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Moving from Ecological Conservation to Restoration: An Example from Central Taiwan, Asia
Yueh-Hsin Lo1, Yi-Ching Lin2, Juan A. Blanco3, Chih-Wei Yu4 and Biing T. Guan1,*
1National 2Tunghai

Taiwan University, University, 3University or British Columbia 4New Taipei City Government 1,2,4Taiwan 3Canada

1. Introduction
The concept of “natural conservation” has been evolving since the beginning of the first efforts to preserve the natural landscape. The creation of the first national parks in the 19th century was originated by the belief that landscapes of exceptional beauty should be preserved from human influence and maintained in their current state for the enjoyment of future generations (Runte, 1997). During the first years of the establishment of national parks around the world, defining these “exceptional landscapes” was usually based on the static beauty of the area: majestic mountains, glaciers, old forests, gorges, canyons, waterfalls, etc. The protection in these areas was basically achieved through the prevention of creating human structures in the sites, reducing and controlling human activity and, in practice, maintaining the areas as they look at the time when they were declared as protected. Therefore, this protection was not based on ecological considerations, but on a human-centered view of natural sites. The type of protected ecosystems varied widely among regions, depending on the history of human impact on them. For example, national parks in North America were created to protect largely untouched, almost pristine landscapes practically unaffected by the low populations of native peoples previous to European contact (Runte, 1997). Similarly, in South America and Africa, large natural areas could still be found during the 19th and 20th centuries were the human impact was thought to be minimal. However, in Europe or Asia, were the history of urban development can be traced back for millennia and the density of population is also higher, it was more difficult to found those untouched areas. As a consequence, national parks were created to protect landscapes of indisputable beauty but usually with a noticeable human influence on them. Ecosystems at this time of early
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Corresponding Author

or the Pyrenean wild goat. The improvement of this species-oriented conservationism from the first days of creation of protected areas is that it recognizes individual species as worth of the preservation. lions. 2010). 2007. with the arrival of modern ecology.. This danger was highlighted by scientists and environmental managers around the world. 1989). Odum. it was just a matter of time that some of the iconic wildlife species of the world started to suffer from fast reductions in their populations. natural conservation was seen by the main public as “avoiding things getting worse”. 2001).5% in total area and 1. such in the case of “ugly” species as it is amphibians.. cacti.. Folch et al. etc. As a consequence of this human-centered vision to define the areas to protect... such as the halt in commercial hunting of whales (Stevick et al. etc. swamps. Some of these activities have achieved important successes. that should not be altered or they would lose their integrity. However. the Yangtze River dolphin. even if they are not in “beautiful landscapes” with some sort of legal protection. gorillas. arriving to a single climax state (Clements. Protected areas overall remain a core element of biodiversity conservation (Andam et al. However.. and as a consequence.4% in numbers of sites. it moves one step from the human-centered conservation of some specific favourite areas to protect species and control the factors that affect their populations. sequoias. Sanderson et al. 2008). 2004. or even facing extinction. 1916. in other cases the protection of the target species was not enough to prevent its decline or extinction (e. Gaston et al. 2009). Pickett and McDonnell. and therefore the efforts were oriented into maintaining it. elephants. was and still remains very real (Laliberte & Ripple.. increasing yearly by an average 2.. Many of these campaigns were supported by the public due to the easy sympathy or spiritual connection with some of these majestic species. 2001) or the increase in numbers of American buffalos (Waldman. However. The danger of losing species such as whales. deserts. This climax state was usually identified with the state that nature reaches in absence of human influence. 2008). regions such as oceans. these regions are still underrepresented in the protected areas around the globe (Noss et al. Therefore. During the 20th century urban development was extended to all the regions of the world. reptiles. 1969. 2008. the main drawback of this type of ecological conservation is that it is targeted to one species. Therefore. and the society responded with the creation of environmentalist groups. insects. brown bears. or just the species were not interesting enough for the public opinion and therefore not the main focus of protection efforts. not to the ecosystem that supports that species.000 designated sites (Butchard et al. panda bears. On the other hand. this static view was gradually substituted by the classic view of gradual linear change along a continuum. Since the Rio Earth Summit in 1992. Even today. shrublands and other similar ecosystems were usually considered “badlands” and unworthy of legal protection. the breeding programs of panda bears (Peng et al.g. 2003). With a booming human population and the intensification of economic development (first in Europe and North America and lately in the rest of the world) practically all the ecosystems in the world were impacted in one way or another.. 2001).. buffalos. Turvey et al.340 Forest Ecosystems – More than Just Trees conservation efforts were seen as static entities. 1995). This was the base to develop programs and activities focused on the protection of individual high-profile species. This species-oriented conservation followed the theory that if the . and by 2006 covering more than 24 million km2 in about 133. tigers. whose social pressure helped to create lists of endangered animal and plant species needing specific actions for conservation. very few protected areas have been established in productive landscapes with clear economical potential for agriculture or forest management (Scott et al. the global network of protected areas has continued to grow steadily.

animal or microorganism species will survive in a given ecosystem as long as the right conditions exist to support the niche that the species inhabits. the target species could survive or even increase its population. harvesting. Asia 341 causes of non-natural mortality are controlled (i..Moving from Ecological Conservation to Restoration: An Example from Central Taiwan. arboretums and other centers where collections of plant and animals are kept under controlled conditions are an important part of this strategy. soil alteration and erosion. practices. any plant. or destroyed. In addition. etc.. clear-cutting. removal of invasive species. The goal of ecological restoration is therefore the reestablishment of the characteristics of an ecosystem. The main implication of this conceptual model is that ecosystems that have been altered by human activity may not revert back to its original state if left alone. these altered ecosystems could just reach a different stable state defined by the actions of human management on them (i. in very long time scales) by the ecosystems if left alone. the idea of the balance of nature has been replaced with the flux of nature (Wu & Loucks. As a consequence. a more holistic approach to conservation has emerged since the last quarter of the 20th century. Ecological restoration involves assisting the recovery of an ecosystem that has been degraded. Within this approach. 1995. it is not a passive conservation effort in which humans are just consider outsiders that should not be “in the way” of Nature. Their dynamics are not only complex but also dependent on the spatial context and the history of natural disturbance and human influence (Hobbs & Cramer. damaged. and the availability of resources increased. 2008). This new approach is in the origin of the last trend in conservation: ecological restoration. Ultimately.). and policies have shifted over time recognizing this need to preserve the ecosystem and not just the target species (Adams. stopping illegal logging. etc. modification of the physic-chemical environment. 2000). Examples of such alternative states are grasslands or forest dominated by invasive species or shrublands that substitute forests. Ecological restoration is different from the earlier protection of specific areas because “restoration” means human intervention to bring the ecosystem back to a state different from the one in which it is currently. and they could be ineffective if the ecosystem is too altered to keep the target species. Quite differently. 1987). Ecological restoration is based on the new view of ecosystems as biological communities established on a geophysical substrate that can develop into alternative stable states rather than into a single climax state (Lewontin 1969).). actions such as banning hunting. plantation of trees and plants. On the contrary. poaching. etc. 1998). and ecosystems are thought to be mostly in non-equilibrium. lost of native species. typically as a result of human activities (Sala et al. and that will not be reached (or if so. ecological restoration needs direct human actions (i. actions that could be beneficial for the target species are not necessary relevant for other species in the ecosystem. hunting. 2005).e. Pickett & Ostfield. Conservation paradigms. invasive species.. introduction of lost species. . As a consequence. changes in hydrological regime. 1995. Therefore.e.). 2009). However. Wallengton et al. as they provide research insights in the biology of the species and they can specially increase the population sizes of plants and animal species (Bagarinao. even after removing the human factors directly affecting it. poisoning.e.. the actions in the conservation effort will be directed to keep the integrity (bio-physical diversity) and the functionality of the ecosystem. ecological restoration is different from conservation of emblematic species in that restoration targets the whole ecosystem.. controlling access to the areas were the species is distributed can be part of this strategy (Folch et al. herbariums. In addition. 2004). zoos. Therefore. such as biodiversity and ecological function that were prevalent before degradation (Jordan et al.

both within and outside protected areas. there is an urgent need to greatly expand the scale of ecosystem restoration for both conservation and production (Naveh. protected nature reserves (Naveh... 1996). 2010). Although small-scale restoration projects can be valuable. Actions such as targeted habitat management. the emblematic species (and their companion species) will be preserved in the restored area. A meta-analysis of 89 restoration assessments in a wide range of ecosystem types across the globe indicated that ecological restoration increased the provision of biodiversity by 44% and ecosystem services by 25%.. Soulé & Terborgh 1999). (Hsu & Agaramoorthy. Hence. it is clear that detailed field research is needed to guide restoration efforts. seed production and species reintroduction have yielded notable successes: among many examples. we seek to evaluate whether the combination of selective cutting.000 vascular plants in six different forest types (Boufford et al.342 Forest Ecosystems – More than Just Trees assuming that if the correct ecological conditions are maintained. and with Jade Mountain (the highest peak). Nelleman & Corcoran. removal of invasive species. 2011). Hobbs & Norton. 2010). In this study. The island has a very complex terrain. Taiwan harbours over 4. site and situation specific (Hobbs & Norton. and understory vegetation control can be a cost-effective method to gradually restore plantation forests to native forests. 1999). 2006). 2007. Ecological restoration. 1996). direct seeding..l. An example from central Taiwan 2..s. captive breeding. the effectiveness of restoration actions in increasing the provision of both biodiversity and ecosystem services has not been evaluated systematically. 400 . Wildlife resources are also abundant with 61 species of mammals. In this chapter we describe the field research done to guide the restoration of native conifer forests in central Taiwan. 1994. 2. an species alien to Taiwan. and separated from the main continent by a strait of 130 km in its narrowest point. Our research has as main objective the identification of the best conditions for seedling establishment of two tree native species: the evergreen Fabaceae Lithocarpus castanopsisifolius and Lithocarpus kwakamii (stone oak) in a former plantation of Japanese cedar (Cryptomeria japonica). 1996). 2006).000 km2 and is located at the fringe of the Asian continental shelf at the western rim of the Pacific Basin. at least 16 bird species extinctions have been prevented by such means between 1994 and 2004 (Butchart et al. reaching 3952 m a. with about two thirds of Taiwan's land area at slopes over 10% and almost half of the island with slopes over 40%. but values of both remained lower in restored versus intact reference ecosystems (ReyBenayas et al. 2009). restoration efforts to date have been criticized for being ad hoc. closeness to the continent and wide gradient of altitudes. Large-scale ecosystem restoration is needed to arrest and reverse the degradation of landscapes around the world (Manning et al. is being increasingly applied worldwide (Clewell & Aronson.1 Historical background Taiwan covers an area of 36. and it is increasingly found as part of natural resource management plans. Ecological restoration has developed quickly since the first meetings of the Society of Ecological Restoration International in the early 1990s (Greipsson. We also seek to identify the potential barriers that hinder seed and seedling survival. As a consequence of its insularity. Field experiments can be a useful guide as to which restoration practices are the most useful for achieving this goal (Kimmins et al. 1994. However. or focusing on small. From these results.

Currently. herbivory.600 m3 harvested from 1965 to 1975. the harvesting-reforestation approach is no longer viable and alternatives need to be devised. Drought. making these stages the bottleneck of ecological restoration (Fenner & Thompson.000 m3 and by 1990. in a gradual manner. 2005). but it is .132 ha) of the island’s total land area (Taiwan Forestry Bureau. at the peak of exploitative management in the island. Lu et al. with plantations of native species and timber harvesting program through the 1950s to the 1970s. seedling planting and direct seeding are two common tools used in forest restoration. Asia 343 species of birds. 1997). timber was harvested mainly from forest plantations with an annual cut of about 100. and camphor. national forest lands are managed almost exclusively for the purposes of streamflow regulation.1% (41. Don) plantations that were established in sites now considered as unsuitable for harvesting. 30 species of amphibians. These levels of harvesting brought petitions from citizens and environmental protection groups urging forest protection. 1997). covering about 1. 99% of Taiwan’s timber supply was imported (Wang. This species was introduced from Japan with the start of the Japanese colonial rule at the end of the 19th century. corresponding to about 18. 2008) To improve seed establishment rates. Till date. To restore and promote biodiversity. primarily cypress. This intensive level of exploitation was essentially halted with the national forestry management policy of 1976 (Wang. Fenner & Thompson. 2005. it is necessary to understand the best ways of promoting native trees establishment.552. Since then. 92 species of reptiles.. most of Taiwan’s Japanese cedar plantations either are approaching or have passed the prescribed rotation age.000 ha cut annually (Lu et al. spruce. 1999). 2000. with an average of 1. and the estimated 50. 1995). the first national park was created in 1984 in Kenting (south Taiwan).. However. mainly due to soil and slope protection concerns. due to increasing production costs and declining timber prices.000 species of insects including 400 species of butterflies (Hsu & Agoramoorthy. Large areas of valuable timber.. After 1977. the emphasis of forest management in Taiwan has shifted almost entirely from timber production to forest protection.. Seed and seedling survival are limited by multiple biotic and abiotic factors (Beckage et al. the current management directives mandate the restoration of plantations no longer serving for timber production back to native forests. 140 species of fresh-water fish. 2001). Economic pressures led to an aggressive management.Moving from Ecological Conservation to Restoration: An Example from Central Taiwan. The protected area covers 12.. The interest on conservation is not limited to natural forests.2% of the total land area of Taiwan (Hsu & Agoramoorthy. were cut and shipped primarily to Japan. and light are the three most important causes for seedling mortality (Leck et al. Under this new approach. 6 national parks. and conservation of biological diversity. erosion control. Among other concerns. It has become the most widely planted tree species in Taiwan. especially in the marginal plantations created during the 50s and 60s. 1997. 2001). One example is the important number of existing Japanese cedar (Cryptomeria japonica D. To protect this rich biodiversity. Leck et al. 2008). but it is also extending into plantation forests. The history of forest conservation and restoration in Taiwan is closely linked to the economic development of the island. 18 nature reserves and 24 nature protected areas have been designated to ensure protection for wildlife and their habitats. Timber harvesting peaked during the Japanese colonial period and immediately following World War II. Knowledge on how to use current forestry practices to accelerate and support the conversion from plantations into native forests is needed to design successful restoration plans in these plantations. The former has the advantage of high success rate.

the creation of an environment that enhances the survival of tree seeds and seedlings is a key element of a successful gradual forest restoration strategy. Leck et al. In addition..s. 2000. On the other hand.344 Forest Ecosystems – More than Just Trees also more expensive than the later (Bullard et al. Brokaw & Busing... Thus. Direct seeding has the advantage in term of cost. 1984. 1.. with a mean annual rainfall of 1500 mm (NTUF.2 Material and methods 2. Fenner & Thompson. Fig.2. partial removal of trees can enhance local light availability and create physical environments similar to natural gaps that are essential for seedling survival (Augspurger. Mean annual temperature of the study site is 19. Selective cutting and thinning are common forestry practices that can also be used for restoration. spatial arrangements of the treatment combinations (b). 2008).1 Experimental site This study was carried out in a 10-ha Japanese cedar plantation in the Heshe District of the National Taiwan University Experimental Forest. 1992).. central Taiwan (120° 52’ E. 2011). During these procedures. 14421602 m a. the site was clear-cut in 1958 and planted with Fig. Therefore. Masaki et al. Therefore. 2005. 2008). 2..8°C. understory vegetation may reduce seedling predation by providing protection (Smit et al. 1). Location of the experimental site.. and the overall stand abiotic environment is not greatly altered. 23° 37’ N. and seed transects (c). but it usually has low success rates. only a portion of trees is removed. . 2006). understory vegetation control may cast both positive and negative effects on the survival of seeds and seedlings (Beckage et al. 2007). 2000. the presence of understory vegetation may reduce seedling survival by reducing light availability or increasing competition between seedlings and understory vegetation (Leck et al. National Taiwan University Experimental Forest – NTUEF (a).l. Originally an evergreen broad-leaf forest dominated by Fagaceae and Lauraceae species.

Species . 20% of the standing volume was harvested to create gaps of different sizes. Due to typhoon damages in 1969.Moving from Ecological Conservation to Restoration: An Example from Central Taiwan. We regard those edge areas as the reference for the restoration project and to set the initial restoration goal: the successful establishment of late succession components of the reference stands in the plantation. Lithocarpus castanopsisifolius (Hayata) Hayata and Pasania kawakamii (Hayata) Schottky were selected as the target species for reintroduction as they are late succession species and relatively abundant in the surrounding areas from were collected seeds. we focused only on the reintroduction of the main Fagaceae species. We established four plots. respectively. remnants of the original vegetation can still be found. 1c). kawakamii were placed along the remaining four transects (Fig. with the hazard defined as the instantaneous mortality risk of a seed (Cox. Thus. we went back on day 140 as the final checking time. For each plot. 2. within the plantation (Fig. As a part of the experiment. Eight infrared automatic cameras. we randomly selected two transects from where we removed the ground vegetation in a strip of 1-m wide along the entire transect (devegetated transects). and they were stored at 4 °C until they were used in January 2009. For each group of transects in each plot (cardinal or inter-cardinal). the plantation was selected as a demonstration site to study how to gradually restore Japanese cedar forests back to native vegetation communities. All fresh seeds used in this study were collected between October and November 2008 from the reference stands in the edges of the Japanese cedar plantation. 2. At day 140. whereas the ground vegetation of other two transects was left untouched (vegetated transects. for a total of 120 seeds per transect and 1920 seeds per species for the entire experiment. one for each species-treatment combination.5 m along each transect. the entire experiment consisted of 4 treatments for each species. After that.2. whereas canopy openness of the two unthinned plots was 13% and 11%. In January 2009 we placed 30 fresh seeds every 2. the number of seeds still present was counted every day during the first 35 days. two thinned and two unthinned. 1c).3 Data analysis Cox regressions were used to analyze the survival and seedling establishment. almost all the seeds were removed or consumed.5-m and 10-m marks.2.2 Materials and experimental design An initial inventory found that while the saplings of late succession Lauraceae species (mainly dispersed by birds) were relatively abundant. starting and ending at the 2. The thinning and understory vegetation removal treatments were used as the explanatory variables. whereas the seeds of P. Within 500-m from the edges of the plantation. only a few saplings of Fagaceae species were present. a 10meter transect was set at each of the 8 cardinal and inter-cardinal directions (Fig. After placing the seeds. Fig. 1c). The seeds of L. therefore the experiment ended at that time. The canopy openness of the two thinned plots was 27% and 29%. In 2005. 1972). Asia 345 Chinese fir (Cunninghamia lanceolata). were also set up to capture how the seeds were removed or consumed and by which animal species under different treatment conditions. the stand was replanted with Japanese cedar in 1971. castanopsisifolius were placed along the cardinal direction transects. 1b). Thus.

. 2. Twenty days since the beginning of the observation.3 Results 2. kwakamii seeds. 2. with survival analysis using the R package Survival (R Core Team.3. and dotted lines represent unthinned treatment. respectively.346 Forest Ecosystems – More than Just Trees were analyzed separately. 82% and 48% of L. Fig. Solid lines indicate thinned treatment. whereas opened squares represent without ground vegetation. castanopsisifolius seeds were missing in the unthinned and thinned plots. These results were similar for the P. respectively. Seed removal rates over a period of 61 days in different thinning and understory vegetation treatments for Lithocarpus castanopsisifolius (upper panel) and Pasania kawakamii (lower panel). 2010). 2). We used R to conduct all statistical analyses. Solid black dots indicate ground vegetation intact. with 95% and 48% seeds disappearing in the unthinned and thinned plots.1 Field observations Seeds were first removed from transects with no ground vegetation cover (Fig.

466 0. castanopsisifolious seeds had disappeared from the vegetated plots after 35 days.3 Seedling establishment Probabilities of successful seedling establishment at day 140 differed among species and treatments. For P.93 P <0. but after 15 days there seemed to be a tendency for a slightly higher probability survival in devegetated plots for both canopy types. removing part of the canopy significantly influenced seed removal (mortality risk) in the study site (Table 1.11 0. seedlings only successfully established in the unthinned plots (Fig. it indicates equal mortality risk.18 0.04 1.509 0.001 0. (Fig.001 0. These two species can be seen as potential dispersers of large seeds in the late succession period.18 0. From the images captured by the automatic cameras.3. seeds in the thinned treatments suffered the highest removal risk (Table 1). kwakamii. These two species were redbellied squirrel (Callosciurus erythraeus) and Owslon's long-nosed tree squirrel (Dremomys pernyi owstoni). However. In addition to the main effects. 3 right panel). Similar results were found for L. 2.65 1. No significant difference was detected between vegetated and devegetated plots. there was a small seedling establishment probability in thinned plots. Ninety-four percent of L.091 <0. For L. Asia 347 Seeds of P.69 -6. castanopsisifolius (Table 1). castanopsisifolious. the treatment producing the biggest differences after 35 days for P. castanopsisifolius seeds. 3) Risk ratio of devegetated relative to vegetation treatment. kwakamii was thinning.2 Seed removal The results from Cox regressions indicated that. Notes: 1) Hazard ratio is defined as the ratio of mortality risk between two factor levels. kwakamii.Moving from Ecological Conservation to Restoration: An Example from Central Taiwan. 2) Risk ratio of thinned relative to unthinned treatment. .2). Effects of thinning and ground vegetation treatments on the seed hazard (instantaneous mortality risk) based on Cox regressions. but the success rate was much higher for unthinned plots. This pattern was not clearly identified for L.09 Z4 -5. 2. the interaction between canopy type and understory vegetation cover was non-significant (Table 1). 3 left panel). for both species. Results indicated that. we identified two mammal species as the acorn consumers/removers during the observation period. If hazard ratio = 1.70 1.3. kwakamii were removed more slowly from the devegetated plots for both thinned and unthinned treatments. with 98% of seeds disappearing from thinned. but this effect disappeared after 23 days of exposure.66 0. versus 91% from devegetated plots. Fig.73 1.351 Table 1. 4) Cox regression coefficient. Treatment Lithocarpus castanopsisifolius Thinning2 Understory removal3 Thinning  Vegetation Pasania kwakamii Thinning Understory removal Thinning  understory removal df 1 1 1 1 1 1 Hazard ratio1 0. a much lower probability of survival than in close canopy plots (84% seeds disappeared after 35 days). for P.05 1.

Probabilities of successful seedling establishment after 140 days of seed planting for Lithocarpus castanopsisifolius (left panel) and Pasania kawakamii (right panel). canopy and ground vegetation conditions were observed. In contrast. The understory vegetation was rather dense in the study site. they suggest that keeping canopy vegetation intact may be more effective than removing understory for the two species studied. Error bars represent the mean ± standard error. The improved light conditions due to . The thinning treatment reduced seedling establishment in both species and significantly reduced seedling survival in both species. castanopsisifolius. seedling establishment was not significantly affected by removing ground vegetation. Survival rates for the unthinned plots were consistently higher than for the thinned plots in either vegetated or devegetated after two weeks of seeding (Figure 2). although understory removal showed a tendency to increase seed survival in L. Despite that our results are species-specific. 3.348 Forest Ecosystems – More than Just Trees Fig. differences in seed survival between the two vegetation types did not show consistent patterns and the difference was rather small. especially in the thinned plots. 2. although statistically not significant. The observed differences in seed survival may arise from changes in foraging behaviours of seed predators as a result of the forest management treatments. In addition. The understory grew rapidly after selective thinning was conducted in 2005.4 Discussion Differential seed survival and seedling establishment among species. Such results suggest that removing the understory layer may not be as effective as keeping the canopy cover.

including red-bellied Squirrel and Owslon's long-nosed tree squirrel. where instead of consuming the seeds. while predation rates by field mice. 2004). 2004. and the species-specific responses of seedling establishment to management activities provide a clear indication of the need to shift from a static conservation in which the “no human action” approach is favoured into a more active restoration strategy. kwakamii. and they could be a future research line at these sites. and Machilus thunbergii. however. Therefore.Moving from Ecological Conservation to Restoration: An Example from Central Taiwan. Hulme & Kollmann. many rodent species. 2004). 2010). 2004). which was mostly composed by broadleaf species. Schnurr et al. which are shadetolerant but can take advantage of increased light levels (C. Seed predation may increase or decrease among different microhabitats depending on the composition of seed predators (Schupp et al. 2004. and in a lesser way P. The effect of both canopy structure or understory vegetation density on seed survival. Our experiments are the first ones of these kind in Taiwan.. For example.. For instance.. the openness of the canopy layer can also influence the behaviour of seed predators (Boman and Casper.-W. the movements of mammals were monitored in a nearby plantation. 2004. 2002. personal observation). such as Schefflera actophylla. Yu. Seed tracking techniques are required to study the details of scatter-hoarding behaviours (Forget et al. thinning could reduce seed survival by favouring understory growth and then providing more protection for seed predators. but in which P. den Ouden. Scatter-hoarding behaviour has been widely observed in squirrels and jays in various forest ecosystems. Zhang et al. Mammalian activities were decreased immediately after thinning treatments (Lin & Bridgman. In addition of these two species. To try to understand the relationship between seed predators and seedling establishment. kwakamii will remain mostly suppressed. 2004).. such as red-bellied squirrels or the Formosan field mouse have been observed to eat seedlings (Young. Hulme & Kollmann.. Peromyscus mexicanus are higher in canopy gaps (Tamura & Katsuki. In addition. 1994. personal observation). seed predation by squirrels is reduced in canopy gaps. 2005). Forget et al. Therefore. and our preliminary results indicate that for other species the effects of thinning and understory removal can be the opposite (data not shown). Fenner and Thompson. Asia 349 thinning were believed to facilitate the rapid growth. Tamura & Katsuki. 1995. castanopsisifolious will successfully regenerate. 2004. the photos taken from our automatic cameras in our plots also suggested that major seed predators in the study sites were granivorous animals. the future tree composition of these stands could be a multi-story canopy in which the Japanese cedar (an alien species) dominates the canopy. Tamura & Katsuki. is somewhat context-dependent (den Ouden. we are carrying similar experiments for other tree species at the same plots. with lower layers of L. 2004. the new state of this forest under a conservation-only strategy would not be . as well as the effects of understory vegetation (Chambers and MacMahon. In addition. As a consequence. This may be the case in our study. Under this scenario. they moved seeds away and cached the seeds (Vander Wall. which showed increases in seed predation in conditions with higher canopy openness after thinning. they can be reducing the seed survival but increasing seed dispersal rates at the same time. 2004). Some of the animals observed by the automatic camera may also function as seedling predators. 2004). The cached seeds may germinate at later dates. castanopsisifolius and other similar native species. These animals may also become seed dispersers via scatter-hoarding behaviour. 1994. it can be expected that L. If no action were taken.

Only after a major natural disturbance (i. soil moisture and species-specific factors for trees (seed size. if this “new ecological state” is to be avoided and the recovery towards the original mixed forest is needed. kwakamii should be implemented in these stands to ensure that P. seed preference) can make the same management have very different results in restoring different sites. 3. We suggest that other similar programs monitoring seed survival and seedling establishment should be enacted in other forest regions around the world. rare in the seed rain of the planted forest after thinning (Sun. to facilitate the transition from planted forest to native forest. . Therefore. Also. a program of seeding combined with additional research to improve the regeneration of P. The interaction between light availability. With extensive plantation forests in Taiwan. Biing T. these activites should be localized and not general through the stands to avoid the inhibitory effect that close canopy or exposed forest floor could have for the regeneration of other important native species. we warn the readers from assuming that the results presented here could be applied to other forests types or regions. Acknowledgements The authors thank Dr. Our study indicated that for the two species studied. seems to be species-specific. the management practices could be widely applied to facilitate the regeneration of native species. our research shows how the shift from passive ecological conservation towards a more active ecological restoration can be successful if enough ecological information on ecosystem structure and function is available. a typhoon or a stand-replacing wildfire) this hybrid stand could be transformed into the original mixed forests. Conclusions The native forests within the elevational range in this experimental area were dominated by Fagaceae and Lauraceae species. The efficiency of these management techniques. seed dispersal) and seed predators (foraging behaviour. 4. however. Such results suggested a high degree of recruitment limitation in the plantation. generating a new “hybrid” state with elements from both the human-altered (plantation) and original (mixed forest) ecosystems. These families were. Therefore. it is essential to develop management strategies to overcome recruitment limitation of native species in the plantation forest. if no active restoration is implemented. The next step is to apply such treatments to a broader area to assess the operational costs of such management techniques. 2010). We also thank National Taiwan University Experimental Forest for their help during the field work period. Other research has shown opposite effects of thinning and vegetation removal in other species at the same sites (data not shown). however. but a hybrid of both. However. Therefore. the ecological barriers for seedling establishment could prevent these ecosystems from recovery for a long time. suggesting an easy and inexpensive mean for forest restoration.350 Forest Ecosystems – More than Just Trees either the Japanese cedar plantation or the original mixed forest. especially in tropical forest where little is known about many of the native tree species. Guan and Dr. Po-Jen Jiang for his assistance in setting up automatic cameras.e. This study was supported by a grant to Dr. All things considered. Yiching Lin by the Taiwan National Science Council (NSC 97-2313-B-002-041-MY3). keeping the canopy cover could be an effective management tool to overcome recruitment limitation. kwakamii finds favourable conditions to regenerate.

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