Functional performance of oak seedlings naturally regenerated across microhabitats of distinct overstorey canopy closure

The extent to which seedling recruitment is limited by summer drought in Mediterranean-type ecosystems depends on the light microsite, yet the relationship between light availability and water status, functional performance, and survival of seedlings in these systems is still unclear. Over a 3-year period, we studied the pattern of survival and functional performance of seedlings of Quercus petraea (Matt.) Liebl. and Quercus pyrenaica Willd. in a montane forest in central Spain, which is the southern edge of the natural range of Q. petraea. After a mast year of the two species, 72 plots were established in six microhabitats spanning a range of overstorey canopy closure: closed, partial and open canopies dominated by either Q. petraea or Q. pyrenaica adult trees. Seedlings of each species naturally emerged beneath the conspecific-dominated canopies. The second and third years of study were extremely dry. Three years after emergence, the greatest seedling survival occurred beneath the partial canopy of Q. pyrenaica trees (8%) and the lowest (0%) beneath the closed canopies of Q. pyrenaica and Q. petraea. Survival for Q. pyrenaica increased linearly with understorey light across the range of 10–35% Global Site Factor. Plant water deficit (estimated by leaf water potential) was high across microhabitats, and increased with light availability for Q. pyrenaica. Potential for photosynthesis (estimated by the electron transport rate of photosystem II) decreased with canopy closure; and potential for light harvesting (e.g. specific leaf area (SLA) and chlorophyll concentration) increased with closure. Extreme water deficit could be the main contributor to seedling death in the more open microhabitats, whereas light level was insufficient to maintain carbon balance under the water-stressful conditions existing beneath the closed tree canopies. Seedling establishment appears to be a limiting factor for the recruitment of both oaks within this forest in a wide range of microhabitats, especially for the more drought-sensitive Q. petraea. Moderate reductions of tree canopy cover can improve seedling establishment, but extreme summer droughts can prevent the success of any silvicultural practice made.     

The role of germination microsite in the establishment of sugar pine and Jeffrey pine seedlings

Mature shrubs can provide microhabitats that are beneficial to tree seedling growth and development. Sugar pine trees (Pinus lambertiana) grow in a narrow zone on the eastern slope of the Carson Range in extreme western Nevada, whereas Jeffrey pine (Pinus jeffreyi) is the dominant tree species in the region, an area extensively disturbed by wild fire. This study compares seedling establishment of sugar pine and Jeffrey pine relative to mature shrubs. In the fall of 2002 (cohort 1) and 2003 (cohort 2), 13,600 seeds of both species were planted in wire mesh enclosures, at three sites, under a variety of microhabitat treatments: under shade and in the open, under two species of shrub cover, and with and without plant litter. Seedlings were monitored for survival through two growing seasons. Even though more sugar pine seedlings emerged, more Jeffrey pine seedlings survived, and Jeffrey pine was the more drought tolerant species, better suited for the xeric climate found in the Carson Range. Litter slightly hindered seedling emergence but had no effect on survival and there was no significant species × litter interaction. Supplemental water facilitated survival in all treatments with highest survival in shade treatments. Sugar pine seedlings showed a significant increase in survival over Jeffrey pine seedlings with the addition of water, particularly in open treatments and more of both species survived under manzanita shrubs with water. The highest seedling mortality occurred when shrub canopy was removed, and seedlings experienced the effect of full sun and competition for soil water. For either species, microhabitat is a significant factor in determining success or failure in rehabilitation efforts after disturbance.     

The unstable fate of seedlings of the small-seeded pioneer tree species, Betula maximowicziana

The dynamics of the seedling population of the pioneer tree, Betula maximowicziana, were studied for 6 years under various site conditions, with special reference to mortality and mortality factors, in order to better understand their seedling bank formation process. We found that current year seedlings suffered high annual mortality of more than 80%, but that mortality fell with age. Canopy covers and cohorts maintained strong influences on seedling mortality during the first 3 years after emergence. The open site with disturbed soil surface was best suited for 3-year seedling survival. Since the seedlings of B. maximowicziana emerged regardless of canopy covers, high seedling mortality was thought to be the main factor that interrupted their regeneration under a canopy. The occurrence of various hazards together caused high seedling mortality. The major mortality factors, regardless of site conditions, were drought, cold hazard and debilitation, though occurrence of the three factors varied, dependent upon the climate of that particular year. Annual mortality and the occurrence of each mortality factor at a fine scale of 0.2 m² quadrate each showed certain spatio-temporal variations within the same site condition. The extremely high mortality of B. maximowicziana seedlings should be attributed to their inherently small, initial size. The poor root system of the small seedlings may be responsible for their low resistance to various hazards, such as drought and frost heaving.     

Effects of sika deer (Cervus nippon) and dwarf bamboo (Sasamorpha borealis) on seedling emergence and survival in cool-temperate mixed forests in the Kyushu Mountains

Effects of sika deer (Cervus nippon) and dwarf bamboo (“sasa;” Sasamorpha borealis) on seedling emergence and survival were investigated in cool-temperate mixed forests in the Kyushu mountain range, Japan. We compared the effects of sika deer between two sites with different sika deer densities. One site (no-sasa forest) has sparse cover of S. borealis and a high density of sika deer, and the other site (sasa forest) has dense cover of S. borealis with a low density of sika deer. In the no-sasa forest, more seedlings emerged and the survival rate of the seedlings was higher compared with the sasa forest. Compared with the sasa forest, the canopy in the no-sasa forest was more open, the organic layer was shallower and drier, the mean daily soil temperature was higher, and soil temperature fluctuated more widely. Those environmental parameters did not differ between unfenced and fenced plots. The comparison between the sasa and no-sasa forests suggested that removal of S. borealis by sika deer had an indirect positive effect on seedling emergence and survival. However, the fenced-plot experiments indicated that sika deer inhibited seedling emergence and survival. These contradictory conclusions could result from the much larger negative effects of S. borealis on seedling emergence and survival compared with the effects of sika deer browsing.     

Refugial forests of the southern Appalachians: photosynthesis and survival in current-year Abies fraseri seedlings

Fraser fir (Abies fraseri (Pursh) Poiret) is an endemic, high-elevation conifer confined to six relict mountaintop communities in the southern Appalachian Mountains, USA. High adult mortality has occurred over the past 50 years, possibly the result of an introduced insect (Adelges piceae Ratzeburg), air pollution, or both. Knowledge of the mechanisms of and limitations to seedling establishment may allow reestablisment and perpetuation of this unique community type, notwithstanding global climate change. We monitored seedling emergence and mortality in relation to photosynthetic performance and water relations in microsites differing in canopy openness (sunlight exposure) over the summer of 2004. Abundance of cotyledonous seedlings in early summer was 2.3 times greater (849 versus 366 seedlings m(-2)) in microsites with lower sky exposure (greater canopy closure) than in microsites with greater sky exposure (greater canopy openness). In contrast, late-season abundance and survival were greater in areas beneath more open canopies than in areas beneath less open canopies (3.3 times and 11.7 times greater, respectively). However, newly emerged seedling survival in a completely open site (no overhead canopy) was zero, despite an initial density of 124 seedlings m(-2). Seedling water status was similar in open- and closed-canopy sites (-0.52 and -0.74 MPa, respectively). Photosynthetic carbon gain was higher in newly emerged seedlings at open canopy than at closed canopy sites, especially during early morning. Based on photosynthetic light response curves and measured sunlight regimes, seedlings in open canopy sites were estimated to assimilate 3.3-4.5 times more carbon than seedlings at closed sites. Reductions in carbon gain of closed-site seedlings, primarily a result of limited sunlight, corresponded to substantial increases in seedling mortality (98 versus 79% in open canopy sites). Thus, sunlight exposure, which reflects overstory canopy structure, appears to be an important factor influencing newly emerged seedling survival and distribution.     

A comparison of seedling emergence and survival between Quercus glauca and Symplocos prunifolia

Seedling emergence and initial survival were compared for two evergreen broad-leaved species, Quercus glauca and Symplocos prunifolia. The relationships between the two seedling dynamics variables and environmental factors for the two species were also investigated. The number of seedlings that emerged in the study period was larger for S. prunifolia than for Q. glauca, while the survival rate was lower for S. prunifolia, presumably due to the closed canopy of this site. Models were selected for each species to determine the combination of variables explaining the most variation in emergence and survival of seedlings. The model selected for seedling emergence of Q. glauca showed that more seedlings emerged in lower hillslope positions and where the canopy in winter was more open. The model selected for S. prunifolia showed that fewer seedlings emerged on steeper slopes and that more emerged under a more open canopy in winter. With respect to seedling survival, models with only the proportion of open canopy in summer were selected for both species. These models showed that the survival rate was higher where the canopy in summer was more open. This is to be expected as both species are regarded as pioneer or mid-successional species. Q. glauca seemed to have the ability to persist as a dominant in the secondary forest at this site for longer than S. prunifolia due to the former’s better seedling survival rates and the rarity of climax species such as Castanopsis cuspidata.     

Tree recruitment in gaps of various size,clearcuts and undisturbed mixed forest of interior British Columbia,Canada

Tree seedling recruitment was monitored after various types of logging in mixed conifer and deciduous forests of northern British Columbia, Canada. Predicting tree seedling recruitment after disturbance is fundamental to understanding forest dynamics and succession and is vital for forest management purposes. Seedling recruitment success in multi-species northern latitude forests varied as a function of mature tree canopy cover, gap size and position in a gap. Recruitment was abundant within canopy gaps across a wide range of gap sizes (20–5000 m²), but recruit numbers dropped off rapidly under the closed forest canopy and in the open conditions of clearcuts. Inside canopy gaps, recruitment was similar by gap position in small gaps (<300 m²) but, in these northern latitude forests, exhibited a trend of increasing density from the sunny north to shady south end of larger gaps. This was true for all tree species regardless of their shade tolerance ranking. There was no evidence of gap partitioning by any of the tree species during the regeneration phase suggesting that adaptation to the subtleties of gap size during early recruitment are not well developed in these tree species. Favorable locations for emergence and early establishment of germinants were less favorable for growth and survival of established seedlings, i.e. the regeneration niches in these forests were discordant. Tree abundance and species diversity appears to be controlled more by differentiation among growth and survival niches than by the regeneration niches. From the perspective of forest management, abundant natural regeneration of all the dominant tree species of these mixed-species forests can be obtained after partial cutting.     

Seedling emergence of Betula maximowicziana following human disturbance and the role of buried viable seeds

We observed seed rain and seedling emergence of a pioneer tree species, Betula maximowicziana Regel., for 5 years at various sites differing in canopy and soil surface conditions, and clarified that a buried seed bank contributed to the recruitment of B. maximowicziana seedlings. More seedlings emerged at sites where the soil surface had been disturbed than at sites where the soil surface had not been disturbed. Within each soil surface condition, the densities of the emerged seedlings were generally proportional to the densities of the seeds dispersed the previous year. However, the seedling densities were disproportionately higher than the dispersed seed densities at the sites where the soil surface had been disturbed in the previous year. Soil samples taken during different seasons indicated that B. maximowicziana maintained a persistent seed bank in the soil below 1 cm deep, while it formed a more transient seed bank in the top 1 cm of the soil. The seed rain in the previous year was the primary contributor to the establishment of B. maximowicziana seedlings in a year after a mast seed year. On the other hand, the soil seed bank had a compensational role in a year after a poor seed year. Canopy cover did not affect seedling emergence. Therefore, the previously reported lack of a persistent seedling bank of B. maximowicziana under a canopy will be partially attributed to a relatively rare soil surface disturbance in a stable stand.     

Effects of nurse shrubs and tree shelters on the survival and growth of two Austrocedrus chilensis seedling types in a forest restoration trial in semiarid Patagonia, Argentina

? Context

Harsh environmental conditions in xeric sites of Andean Patagonia, affect the emergence, survival, and growth of either naturally grown or planted Austrocedrus seedlings.

? Aims

We evaluated the effects of nurse shrubs and tree shelters on the survival and growth of Austrocedrus seedlings as compared to unprotected (control) areas and how differently produced seedling types perform under these treatments.

? Methods

In 2006, two Austrocedrus seedling types (1?+?2?=?S1 and Plug 2?+?1?=?S2) were planted under shrub cover (C1), tree shelter (C2), and control (C3). Soil surface temperature and moisture were measured for each treatment during the first growing season, while seedling survival and height were recorded during 5 years.

? Results

Survival was not affected by cover type the first year, but it was affected by seedling type (S1?>?S2). After an extremely dry second growing season, seedling survival significantly decreased in relation to either cover or seedling type. Five years after plantation, survival was significantly higher for C1 and C2 (40?C60 %) as compared to C3-S2 (18 %). Seedling height was significantly affected by cover, but not by seedling type.

? Conclusions

The use of nurse shrubs and/or tree shelters is useful in Austrocedrus seedling restoration trials. Seedling morphology appears as relevant for survival in semiarid environments.     

Impact of vertebrate acorn- and seedling-predators on a Mediterranean Quercus pyrenaica forest

We have experimentally investigated the impact of biotic factors, acting at the seed and seedling stages, on a Quercus pyrenaica forest in the Sierra Nevada mountains (SE Spain). We monitored the natural establishment of the oak for 3 years in two forest plots and two shrubland plots, by counting seedlings and juveniles. In addition, we established several experiments in these plots to examine acorn and seedling survival, while also considering the microhabitat effect on survival probability. Dispersed acorns were quickly consumed by several species of predators, particularly wild boar (Sus scrofa) and woodmouse (Apodemus sylvaticus). Less than 4% of the experimental acorns survived to produce seedlings, even when they were buried 4 cm in soil, simulating caches. No effect of microhabitat was found on predation, and thus no safe site appears to exist for Q. pyrenaica acorns in the study area. Some 98% of the 1000 experimental seedlings were killed by herbivores, notably woodmice, wild boar, and domestic and wild ungulates. Seedling survival varied spatially, being significantly higher under shrubs (4%) than in any other microhabitat (less than 0.5%). Both acorn and seedling survival were much lower in the shrublands than in the forests. In shrubland plots, the main agent of seedling mortality was trampling by domestic ungulates foraging in herds. This study suggests that the regeneration of Q. pyrenaica forests in Mediterranean mountains can be limited by herbivores acting at several life-history stages.     

Establishment limitation of holm oak (Quercus ilex subsp. ballota (Desf.) Samp.) in a Mediterranean savanna — forest ecosystem

? Tree recruitment in Mediterranean savannas is generally hampered, in contrast with the original oak forests where these savannas are derived from. We asked whether this difference in recruitment success can be explained by differential post-dispersal survival. For one year we monitored experimentally cached holm oak acorns in a savanna — forest ecosystem in Central Spain, and recorded cache pilferage, type of pilferer, boar rooting, seedling emergence, seedling survival and the cause of mortality.

? Cache pilferage was significantly lower in savanna (8%) than in forest (21%). However, the higher cache survival was more than offset by lower seedling emergence and, particularly, by nine times higher seedling mortality in savanna, mainly due to desiccation. Wild boar rooting did not differ between experimental caches and controls without acorns, indicating that individual cached acorns do not trigger rooting activity.

? Our results indicate that the difference in post-dispersal survival between savanna and forest is due to lower emergence and, primarily, higher seedling mortality in savanna, not to higher cache pilferage. Absence of safe sites such as shrubs, abundantly present in the forest, may well explain the lack of recruitment in the savanna. Management measures appear necessary for long-term persistence of Mediterranean savannas in general.

     

Methods to improve the recruitment of holm-oak seedlings in grazed Mediterranean savanna-like ecosystems (dehesas)

Context

??Dehesas?? are savanna-like ecosystems of human origin that extend broadly in the Mediterranean area of the Iberian Peninsula. They consist of scattered oaks (mainly Quercus ilex subsp. ballota L. holm-oak), an annual grassland layer and interspersed shrubs. These ecosystems, used for grazing and wild game, support high plant and animal biodiversity and provide important environmental services. At present, Mediterranean ??dehesas?? are endangered by the lack of oak regeneration.

Aims

This paper analyses the efficiency of: (1) using shrubs as nurse plants; (2) drip irrigation of seedlings during summer; and (3) a combination of the two methods for the restoration of a ??dehesa?? in a mid-mountain Mediterranean area of southern Spain.

Methods

Different techniques were tested to improve the recruitment of holm-oak seedling during a 3-year field experiment: (1) acorn plantation in open spaces, irrigating seedlings during the first dry season; (2) acorn plantation beneath the canopy of Myrtus communis L. and (3) both methods combined.

Results

There was a large facilitative effect of myrtle for the recruitment of holm-oak seedlings, regardless of the supply of irrigation. This effect was associated with a large decrease in air temperature and photosynthetically active radiation beneath myrtle canopies. By contrast, summer irrigation of seedlings planted in open spaces did not improve seedling survival after 3 years despite a small and transient positive effect on seedling survival during the 1st year.

Conclusion

The use of evergreen shrubs, such as myrtle, as nurse plants may be considered to restore ??dehesas?? instead of expensive seedling irrigation techniques. Several studies have promoted abandoning grazing to increase holm oak self-regeneration in ??dehesas??. However, creating closed patches of naturally occurring evergreen shrubs could provide suitable sites for oak planting when necessary, thus enhancing seedling recruitment without damaging the environmental and economic value of these ecosystems.     

Interacting effects of canopy gap, understory vegetation and leaf litter on tree seedling recruitment and composition in tropical secondary forests

We experimentally investigated interacting effects of canopy gaps, understory vegetation and leaf litter on recruitment and mortality of tree seedlings at the community level in a 20-year-old lowland forest in Costa Rica, and tested several predictions based on results of previous studies. We predicted that experimental canopy gaps would greatly enhance tree seedling recruitment, and that leaf litter removal would further enhance recruitment of small-seeded, shade-intolerant seedlings in gaps. We created a large (320–540 m²) gap in the center of 5 out of 10 40 m × 40 m experimental plots, and applied the following treatments bimonthly over a 14-month-period in a factorial, split–split plot design: clipping of understory vegetation (cut, uncut), and leaf litter manipulations (removal, addition, control). As expected, experimental gaps dramatically increased tree seedling recruitment, but gap effects varied among litter treatments. Litter addition reduced recruitment in gaps, but enhanced recruitment under intact canopy. Species composition of recruits also differed markedly between gap treatments: several small-seeded pioneer and long-lived pioneer species recruited almost exclusively in gaps. In contrast, a few medium-to-large-seeded shade-tolerant species recruited predominantly under intact canopy. Leaf litter represents a major barrier for seedling emergence and establishment of small-seeded, shade-intolerant species, but enhances emergence and establishment of large-seeded, shade-tolerant species, possibly through increased humidity and reduced detection by predators. Periodic clipping of the understory vegetation marginally reduced tree seedling mortality, but only in experimental gaps, where understory vegetation cover was greatly enhanced compared to intact canopy conditions. Successful regeneration of commercially valuable long-lived pioneer trees that dominate the forest canopy may require clear-cutting, as well as weeding and site preparation (litter removal) treatments in felling clearings. Management systems that mimic natural canopy gaps (reduced-impact selective logging) could favor the regeneration of shade-tolerant tree species, potentially accelerating convergence to old-growth forest composition. In contrast, systems that produce large canopy openings (clear-cutting) may re-initiate succession, potentially leading to less diverse but perhaps more easily managed “natural plantations” of long-lived pioneer tree species.     

Patterns of Abies cephalonica seedling recruitment in Mount Aenos National Park, Cephalonia, Greece

Patterns of Abies cephalonica Loudon seedling recruitment at its westernmost geographical limit (western Greece), were studied by monitoring their emergence, survival, and establishment for up to four years in the microhabitats to which seeds are dispersed. Seedlings were monitored on 200 permanent sampling plots established in 11 different locations covering the entire strictly protected Mount Aenos National Park (Cephalonia). Annual seedling mortality rate was estimated in relation to habitat characteristics as well as to their number of cotyledons. A. cephalonica presented the typical behaviour of a masting species in the number of seedlings produced over the four years of study. Of the habitat characteristics studied, ground cover, elevation, soil depth, slope and canopy openness were the main parameters that classified seedling habitat with respect to their emergence. Seedling survival was higher in years of low seedling production. Seedlings with higher cotyledon numbers exhibited higher survival. Our data indicate that A. cephalonica is characterized by remarkable reproductive plasticity, which is expressed in the number of seedlings produced each year and those surviving with respect to its habitat characteristics. Preserving the species implies preserving its habitat. In this sense, Mount Aenos National Park may be the most important protected area for the endemic fir in view of preserving its integrity through pre-emptive management, in the light of current and future threats.     

Effects of artificial shade on early performance of white spruce seedlings planted on clearcuts

Chlorophyll fluorescence, chlorophyll content, growth, and mortality of white spruce (Picea glauca [Moench] Voss) seedlings were monitored for 2 years after planting under three scenarios of artificial shade: no-shade (control), shade in summer only, and shade all year. The shade frames allowed 50–60% light transmission, with limited effects on air temperature, relative humidity, soil temperature, and soil moisture around seedlings. Based on fluorescence yield and chlorophyll content measurements, summer-only shade reduced photoinhibition and photooxidation, especially in summer and fall; extending to all year shading did not further reduce either photoinhibition or photooxidation. Shade tended to reduce seedling diameter and mortality, but after 2 years the cumulative effect on mortality was not statistically significant. Study results support the establishment of white spruce seedlings under partial forest canopy, especially on sites with harsh environmental conditions.     

Seedling demography and establishment of Cryptomeria japonica in a cool-temperate, old-growth, conifer hardwood forest in the snowy region of Japan

To clarify the establishment pattern of Cryptomeria japonica seedlings, the demography of current-year seedlings was studied under various micro-environmental conditions for 2 years in a cool-temperate old-growth forest in the snowy region of Japan. In both years, more than 70% of seedlings that emerged died within a year, and differences in the survival rate of seedlings with respect to micro-environmental conditions were similar. While a number of seedlings survived at the sites on fallen logs and in mineral soil, almost no seedlings survived at the sites with either broad-leaved or coniferous litters at the end of the first growing season. The negative litter effect may kill most small seedlings of C. japonica. In addition, we found that survival rates of the seedlings were greater as the distance from the nearest adult and canopy openness increased. An analysis using Quantification Theory I showed that, of the possible micro-environmental factors, substrate conditions had the strongest effect on the survival of seedlings. The analysis also indicated that distance from the nearest adult and canopy openness also affected seedling survival significantly and to almost the same extent, although the effects were weaker than those for the substrate condition. Given the limited establishment sites for C. japonica seedlings, layering is suggested to be the main mode of regeneration in this forest.     

Seedling survival and growth of Aquilaria malaccensis in different microclimatic conditions of northeast India

We studied seedling survival and growth of Aquilaria malaccensis in three different environmental conditions(homegardens,green house and in different canopy conditions) of northeast India.Results show that mean seasonal survival was highest in green house(95.53%±2.33),followed by homegardens(89.3%±1.89) and different canopy conditions(77.62%±6.73);the highest values were found during February to April for both the homegardens(96%±1.68) and green house(98% ± 0.88) and lowest during November to January(78%±2.99) in homegardens and May to July(90%±4.53) in green house.In case of transplanted seedlings in different canopy conditions,mean seasonal survival was highest during May to July(98%±1.92) and lowest during August to October(66%±12.81).However,mean seasonal growth of collar diameter was highest in different canopy conditions(23.99%±1.76) compared to green house(21.52%±2.70) and homegardens(12.44%±1.33) and it was highest during rainy season(May to July) and lowest during dry winter(November to January) in all the three experimental sites.These variations may be due to the different microclimatic conditions as well as nutrient status of the soil in all the three experimental sites.Although,seedling quality plays a great role in their survival and growth,based on the result of green house experiment,it can be concluded that maintenance of seedlings in green house conditions during their early growth period may improve both the survival and growth for large scale plantation of the species.Thus,the species can be reintroduced in its natural forest range to compensate the loss of natural population of this precious species in northeast India.     

Canopy tree development and undergrowth bamboo dynamics in old-growth Abies–Betula forests in southwestern China: a 12-year study

Interactions between forest canopy characteristics and plants in the forest understory are important determinants of forest community structure and dynamics. In the highlands of southwestern, China the dwarf bamboo Bashania fangiana Yi is an understory dominant beneath a mixed canopy of the evergreen Abies faxoniana (Rheder & Wilson) and the deciduous Betula utilis (D. Don). The goal of this study was to better understand the role of bamboo dominance, canopy characteristics, and periodic bamboo dieback on forest development. To achieve this goal, we measured tree seedling, tree saplings, and trees, forest canopy characteristics, and bamboo cover in permanent forest (n = 4) and gap plots (n = 31) in a mixed A. faxoniana and B. utilis forest in Sichuan, China. Dwarf bamboos died off in 1983 in the gap plots, and in three of the four forest plots. Forest development was assessed for the period 1984–1996. The seedling bank in forest and gap plots increased after bamboo die-off. A. faxoniana seedlings increased more than B. utilis in forest plots; the opposite pattern characterized gap plots. The proportion of seedlings on raised micro-sites on the forest floor also changed and new seedling were more abundant on the forest floor. By 1996, bamboo seedling cover and biomass had recovered to ca. 45% or their pre-flowering values. Rates of bamboo seedling recovery were faster beneath canopy gaps and deciduous trees than beneath forest or evergreen trees. Tree mortality exceeded recruitment in plots with dense bamboo; the opposite pattern was found in the plot with little bamboo. The mortality rate for B. utilis trees (2.4% year⁻¹) was higher than that for A. faxoniana (0.8% year⁻¹) and forests with dense bamboos became more open over the census period. Tree mortality was size-dependent and intermediate sized trees had the lowest rates of mortality. Stand basal area increased mainly due to greater basal area gain than loss for A. faxoniana. Interactions between tree species life history, canopy type, and bamboo life-cycles create heterogeneous conditions that influence tree and bamboo regeneration and contribute to the coexistence of A. faxoniana and B. utilis in old-growth forests in southwestern China.     

Season and substrate effects on the first-year establishment of current-year seedlings of major conifer species in an old-growth subalpine forest in central Japan

Seasonal and substrate effects on first-year seedling establishment of major conifer species (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis, and Tsuga diversifolia) were assessed in an old-growth subalpine forest in central Japan. Older seedlings (≥1 year) were more abundant on woody debris (WD), such as fallen logs, than on soil. It is known that this preferential seedling establishment on WD is determined by the first-year seedling establishment. The present results indicate that first-summer seedling survivorship clearly contributes to the creation of such seedling–substrate associations, because mortality during the first-summer was greater on soil than WD but that during the first-autumn/winter was not different. Although the standardized mortality rate of current-year seedlings on WD was not so different among the three seasons, that on soil significantly decreased in the winter; indicating that soil is not an unsuitable substrate for seedling survivorship during snow-covered winter. Thus, it can be concluded that biotic and abiotic factors enhancing seedling mortality on soil are most active during summer, and seedling–substrate associations seem to be determined mainly during the first-summer soon after seedling emergence. Furthermore, the first-year survivorship of P. jezoensis var. hondoensis, which has smaller-sized seeds/seedlings that are disadvantageous for early survival, decreased with progressing WD decay toward soil. This suggests that the properties of WD also affect current-year seedling survivorship, especially for species sensitive to substrate properties. In conclusion, initial survivorship affects seedling bank dynamics, and such early dynamics are greatly regulated by the highly heterogeneous substrate availability on forest floors.     

Shrubs facilitating seedling performance in ungulate-dominated systems: biotic versus abiotic mechanisms of plant facilitation

Shrubs are important microsites that facilitate seedling performance in abiotically stressed environments. However, shrub facilitation might work differently in ecosystems with high biotic stress (e.g., ungulate-dominated environments). We wonder whether ungulates through their preferences for different shrubs affect the facilitation process of seedlings, and whether this process remains consistent across different levels of abiotic stress. We study seedling performance (survival and growth) of a Mediterranean oak (Quercus pyrenaica) in three microsites (open, preferred and non-preferred shrubs by ungulates) for two contrasting climatic environments and throughout a complete growing season (increasing abiotic stress). Overall, there was a positive effect of shrubs in seedling survival. We found a microsite effect on seedling growth but this effect differed depending on the abiotic environment, with seedlings growing better in open microsites in the less stressful site. The larger abiotic context (climate) played an important role in determining which microsite is the most favorable for seedling growth but not necessarily for the eventual survival of seedlings. Wild ungulates started to damage seedlings early in the growing season, whereas abiotic stress (desiccation) affected mostly those plants that were not previously attacked by animals, showing the hierarchical effect (earlier and longer effect) of the biotic over the abiotic mortality agents. Non-preferred shrubs worked better as nurse shrubs only under high ungulate pressure. From the abiotic perspective, preferred and non-preferred shrubs did not apparently differ in the facilitation effect. Importantly, we found a greater facilitative effect of shrubs on the survival than on the growth of seedlings.