八仙山不同类型松栎林群落主要特征分析

为探究八仙山保护区不同类型森林群落的更新潜力、多样性程度以及稳定性水平,阐明三者间的关系,该文以保护区内油松林、蒙古栎林和油松-栓皮栎混交林(松栎混交林) 3种不同类型天然次生林为对象,调查建群种径级结构和更新潜力,计算不同层次群落多样性,测定M. Godron稳定性,并采用主成分法建立评价模型。结果表明:(1)油松种群径级结构近似正态分布,处于成熟期,但幼苗较少;栓皮栎、蒙古栎以及阔叶杂木的幼苗、幼树较多,更新潜力较大。(2)松栎混交林的乔木层多样性较高,而灌木层多样性最低,油松林的草本层多样性最低;松栎混交林群落总体物种丰富度最低而均匀度最高。(3) M. Godron稳定性表明,蒙古栎林距离稳定点较近,而油松林偏离较远。(4) PCA双序图表明,M. Godron稳定性与种群更新潜力、Alatalo均匀度呈较强正相关,综合排序依次为松栎混交林、蒙古栎林和油松林。综上结果表明,建群种更新潜力和物种均匀度对群落稳定性影响较大,林地管理应注重对幼苗幼树的保护。     

Comparative studies on acorn and seedling dynamics of fourQuercus species in an evergreen broad-leaved forest

The dynamics and regeneration probabilities ofQuercus acuta, Q. gilva, Q. salicina, andQ. sessilifolia were studied in a 0.48 ha plot in an old-growth evergreen broad-leaved forest, Kagoshima prefecture, southwestern Japan. The canopy was well closed and had only one small gap. Seventeen tree species occurred in the canopy. The fourQuercus species ranked second to fifth in basal area. Excluding the trees originated from sprouts,Q. acuta and Q. gilva lacked trees at the subcanopy layer, andQ. salicina andQ. sessilifolia had some trees at the same layer. For eachQuercus species, the amount of fallen acorns fluctuated from year to year. The number of current year seedlings was positively correlated with the number of fallen acorns in the previous year. A disproportion between the spatial distribution of fallen acorns and that of established seedlings has been shown. The establishment of seedlings was frequently observed where no acorns had fallen in the previous year. Saplings of four species survived for several years and formed sapling banks under the closed canopy. During the three years of the study, the mortality ofQ. salicina andQ. sessilifolia saplings was lower than that ofQ. acuta andQ. gilva. However, the differences in the average annual height growth of saplings between four species were not significant, and the most of them did not grow much. These results suggest that the saplings of the fourQuercus, especiallyQ. gilva andQ. acuta, have no chance to recruit to the canopy or subcanopy layer within a closed stand, and that some changes in the environment caused by gap formation are required for their recruitment.     

Comparative ecophysiology of two representativeQuercus species appearing in different stages of succession

Ecophysiological comparisons were made of the growth and photosynthetic characteristics between seedlings of deciduousQuercus serrata and evergreenQuercus myrsinaefolia. Q. myrsinaefolia seedlings naturally occurring in secondary coppice forests showed exponential-like growth in height with age, while sympatricQ. serrata seedlings were considerably smaller in height, their growth being limited by shortage of light. The photosynthetic characteristics measured under laboratory conditions showed no bases for the differences in growth between the two species on the forest floor: Light compensation points of the seedlings raised under 5% daylight were almost identical for the two species, being about 6.0 μE·m⁻²·s⁻¹. Growth analysis of seedlings planted in a coppice forest showed that bothQ. serrata andQ. myrsinaefolia could hardly grow during the summer under the shrub layer, when relative photon flux density (RPFD) was 0.9±0.5%. In the winter, when RPFD under the leafless canopy increased to 29.3±2.7%, the dry matter production of the evergreen seedlings ofQ. myrsinaefolia was much improved. Current-year seedlings of the species showed NAR of 0.102±0.021 g·dm⁻²·mo⁻¹ during the winter. Temperature dependency of photosynthesis and increment of leaf temperature by direct solar beam also indicated active photosynthesis ofQ. myrsinaefolia on the forest floor during the winter.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Nutrient utilization in Quercus leucotrichophora and Pinus roxburghii seedlings at five soil fertility levels

Abstract. Nutrient utilization and growth performance of Pinus roxburghii Sarg. and Quercus leucotrichophora A. Camus, the two major forest trees of the Central Himalaya Mountains between 1000 and 2000 m elevation, were analysed at different nutrient levels. The early successional P. roxburghii occurs naturally on nutrient-poor sites and the late successional Q. leucotrichophora on nutrient-rich sites. Seedlings of these two species were grown both in monoculture and mixed, in plastic bags containing 1 kg soil and representing five levels of nutrients (NPK) in a 12 : 32 : 16 ratio. The P. roxburghii seedlings showed significantly greater nutrient extraction efficiency (nutrient unit extracted by seedlings per nutrient unit in the soil) than the Q. leucotrichophora seedlings, and this difference was increased in interspecific competition, particularly at the higher nutrient levels. In interspecific competition the extraction efficiencies of P. roxburghii for N, P and K were distinctly higher than those of Q. leucotrichophora. This we consider to be the main reason for the replacement of Q. leucotrichophora by P. roxburghii, even on nutrient-rich soils, after disturbance and subsequent increase in light availability. In both the species, re-translocation of nutrients from senescing leaves declined with increasing soil fertility. For example, re-translocation for P. roxburghii was 49% N, 30% P and 32% K at the lowest soil fertility, and 20% N, 8% P and 13% K at the highest soil fertility.     

Anthropogenic disturbances and plant biodiversity in forests of Uttaranchal,central Himalaya

Eight forest types varying in disturbance frequencies were identified along an elevational gradient in Uttaranchal, central Himalaya. Low elevation forests were close to human habitation and had high disturbance frequency, while high elevation forests were situated far from the human habitation and had low disturbance. The dominant tree species at low elevation were Pinus roxburghii and Quercus leucotrichophora, while Q. floribunda and Q. semecarpifolia dominated the high elevation forests. Pyracantha crenulata was the shrub present in all the forests except in Q. semecarpifolia forest and Anaphalis contorta, a herb species, was present in all the forests. Disturbance decreased the dominance of single species and increased the plant biodiversity by mixing species of different successional status. Species richness and diversity for all the vegetation layers were higher in low elevation–high disturbance forests. Mean tree density decreased from high to moderate and increased in low disturbance. The shrub density decreased from high to low disturbance while the reverse occured for herbs. High proportion of early successional species in disturbed forests indicated that disturbance induces succession. The mean number of young individuals increasing from high to low disturbance indicates that disturbance adversely affects regeneration. But, however, the high number of young individuals of Coriaria nepalensis, a small non-leguminous nitrogen fixing tree, in disturbed forests shows that the forest is regenerating. This species could be helpful in the re-establishment of original vegetation through triggering the regeneration of these forests. High elevation–low disturbed forests separated from low elevation–high disturbed forests. Forest type and elevation may have more influence on tree richness while shrub and herb richness may be more sensitive to disturbance and forest types.     

Forest fire in the central Himalaya: climate and recovery of trees

 A forest fire event is influenced by climatic conditions and is supported by accumulation of fuel on forest floor. After forest fire, photosynthetically active solar radiation was reduced due to accumulation of ash and dust particles in atmosphere. Post-fire impacts on Quercus leucotrichophora, Rhododendron arboreum and Lyonia ovalifolia in a broadleaf forest were analysed after a wild fire. Bark depth damage was greatest for L. ovalifolia and least for Q. leucotrichophora. Regeneration of saplings was observed for all the tree species through sprouting. Epicormic recovery was observed for the trees of all the species. Young trees of Q. leucotrichophora (<40 cm circumference at breast height) were susceptible to fire as evident by the lack of sprouting. Under-canopy tree species have a high potential for recovery as evident by greater length and diameter of shoots and numbers of buds and leaves per shoot than canopy species. Leaf area, leaf moisture and specific leaf area were greater in the deciduous species, with few exceptions, than in evergreen species. Received: 26 July 1995 / Revised: 1 March 1996 / Accepted: 9 September 1996     

Stand dynamics during a 12-year period in a second-growth stand in a cool temperature forest in northern Japan

In 1981 and 1993, trees over 2 m high were measured and mapped to clarify stand dynamics in two permanent plots of 0.1 ha in a secondary, cool temperate, mixed broadleaf/conifer forest after logging in the Tomakomai Experiment Forest, central Hokkaido, northern Japan. The species could be placed in two groups according to the change in basal area and density, and annual height growth:Quercus mongolica var.grosseserrata, Phellodendron amurense, Prunus sargentii andTilia japonica (Group I: GI) showed episodic regeneration and/or fast height growth;Acer mono, Acer palmatum var.matsumuae andSorbus alnifolia (Group II: GII) showed continuous regeneration and/or slow height growth. The age distribution of stems over 10 cm in d.b.h. suggests synchronous regeneration of GI and GII species. Vertical stratification was promoted during the 12-year period by the difference in annual height growth between the two species groups. Additional tree censuses in both remnant old-growth stands and second-growth stands after large scale blowdowns demonstrated thatQ. mongolica var.grosseserrata is most dominant in stands varying in the stages of development. Disturbance history and successional trends in dominant species implied that stand-devastating disturbances were responsible for the regeneration ofQ. mongolica var.grosseserrata in the study forest.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

Dynamics and conservation of the gwangneung national forest in central Korea: A national model for forest restoration

Forests in Gwangneung National Arboretum District (GNAD) have been protected since the 15^th century. Consequently, these forests support more than 20% of all plant species in Korea. We constructed vegetation maps for landscape analysis, and forest dynamics, species diversity, and sustainable management were discussed. Secondary forests compose 51.0% of the whole vegetation, while plantations compose 45.2%.Quercus serrata dominates the forest, and the plantations are comprised mainly ofPinus koraiensis. Although dominated by plantations and human installations, the presence of a rare riparian hardwood forest, composed primarily of Q.aliena, was notable. Species diversity of the riparian (H′ = 3.38) was significantly (p<0.0001) higher than the upland (H′ = 1.56). Species turnover rate as a spatial heterogeneity was also higher. Such high species diversity and heterogeneity are justification to conserve the riparian and lowland forests in GNAD. Extensive recruitments of their own seedlings and saplings suggest a sustainable regeneration of Q.serrata and Q.aliena stands in the lower elevations, and the opposite is true for the Q.mongolica andP. rigida stands in higher elevations. GNAD contains diverse natural landscape elements that range from riparian to upland vegetation, which may well serve as a national model for forest restoration.     

Deer Browsing,Forest Edge Effects,and Vegetation Dynamics Following Bottomland Forest Restoration

Vegetation composition and forest stand development are frequently mediated by browsing herbivores. These relationships have received little attention in a forest restoration context even though White‐tailed deer (Odocoileus virginianus) is likely to influence these agriculture, forest, and restored ecosystem mosaic landscapes. Tree species composition, herbaceous vegetation, and deer browsing patterns were assessed 5 and 7 years following bottomland hardwood forest restoration on a 526‐ha site in the Cache River watershed in southern Illinois, United States. Light‐seeded tree species (Fraxinus pennsylvanica, Acer negundo, Liquidambar styraciflua, and Platanus occidentalis) of volunteer origin dominated the woody vegetation component, with especially high stocking and density near existing forest cover and potential seed sources. At more distant locations, presumably planted Quercus spp. were more likely to dominate and were the only tree species found in 15% of plots in year 7. Quercus stocking increased over the course of the study, constituting 7% of trees during year 7. Deer herbivory was associated with reduced stem height and disproportionately impacted seedlings of Quercus palustris and Celtis spp. Our results suggest that deer browsing influences forest stand composition and density as a function of distance from the nearest forest edge. Herbaceous vegetation had little impact on early stand development. Continued spread of the exotic and invasive Lonicera japonica and potential mortality of F. pennsylvanica due to an anticipated Emerald ash borer (Agrilus planipennis) epidemic, combined with low stand density and delayed canopy closure, may result in persistent overstory gaps and compromise long‐term restoration success.     

Tree structure,regeneration and survival of seedlings and sprouts in highmontane forests of the Garhwal Himalayas,India

The tree structure of some timberline and sub-alpine forests of the Garhwal Himalayas was studied with reference to the regeneration status and survival of tree seedlings and sprouts. Nine tree and shrub species were chosen. The dominating tree species wereAbies pindrow andQuercus semicarpifolia. Sprouts were found to have a higher survival rate than seedlings, but both suffered from winter conditions. Girth class measurements showed that trees between 1.6 and 2.5 m were best represented.     

Differences in sapling architecture betweenFagus crenata andFagus japonica

The ecological significance of architectural patterns for saplings ofFagus crenata andFagus japonica co-occurring in a secondary oak forest were evaluated by comparing the size and shape of leaves, trunks and crowns.Fagus japonica saplings were different fromF. crenata saplings in some architectural properties: (i) the leaf area and specific leaf area were larger; (ii) the ratio of sapling height to trunk length was lower, indicating greater leaning of the trunk; and (iii) the projection area of the crown was larger and the leaf area index lower indicating less mutual shading of leaves. These architectural features indicated thatF. japonica saplings were more shade tolerant thanF. crenata andF. crenata saplings were superior toF. japonica for growth in height and could, therefore, utilize sunlight in the upper layer. An erect trunk inF. crenata and a leaning trunk inF. japonica may be important characteristics associated with the regenerations patterns of each species; regeneration from seedlings under canopy gaps in the former and vegetative regeneration by sprouting in the latter.     

Decomposition of woody branch litter on an altitudinal transect in the Himalaya

Decomposition of branch litter of four angiosperm and one conifer species was studied over a two-year period. Litter species and the corresponding forest type are: (i) Shorea robusta, sal forest at 329 m; (ii) Lyonia ovalifolia, mixed-pine broadleaf forest at 1 350 m; (iii) Pinus roxburghii, pine forest at 1 750 m; (iv) Quercus leucotrichophora, mixed oak-pine forest at 1 850 m; and (v) Quercus lanuginosa, mixed oak forest at 2 150 m. The weight loss ranged from 44–89%. Litter moisture and air temperature had significant positive effect on decomposition. The decomposition rate decreased with an increase in altitude and was inversely related with lignin content. Linear combinations of lignin content with rainfall and with temperature indicated significant interactive influence on decomposition.Authorities for plant names are given in Table 1.We gratefully acknowledge financial support from the Department of Science and Technology, Government of India.     

Seedling recruitment ofChamaecyparis obtusa andSciadopitys verticillata in different microenvironments in an old-growthSciadopitys verticillata forest

Seedling recruitment ofChamecyparis obtusa andSciadopitys verticillata in different microenvironments in an old-growthS. verticillata forest was studied for five years. Exposed mineral soil beneath a gap, which was newly created by a typhoon, supported the best seedling emergence forC. obtusa. S. verticillata seemed to require no specific microenvironment for its seedling emergence. Mortality of seedlings was higher for both species beneath a closed canopy, where all new seedlings ofC. obtusa died within a year after their emergence. Beneath the gap, mortality of seedlings was generally lower for both species, regardless of the presence of litter on the ground. The occurrence of saplings was more frequent forC. obtusa beneath the gaps and forS. verticillata beneath a closed canopy. Difference of seedling recruitment behaviour could maintain the coexistence ofC. obtusa andS. verticillata in the canopy layer of this forest in a non-equilibrium state caused by occasional disturbance.     

Distribution of Quercus leucotrichophora at Different Location and Elevation in Jandool Valley,Northern Pakistan

Quercus leucotrichophora is one of the favorite fuelwood trees of Jandool valley District Dir (lower). The tree is of prime importance in the area, therefore, a phytosociological study was carried out to know the current status of Quercus leucotrichophora at different sites of two locations (Maskini & Zardari Banda) during September to November 2017. During the survey, 28 quadrates of size (10 × 10) were randomly constructed, 14 in each site. The survey data showed that plant density of Quercus leucotrichophora was higher in Zardari Banda than Maskini at the middle site as compared to base and top. The results showed that plant of Maskini and Zardari Banda had little difference in term of height, while in case of sites the maximum plant height was recorded at top of Zardari Banda and base site of site Maskini. Plant of Maskini and Zardari Banda had different girth at different sites. More number of cut plants was recorded at the base and middle of both locations, while less number of cut plants was noted at the top. The number of seedlings and soil physiochemical attributes like soil texture, class, electrical conductivity, organic matter, lime, nitrogen, phosphorus, and potassium content was better in Zardari Banda as compared to Maskini. Due to large variations in both locations and sites, the study provided very helpful knowledge to the scientific community about forest trees to design a solid integrated approach for proper management and protection from overuse of the species in the target area. In future, similar survey studies should be carried out for timely information and resolution of issues related to forest trees.     

The response of forest floor vegetation and tree regeneration to deer exclusion and disturbance in a riparian deciduous forest,central Japan

The response of forest floor vegetation and regeneration of major treespecies to deer exclusion in a riparian forest were studied for 3 years withtheinteractive effects of natural disturbances. At the start of this study, deerdensity had quickly increased to a fairly high level (29–31 individualskm^–2) during the last decade and had severely reduced theamount of forest floor vegetation in the study area. Dwarf bamboos, which weredominant before, had almost diminished from the forest floor. During the studyperiod, aboveground biomass increased steadily but species diversity did notchange much in the exclosures. Outside the exclosures, the seedlings of alltreespecies were damaged greatly by deer browsing, especially the taller ones. Deerbrowsing had little effect on the emergence of tree seedlings, but deertrampling might have accelerated emergence indirectly by disturbing the soilsurface for some species. Differences in plant responses were also observed fordeer browsing and the presence of dwarf bamboo that strongly inhibits therecruitment of tree seedlings. The nine major species were classified intothreegroups according to the response of seedlings to deer browsing and disturbance.Group 1 consists of the species whose seedling survival is affected bybrowsing,but seldom by disturbances (Phellodendron amurense,Kalopanax pictus, Quercus crispulaandMalus toringo). Groups 2 and 3 consist of species adaptedto tree-fall disturbance (Betula spp.) and ripariandisturbance (Alnus hirsuta var.sibirica, Ulmus davidiana var.japonica, Populus maximowiczii andSalix sachalinensis), respectively, and seedling survivalof these two groups is principally affected by light conditions. The effect ofdeer browsing on seedling survival and growth was greater for Group 2 than forGroup 3.     

Comparative structural study ofQuercus serrata andQ. acutissima formed byPisolithus tinctorius andHebeloma cylindrosporum

Ectomycorrhizas were synthesized in pots and growth pouches betweenQuercus serrata, Q. acutissima, and two ectomycorrhizal fungi,Pisolithus tinctorius andHebeloma cylindrosporum. Root morphology and the structure of the mantle and Hartig net were compared using light, fluorescence, scanning and transmission electron microscopy.P. tinctorius initially colonized root cap cells, and eventually produced a highly branched lateral root system with a complete mantle, whereasH. cylindrosporum promoted root elongation with few hyphae on the root apex surface indicating that interaction between roots differs with fungal species. Hartig net structure and hyphal inclusions varied between all the combinations tested. There were structural differences between mycorrhizas ofH. cylindrosporum/Q. acutissima grown in soil and growth pouches, which indicate that the growth pouch environment can induce artefacts in roots. Fruit bodies ofH. cylindrosporum developed in pots withQ. acutissima. AlthoughP. tinctorius has been used to inoculate oak seedlings in the nursery, results of this study indicate thatH. cylindrosporum may also be an effective ectomycorrhizal fungus forQ. serrata andQ. acutissima.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

Large-Scale Patterns of Quercus ilex, Quercus suber, and Quercus pyrenaica Regeneration in Central-Western Spain

In Central-Western Spain, forests and woodlands composed of Quercus sp. support outstanding levels of biodiversity, but there is increasing concern about their long-term persistence due to a lack of regeneration. We hypothesize that this regenerative lack is operating on a large geographic scale; that there are differences in the abundance of regeneration between three oak species; that oak regeneration is governed mainly by forest management and structure; and that shrubs act as important physical protectors of seedlings and saplings. We analyzed whether densities of oak seedlings and saplings in several size classes were related to stand-structure, understory, and physiographic variables potentially affecting regeneration. Data collected at a regional level (1 km × 1 km grid) by the Spanish Forest Inventory were evaluated from 2,816 plots. Results revealed that regeneration failure was common for all size categories, from small seedlings to large saplings, and for the three oak species studied, especially the evergreens. Of the Quercus ilex, Q. suber, and Q. pyrenaica plots studied, 49%, 62%, and 20% were lacking any small seedlings, and 82%, 96%, and 56% did not have any large saplings, respectively. Regeneration was positively correlated with tree cover and density, especially of small and medium-sized trees, and negatively correlated with the presence of large trees, indicating that regeneration failure is mostly associated with more open, uniform, and/or aged woodlands. Regeneration densities of Q. ilex and Q. suber were positively correlated with all understory variables, suggesting that the presence of pioneer shrubs represent a major safe site for early tree recruitment, independent from specific shrub species.