Phenology of plants in relation to ambient environment in a subalpine forest of Uttarakhand,western Himalaya

Observations on phenology of some representative trees, shrubs, under-shrubs and herbs in a subalpine forest of Uttarakhand, western Himalaya were recorded. With the commencement of favorable growth season in April, occurrence of leaf fall was indicatory growth phenomenon in Quercus semecarpifolia, Q. floribunda and Abies spectabilis. However, active vegetative growth in herbaceous species starts onward April and fruit maturation and seed dehiscence are completed from mid of September to October. In general, vegetative growth and reproductive stages in majority of the studied species seems to be dependent on adequate moisture content and also flowering and fruiting in subalpine plants correlate ambient temperature.     

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.     

Ecological and biogeographical analyses of Mediterranean forests of northern Morocco

Mediterranean forests in northern Morocco have been studied. Tree species composition and abundance were investigated in 84 forest sites, distributed throughout the sandstone formation of the Tangier region. The relative abundance of 15 tree and arborescent shrub species was related, by ordination analysis, to altitude, disturbance by fire, and soil fertility. High-elevation forests were dominated by Cedrus atlantica, Quercus pyrenaica and Pinus pinaster. Sacred forests (protected as holy burial places) had a distinct stand structure, and were considered as refuges, for maintaining biodiversity at landscape scale. The evergreen oak Quercus suber and the semideciduous Quercus canariensis co-dominated the lowland forests. The mean basal area of the studied forests was 34 m² ha^–1. The mean trunk size (dbh) for Q. suber was 24 cm and for Q. canariensis 36.5 cm; in both cases there were indications of declining regeneration at population level. A comparative biogeographical analysis with the equivalent region in southern Spain (separated by the Strait of Gibraltar; 14 km wide), indicated a possible biodiversity loss in the Moroccan forests.     

The effect of environmental factors on the regeneration of Quercus semecarpifolia Sm. in Central Himalaya,Nepal

Poor regeneration of oak forest has been observed in north America and Eurasia. In the Himalaya it has mainly been explained by anthropogenic pressures. This study analyses the regeneration of two Quercus semecarpifolia Sm. forests (2–3000 m a.s.l.), where one forest has almost nil disturbance and the other has different degrees of canopy disturbance. The relationships between biotic and abiotic environmental variables and the number of Quercus semecarpifolia seedlings and saplings (i.e., recruits <10 cm Diameter at Breast Height (DBH)) were analysed by means of Generalized Additive Models (GAM). The two forests were compared with respect to size-class distributions of mature trees (i.e., >10 cm DBH), and the number of recruits.The size-class distributions indicate that regeneration is most reliable in the nearly undisturbed forest. Most recruits were found under high canopy cover and high potential radiation. Canopy disturbance has a negative effect on the number of seedlings. Seedlings seem to prefer a pH of around 6, Loss-on-Ignition between 20 and 30 %, and total nitrogen between 2 and 3 %. In general, there were very few plots with saplings, and there was not a clear relationship between the number of saplings and the soil variables. The size-class diagrams indicate a lack of young trees in the disturbed forest, and it is hypothesized that fire may inhibit recruits reaching the canopy phase.     

A Neighborhood Analysis of the Consequences of Quercus suber Decline for Regeneration Dynamics in Mediterranean Forests

In forests, the vulnerable seedling stage is largely influenced by the canopy, which modifies the surrounding environment. Consequently, any alteration in the characteristics of the canopy, such as those promoted by forest dieback, might impact regeneration dynamics. Our work analyzes the interaction between canopy neighbors and seedlings in Mediterranean forests affected by the decline of their dominant species (Quercus suber). Our objective was to understand how the impacts of neighbor trees and shrubs on recruitment could affect future dynamics of these declining forests. Seeds of the three dominant tree species (Quercus suber, Olea europaea and Quercus canariensis) were sown in six sites during two consecutive years. Using a spatially-explicit, neighborhood approach we developed models that explained the observed spatial variation in seedling emergence, survival, growth and photochemical efficiency as a function of the size, identity, health, abundance and distribution of adult trees and shrubs in the neighborhood. We found strong neighborhood effects for all the performance estimators, particularly seedling emergence and survival. Tree neighbors positively affected emergence, independently of species identity or health. Alternatively, seedling survival was much lower in neighborhoods dominated by defoliated and dead Q. suber trees than in neighborhoods dominated by healthy trees. For the two oak species, these negative effects were consistent over the three years of the experimental seedlings. These results indicate that ongoing changes in species’ relative abundance and canopy trees’ health might alter the successional trajectories of Mediterranean oak-forests through neighbor-specific impacts on seedlings. The recruitment failure of dominant late-successional oaks in the gaps opened after Q. suber death would indirectly favor the establishment of other coexisting woody species, such as drought-tolerant shrubs. This could lead current forests to shift into open systems with lower tree cover. Adult canopy decline would therefore represent an additional factor threatening the recruitment of Quercus forests worldwide.     

Establishment and growth of Quercus floribunda seedlings after a mast year

Abstract. Population dynamics, mortality factors, growth and dry weight changes in Quercus floribunda seedlings were studied to determine the implication of sun and shade microsites and the role of mast years on regeneration and maintenance of oak forests in Kumaun Himalaya. The number of seedlings was significantly greater at the sun microsites, but seedling survival was not, as ground herbage clearance at sun microsites accounted for 53.5% mortality. Insect herbivores (e.g. acorn worm) were also one of the major mortality factors. At both microsites, most mortality was encountered between June and August. The initial seedling height at sun microsites was significantly lower compared to shade microsites. Seedlings at sun microsites were superior in terms of number of leaves and peak dry weight. Dry mass allocation in seedling components and root:shoot ratio was similar at both microsites. The study shows the importance of sun microsites and of masting in the potential for regeneration and expansion of Q. floribunda forests.     

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.     

Regeneration Patterns of European Oak Species (Quercus petraea (Matt.) Liebl., Quercus robur L.) in Dependence of Environment and Neighborhood

Quercus robur L. (pedunculate oak) and Quercus petraea (Matt.) Liebl. (sessile oak) are two European oak species of great economic and ecological importance. Even though both oaks have wide ecological amplitudes of suitable growing conditions, forests dominated by oaks often fail to regenerate naturally. The regeneration performance of both oak species is assumed to be subject to a variety of variables that interact with one another in complex ways. The novel approach of this research was to study the effect of many ecological variables on the regeneration performance of both oak species together and identify key variables and interactions for different development stages of the oak regeneration on a large scale in the field. For this purpose, overstory and regeneration inventories were conducted in oak dominated forests throughout southern Germany and paired with data on browsing, soil, and light availability. The study was able to verify the assumption that the occurrence of oak regeneration depends on a set of variables and their interactions. Specifically, combinations of site and stand specific variables such as light availability, soil pH and iron content on the one hand, and basal area and species composition of the overstory on the other hand. Also browsing pressure was related to oak abundance. The results also show that the importance of variables and their combinations differs among the development stages of the regeneration. Light availability becomes more important during later development stages, whereas the number of oaks in the overstory is important during early development stages. We conclude that successful natural oak regeneration is more likely to be achieved on sites with lower fertility and requires constantly controlling overstory density. Initially sufficient mature oaks in the overstory should be ensured. In later stages, overstory density should be reduced continuously to meet the increasing light demand of oak seedlings and saplings.     

Tree population structure of certain Himalayan forest associations and implications concerning their future composition

This study deals with the regeneration status of certain tree components of forests in the northwestern catchment of the river Gola in Kumaun Himalaya and explores possible compositional changes in time.Pinus roxburghii, Quercus floribunda andQuercus lanuginosa in their respective forest types had a stable population structure with a large percentage of young individuals (seedlings) and relatively few old ones, indicating that these species are frequent reproducers. The girth class distribution of the individuals ofQuercus leucotrichophora in theQuercus leucotrichophora forest suggests that this species is likely to disappear gradually. In this forest, although forest composition will remain unchanged for some time, a complete absence of seedlings of any species shows the possibility of its complete replacement by a scrub or a grassland vegetation. In the mixed forest, the dominant species, viz.,Quercus leucotrichophora, Persea odoratissima andBauhinia retusa did not have saplings/seedlings, indicating a possible change in forest composition in the near future whenSapium insigne, Cocculus laurifolius, Celtis eriocarpa andAcer oblongum are likely to gain dominance. Finally, suggestions are made to explain the poor regeneration ofQuercus leucotrichophora.Nomenclature: see Figure 3.Financial support from the Indian Space Research Organisation, Bangalore, is gratefully acknowledged.     

More than just drought: complexity of recruitment patterns in Mediterranean forests

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species (Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems.     

Effects of <Emphasis Type="Italic">Quercus suber</Emphasis> Decline on Woody Plant Regeneration: Potential Implications for Successional Dynamics in Mediterranean Forests

In the last two decades, widespread tree decline and mortality have been documented in forests worldwide. These mortality events usually show certain level of host-specificity, translating into rapid changes in the relative abundance of the adult community. Despite these short-term changes, it is poorly understood whether the decline and mortality of certain tree species are likely to result in long-term vegetation shifts. Trajectories of forest recovery and the probability of occurrence of permanent vegetation shifts are to a large extent determined by post-mortality regeneration dynamics. Using a spatially explicit neighborhood approach, we evaluated the spatial patterns of natural regeneration of the woody plant community in mixed Mediterranean forests affected by the decline of their dominant tree species, Quercus suber. We predicted the abundance, survival, and richness of the seedling and sapling bank as a function of the distribution and health status of the tree and shrub community. Results indicated that Q. suber decline had detectable effects on seedlings and saplings of coexistent woody species from very different functional groups (trees, shrubs, and lianas). The sign and magnitude of these effects varied substantially among coexistent species, which could imply shifts in the species ranking of seedling and sapling abundance, affecting successional trajectories and potentially leading to vegetation shifts. Because most of these changes pointed towards a loss of dominance of Q. suber, management strategies are urgently needed in order to attenuate adult mortality or promote its regeneration, counteracting the negative effects of global change drivers (exotic pathogens, climate change) on these valuable forests.     

Long-term radial growth and climate-growth relationships of Quercus petraea (Matt.) Liebl. and Quercus cerris L. in a xeric low elevation site from Hungary

Sessile oak (Quercus petraea [Matt.] Liebl.) and Turkey oak (Quercus cerris L.) dominated mixed forests are common in low montane and hilly regions in Hungary. Here, we aimed to describe the long-term pattern and climatic responses of the radial growth of Q. petraea and Q. cerris in a xeric low-elevation forest, using retrospective tree-ring analysis for the period 1910–2019. We performed separate analyses with time series of full tree-ring (TRW), earlywood (EW) and latewood (LW) widths. Our results showed that the radial growth of the two species was largely synchronous over time, but became transiently divergent for a 20-year period after a drought in 1968, due to the greater-than-expected growth of Q. cerris and the supressed growth of Q. petraea. Precipitation was the major growth-limiting factor for both species, with a strong positive influence on LW particularly during the current early growing season (March-June), on EW in the previous late summer (August-September) and in the current early spring (March), and on all tree-ring traits in the previous December. The radial growth of both species was negatively related to temperature in the spring (May) and late summer (August) of both current and previous years. The climate-growth relationships showed general instability over time: the most striking temporal change was a gradual shift of positive correlations with precipitation and SPEI during the growing season from spring (March-May) to summer (June-August) since the 1980s over the analysed period. The two species had similarly low growth resistance to droughts in four studied pointer years (1968, 1993, 2002 and 2012), but Q. cerris exhibited a greater capacity to recover over the four post-drought years, and thus higher growth resilience, particularly after the drought of 1968. Our results contribute to the better understanding of the role of climate variability and droughts in the growth of the two co-existing species in transitional locations between closed forests and forest-steppes.     

Diversity and specificity of ectomycorrhizal fungi retrieved from an old-growth Mediterranean forest dominated by Quercus ilex

We analysed the ectomycorrhizal (ECM) fungal diversity in a Mediterranean old-growth Quercus ilex forest stand from Corsica (France), where Arbutus unedo was the only other ECM host. On a 6400 m2 stand, we investigated whether oak age and host species shaped below-ground ECM diversity. Ectomycorrhizas were collected under Q. ilex individuals of various ages (1 yr seedlings; 3-10 yr saplings; old trees) and A. unedo. They were typed by ITS-RFLP analysis and identified by match to RFLP patterns of fruitbodies, or by sequencing. A diversity of 140 taxa was found among 558 ectomycorrhizas, with many rare taxa. Cenococcum geophilum dominated (35% of ECMs), as well as Russulaceae, Cortinariaceae and Thelephoraceae. Fungal species richness was comparable above and below ground, but the two levels exhibited < 20% overlap in fungal species composition. Quercus ilex age did not strongly shape ECM diversity. The two ECM hosts, A. unedo and Q. ilex, tended to share few ECM species (< 15% of the ECM diversity). Implications for oak forest dynamics are discussed.     

Regeneration of oaks (Quercus robur/Q. petraea) and three other tree species during long-term succession after catastrophic disturbance (windthrow)

In broadleaved temperate forests in Europe, oak (Quercus robur/Q. petraea) regeneration is reported to be weak or absent. However, most work concern seedlings or saplings, studied relatively few years. We studied a Picea abies stand in Sweden, windthrown and logged (all stems harvested) in 1969, testing the hypothesis that oaks regenerate in the long term among competing tree species after catastrophic disturbance. In 2011, after 40 years/growth seasons, we recorded live and dead trees in the new stand and investigated the surroundings, competition, and succession. The following trees, up to 26 m tall, colonized: Sorbus aucuparia, Betula pendula/B. pubescens, Fagus sylvatica, Q. robur/Q. petraea, and Corylus avellana (a shrub). Betula dominated, and only Fagus was regenerating in 2011. Sorbus had produced most of the dead trees, mainly or partly through intraspecific competition. In the stand, compared to the surroundings, Quercus, Picea, and Alnus glutinosa were under-represented, and Sorbus, Betula, and Fagus were over-represented. Yet, the density of Quercus was far from negligible; 48 large trees/ha. Most of the oaks (74 %) were co-dominant trees and many grew near Sorbus. Thus, oaks can survive and grow fast among pioneer trees and browsing animals, a conclusion which is supported by the literature (nine studies identified). High mortality of Sorbus is part of a long-term succession, where Fagus might come to dominate. However, Quercus likely will persist, in low density. We propose three key traits contributing to long-term persistence of Q. robur/Q. petraea in European temperate forests: long life span, ecological plasticity, and resistance to disturbances.     

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。     

Comparison between the soil seed banks of a burnt and an unburnt Quercus pyrenaica Willd. forest

Soil samples from burnt and unburnt sites dominated by Quercus pyrenaica Willd. forest in León province (NW Spain) were collected, separated into an upper layer (0–2 cm depth) from a lower layer (2–5 cm), put in a greenhouse, and seeds allowed to germinate.A total of 670 identified seedlings comprising 56 species and belonging to 24 families were recorded. Most seedling were herbaceous perennials (hemicryptophytes: 59%), followed by annuals (therophytes: 23%). Many species that germinated from buried seeds were not observed as plants at any site, and came from exogenous communities. The principal means of seed dissemination were anemochory (45%) and autochory (23%).The number of species as well as seedlings was not significantly different between soils from the burnt and unburnt sites, but seedlings were more numerous in samples from the unburnt sites, as the number of seedlings was always highest in the upper soil layer. Germination behaviour of four particular species was characterized.Seedlings from the seed bank temporarily were found in the early stages of recovery of the burnt Quercus pyrenaica forests.     

Germination of Quercus ilex and Q. pubescens in a Q. ilex coppice

Quercus ilex L. (holm oak) coppices, widespread around the Mediterranean basin, are probably the result of 5 000 years of prolonged human disturbance of the original Quercus pubescens Willd. (downy oak) forests. Since disturbance has almost ceased in recent years, a question arises as to the development of these coppices: will the Q. pubescens forests return, or will Q. ilex remain the dominant species? To investigate the phenomenon, we analyzed the first stages, i.e. germination of the two species in holm oak coppices. Our experiments show that both species germinated better in coppices than in clearings or clear-cuts. Moreover, Q. pubescens appears to be slightly favored above Q. ilex and it is suggested auto-allelopathy is involved, at least partially inhibiting the germination of Q. ilex.     

Long‐term directional changes in upland Quercus forests throughout Oklahoma,USA

Questions: (1) How have the composition and structure of undisturbed upland Quercus forests changed over 50 years across a large region and moisture gradient; (2) What factors are associated with long‐term and broad‐scale changes in these forests? Location: Oklahoma, USA. Methods: We re‐sampled 30 forest stands originally sampled in the 1950s across a large geographical area and compared basal area, tree density, and sapling density between the sampling periods using paired t‐tests, CCA, and DCA. We examined vegetation dynamics in the context of drought indices compiled for the sample period. Results: Total and Quercus stellata basal area and tree density increased, but Q. stellata and Q. marilandica sapling density decreased. Juniperus virginiana and woody species richness increased for all measures. DCA indicated that re‐sampled stands generally changed from Q. stellata–Q. marilandica‐dominated forests to forests with greater woody species richness and more J. virginiana. Q. stellata remained a dominant tree species; otherwise, composition shifted towards mesophytic and invasive woody species. Measurements taken in the 1950s immediately followed a major drought; whereas subsequent decades were significantly moister. Conclusions: Fire exclusion and drought may have played an important role in driving changes towards lower dominance by Quercus, increased importance of mesophytic and invasive species, and greater woody species richness. These phenomena are similar to those found in Quercus‐dominated forests throughout the northern hemisphere.     

Subalpine forest development following a blowdown in the Mount Zirkel Wilderness,Colorado

Abstract. We documented the occurrence of a 1934 blowdown in a subalpine forest in northwestern Colorado, USA. Prior to the blowdown, the stand was dominated by old‐growth Picea engelmannii ‐ Abies lasiocarpa forests. Although blowdowns are believed to trigger outbreaks of Dendroctonus rufipennis (spruce beetle), we found no detectable increase in beetle caused mortality. Forest recovery was by both release of the previously suppressed regeneration and by new seedling establishment. Both recovery pathways were dominated by Abies. The blowdown thus caused a shift in species dominance from Picea to Abies; 65 yr after the blowdown, the fallen logs and tip up mounds continue to provide favourable habitat for seedling establishment of both species. The present study shows that the legacy of blowdowns can influence forest dynamics for decades following the disturbance event.