Forest gaps regulate seed germination rate and radicle growth of an endangered plant species in a subtropical natural forest

The survival rate of Castanopsis kawakamii from seed to seedling is relatively low,leading to difficulties in the regeneration of its natural forests.Forest gaps play a vital role in plant regeneration and biodiversity maintenance in forest ecosystems.Unfortunately,our understanding of the effects of gap size and within-gap position on the seed germination and radicle growth of C.kawakamii is still limited.In particular,our knowledge on the relationship between gap size and environmental factors and their influence on seed germination and radicle growth is incomplete.In the present study,we studied the influences of forest gaps and within-gap position on seed regeneration on the germination and radicle growth of an endangered species C.kawakamii in a subtropical natural forest in China.We selected three large gaps(LG,gap size above 200 m^(2)),three medium gaps(MG,gap size 50-100 m^(2)),three small gaps(SG,gap size 30-50 m^(2)),and non-gap(NG),and planted the seeds of C.kawakamii in five positions within each gap.The results showed that(1)the influence of forest gaps on seed germination rate was,from highest to lowest,medium gaps(51%),non-gap(47%),small gaps(40%)and large gaps(17%),and the seed germination rate was the highest in all positions in medium gaps,with the exception of the east position.(2)Radicle length in forest gaps was,from highest to lowest,medium gaps,small gaps,large gaps and non-gap,and it was the highest in the east,south,west and north positions of medium gaps.(3)Canopy openness(gap size)and air temperature were the main factors influencing seed germination and radicle growth of C.kawakamii.We concluded that medium-sized gaps were the most suitable for seed germination and radicle growth of C.kawakamii,and they promote the regeneration of this endangered species in the investigated natural forest.

Response of ground beetles(Coleoptera:Carabidae)to forest gaps formed by a typhoon in a red pine forest at Gwangneung Forest,Republic of Korea

In Korea, damaging typhoons related to climate change have increased steadily since the 1990s. Red pine （Pinus densiflora） forests in Gwangneung Forest were greatly disturbed by typhoon Kompasu in 2010. A survey was carried out to clarify differences in ground beetle （Coleoptera： Carabidae） communities between forest gaps and undamaged forests. Ground beetles were sampled using pitfall traps from early May to late October 2011. Vegetation changes, litter layer, organic matter layer, and soil conditions were also measured. A total of 1035 ground beetles of 32 species were collected. Contrary to our expectation, species richness, abundance, and community structure of the ground beetles in forest gaps were similar to those in undamaged forests. Species richness and abundance of habitat type were also similar. However, species diversity and estimated species richness in forest gaps were significantly higher than in undamaged forests. These findings suggest that forest gaps formed by a typhoon did not lead to great change in ground beetle communities.

The effect of low-temperature event on the survival and growth of Juglans mandshurica seedlings within forest gaps

In forest ecosystems, gap formation changes the allocation of abiotic resources and thus affects the survival and growth of understory plants. However, how tree seedling survival and growth respond to low-temperature events and the influencing mechanisms remain unclear. To clarify how low-temperature event limits the survival and growth of tree seedlings in the montane regions of eastern Liaoning Province, northeast China, we investigated temperature and light intensity within secondary forest gaps, and the survival and growth of Juglans mandshurica seedlings after a low-temperature event in the spring of 2014. Damage to seedlings due to low temperature sig- nificantly varied in different aspects. Seedlings in gaps on southeast-facing slopes were the most seriously damaged, followed by those in gaps on northeast-facing slopes. In contrast, seedlings in west-facing gaps and in control plots without slope aspect were not damaged. The freezing injury index for seedlings was negatively correlated with minimum temperature （r = - 0.608, P 〈 0.01）, but it was positively correlated with light intensity （r= 0.818, P 〈 0.01）. In addition, height and root collar diameter of damaged seedlings were significantly lower than those of the undamaged seedlings （P 〈 0.01） during the early growing season （April-July）, but no significant difference were observed during the late growing season （July-Oc- tober） （P 〉 0.05）. The extent of seedling damage was directly related to slope aspect. Low temperature and high light intensity were found to be the dominant factors affecting extent of damage to seedlings on southeast- and northeast-facing slopes.

Distribution characteristics and succession regulation of the forests in alpine and canyon region of western Sichuan Province, P.R.China

Since 1950, 700 plots were established in the alpine and canyon region of western Sichuan. The distribution charac-teristics and the relationships between forest succession and environmental gradients were studied. The results showed that the main tree species were Picea and Abies in this region, and there were more than 90 forest types. Abies forests mainly dis-tributed in the middle and upper reaches of rivers and their branches, and Picea forests mainly distributed in wide valleys and on half-shaded and half-sunny slopes. The natural regeneration was poor under primitive spruce and fir forest canopy, but was good in the spruce and fire forest gap. The relationship between forest succession and vertical gradient was closely related to the relationship between forest succession procession and plant synusia under primary forests. Human activities could promote and postpone succession process. The results of expanding regeneration were often influenced by topography, vegetation and wind direction.

Microclimate regimes following gap formation in a montane secondary forest of eastern Liaoning Province, China

In order to improve the understanding of the role of a canopy opening/gap on the physical environments in a secondary forest in Northeastern China, a case study was conducted in and around a small irregular gap in a montane secondary forest. The secondary forest, which was severely disturbed by human beings about 50 years ago, was dominated by Quercus mongolica and Fraxinus rhynchophyllaan. Temporal variation in photosynthetic photon flux density （PPFD）, air temperature （TA） at 10 cm above the ground, soil temperature （Ts） and soil water content （SWC） at top-layer （0-15 cm） and sub-layer （15-30 cm） were measured from May to September after the second year since the formation of the small gap （the ratios of gap diameter to stand height were less than 0.5） in 2006 respectively. Results indicated that the highest value of PPFD occurred at the northern edge of the gap, particularly at the beginning of the growing season in May. On sunny days, the highest value of PPFD appeared earlier than that on overcast days. Maximum and mean values of TAwere higher in the northern part of the gap, and the minimum values of TAwere at the southern edge of the gap. Soil temperature varied obviously in the gap with the range from 1 to 8 ℃. Maximum values of Ts occurred at the northern part of the gap, which was significantly correlated with the maximum values of TA （R = 0.735, P〈0.05）. SWC was higher in the top-layer （0-15 cm） than that in sub-layer （15-30 cm）, but the difference of them was not significant （p〉0.05）, which might be attributed to the small gap size and the effects of aboveground vegetations. From these results, the maximum of PPFD in the study area occurred at the northern part of the gap, which was consistent with the results observed in north hemisphere, but the occurrence time varied with the differences of the latitudes. The highest values of air and soil temperatures also occurred in the northern part of the gap because they were affected by the radiation. However, the variation of temperature in July was different from other months due to the influence of gap size. And the values of soil water content were neither higher in the gap in the wet season nor lower in the dry season, which might be affected by the gap size and topography the gap located. The variations of light, soil and air temperatures, and soil moisture in this small irregular gap might be related to the effects of the micro-site, which affects the regeneration of plant species.

Natural regeneration characteristics of Pinus sylvestris var. mongolica forests on sandy land in Honghuaerji, China

Natural regeneration in Mongolian pine, Pinus sylvesttis var. mongolica, forest at Honghuaerji of China （the original of the natural Mongolian pine, forest on sandy land） was studied in 2004. The total mean values of regeneration indexes were higher in mature stands （more than 80% individual stems were older than 50 years）, the maximum of regeneration index reached 29 seedlings, m^ 2, with lowest values in the younger stand, e.g., in 32-year old and 43-year old stands. The stand age was an important factor determining the natural regeneration, which was the best in the older stands in this investigation （e.g. about 80-year old）. The regeneration index seemed not to be closely in relation to canopy openness although Mongolian pine is a photophilic tree species. In each type of gaps, natural regeneration was very well. Regeneration indexes were satisfactory at the south and east edges in the circle gaps; and at the east edge of the narrow-square gaps. Results indicated that Mongolian pine, seedlings could endure shading understory, but it would not enter the canopy layer without gap or large disturbance, e.g., fire, wind/snow damage or clear cutting etc. These results may provide potentially references to the management and afforestation of Mongolian pine, plantations on sandy land in arid and semi-arid areas. Researches such as the comprehensive comparisons on regeneration, structure and ecological conditions and so on between natural Mongolian pine, forests and plantations should be conducted in the future.

Effects of gaps on regeneration of woody plants: a meta-analysis

Forest gaps, openings in the canopy caused by death of one or more trees, have a profound effect on forest regeneration and drive the forest growth cycle. It is therefore necessary to understand the effects of forest gaps on regeneration for modern forest management. In order to provide a quantitative assessment of the effects of forest gaps on regen-eration of woody plants, we conducted this review of gap effects on woody plant regeneration on the basis of 527 observations from 42 indi-vidual papers, and reported the results of these data in a meta-analysis. Overall, densities of regenerated woody plants were significantly greater （359%） in forest gaps than on the closed-canopy forest floor. The regen-eration density in gaps of plantation forests was significantly greater （P＆lt;0.05） than that of natural forest because the regeneration in gaps of plan-tation forests was improved by both gap effects and experimental meas-ures. Similarly, in comparison to natural gaps, regeneration was better enhanced in artificial gaps. Regeneration density exhibited a significantly positive correlation with gap size, but a negative correlation with gap age because the gap size decreased with increasing gap age. Shade tolerance of woody plants affected regeneration density in gaps and understory. Average regeneration density of shade-tolerant species exhibited a sig-nificantly positive response to gaps but densities remained lower in total than those of intermediate and shade-intolerant species. Gap effects on regeneration decreased in response to increasing temperature and pre-cipitation because of the limiting effects of lower temperature and moisture on woody plant regeneration. In summary, forest gaps enhance woody plant regeneration, and the effects of gaps varied by forest type, gap characteristics, environmental factors and plant traits. The results of this meta-analysis are useful for better understanding the effects and roles of gaps on forest regeneration and forest management.

Succession Features and Dynamic Simulation of Subalpine Forest in the Gongga Mountain, China

The Gongga Mountain of eastern Tibet Plateau is a representative of the alpine regions with high peaks and deep valleys. Climate change over the last thousand years has controlled the dynamics of glacier and debris flow occurrence, which resulted in substantial changes in the mountainous environment. The authors surveyed the community structure of primary forests in Gongga Mountain and forest succession processes in woodland plots. The changing features in the subalpine environment are discussed in this paper. Tree species and sizes between the glacier shrinking areas and debris flow fans in Hailuogou Valley are compared. The pioneer species that settle in debris flow fans and the glacier shrinking areas are Salix spp. and Populus purdomii. Abies fabri and Picea brachytyla are the climax tree species. The succession process of primary vegetation in Hailuogou (2700 ~ 3200 m) can be divided into four stages: Slash surface (20~ 200 yr) Salix-Populus seeding community (10 ~ 30 yr) Populus-Salix sapling community (30 ~ 100 yr) Populus-Abies mixed community (50 ~ 100 yr) Abies-Picea climax In a natural and undisturbed environment, trees compete for light, water and nutrients. Disaster disturbance in mountains is a very important driving factor for regeneration of woody plants. Repeated destruction of forests by glacier movement or debris flows generated additional forest gaps that allow young plants to grow. In this study the Gongga Forest Succession Model (GFSM) was developed for simulation of forest community succession processes on different scales in Gongga Mountain. A soil succession module was added to the GFSM model to simulate soil formation and chemical element change of woodland. In order to represent major features of forests in Southwestern China, many field works has been done to identify ecological parameters of various trees in the subalpine region. On the basis of simulation of tree life history, the GFSM combines forest succession with soil change in both material components and nutrition content. The Monte-Carlo method was applied to simulate random weather fluctuation and the uncertainty of tree death. These modeled processes agreed with the field investigation results in this region. The elevation distribution of different tree species was also simulated; and the results are consistent with field observations of ecological features of tree species. The modeling approach reflects well the succession dynamics of primary forests in Southwestern China. These results are very useful for improving the management policies and prediction technology for restoration and conservation of primary forests in Southwestern China.

The Altitudinal Belts of Subalpine Virgin Forest on Mt.Gongga Simulated by a Succession Model

How to accurately simulate the distribution of forest species based upon their biological attributes has been a traditional biogeographical issue.Forest gap models are very useful tools for examining the dynamics of forest succession and revealing the species structure of vegetation.In the present study,the GFSM(Gongga Forest Succession Model) was developed and applied to simulate the distribution,composition and succession process of forests in 100 m elevation intervals.The results indicate that the simulated results of the tree species,quantities of the different types of trees,tree age and differences in DBH(diameter at breast height) composition were in line with the actual situation from 1400 to 3700 MASL(meters above sea level) on the eastern slope of Mt.Gongga.Moreover,the dominant species in the simulated results were the same as those in the surveyed database.Thus,the GFSM model can best simulate the features of forest dynamics and structure in the natural conditions of Mt.Gongga.The work provides a new approach to studying the structure and distribution characteristics of mountain ecosystems in varied elevations.Moreover,the results of this study suggest that the biogeochemistry mechanism model should be combined with the forestsuccession model to facilitate the ecological model in simulating the physical and chemical processes involved.

Species Richness, Diversity and Density of Understory Vegetation along Disturbance Gradients in the Himalayan Conifer Forest

We investigated whether species richness, diversity and density of understory herbaceous plants differed along logging(gap) and grazing(primarily by cattle) disturbance gradients, and sought to identify drivers of richness, diversity and density of understory vegetation of logged sites. A factorial experiment was conducted in the mixed conifer forest of Gidakom in Western Bhutan. Levels of the logging treatment included small(0.15 – 0.24 ha), medium(0.25 – 0.35 ha) and large(0.36 – 1.31 ha) gaps. The grazing treatment included grazed(primarily by cattle) and ungrazed(where herbivores were excluded by a fence) plots nested within each gap. Data were collected from 12 gaps(4 replicates at each level of logging) using the point intercept method. Shannon Weaver Diversity and Margalef's indices were used to estimate species diversity and describe species richness, respectively. Soil samples were analyzed for pH and nutrients. The interaction effect of logging and grazing was significant(p≤0.001) only on species diversity. Relative to ungrazed areas, species diversity was significantly higher(0.01≤p≤0.05) in medium grazed gaps. Under grazed conditions, soil P was negatively correlated with gap size and species diversity. While species diversity was positivelycorrelated(0.01≤p≤0.05) with soil N in grazed plots species richness was positively correlated(0.001≤p≤0.01) with soil N in ungrazed plots. Relative density of Yushania microphylla and Carex nubigena were higher under ungrazed conditions. Our study suggests that the combined effect of cattle grazing and logging results in higher species diversity of understory vegetation in medium and grazed gaps in mixed conifer forests of Bhutan,whereas increase or decrease in relative density of major species is determined primarily by the independent effects of grazing and logging. From management perspective, forest managers must refrain from creating large gaps to avoid loss of nutrients(mainly P and N), which may eventually affect tree regeneration. Managers intending to maintain understory vegetation diversity must consider the combined effects of grazing and logging, ensuring low to moderate grazing pressure.

冬季森林鸟类对林窗的响应分析

Although forest gap ecology is an important field of study, research remains limited. By plot setting and point counted observation, the response of birds to forest gaps in winter as well as bird distribution patterns in forest gaps and intact canopies were studied in a north tropical monsoon forest of southwestern China from November 2011 to February 2012 in the Fangcheng Golden Camellia National Nature Reserve, Guangxi. The regression equation of bird species diversity to habitat factor was Y1=0.611＋0.002 X13＋0.043 X2＋0.002 X5-0.003 X8＋0.006 X10＋0.008 X1 and the regression equation of bird species dominance index to habitat factor was Y3=0.533＋0.001 X13＋0.019 X2＋0.002 3（3-0.017 X4＋0.002 X1. There were 45 bird species （2 orders and 13 families） recorded in the forest gap, accounting for 84.9% of all birds （n=45）, with an average of 9.6 species （range： 2-22）. Thirty-nine bird species （5 orders and 14 families） were recorded in non-gap areas, accounting for 73.6% of all birds （n=39）, with an average of 5.3 species （range： 1-12）. These results suggested that gap size, arbor average height （10 m from gap margin）, arbor quantity （10 m from gap margin）, shrub quantity （10 m from gap margin）, herbal average coverage （1 m from gap margin） and bare land ratio were the key forest gap factors that influenced bird diversities. On the whole, bird diversity in the forest gap was greater than in the intact canopy. Spatial distributions in the forest gaps were also observed in the bird community. Most birds foraged in the ＂middle＂ and ＂canopy＂ layers in the vertical stratification. In addition, ＂nearly from＂ and ＂close from＂ contained more birds in relation to horizontal stratification. Feeding niche differentiation was suggested as the main reason for these distribution patterns.

Effects of gap size on diversity of soil fauna in a Cunninghamia lanceolata stand damaged by an ice storm in southern China

Cunninghamia lanceolata （Lamb.） Hook. is economically the most important tree species in southern China and has been cultivated in plantations on a large scale. This species is widely used in construction, furniture, utensils and shipbuilding. Soil fertility of C. lanceolata stands affects soil structure, porosity and nutrient availability, which causes changes in fauna activity. During January to February 2008, an ice storm caused extensive damage to C. lanceolata stands. Despite the environmental importance of soil fauna, basic information on the distribution and diversity of soil fauna in C. lanceolata stands after ice storm damage is lacking. To assess the response of soil fauna diversity and distribution to forest gaps following the ice storm, five small gaps （each 30-40 m^2）, five large gaps （each 80-100 m^2） and five canopy cover plots were selected within a 2-ha C. lanceolata stand. Soil samples were collected from 0 to 10 cm depth in March 2011 to measure soil fauna diversity and abundance. The abundance and community composition of the soil fauna varied with gap size. In canopy cover sites, the number of individuals was 2.0 and 5.2 times greater than in the small gaps and large gaps. Three taxa （Nematoda, Oribatida and lnsecta） of soil inver- tebrates occurred commonly, and Nematoda dominated the communities in all three habitat types. The Shannon-Wiener diversity index, Margalef diversity index, and Pielou evenness index were high in the small gaps, indicating that they harbored the most species, with the most even distribution, and the highest diversity. Our results indicated that gap size apparently affected abundance and community composition of the soil fauna.

Loss of total phenols from leaf litter of two shrub species: dual responses to alpine forest gap disturbance during winter and the growing season

Aims Alpine forest gaps can control understory ecosystem processes by manipulating hydrothermal dynamics.Here,we aimed to test the role of alpine forest gap disturbance on total phenol loss(TPL)from the decomposing litter of two typical shrub species(willow,Salix paraplesia Schneid.,and bamboo,Fargesia nitida(Mitford)Keng f.).Methods We conducted a field litterbag experiment within a representative fir(Abies faxoniana Rehd.)forest based on‘gap openness treatments’(plot positions in the gap included the gap center south,gap center north,canopy edge,expanded edge and closed canopy).The TPL rate and litter surface microbial abundance(fungi and bacteria)of the two shrub species were measured during the following periods over 2 years:snow formation(SF),snow cover(SC),snow melting(ST),the early growing season(EG)and the late growing season(LG).Important Findings At the end of the study,we found that snow cover depth,freeze–thaw cycle frequency and the fungal copies g−1 to bacterial copies g−1 ratio had significant effects on litter TPL.The abundances of fungi and bacteria decreased from the gap center to the closed canopy during the SF,SC,ST and LG periods and showed the opposite trend during the EG periods.The rate of TPL among plot positions closely followed the same trend as microbial abundance during the first year of incubation.In addition,both species had higher rates of TPL in the gap center than at other positions during the first winter,first year and entire 2-year period.These findings suggest that alpine forest gap formation accelerates litter TPL,although litter TPL exhibits dual responses to gap disturbance during specific critical periods.In conclusion,reduced snow cover depth and duration during winter warming under projected climate change scenarios or as gaps vanish may slow litter TPL in alpine biomes.

Changes in carbon,nitrogen and phosphorus stoichiometry in decaying logs with gap positions in a subalpine forest

Aims The redistribution of forest gaps based on solar radiation and precipitation(mainly rainfall and snowfall)can create heterogenous microenvironments inside and outside the gaps.Here,we investigated the effects of forest gaps on carbon(C),nitrogen(N)and phosphorus(P)stoichiometry in decaying logs by altering the microenvironments.Methods Minjiang fir(Abies faxoniana)logs of decay classes I–V were incubated in situ on the floor under the gap center(GC),gap edge(GE)and closed canopy(CC)in a subalpine forest in the eastern Qinghai-Tibet Plateau.The C,N and P concentrations in decaying bark,sapwood and heartwood were measured in August from 2013 to 2016.Important Findings Our results showed that the N concentration in bark decreased from the CC to the GC,while the opposite trend was found in sapwood and heartwood.Additionally,the C/N ratio in sapwood and heartwood of decay classes IV and V decreased from the CC to the GC,and the N/P ratio in heartwood of decay class V increased from the CC to the GC,implying that forest gaps have strong effects on C/N/P stoichiometry in highly decayed logs.Briefly,highly decayed logs are more susceptible to the microenvironment,and the proportion of highly decayed logs under the CC or the GC should be appropriately adjusted if necessary.

STATISTICAL DETECTION OF SPATIAL PLANT PATTERNS UNDER THE EFFECT OF FOREST USE

The analysis of the consequences of land use （in particular forest use） may be considered as a partial step towards an integrated modeling of a land system. In the paper a forest territory is considered, where a gapcut is made, and after a given time period the eventual change in the spatial distribution of undergrowth plants and tree seedlingsis to be detected. Floristic data are collected along a line transect. A method for the detection of the change in the plant distributions along the transect is proposed to see whether this occurs at the geometric frontier of the human intervention. Since in the considered case the distribution of the changepoint estimate is not known, as a substitute of its confidence interval, the socalled changeinterval is calcu lated, using an adaptation of the bootstrap method. As an illustration, for a concrete plant species, the maximum likelihood estimation of the changepoint and the calcu lation of the above mentioned changeinterval is presented. Finally, the validation of the proposed method against some typical ecological situations is also presented, which provides a justification of the used algorithms.