type="Italic">Castanopsis cuspidata</Emphasis> var. <Emphasis Type="Italic">sieboldii</Emphasis> population in an old-growth,evergreen, broad-leaved forest: The importance of sprout regeneration

The population structure and dynamics of Castanopsis cuspidata var. sieboldii were studied to evaluate vegetative and sapling regeneration in an old-growth, evergreen broad-leaved forest exposed to low-severity typhoon disturbances by annual typhoons on the Tsushima Islands in Japan. The density of individuals 5cm d.b.h. was 38.0ha^–1 in 1990; 7.9% were multiple-stemmed individuals. Over the 7-year study period (1990–1997), the number of individuals decreased, although the number of stems increased. Over 30% of apparently dead individuals were reconstituted by sprouting stems. Compared with sprout regeneration, sapling regeneration rarely occurred and was only observed in canopy gaps. Most individuals had at least one sprout shoot (H 30cm, d.b.h. <5cm), and the number and size of sprout shoots increased as the size of the individuals increased. During the study period, larger individuals with stem breakage tended to produce sprout stems. The density of saplings was 1074ha^–1 and they were more abundant in canopy gaps than under closed canopies, but large saplings were very rare even in canopy gaps. The population of C.cuspidata var. sieboldii consisted primarily of single-stemmed individuals with a few multiple-stemmed individuals providing a sprout bank. Larger individuals responded to the low severity typhoon disturbances and formed sprout stems. Although many saplings were observed, regeneration occurred more often by sprout formation than by growth of saplings. Thus, sprout regeneration is an important mode of regeneration, which allows this pioneer-like species to maintain its population in this forest.     

粤北山地常绿阔叶林自然干扰后冠层结构与林下光照动态

以粤北车八岭2008年受冰灾破坏的山地常绿阔叶林为研究对象,设置2 hm2固定样地开展连续3a(2008—2010年)的群落调查,并采用半球面影像技术(Hemispherical photography)获取冠层结构和林下光照指标,分析灾后森林演替过程中冠层结构和林下光照的动态。研究发现:1)灾后森林恢复过程中,样地林下光照(直射光、散射光和总光照)均随林冠开度的减少、叶面积指数的增加而减少;2)从2008到2010年,各年度冠层结构和林下光照的差异均极显著(P<0.0001),但年间差异程度有逐年减少的趋势;3)灾后森林恢复前3a,林下直射光对林下总光照的贡献大于散射光,其时空波动性也大于散射光;4)林冠开度对冠层结构的反映程度比叶面积指数高,冠层结构对林下散射光的影响比对直射光大。灾后林木先是迅速生长然后生长速度缓慢下来并逐渐稳定,随森林逐渐郁闭林下光照也随之减少,其中林冠开度用于评价冠层结构动态的效果更佳,林下直射光比散射光的时空变化更复杂。     

Canopy disturbance and gap partitioning promote the persistence of a pioneer tree population in a near‐climax temperate forest of the Qinling Mountains,China

An unresolved question of temperate forests is how pioneer tree species persist in mature forests. In order to understand the responsible mechanisms, we investigated a near‐climax mixed temperate forest dominated by Betula albosinensis in the Qinling Mountains of China. Through establishing four 50 m × 50 m plots, we examined the canopy disturbance characteristics and its effects on tree recruitments. We further test the intra‐ and interspecific effects on the recruitment of B. albosinensis. The obtained data demonstrated canopy disturbance was frequent but most small‐sized. The canopy gaps are caused mainly by adult B. albosinensis by snapping. The regeneration of coexistent tree species shows a distinct preference for gap size. B. albosinensis were clumped at the juvenile stage and small scales. B. albosinensis juveniles were positively associated with B. utilis juveniles and negatively associated with the conspecific and B. utilis large trees. In addition, B. albosinensis juveniles showed negative associations with contemporary other tree species. Our results suggested that canopy disturbance caused by canopy trees and gap partitioning among the coexistent tree species are important for the persistence of the mixed forest. As a main gapmaker, B. albosinensis appear to develop a self‐perpetuating life‐history trait and allow them to persist.     

古田山常绿阔叶林凋落量时间动态及冰雪灾害的影响

2006年10月至2009年12月期间,我们通过对古田山24 ha常绿阔叶林动态样地169个种子雨收集器的凋落物进行烘干、分类、称量和数据分析,研究了凋落量的组成特征和时间动态,以及受2008年2月特大冰雪灾害的影响.2007年和2009年凋落量分别为532.05 g/m2和375.17 g/m2,年际变化显著,这与2008年的冰雪灾害有关.2007年古田山常绿阔叶林各组分在凋落量中所占比例依次为:叶(78.99%)>枝(14.69%)>皮(3-33%)>其他(2.99%);叶凋落量中各组分所占比例依次为:常绿阔叶树种叶(78.70%)>落叶阔叶树种叶(12.37%)>针叶树种叶(8.92%),其中甜槠(Castanopsis eyrei)、木荷(Schima superba)、马尾松(Pinus massoniana)和短柄袍(Quercus serrata vat.brevipetiolata)4个优势种年凋落量合计占叶凋落总量的71.36%,它们的动态直接影响着凋落总量的变化格局.凋落量高峰发生在春季(4月)和秋冬季(10月末至12月初),其中总凋落量、叶凋落量动态呈双峰型,枝条凋落量和树皮凋落量动态为不规则峰型,其他凋落量动态为单峰型.冰雪灾害后总凋落量、叶凋落量、枝条凋落量显著减少(P<0.05),其中甜槠、杨梅叶蚊母树(Distylium myricoides)叶凋落最显著减少(P<0.05)、木荷叶凋落量减少达到边缘显著水平(P<0.1),这也反映了冰亏灾害期间森林群落的受损情况.其他凋落量(主要成分为虫粪)在4B份增加极其显著(P<0.01),说明灾后植物的枝叶出现了补偿性生长.     

Canopy cover and tree regeneration in old-growth cove forests of the Appalachian Mountains

Relationships between canopy cover and tree regeneration were determined for various species in cove forests of the Great Smoky Mountains. Old-growth stands were sampled with six plots covering a total area of 4.8 ha. Each plot was subdivided into contiguous 10×10 m quadrats. Canopy cover overlying each of the 480 quadrats was characterized with three different indices based on visual estimates of cover. Influences of: (1) overlying cover, (2) proximate openings, and (3) total area of proximate openings on quadrat regeneration densities were determined. Most species reproducing by seed and some species reproducing by vegetative means had higher densities in quadrats with openings, but only the intolerants were highly dependent on gaps. Tsuga canadensis, a very shade-tolerant species, was one of the few species with abundant regeneration beneath dense canopy cover. In general, understory areas near gaps had somewhat higher regeneration densities than other areas with overlying cover. Several shade-tolerant species showed a positive regeneration density response to canopy openings and an ability to regenerate in gaps 0.01–0.03 ha in area. These openings were too small for intolerant species. Many species exhibited a positive response to total size of the proximate opening(s). A sharp increase in regeneration density with area of the opening(s) was evident at approximately 0.04 ha for the shade-intolerant species.     

Effect of forest canopy composition on soil nutrients and dynamics of the understorey: mixed canopies serve neither vascular nor bryophyte strata

Question: The effect of overstorey composition on above‐ground dynamics of understorey vegetation is poorly understood. This study examines the understorey biomass, production and turnover rates of vascular and non‐vascular plants along a conifer–broadleaf gradient of resource availability and heterogeneity. Location: Canadian boreal forests of northwest Quebec and Ontario. Methods: We sampled mature stands containing various proportions of black spruce (Picea mariana (Mill.) BSP), trembling aspen (Populus tremuloides Michx.) and jack pine (Pinus banksiana Lamb.). Above‐ground biomass of the understorey vegetation was assessed through harvesting; annual growth rates were calculated as the differences between biomass in 2007 and 2008, as estimated by allometric relationships, and turnover rates were estimated as net primary production divided by the biomass in 2007. Results: Higher aspen presence, linked to greater nutrient availability in the forest floor, was generally associated with higher vascular biomass and production in the understorey. This effect was less pronounced in sites of high intrinsic fertility. In contrast, bryophyte biomass was positively associated with conifer abundance, particularly in wet sites of the Quebec study area. Non‐linear responses resulted in total understorey biomass being lower under mixed canopies than under pure aspen or pure conifer canopies. Turnover rates did not differ with overstorey composition. Conclusions: While resource availability is a main driver of understorey productivity, resources as drivers appear to differ with differences in understorey strata components, i.e. vascular versus non‐vascular plants. Resource heterogeneity induced by a mixed canopy had overall negative effects on understorey above‐ground productivity, as this productivity seemed to rely on species adapted to the specific conditions induced by a pure canopy.     

Regeneration and coexistence of two subalpine conifer species in relation to dwarf bamboo in the understorey

Abstract. Microhabitats for seedling establishment and gap regeneration in subalpine forests of northern Japan were studied for two conifers, Abies sachalinensis and Picea glehnii. The abundance of understorey dwarf bamboo (Sasa spp.) was different for the four plots examined. Two types of micro-habitats were recognized for the two conifers: ground and elevated woody substrates (fallen logs and buttresses). Picea regenerated mostly on elevated sites, while Abies regenerated on both ground and elevated sites. The densities of Picea were independent of those of Sasa, but Abies densities decreased with increasing abundance of Sasa because Sasa reduced regeneration on the ground. Density of Abies on elevated sites was higher than that of Picea, irrespective of Sasa and of the density of adult trees. There was no significant difference in growth in sapling trunk height between the two conifers, but Picea grew more slowly under the canopy than Abies and was aggregated into gaps. Thus, in forests with less Sasa, the recruitment capacity of Abies was greater than that of Picea. The long life span of Picea compensated for its low density on elevated sites. Examination of a dynamic system model showed that Picea was excluded by Abies in forests without Sasa because regeneration on the ground is more advantageous than on elevated sites, but the two conifers could coexist in forests with Sasa because of the increased relative success of regeneration on elevated sites by Picea saplings.