Biodiversity Conservation in Sacred Groves of Manipur, Northeast India: Population Structure and Regeneration Status of Woody Species

Sacred groves are forest patches conserved by the local people intertwined with their socio-cultural and religious practice. These groves harbour rich biodiversity and play a significant role in the conservation of biodiversity. Population structure and regeneration status of woody species were studied during 2001–2002 in the four sacred groves of Manipur, a state in north east India. A total of 96 woody species was recorded from the four groves, the highest being Konthoujam Lairembi sacred grove (55 species) and lowest in Heingang Marjing sacred grove having 42 species. The density–diameter distribution of woody species in the four groves showed highest stand density and species richness in the lowest girth class (30–60 cm) and decreased in the succeeding girth classes. Overall population structure of the groves based on the number of tree seedlings, saplings and adults, displayed a greater proportion of seedlings followed by saplings and adults while for the selected tree species it varied seasonally and recruitment of species increased during rainy season attaining peak during June. Regeneration status of the four sacred groves based on strength of different age groups in their population showed good regeneration. High occurrence of ‘additional species’ to the groves may be due to the invasion through dispersal from other areas. Possibly, the prevailing favourable microenvironmental conditions contributed to their establishment and growth in the groves. Absence of seedlings and saplings of some of the species in the groves may be due to their poor seed germination and establishment of seedlings in the forest.     

Comparing evaluations of forest health based on aerial surveys and field inventories: Oak forests in the Northern United States

Aerial sketch-map surveys and systematic forest field inventories may be used separately or in combination to indicate the status of regional forest health. During recent decades, aerially conducted sketch-maps of forest damage and forest inventories have been used to assess oak (Quercus spp) forest health across a 24-state region spanning the northern U.S. In order to more fully inform the monitoring of oak forest health and integrate these independent datasets, the effect of the quality, timing, and repeated sampling of aerial data on correlations with field-based oak forest assessments was assessed. Study results indicated that aerial damage surveys were weakly correlated with indicators of oak forest sustainability (e.g., oak seedlings and saplings), but more highly correlated with overstory attributes such as tree mortality and standing dead. The highest correlations between aerial damage surveys and oak mortality/standing dead were found when the time between the aerial survey and subsequent forest inventory was 4–6 years. Aerial surveys may have their greatest efficacy in supplementing field inventories of oak forest health when they are conducted in a high quality manner with bi-annual or longer remeasurement periods (due to rare pest damage events).     

云南楚雄彝族的“神树林”与生物多样性保护

所述的“神树林”是广泛意义上的概念,包括多种因文化信仰而保护的各种森林块.这些森林块不仅有多种的生态功能,也是村社水平生物多样性相对集中的地方.通过对云南楚雄彝族的“神树林”进行了广泛调查,取样比较了自然保护区、村社集体林和“神树林”3种不同管理模式下的森林群落的植物物种多样性,结果表明“神树林”群落的物种总数(67)、样方特有种(17)、Shannon-Wiener物种多样性指数(2.96)都显著高于自然保护区群落(分别为44、8、2.17)和集体林群落(分别为34、4、2.39),表明云南楚雄彝族的“神树林”在当地生物多样性保护和管理中有着不可低估的作用.     

Environmental change in Garry oak (Quercus garryana) ecosystems: the evolution of an eco-cultural landscape

Globally, colonialism resulted in the suppression of aboriginal land management practices, abetted by the concept of terra nullius, “belonging to no one”; the belief that aboriginal people had little influence on or ownership of the land. Until recently, this ideology was entrenched in resource management and policy. Traditional ecological knowledge, historical ecology, archaeology, and palaeoecological research have shown these assumptions to be wrong. In this paper we take a multidisciplinary approach (biogeography, paleoecology, dendrochronology, and bioclimatic envelope modeling) to better understand the role of climate and fire in the formation of eco-cultural landscapes. We synthesize results from pollen and charcoal analysis in Garry oak ecosystems that indicate there were continuous and frequent prescribed burning events, with more severe fires occurring every 26–41 years in southwest British Columbia throughout the Anthropocene (~last 250 years) that substantially altered forest structure and composition. These results are consistent with stand age reconstructions in BC and Washington with Garry oak establishment beginning ~1850 AD, corresponding with modern fire exclusion, aboriginal population decline, and end of the Little Ice Age. Douglas-fir recruitment has been continuous since ~1900, with succession of oak woodland to closed conifer forest at most sites. These findings indicate that the structure of many Garry oak ecosystems have been profoundly influenced by eco-cultural practices. Overwhelming evidence indicates that in many cases these ecosystems are dependent on prescribed fire for their open structure. In the absence of aboriginal land-management practices, active management will be necessary to maintain Garry oak woodland.     

云南楚雄彝族的"神树林"与生物多样性保护

所述的“神树林”是广泛意义上的的概念,包括多种因文化信仰面保护的各种森林块,这些森林块不仅有多种的生态功能,也是村社水平生物多样性相对集中的地方,通过对云南楚雄彝族的“神树林”进行了广泛调查,取样比较了自然保护区,村社集体林和“神树林”３种不同管理模式下的森林群落的植物物种多样性,结果表明“神树林”群落的物种总数的植物物种多样性,结果表明“神树林”群落的物种总数（６７）、样方特有种（１７）、Ｓｈａ     

Field study of beech (t Fagus sylvatica L.) and melojo oak (t Quercus pyrenaica Willd) leaf litter decomposition in the centre of the Iberian Peninsula

The effect of the canopy on leaf decomposition of beech (t Fagus sylvatica) and melojo oak (t Quercus pyrenaica) was studied during a period of 660 days in a mountain forest of central Spain; response of leaves to leaching was also studied to determine the effects of rainfall after leaf fall. Beech leaves lost 5.8% of their weight by leaching, whereas melojo oak leaves lost 13%. Under both types of canopy, beech leaves showed almost no difference in their decomposition patterns, with decay constants of 0.31 and 0.32 respectively. Melojo oak leaves showed quite a different behaviour under both canopies; decay constant was 0.47 under t Quercus pyrenaica and 0.77 under beech canopy. Total immobilization of nitrogen was less in the melojo oak forest. Effects of summer dryness were sharper in the melojo oak forest, where decomposition stopped during the summer. This delay in the decomposition might have been due to the lower canopy density in the melojo oak forest. The decomposition patterns of the leaves of both species, under the canopy of the other species, suggests what might happen in mixed stands. Thus, the presence of melojo oaks in beech forest would increase decay and decrease nitrogen immobilization. The presence of beech trees in melojo oak forests would improve microclimatic conditions and increase decay, whereas beech litter on the soil would immobilizate more nitrogen.     

Effect of agricultural land‐use change on the structure of a temperate forest ant–plant interaction network

Studies on the responses of ant–plant interactions to land‐use change have mainly focused on tropical habitats, usually without considering the impacts on the structure of interaction networks. Here we show that land‐use modifies the structure of the ant–plant interaction networks in a temperate habitat. Ant–plant interactions and plant diversity were recorded in an oak forest and agricultural land in central Mexico. We registered five ant species in the oak forest, and four ant species in the agricultural land. Plant diversity was higher in the agricultural land than in the oak forest. In the ant–plant networks of both sites, our results showed a higher dependence of ants on the plants on which they feed than vice versa, and the ants Formica spp. and the plants Barkleyanthus salicifolius were the species with the most strength and greatest influence in the network structure. The ant–plant network in the oak forest showed a nested structure. However, the network at the agricultural land site showed non‐nestedness; the identity of both ants and plants with the highest values of specialization was different and the number of ant species in the network was decreased, but the number of plant species with which they interacted significantly increased. Both ant–plant networks were equally tolerant to simulated extinction of individual species. We conclude that temperate forest ant–plant networks can be inherently fragile and susceptible to the effects of agricultural land‐use change, not on the number of interacting species but on their identity.     

Evaluating the benefits of CAP reforms: Can afforestations restore bird diversity in Mediterranean Spain?

Current Common Agricultural Policy (CAP) subsidies in the Mediterranean region tend to prioritize afforestation on former arable land with oaks rather than pines because pine plantations would maintain lower biological diversities than native forests. Nevertheless, no thorough evaluations of the conservation values of pine plantations as compared to oak remnants have been carried out to date. We analyze the diversity and conservation value of bird assemblages breeding in 200 remnants of Holm oak Quercus ilex woodlands and 82 mature (>50-year-old) pine plantations in central Spain, a Mediterranean region mostly devoted to arable farming. Species–area relationships were compared between forest types. The conservation value of bird assemblages was assessed using the “Species of European Conservation Concern” (SPEC) classification of Burfield and van Bommel [(2004). Birds in Europe: Populations estimates, trends and conservation status. Cambridge: BirdLife International]. Overall numbers of bird species maintained by oak and pine archipelagoes were rather similar, but species–area relationships differed between forest types. Intercepts were higher in oak fragments, whereas slopes were steeper in pine plantations. Small oak fragments held more species (mainly Mediterranean Sylvia warblers) than plantations, whereas large plantations held more species than large oak remnants. Differences in species–area relationships seemed to be due to differences in vegetation structure, especially understorey shrub cover and tree height and cover. We recorded nine SPECs, all exclusive (6) or near-exclusive (3) to oak woodlands, although such woodlands do not appear to be critical for their conservation. Hence, we conclude that pine afforestations have played a role for maintaining and restoring forest bird communities in the farming landscapes of central Spain. Promoting large and shrubby plantations would enhance their conservation value for breeding birds, together with promoting growth, regeneration and expansion of Holm oak remnants by means of set-aside measures previous or alternative to oak reafforestation. The increasing importance of non-commercial as compared to commercial values of Mediterranean forests would justify subsidizing the proposed policy.     

Forest community structure and composition following containment treatments for the fungal pathogen oak wilt

In temperate oak forests in Ohio, USA, we examined variability in forest communities within containment treatment sites for oak wilt (Bretziella fagacearum), a fungal pathogen lethal to susceptible oak species. Containment treatments included quarantine lines in soil for limiting belowground fungal spread and sanitation cutting of 1–3 mature black oak (Quercus velutina) trees within oak wilt infection patches. At 28 sites, we compared tree structure and understory plant communities across a gradient of 1- to 6-year-old treatments and reference forest (untreated and without evidence of oak wilt). While oak seedlings were abundant, oak saplings (1–10 cm in diameter) were absent. In contrast, many native understory plant community measures were highest in oak wilt treatments. Plant species richness 100 m^?2 doubled in treatments, regardless of age, compared with reference forest. Plant cover increased with treatment age, with 6-year-old treatments exhibiting 5?×?more cover than reference forest. Non-native plants averaged only a small proportion (<?0.12) of cover across treatments and reference forest. Variability in understory communities was mostly predictable using treatment age, tree canopy cover, and geographic location, as 20 of 25 understory measures had at least 72% of their variance modeled. While oak wilt treatments did not facilitate oak regeneration nor many conservation-priority species of open savanna-woodland habitats, the treatments did diversify and increase cover of native understory communities with minimal invasion of non-native plants.

     

黑龙江省不同地点蒙古栎林生态特点研究

通过对黑龙江省６个地点的天然蒙古栎林的结构和更新特点的分析,蒙古栎林可划分为不同特点的蒙古栎群落,即纯蒙古栎群落、蒙古栎桦树群落、蒙古栎槭树群落、蒙古栎红松群落和蒙古栎松混交林群落,其演替趋势如下：红蒙古栎群落、蒙古栎桦树群落至蒙古栎槭树落、蒙古栎红松群落,再至蒙古栎红松混交林群落,蒙古栎群落类型的多样性主要反应群落不同的演替阶段,造成蒙古栎群落多样的原因是人类活动和自然因素作用的结果,随着群落的演替,蒙古栎的优势地位逐渐被消弱,乔木种类丰富度增多,草本种类丰富度增多;蒙栎的相对密度下降,林内环境由于干燥逐渐变中性至较湿润,蒙古栎幼苗和幼树在总幼苗和幼树中所占的比例下降,耐荫物中色木槭等的幼苗和幼树所占比例上升;在演替过程中,蒙古栎分布格局－聚集分布的聚集程度逐渐降低,并向随机分布的方向发展。     

The Impacts of Global and National Policy on the Management and Conservation of Sacred Groves of India

India is home to thousands of community-protected forests, called sacred groves. Sacred forests or groves are sites that have cultural or spiritual significance to the people who live around them. These areas may also be key reservoirs of biodiversity. In India, most sacred groves are managed by a community group, not by a government agency. They are often private or community land, not formal protected areas or parks. This poses an interesting challenge in terms of future management and possible policy relating to the sacred groves. On the international level, organizations such as the International Union for the Conservation of Nature and UNESCO have created guidelines for management of sacred sites. On the national level, India’s past Forest Acts and recent Forest Rights Act have relevance to the sacred groves. Local differences in land tenure also affect the groves. Ethnographic research conducted in 2009 and 2010 in the states of Meghalaya and Karnataka, India, evaluated the historic and current management and beliefs associated with sacred forests. Cultural change and pressure to use natural resources within the groves is leading to reduction of these forest areas. In the future, a creative combination of policy approaches to conserve groves that respects their spiritual values is recommended.     

中国北方蒙古栎林起源和发展的初步探讨

古生态学的研究显示 ,栎林 (包括蒙古栎林 )在华北地区和东北地区的分布在地质历史时期不是一成不变的 ,其优势度也呈现上下波动 ,栎树优势度的增减是由于地质历史时期气候的变化造成的 ;蒙古栎的起源时间可能不早于上新世中期 ,蒙古栎林的起源也应不早于上新世中期。蒙古栎林的起源原因有 2种 ,即火与人类的反复砍伐。火是原生性质蒙古栎林在地质历史时期存在和扩散的根本原因 ,过度的人类活动 (主要是反复砍伐 )是促成现在蒙古栎次生林占优势地位的主要原因 ,这是由蒙古栎这个树种的生物学特性决定的 ,在没有外界因素的干扰和火减少的情况下 ,蒙古栎在演替过程中逐渐被耐荫的树种所取代 ,但是在中国北方 ,由于人类活动的加剧 ,蒙古栎林的优势度有增加的趋势。     

北京东灵山区两种生境条件下辽东栎幼苗补充与建立的比较

 北京东灵山区的人工油松（Pinus tabulaeformis）林中常常混生有萌生的辽东栎（Quercus wutaishanica）种群。局部地带辽东栎与油松形成混交林,在山脊及山坡上部一些特殊生境辽东栎甚至取代油松林成为优势种。为了探讨北京东灵山区辽东栎林的天然更新机制及辽东栎幼苗在人工油松林中的天然更新,在油松人工林的林下和林缘两种生境条件下将辽东栎种子播种于2 cm深的土中,对辽东栎幼苗的补充和建立进行了对照试验。结果显示,在辽东栎结实丰年,两种生境条件下播种后辽东栎种子丢失差异并不显著,而辽东栎幼苗补充和建立方面的差异极显著。由于森林动物特别是啮齿类动物的活动对辽东栎幼苗建立的影响具有二重性：消耗种子与促进萌发,本试验特别关注辽东栎幼苗子叶丢失现象及其丢失后果。由于辽东栎种子萌根较早,在幼苗出土前其粗大的主根长达10～20 cm,并已初步形成根系,表明已有相当比例的营养物质从子叶转移到根部。因此当幼苗出土后如果仅仅只是子叶的丢失对辽东栎幼苗的成活和生长均无明显的影响。通过研究发现辽东栎幼苗期子叶丢失有3种形式：1）仅丢失子叶,幼苗其它部位未受伤害,多发生在林缘生境中,占发芽坚果的29.69%;2）整个幼苗连根被拖出地面,多发生在林下生境,占62.43%;3）在子叶与根颈的连接处主干被咬断,林缘为50.88%,显著高于林下的8.41%。但只有后两种形式才导致幼苗死亡。导致这些现象的原因是由于林缘土壤腐殖质含量低,比较干燥,土质较坚实,以及幼苗出土前坚果主根生长得较长等缘故,当幼苗遭受动物捕食时不会轻易地被拖出地面,拖走的往往仅仅是子叶。至生长季节结束,林缘样方辽东栎幼苗的成活率接近20%,茎干平均高度7.94 cm,芽数11.24·株-1,叶片平均干重为81.14 mg·株-1,且几乎全部由丢失子叶的幼苗所构成;而林下样方辽东栎幼苗的成活率不足2%,茎干平均高度4.74 cm,芽数7.52·株-1,叶片平均干重仅为42.27 mg·株-1,以未丢失子叶的幼苗为主。试验结果表明,林缘及类似林缘的环境条件更有利于辽东栎的实生更新。     

巨桉人工林土壤动物群落结构特征

为深入了解巨桉人工林生物养分循环及土壤肥力变迁,客观全面地评价巨桉人工林的生态效益,对比研究了四川省洪雅县巨桉人工林和青冈次生林土壤动物群落结构特征。研究方法,大型土壤动物采用野外分层手捡计数,中小型土壤动物用取土器分层取土,带回室内分别用干湿漏斗分离并镜检。结果表明：（1）洪雅巨桉人工林共获1312头土壤动物,分属6门10纲21目,其优势类群分别为蜱螨目、弹尾目、线虫纲、膜翅目;（2）巨桉人工林土壤动物群落的类群数和个体数均低于青冈次生林．其Shannon—Wiener指数和密度一类群指数分别为1．818和4．002,低于青冈次生林的1．963和8．091;（3）巨桉人工林土壤动物群落中腐食性同功能种团比例为81．78％高于青冈次生林的64．18％;（4）巨桉人工林与青冈次生林土壤动物密度剖面分布具明显表聚性,二者经F-检验无显著差异;（5）巨桉人工林与青冈次生林土壤动物群落有比较高的Jaccard相似性系数和Gower,系数,表明两类林分在土壤动物类群和个体数量分布上有比较大的相似度。     

Plant biodiversity of two tropical dry evergreen forests in the Pondicherry region of South India and the role of belief systems in their conservation

Natural vegetation on the south-eastern coast of Peninsular India has now been reduced to patches, some of which are preserved as sacred groves. The plant biodiversity and population structure of woody plants (>20 cm girth at breast height; gbh) in two such groves, Oorani and Olagapuram, occurring on the north-west of Pondicherry have been analyzed. A total of 169 angiosperms have been enumerated from both sites. The Oorani grove (3.2 ha) had 74 flowering plant species distributed in 71 genera and 41 families; 30 of them are woody species, 8 are lianas and 4 are parasites. The Olagapuram grove (2.8 ha) was more species-rich with 136 species in 121 genera of 58 families; woody species were fewer (21) while 9 lianas and 3 parasites occurred. The vegetation structure indicates that the Oorani grove is a relic of tropical dry evergreen forest, whereas Olagapuram is reduced to a thorny woodland. The latter is heavily degraded as it has lost the status of a sacred grove because of its conversion to Eucalyptus plantations. The Oorani grove has an Amman temple in the centre. The attendant cultural rites and religious rituals have perpetuated the status of a sacred grove which has ensured the protection of the grove.     

Ecological succession following forest decline in a xeric oak forest of south-central United States

Aims The loss of canopy trees associated with forest decline can greatly influence the species composition and structure of a forest and have major impacts on the ecosystem. We studied the changes in forest composition and structure 1 and 5 years following nearly total canopy mortality on several hundreds of hectares of xeric oak forests in south-central United States. Because the forests were within an ecotonal vegetation type composed of a mosaic of forest, savanna and grassland, we sought to learn whether forest decline areas would recover to forest or change to more open savanna and grassland conditions in the landscape pattern of vegetation. Because low intensity fire shaped the vegetation type, we sought to learn whether fire would keep the decline areas open.Methods The study was conducted in a xeric oak forest in east-central Oklahoma, USA. Randomly located vegetation and regeneration surveys were conducted in decline and non-decline stands 1 and 5 years following nearly total canopy mortality. Diameter at breast height (DBH), regeneration and sprout origin were recorded for all woody species.Important findings The major canopy species post oak (Quercus stellata Wangenh.), blackjack oak (Quercus marilandica Muenchh.) and black hickory (Carya texana Buckl.) suffered 85–92% mortality; however, minor canopy components experienced limited mortality. Mortality affected all size classes of canopy trees except those below 5cm breast height diameter. There was abundant regeneration of all species and fire seemed to maintain a high level of sprouting. Decline appeared to decrease the relative importance of stump sprouting and increase other types including root sprouts. Decline areas had abundant true seedlings, with stem origin from a root with the same diameter as the stem, which is very unusual for xeric oak forests. Regeneration height in decline areas was twice that of non-decline forests. Our findings suggest that forest decline may lead to: (i) reduced oak dominance and species change in the canopy, (ii) change in reproduction type to increase success of true seedlings and maintain genetic diversity of oaks.     

Estimation of Mediterranean forest transpiration and photosynthesis through the use of an ecosystem simulation model driven by remotely sensed data

Aim This paper investigates the use of an ecosystem simulation model, FOREST‐BGC, to estimate the main ecophysiological processes (transpiration and photosynthesis) of Mediterranean coastal forest areas using remotely sensed data. Location Model testing was carried out at two protected forest sites in central Italy, one of which was covered by Turkey oak (Circeo National Park) and the other by holm‐oak (Castelporziano Estate). Methods At both sites, transpiration and photosynthesis measurements were collected in the field during the growing seasons over a four‐year period (1999 and 2001 for the Turkey oak; 1997, 1999 and 2000 for the holm‐oak). Calibration of the model was obtained through combining information derived from ground measurements and remotely sensed data. In particular, remote sensing estimates of the Leaf Area Index derived from 1 × 1‐km NOAA AVHRR Normalized Difference Vegetation Index data were used to improve the adaptation of the model to local forest conditions. Results The results indicated different strategies regarding water use efficiency, ‘water spending’ for Turkey oak and ‘water saving’ for holm‐oak. The water use efficiency for the holm‐oak was consistently higher than that for the Turkey oak and the relationship between VPD and WUE for the holm‐oak showed a higher coefficient of determination (R² = 0.9238). Comparisons made between the field measurements of transpiration and photosynthesis and the model estimates showed that the integration procedure used for the deciduous oak forest was effective, but that there is a need for further studies regarding the sclerophyllous evergreen forest. In particular, for Turkey oak the simulations of transpiration yielded very good results, with errors lower than 0.3 mm H₂O/day, while the simulation accuracy for photosynthesis was lower. In the case of holm‐oak, transpiration was markedly overestimated for all days considered, while the simulations of photosynthesis were very accurate. Main conclusions Overall, the approach offers interesting operational possibilities for the monitoring of Mediterranean forest ecosystems, particularly in view of the availability of new satellite sensors with a higher spatial and temporal resolution, which have been launched in recent years.     

Dense understory dwarf bamboo alters the retention of canopy tree seeds

Tree seed retention is thought to be an important factor in the process of forest community regeneration. Although dense understory dwarf bamboo has been considered to have serious negative effects on the regeneration of forest community species, little attention has been paid to the relationship between dwarf bamboo and seed retention. In a field experiment we manipulated the density of Fargesia decurvata, a common understory dwarf bamboo, to investigate the retention of seeds from five canopy tree species in an evergreen and deciduous broad-leaved mixed forest in Jinfoshan National Nature Reserve, SW China. We found that the median survival time and retention ratio of seeds increased with the increase in bamboo density. Fauna discriminately altered seed retention in bamboo groves of different densities. Arthropods reduced seed survival the most, and seeds removed decreased with increasing bamboo density. Birds removed or ate more seeds in groves of medium bamboo density and consumed fewer seeds in dense or sparse bamboo habitats. Rodents removed a greater number of large and highly profitable seeds in dense bamboo groves but more small and thin-husked seeds in sparse bamboo groves. Seed characteristics, including seed size, seed mass and seed profitability, were important factors affecting seed retention. The results suggested that dense understory dwarf bamboo not only increased seeds concealment and reduced the probability and speed of seed removal but also influenced the trade-off between predation and risk of animal predatory strategies, thereby impacting the quantity and composition of surviving seeds. Our results also indicated that dense understory dwarf bamboo and various seed characteristics can provide good opportunities for seed storage and seed germination and has a potential positive effect on canopy tree regeneration.     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.