Impact of land-use type and bark- and leaf-harvesting on population structure and fruit production of the baobab tree (Adansonia digitata L.) in a semi-arid savanna,West Africa

Non-timber forest products (NTFPs) strongly contribute to livelihood security in the semi-arid tropics. There is increasing concern about the population status of NTFP-providing trees and therefore a need for their sustainable use. Thus, this study examines the impact of land-use type on the multipurpose baobab tree (Adansonia digitata L.) in Burkina Faso, combined with rates and patterns of bark- and leaf-harvesting, and their impact on fruit production. We compared stands in a protected area (W National Park of Burkina Faso) with those of surrounding communal area (fallows, croplands and villages) to obtain an indication on the status of the baobab population, to assess its harvesting tolerance and to estimate to what extent their actual use is sustainable. Our results reveal that land-use type has an impact on the population structure of the baobab. The size class distribution curve of park stands was inverse J-shape which indicates good rejuvenation, while the curve of fallows, croplands and villages stands was bell-shaped, indicating a lack of recruitment. However, a high number of seedlings were recorded in villages. Nearly all baobabs were pruned and debarked in villages, croplands and fallows while half of the individuals were harvested in the park. Most of the trees were pruned and debarked moderately. Debarking and pruning were slightly size specific. Pruning in interaction with tree-size had a significant impact on fruit production. In contrast, debarking had no effect on fruit production. We conclude that despite the land-use impact and the intense harvesting, baobabs are still well preserved in the communal area due to their longevity, extremely low adult mortality rates and traditional management practices. However, land-use intensifications may lead to increasing pressure on baobab populations in the future. Therefore, adapted management strategies are needed to guarantee the persistence of this important species and to avoid a shortage of baobab products.     

Baobab (Adansonia digitata L.) density,size-class distribution and population trends between four land-use types in northern Venda,South Africa

Baobabs are important to people's livelihoods as a source of food, fibre and medicine. Baobab products are increasingly being commercialised and exported around the world, and pressure on the resource is growing (Sidibe and Williams, 2002). This paper examines baobab population structure and trends in northern Venda, South Africa.     

Growth and fruit production of Sclerocarya birrea in the South African lowveld

Sclerocarya birrea (A.Rich.) Hochst. subsp. caffra (Sond.) Kokwaro. is a common species throughout the semiarid, deciduous savannas of much of sub-Saharan Africa. It is a favoured species and is frequently maintained in homestead plots and arable fields in an agroforestry situation. Although the abundance and popularity of this species has led to several initiatives to commercialise a number of marula products, the sustainability of the resource base with respect to fruit production has not been considered. This paper reports on a field experiment that monitored growth rates and fruit production of a sample of adult trees from several wild populations. Mean fruit production was 36.8 kg per tree in the first year, and negligible in the second. This was considerably less than previous estimates, which were based largely on small samples or anecdotal reports. The maximum recorded yield was 416.6 kg per tree. Fruit production was positively related to the size of the tree. Growth rates of adult trees were slow. There was a strong positive relationship between mean annual diameter increment and stem diameter. The slow growth rates and low fruit yields indicate that more attention is required regarding the sustainability of there source and its ability to provide sufficient fruit for the growing subsistence and commercial demand at both local and national scales. This revised version was published online in June 2006 with corrections to the Cover Date.     

Individual growth, reproduction and population dynamics of Dioon merolae (Zamiaceae) under different leaf harvest histories in Central Chiapas, Mexico

Leaves of the long-lived Dioon merolae have been harvested intensively for decades (possibly for centuries) for ceremonial purposes by Zoque and mestizo groups inhabiting the Central Depression of Chiapas, Mexico. Over a period of four years, we evaluated vital rates (stem growth, leaf production, reproductive performance, and survival) and projected population growth rates in three populations (250 plants each, divided into eight size classes: new germinants, seedlings, saplings (S1, S2), and adults, A1-A4) with different leaf harvesting histories: non-defoliated by humans for at least 55 years (or very old harvest), defoliated annually until 15 years ago (recovering from harvest), and defoliated annually for at least the past 25 years (currently being harvested intensely). Population structure was affected by leaf harvest history. Stem growth was negatively affected by the annual harvest of leaves in size classes from seedlings up to A4 (ANOVA, P < 0.003); fewer leaves were produced by seedlings, saplings and adults at the annually harvested site (ANOVA, P < 0.027). Survival was high at all sites across all size classes; in the annually harvested site, A4 plants showed a decrease in survival (one dead out of four plants). Sex ratio of adults that produced cones during the four years of study was 61% males to 39% females. At the non-defoliated site, adult classes A2 and A4 produced >80% of the cones; no cones were produced by the A3 and A4 adult size classes at the annually harvested site. Asymptotic estimates of population growth indicated growing populations (λ ≥ 1); the highest mean values of finite population growth rate were obtained in the non-harvested site (λ = 1.0202). Elasticity analysis with population projection matrices indicated that stasis (L, 9-38%) was the component that most contributed to λ, followed by growth (G, 1.2-2.9%), and fecundity (F, 0.2-1.1%). We observed detrimental effects on several vital rates due to continued long-term defoliation, although population growth parameters do not currently suggest a decreased trend as a result of the annual harvest of leaves. The duration of this study of a very long-lived plant species suggests caution when setting levels and frequency of leaf harvest. The results help pinpoint practical recommendations that could be implemented in a sustainable management plan for this species, particularly to increase seed production in the annually harvested site, and recruitment of new germinants and seedlings at all sites. However, sound practices will need to consider the interests of involved stakeholders (landowners, pilgrims, conservation organizations and authorities) to effectively reduce anthropogenic pressure on this endangered species.     

Foliar morphological variation in the white oak Quercus rugosa Née (Fagaceae) along a latitudinal gradient in Mexico: Potential implications for management and conservation

Quercus rugosa Nee (section Quercus) has a distribution from the southern United States to Honduras. This study characterized leaf variation across the whole distribution of the species in Mexico. Ten foliar morphometric characters were scored in each of 241 individuals from 25 localities. A principal component analysis resolved four principal components (PCs) that explained 76.4% of the total variation. A nested analysis of variance revealed significant differences among populations (29.2% average contribution to total variance for the four PCs) and among-trees within populations (39.2%), while 31.6% was due to intra-individual variation. For the first PC (related to leaf size), 52.1% resulted from among population differences. This variation was negatively correlated with latitude (r = −0.86; P < 0.0001), indicating a steep clinal reduction in leaf size from south to north. Mean annual precipitation and an annual aridity index also significantly decreased and increased with latitude, respectively. It is suggested that the morphological cline is the result of plastic and/or adaptive responses to environmental conditions, and indicative of further ecophysiological latitudinal differences among Q. rugosa populations. Additionally, we estimated the magnitude of the least significant difference among populations for the first PC and translated it into a delineation of six latitudinal zones (each with a width of 2°30′), to be considered as preliminary zones for the movement of Q. rugosa seeds with management and conservation purposes, including management in response to global warming.     

Tree rings indicate different drought resistance of a native (Abies alba Mill.) and a nonnative (Picea abies (L.) Karst.) species co-occurring at a dry site in Southern Italy

Climate changes induced by the anthropogenic alteration of the atmospheric radiative balance are expected to change the productivity and composition of forest ecosystems. In Europe, the Mediterranean is considered one of the most vulnerable regions according to climatic forecasts and simulations. However, although modifications in the inter-specific competition are envisaged, we still lack a clear understanding of the ability of the Mediterranean vegetation to adapt to climate changes. We investigated how two co-occurring tree species commonly used in afforestation programmes, the native Abies alba Mill. and the nonnative Picea abies L. Karst., adapt to climate change by assessing their growth performance and physiological responses in relation to past climate variability. Growth was addressed by analysing tree-ring width and carbon and oxygen stable isotopes. Statistical relationships between isotopic value and monthly climate data suggest that the two species underwent ecophysiological adaptation to Mediterranean climatic constraints. These adaptations are also expressed in the ring-width data. Based on the carbon isotope ratio reflecting the stomatal response to drought, we found that the precipitation in the first period of the growing season, i.e. early spring, is a major factor influencing the annual growth of A. alba, which although native, proved to be sensitive to drought. P. abies, on the other hand, showed a higher tolerance to summer drought stress. These findings should help define criteria for sustainability and effective forest conservation in the Mediterranean region.     

Diversity of the calabash tree (<Emphasis Type="Italic">Crescentia cujete</Emphasis> L.) in Colombia

Germplasm of the calabash tree (Crescentia cujete L.) was collected in five major regions of Colombia, i.e. the Andes, Caribbean, Amazon, Orinoco, and Pacific regions. Collecting this multipurpose tree was guided by the indigenous knowledge of farmers and artisans in each region. Large variation in fruit shapes and sizes was found, of which some forms were typical for certain regions. Overall 56 accessions were collected and roughly classified into 22 types by eight fruit shapes and eight sizes. Molecular markers (Amplified fragment length polymorphisms) were applied to leaf tip tissue originating from vegetatively propagated plants in order to assess the diversity available in the germplasm collected as well as to detect patterns of geographical or morphological similarity. One accession each of C. alata H.B.&K. and C. amazonica Ducke were used as outgroups. Overall, genetic diversity was high (mean Nei and Li’s coefficient of 0.43). No relations could be established between either geographical provenance or fruit morphology and patterns of genetic diversity. Concerning the outgroups, the C. amazonica accession appeared to be a distinct species. The C. alata accession, however, did not seem to be sufficiently distinct from C. cujete to merit species status. The latter material may in fact be a hybrid or serve to challenge the validity of interspecific organization of the genus Crescentia.

Annual variation in predation and dispersal of Arolla pine (Pinus cembra L.) seeds by Eurasian red squirrels and other seed-eaters

Seed predation and dispersal are key processes in the survival and distribution of plant species. Many animals cache seeds for later consumption, and, failing to recover some of these seeds, act as seed dispersers, influencing post-dispersal seed and seedling survival. Both animal and plant benefit from scatterhoarding and natural selection of seed characteristics and adaptations of seed predators (and dispersers) is one of the most important examples of co-evolution and mutualism. We studied the producer–consumer Arolla pine (Pinus cembra)–red squirrel (Sciurus vulgaris) system in a subalpine forest in the Italian Alps. Arolla pine produced large seed-crops (masting) at irregular intervals, followed by years with poor or moderate seed production. Squirrel density fluctuated in synchrony with the food resource, eliminating the time-lag normally present when resources are produced in pulses. In all years except 2009 (a mast-crop year), all Arolla pine cones were harvested (their seeds consumed and/or cached) by September to late October by different species. Both squirrels and nutcrackers (Nucifraga caryocatactes) fed on seeds, and their relative pre-dispersal predation rates (on cones in the canopy) differed between years. Overall, nutcrackers consumed more seeds between July and October than squirrels, but in 1 year squirrels took the largest number of seeds. Pre-dispersal seed predation by squirrels tended to be lower in years with large seed-crop size and there was a positive correlation, over the entire study period, between density of recovered hoards and Arolla pine seed density of the previous year. We conclude that (i) squirrels and nutcrackers are important pre-dispersal seed predators and seeds dispersers; (ii) squirrels are also post-dispersal seed predators, and (iii) the proportion of cached seeds consumed by squirrels increased with the size of the Arolla pine seed-crop, suggesting that red squirrel is a conditional mutualistic scatterhoarder of Arolla pine seeds.     

Driving factors for natural tree rejuvenation in anthropogenic pine (Pinus sylvestris L.) forests of NE Germany

The rejuvenation ecology of three main tree species in anthropogenic pine (Pinus sylvestris L.) forests is explored in our study. We focus on the scale of micro-plots, which provide the safe sites for tree rejuvenation. We thrive on the multi-factorial relationship of tree establishment and driving ecological factors using a large dataset from pine stands in NE Germany and applying multivariate analyses. The success of the establishment of the investigated focal tree species Fagus sylvatica L., Quercus petraea Liebl. and Pinus sylvestris L. is, on general, mostly affected by three factors, i.e. water balance of the upper soil layers, browsing pressure, and diaspore sources. Our investigations on the micro-plot scale revealed species-specific differences. For beech saplings <50 cm growth height, primarily the availability of water, indicated by available water capacity (AWC), thickness, quality, and structure of the organic layer, silt and humus content in the topsoil, and the lack of a dense competitive herb layer, were identified as most important factors. On the contrary, oak seems hardly be restricted by hydrologic and/or trophic deficits in the topsoil or humus layer. In conclusion and comparison to Fagus sylvatica L., we assume for Quercus petraea Liebl. advantages in natural regeneration processes under sub-continental climate conditions and thus under the scenarios of climate change. Pinus sylvestris L. regeneration in our investigation area occurs only in a narrow niche. We conclude with regard to future forest development and the objective of stand conversion with low management intensity that oak should be favoured within natural stand regeneration.     

Senescence of Manilkara zapota trees and implications for large frugivorous birds in the Southern Yucatan Peninsula,Mexico

It has long been established that mature forests are mosaics of patches in different development phases but it has seldom explicitly been taken into account in ecological studies. We demonstrate here that these development phases, which are related to the population dynamics of trees, play an important role in the distribution of fauna based on observations on frugivorous birds. In an area close to the Calakmul Biosphere Reserve in Mexico, we studied the abundance of large forest bird species in relation to forest development phases, with a methodology that seems promising for ecological diagnosis and prognosis in forest management planning. Fine-scale forest mapping and bird counts were carried out in two block-transects of 40 m × 3000 m. Tree sampling in a sub-transect was used to generate population characteristics of trees. Large bird species preferred mature or senescent forest patches, whereas relatively young, growing forest patches were avoided. Important large tree species such as Manilkara zapota, Thouinia paucidentata, Guaiacum sanctum and Esenbeckia pentaphylla, characteristic of older forest patches, showed skewed size distributions indicating stress or overexploitation. The population of M. zapota, a key fruiting species that accounted for 26.5% of the total woody biomass, was most heavily affected by stress. A future collapse in the population of M. zapota, a decrease of the total area of older forest, and a decline in the abundance of large birds is likely if stress on the system continues at this level.     

Seasonal patterns of photochemical capacity and spring phenology reveal genetic differentiation among native Scots pine (Pinus sylvestris L.) populations in Scotland

Environment-driven genetic differentiation among populations is a common feature among forest trees, and an understanding of how populations have adapted to their home site conditions is essential for management and conservation practices. In Scotland, 84 native Scots pine (Pinus sylvestris L.) woodlands are recognised by the Forestry Commission and they occupy highly diverse environments from the maritime west coast to continental sites in eastern Scotland. However, it is not known whether adaptations to local environments along sharp temperature and rainfall gradients have occurred in different populations and as a result, the seed transfer guidelines of the species are based only on data from isozymes and monoterpenes. In this study of an outdoor common-garden trial, we used chlorophyll fluorescence to examine whether seedlings from 32 open-pollinated families and eight populations from sites experiencing contrasting annual temperature regimes differed in their response to variation in natural outdoor temperatures between September 2009 and May 2010. In addition, growth initiation in spring was recorded. Photochemical capacity at photosystem II F_v/F_m showed a distinct seasonal trend and remained at relatively high levels (∼0.7) until November. Following a period of over 2 weeks with temperatures below or close to 0 °C, F_v/F_m started decreasing towards its minimum values recorded in early March when population means varied between 0.35 and 0.45. By early May and along with rising temperatures, photochemical capacity had recovered to the same level as observed in early November. Populations were found to respond differently to the cold period starting in December. The largest drop in photochemical capacity was observed in seedlings from a low-altitude population located in the maritime western Scotland, while in seedlings from higher-altitude locations in the cooler eastern Scotland, the response was smaller. In March, the recovery of photochemical capacity was slowest in seedlings from the mildest and coolest sites. Evidence of adaptive genetic differentiation was also found in spring phenology. Initiation of shoot elongation and needle flush were earlier in families from higher altitudes (cooler areas), but population differences were not significant at the α = 0.05 level. These results suggest that adaptation to the spatially heterogeneous environment in Scotland has taken place in Scots pine and that in order to minimise the risk of planting maladapted seed stock, the patterns of environmental and adaptive genetic variation should be taken into account in the management of genetic resources in this species.     

Seed and seedling survival of African mahogany (Entandrophragma spp.) in the Central African Republic: Implications for forest management

African mahoganies of the genus Entandrophragma are among the most valuable and important timber species harvested in Central Africa, representing more than 70% of total export volume from many areas. In spite of the importance of these species, relatively little is known about their regeneration ecology and little effort has gone into understanding the reasons for the consistently reported regeneration failures after logging. I assessed seed survival to germination (Entandrophragma angolense) and seedling survivorship (E. cylindricum) in three different forest types – monodominant Gilbertiodendron, mixed species, and fallow forest – under three different treatments – control, small mesh chicken wire, and large mesh chicken wire – to evaluate the relative importance of different causes of mortality. All seeds were eaten in controls and in both exclosure treatments within Gilbertiodendron forest in a matter of days. Seed survivorship to germination within exclosures in mixed species and fallow forest increased by approximately 10 and 25%, respectively, compared to Gilbertiodendron forest. Six-month seedling survivorship in controls was 37, 12, and 9% in Gilbertiodendron, mixed species and fallow forest, respectively. Seedling mortality was due to different causes in each forest type. In Gilbertiodendron forest controls, an equivalent percentage of seedlings died due to fungal and insect attack (27 and 28%, respectively), while in mixed species forest controls 28 and 55% of seedlings died of these causes, respectively. In fallow forest controls, 48% of seedlings died from predation and/or uprooting by small mammals, all in the first few weeks post-sowing; insect attack (26%) and drought (13%) were other important causes of seedling deaths. Protecting seedlings with exclosures had a dramatic effect on seedling survivorship within the fallow forest, increasing to over 50%.     

Impact of bark beetle (Ips typographus L.) disturbance on timber production and carbon sequestration in different management strategies under climate change

The likely environmental changes throughout the next century have the potential to strongly alter forest disturbance regimes which may heavily affect forest functions as well as forest management. Forest stands already poorly adapted to current environmental conditions, such as secondary Norway spruce (Picea abies (L.) Karst.) forests outside their natural range, are expected to be particularly prone to such risks. By means of a simulation study, a secondary Norway spruce forest management unit in Austria was studied under conditions of climatic change with regard to effects of bark beetle disturbance on timber production and carbon sequestration over a time period of 100 years. The modified patch model PICUS v1.41, including a submodule of bark beetle-induced tree mortality, was employed to assess four alternative management strategies: (a) Norway spruce age-class forestry, (b) Norway spruce continuous cover forestry, (c) conversion to mixed species stands, and (d) no management. Two sets of simulations were investigated, one without the consideration of biotic disturbances, the other including possible bark beetle damages. Simulations were conducted for a de-trended baseline climate (1961–1990) as well as for two transient climate change scenarios featuring a distinct increase in temperature. The main objectives were to: (i) estimate the effects of bark beetle damage on timber production and carbon (C) sequestration under climate change; (ii) assess the effects of disregarding bark beetle disturbance in the analysis.Results indicated a strong increase in bark beetle damage under climate change scenarios (up to +219% in terms of timber volume losses) compared to the baseline climate scenario. Furthermore, distinct differences were revealed between the studied management strategies, pointing at considerably lower amounts of salvage in the conversion strategy. In terms of C storage, increased biotic disturbances under climate change reduced C storage in the actively managed strategies (up to −41.0 tC ha⁻¹) over the 100-year simulation period, whereas in the unmanaged control variant some scenarios even resulted in increased C sequestration due to a stand density effect.Comparing the simulation series with and without bark beetle disturbances the main findings were: (i) forest C storage was higher in all actively managed strategies under climate change, when biotic disturbances were disregarded (up to +31.6 tC ha⁻¹ over 100 years); and (ii) in the undisturbed, unmanaged variant C sequestration was lower compared to the simulations with bark beetle disturbance (up to −69.9 tC ha⁻¹ over 100 years). The study highlights the importance of including the full range of ecosystem-specific disturbances by isolating the effect of one important agent on timber production and C sequestration.     

Understorey plant community characteristics and natural hardwood regeneration under three partial harvest treatments applied in a northern red oak (Quercus rubra L.) stand in the Great Lakes-St. Lawrence forest region of Canada

Throughout eastern North America, stands of northern red oak (Quercus rubra L.) are undergoing successional replacement by shade-tolerant competitors. In the Great Lakes-St. Lawrence (GLSL) forest region, Q. rubra approaches the northern limit of its distribution, and ecosystem-specific silvicultural directives are needed to promote regeneration. We used an inductive, ordination-based approach to explore patterns in understorey plant community composition and microenvironment under different partial harvest treatments applied in a GLSL hardwood stand, and related these to characteristics of natural seedlings of Q. rubra and its competitors Acer rubrum and Acer saccharum.Two years after harvest, we established 2 m × 2 m plots in a stratified random design under 70% (n = 20) and 50% (n = 19) crown closure uniform shelterwood, group selection (n = 15), and uncut upper slope (n = 10) and lower slope (n = 10) areas. Percent cover of understorey vascular plant species, and a suite of microclimatic and edaphic variables were measured in each plot. Density, mean diameter and mean height of seedlings in the understorey (height <1 m) were determined in each plot for Q. rubra, A. rubrum and A. saccharum.Correspondence analysis (CA) ordination extracted two major axes explaining 21.6% of the total inertia in the species cover by plot matrix. Axis one separated uncut plots from the 50% shelterwood along a gradient of canopy cover associated with partial harvest treatments. Plot scores on axis one (13.2%) reflected a shift in dominance of the understorey from shade-tolerant Acer spp. to shade-intolerant colonizers, Rubus idaeus and Carex spp. Plot scores on axis one were directly (p < 0.05) associated with total understorey plant cover, litter depth, soil temperature and pH, but not with measures of plant diversity. Axis two (8.4%) separated plots from upper slope and lower slope areas, and plot scores were inversely associated (p < 0.05) with soil pH, phosphorus and nitrogen levels. Along axis two there was a shift in dominance from competitive (e.g. A. saccharum) to stress-tolerant (e.g. A. rubrum) species as soil fertility declined. Stepwise linear regression indicated seedling diameter in Q. rubra, A. rubrum and A. saccharum was inversely related to canopy cover. This suggests all three species benefited from partial harvest, although the relationship was strongest in Q. rubra. Patterns in understorey composition, microenvironment and seedling characteristics provide the basis to identify the main competitors of Q. rubra seedlings and adjust regeneration efforts along gradients of canopy closure and soil fertility under partial harvest systems within the GLSL forest region.