The role of tree size in the leafing phenology of a seasonally dry tropical forest in Belize, Central America

Leafing phenology of two dry-forest sites on soils of different depth (S = shallow, D = deep) at Shipstern Reserve, Belize, were compared at the start of the rainy season (April-June 2000). Trees greater than or equal to 2.5 cm dbh were recorded weekly for 8 wk in three 0.04-ha plots per site. Ten species were analysed individually for their phenological patterns, of which the three most common were Bursera simaruba, Metopium brownei and Jatropha gaumeri. Trees were divided into those in the canopy (> 10 cm dbh) and the subcanopy (less than or equal to 10 cm dbh). Site S had larger trees on average than site D. The proportion of trees flushing leaves at any one time was generally higher in site S than in site D, for both canopy and subcanopy trees. Leaf flush started 2 wk earlier in site S than site D for subcanopy trees, but only 0.5 wk earlier for the canopy trees. Leaf flush duration was 1.5 wk longer in site S than site D. Large trees in the subcanopy flushed leaves earlier than small ones at both sites but in the canopy just at site D. Large trees flushed leaves earlier than small ones in three species and small trees flushed leaves more rapidly in two species. Bursera and Jatropha followed the general trends but Metopium, with larger trees in site D than site S, showed the converse with onset of flushing I wk earlier in site D than site S. Differences in response of the canopy and subcanopy trees on each site can be accounted for by the predominance of spring-flushing or stem-succulent species in site S and a tendency for evergreen species to occur in site D. Early flushing of relatively larger trees in site D most likely requires access to deeper soil water reserves but small and large trees utilize stored tree water in site S.

Ecosystem service trade-offs and stakeholder claims - a meso-scale study from a Madagascar biodiversity hotspot

The north-eastern escarpment of Madagascar has been labelled a global biodiversity hotspot due to its extremely high rates of endemic species which are heavily threatened by accelerated deforestation rates and landscape change. The traditional practice of shifting cultivation or "tavy" used by the majority of land users in this area to produce subsistence rice is commonly blamed for these threats. A wide range of stakeholders ranging from conservation to development agencies, and from the private to the public sector has therefore been involved in trying to find solutions to protect the remaining forest fragments and to increase agricultural production. Consequently, provisioning, regulating and socio-cultural services of this forest-mosaic landscape are fundamentally altered leading to trade-offs between them and consequently new winners and losers amongst the stakeholders at different scales. However, despite a growing amount of evidence from case studies analysing local changes, the regional dynamics of the landscape and their contribution to such trade-offs remain poorely understood. This study therefore aims at using generalised landscape units as a base for the assessment of multi-level stakeholder claims on ecosystem services to inform negotiation, planning and decision making at a meso-scale. The presented study applies a mixed-method approach combining remote sensing, GIS and socio-economic methods to reveal current landscape dynamics, their change over time and the corresponding ecosystem service trade-offs induced by diverse stakeholder claims on the regional level. In a first step a new regional land cover classification for three points in time (1995, 2005 and 2011) was conducted including agricultural classes characteristic for shifting cultivation systems. Secondly, a novel GIS approach, termed “landscape mosaics approach” originally developed to assess dynamics of shifting cultivation landscapes in Laos was applied. Through this approach generalised landscape mosaics were generated allowing for a better understanding of changes in land use intensities instead of land cover. As a next step we will try to use these landscape units as proxies to map provisioning and regulating ecosystem services throughout the region. Through the overlay with other regional background data such as accessibility and population density and information from a region-wide stakeholder analysis, multiscale trade-offs between different services will be highlighted. The trade-offs observed on the regional scale will then be validated through a socio-economic ground-truthing within selected sites at the local scale. We propose that such meso-scale knowledge is required by all stakeholders involved in decision making towards sustainable development of north-eastern Madagascar.

Revealing regional deforestation dynamics in north-eastern Madagascar - Insights from multi-temporal land cover change analysis

The north-eastern escarpment of Madagascar harbours the island’s last remaining large-scale humid forest massifs surrounded by a small-scale agricultural mosaic. There is high deforestation, commonly thought to be caused by shifting cultivation practiced by local land users to produce upland rice. However, little is known about the dynamics between forest and shifting cultivation systems at a regional level. Our study presents a first attempt to quantify changes in the extent of forest and different agricultural land cover classes, and to identify the main dynamics of land cover change for two intervals, 1995–2005 and 2005–2011. Over the 16-year study period, the speed of forest loss increased, the total area of upland rice production remained almost stable, and the area of irrigated rice fields slightly increased. While our findings seem to confirm a general trend of land use intensification, deforestation through shifting cultivation is still on the rise. Deforestation mostly affects the small forest fragments interspersed in the agricultural mosaic and is slowly leading to a homogenization of the landscape. These findings have important implications for future interventions to slow forest loss in the region, as the processes of agricultural expansion through shifting cultivation versus intensified land use cannot per se be considered mutually exclusive.

Late-Holocene climate variability and ecosystem responses in Alaska inferred from high-resolution multiproxy sediment analyses at Grizzly Lake

The late-Holocene shift from Picea glauca (white spruce) to Picea mariana (black spruce) forests marked the establishment of modern boreal forests in Alaska. To understand the patterns and drivers of this vegetational change and the associated late-Holocene environmental dynamics, we analyzed radiocarbon-dated sediments from Grizzly Lake for chironomids, diatoms, pollen, macrofossils, charcoal, element composition, particle size, and magnetic properties for the period 4100–1800 cal BP. Chironomid assemblages reveal two episodes of decreased July temperature, at ca. 3300–3150 (ca −1 °C) and 2900–2550 cal BP (ca −2 °C). These episodes coincided with climate change elsewhere in the Northern Hemisphere, atmospheric reorganization, and low solar activity. Diatom-inferred lake levels dropped by ca. 5 m at 3200 cal BP, suggesting dry conditions during the period 3200–1800 cal BP. P. glauca declined and P. mariana expanded at ca. 3200 cal BP; this vegetational change was linked to diatom-inferred low lake levels and thus decreased moisture availability. Forest cover declined at 3300–3100, 2800–2500 and 2300–2100 cal BP and soil erosion as inferred from increased values of Al, K, Si, Ti, and Ca intensified, when solar irradiance was low. Plant taxa adapted to disturbance and cold climate (e.g. Alnus viridis, shrub Betula, Epilobium) expanded during these periods of reduced forest cover. This open vegetation type was associated with high fire activity that peaked at 2800 cal BP, when climatic conditions were particularly cold and dry. Forest recovery lagged behind subsequent climate warming (≤+3 °C) by ca. 75–225 years. Our multiproxy data set suggests that P. glauca was dominant under warm-moist climatic conditions, whereas P. mariana prevailed under cold-dry and warm-dry conditions. This pattern implies that climatic warming, as anticipated for this century, may promote P. glauca expansions, if moisture availability will be sufficiently high, while P. mariana may expand under dry conditions, possibly exacerbating climate impacts on the fire regime.

Beyond deforestation monitoring in conservation hotspots: Analysing landscape mosaic dynamics in north-eastern Madagascar

Due to its extraordinary biodiversity and rapid deforestation, north-eastern Madagascar is a conservation hotspot of global importance. Reducing shifting cultivation is a high priority for policy-makers and conservationists; however, spatially explicit evidence of shifting cultivation is lacking due to the difficulty of mapping it with common remote sensing methods. To overcome this challenge, we adopted a landscape mosaic approach to assess the changes between natural forests, shifting cultivation and permanent cultivation systems at the regional level from 1995 to 2011. Our study confirmed that shifting cultivation is still being used to produce subsistence rice throughout the region, but there is a trend of intensification away from shifting cultivation towards permanent rice production, especially near protected areas. While large continuous forest exists today only in the core zones of protected areas, the agricultural matrix is still dominated by a dense cover of tree crops and smaller forest fragments. We believe that this evidence makes a crucial contribution to the development of interventions to prevent further conversion of forest to agricultural land while improving local land users' well-being.

Mast fruiting of large ectomycorrhizal African rain forest trees: importance of dry season intensity, and the resource limitation hypothesis

Mast fruiting is a distinctive reproductive trait in trees. This rain forest study, at a nutrient-poor site with a seasonal climate in tropical Africa, provides new insights into the causes of this mode of phenological patterning. •  At Korup, Cameroon, 150 trees of the large, ectomycorrhizal caesalp, Microberlinia bisulcata, were recorded almost monthly for leafing, flowering and fruiting during 1995–2000. The series was extended to 1988–2004 with less detailed data. Individual transitions in phenology were analysed. •  Masting occurred when the dry season before fruiting was drier, and the one before that was wetter, than average. Intervals between events were usually 2 or 3 yr. Masting was associated with early leaf exchange, followed by mass flowering, and was highly synchronous in the population. Trees at higher elevation showed more fruiting. Output declined between 1995 and 2000. •  Mast fruiting in M. bisulcata appears to be driven by climate variation and is regulated by internal tree processes. The resource-limitation hypothesis was supported. An ‘alternative bearing’ system seems to underlie masting. That ectomycorrhizal habit facilitates masting in trees is strongly implied.

An ice-core based history of Siberian forest fires since AD 1250

Forest fires play a key role in the global carbon cycle and thus, can affect regional and global climate. Although fires in extended areas of Russian boreal forests have a considerable influence on atmospheric greenhouse gas and soot concentrations, estimates of their impact on climate are hampered by a lack of data on the history of forest fires. Especially regions with strong continental climate are of high importance due to an intensified development of wildfires. In this study we reconstruct the fire history of Southern Siberia during the past 750 years using ice-core based nitrate, potassium, and charcoal concentration records from Belukha glacier in the continental Siberian Altai. A period of exceptionally high forest-fire activity was observed between AD 1600 and 1680, following an extremely dry period AD 1540-1600. Ice-core pollen data suggest distinct forest diebacks and the expansion of steppe in response to dry climatic conditions. Coherence with a paleoenvironmental record from the 200 km distant Siberian lake Teletskoye shows that the vegetational shift AD 1540-1680, the increase in fire activity AD 1600-1680, and the subsequent recovery of forests AD 1700 were of regional significance. Dead biomass accumulation in response to drought and high temperatures around AD 1600 probably triggered maximum forest-fire activity AD 1600-1680. The extreme dry period in the 16th century was also observed at other sites in Central Asia and is possibly associated with a persistent positive mode of the Pacific Decadal Oscillation (PDO). No significant increase in biomass burning occurred in the Altai region during the last 300 years, despite strongly increasing temperatures and human activities. Our results imply that precipitation changes controlled fire-regime and vegetation shifts in the Altai region during the past 750 years. We conclude that high sensitivity of ecosystems to occasional decadal-scale drought events may trigger unprecedented environmental reorganizations under global-warming conditions.

High Bee and Wasp Diversity in a Heterogeneous Tropical Farming System Compared to Protected Forest

It is a globally important challenge to meet increasing demands for resources and, at the same time, protect biodiversity and ecosystem services. Farming is usually regarded as a major threat to biodiversity due to its expansion into natural areas. We compared biodiversity of bees and wasps between heterogeneous small-scale farming areas and protected forest in northern coastal Belize, Central America. Malaise traps operated for three months during the transition from wet to dry season. Farming areas consisted of a mosaic of mixed crop types, open habitat, secondary forest, and agroforestry. Mean species richness per site (alpha diversity), as well as spatial and temporal community variation (beta diversity) of bees and wasps were equal or higher in farming areas compared to protected forest. The higher species richness and community variation in farmland was due to additional species that did not occur in the forest, whereas most species trapped in forest were also found in farming areas. The overall regional species richness (gamma diversity) increased by 70% with the inclusion of farming areas. Our results suggest that small-scale farming systems adjacent to protected forest may not only conserve, but even favour, biodiversity of some taxonomic groups. We can, however, not exclude possible declines of bee and wasp diversity in more intensified farmland or in landscapes completely covered by heterogeneous farming systems.

Changes in biodiversity and vegetation composition in the central Swiss Alps during the transition from pristine forest to first farming

Aim: We investigate the response of vegetation composition and plant diversity to increasing land clearance, burning and agriculture at the Mesolithic–Neolithic transition (c. 6400–5000 bc) when first farming was introduced. Location: The Valais, a dry alpine valley in Switzerland. Methods: We combine high-resolution pollen, microscopic charcoal and sedimentological data to reconstruct past vegetation, fire and land use. Pollen evenness, rarefaction-based and accumulation-based palynological richness analyses were used to reconstruct past trends in plant diversity. Results: Our results show that from c. 5500 cal. yr bc, slash-and-burn activities created a more open landscape for agriculture, at the expense of Pinus and Betula forests. Land clearance by slash-and-burn promoted diverse grassland ecosystems, while on the long term it reduced woodland and forest diversity, affecting important tree species such as Ulmus and Tilia. Main conclusions: Understanding the resilience of Alpine ecosystems to past disturbance variability is relevant for future nature conservation plans. Our study suggests that forecasted land abandonment in the Alps will lead to pre-Neolithic conditions, with significant biodiversity losses in abandoned grassland ecosystems. Thus, management measures for biodiversity, such as ecological compensation areas, are needed in agricultural landscapes with a millennial history of human impact, such as the non-boreal European lowlands. Our study supports the hypothesis that species coexistence is maximized at an intermediate level of disturbances. For instance, species richness decreased when fire exceeded the quasi-natural variability observed during the Mesolithic times. Under a more natural disturbance regime, rather closed Pinus sylvestris and mixed oak forests would prevail.

More efficient aboveground nitrogen use in more diverse Central European forest canopies

We hypothesized that biodiversity improves ecosystem functioning and services such as nutrient cycling because of increased complementarity. We examined N canopy budgets of 27 Central European forests of varying dominant tree species, stand density, and tree and shrub species diversity (Shannon index) in three study regions by quantifying bulk and fine particulate dry deposition and dissolved below canopy N fluxes. Average regional canopy N retention ranged from 16% to 51%, because of differences in the N status of the ecosystems. Canopy N budgets of coniferous forests differed from deciduous forest which we attribute to differences in biogeochemical N cycling, tree functional traits and canopy surface area. The canopy budgets of N were related to the Shannon index which explained 14% of the variance of the canopy budgets of N, suggesting complementary aboveground N use of trees and diverse understorey vegetation. The relationship between plant diversity and canopy N retention varied among regional site conditions and forest types. Our results suggest that the traditional view of belowground complementarity of nutrient uptake by roots in diverse plant communities can be transferred to foliar uptake in forest canopies.

Structure and inferred dynamics of a large grove of Microberlinia bisulcata trees in central African rain forest: the possible role of periods of multiple disturbance events

A 272-ha grove of dominant Microberlinia bisulcata (Caesalpinioideae) adult trees greater than or equal to 50 cm stem diameter was mapped in its entirety in the southern part of Korup National Park, Cameroon. The approach used an earlier-established 82.5-ha permanent plot with a new surrounding 50-m grid of transect lines. Tree diameters were available from the plot but trees on the grid were recorded as being greater than or equal to 50 cm. The grove consisted of 1028 trees in 2000. Other species occurred within the grove. including the associated subdominants Tetraberlinia bifoliolata and T. korupensis. Microberlinia bisulcata becomes adult at a stein diameter of c. 50 cm and at an estimated age of 50 y. Three oval-shaped subgroves with dimensions c. 8 50 in x 13 50 in (90 ha) were defined. For two of them (within the plot) tree diameters were available. Subgroves differed in their scales and intensities of spatial tree patterns, and in their size frequency distributions, these suggesting differing past dynamics. The modal scale of clumping was 40-50 m. Seed dispersal by pod ejection (to c. 50 in) was evident from the semi-circles of trees at the grove's edge and from the many internal circles (100-200 m diameter). The grove has the capacity. therefore, to increase at c. 100 m per century. To form its present extent and structure. it is inferred that it expanded and infilled from a possibly smaller area of lower adult-tree density. This possibly happened in three waves of recruitment, each one determined by a period of several intense disturbances. Climate records for Africa show that 1740-50 and 1820-30 were periods of drought, and that 1870-1895 was also regionally very dry. Canopy openings allow the light-demanding and fast-growing ectomycorrhizal M. bisulcata to establish, but successive releases are thought to be required to achieve effective recruitment. Nevertheless, in the last 50 y there were no major events and recruitment in the grove was very poor. This present study leads to a new hypothesis of the role of periods of multiple extreme events being the driving factor for the population dynamics of many large African tree species such as M. bisulcata.

Rainfall input, throughfall and stemflow of nutrients in a central African rain forest dominated by ectomycorrhizal trees

Incident rainfall is a major source of nutrient input to a forest ecosystem and the consequent throughfall and stemflow contribute to nutrient cycling. These rain-based fluxes were measured over 12 mo in two forest types in Korup National Park, Cameroon, one with low (LEM) and one with high (HEM) ectomycorrhizal abundances of trees. Throughfall was 96.6 and 92.4% of the incident annual rainfall (5370 mm) in LEM and HEM forests respectively; stemflow was correspondingly 1.5 and 2.2%. Architectural analysis showed that ln(funneling ratio) declined linearly with increasing ln(basal area) of trees. Mean annual inputs of N, P, K, Mg and Ca in incident rainfall were 1.50, 1.07, 7.77, 5.25 and 9.27 kg ha(-1), and total rain-based inputs to the forest floor were 5.0, 3.2, 123.4, 14.4 and 37.7 kg ha-1 respectively. The value for K is high for tropical forests and that for N is low. Nitrogen showed a significantly lower loading of throughfall and stemflow in HEM than in LEM forest, this being associated in the HEM forest with a greater abundance of epiphytic bryophytes which may absorb more N. Incident rainfall provided c. 35% of the gross input of P to the forest floor (i. e., rain-based plus small litter inputs), a surprisingly high contribution given the sandy P-poor soils. At the start of the wet season leaching of K from the canopy was particularly high. Calcium in the rain was also highest at this time, most likely due to washing off of dry-deposited Harmattan dusts. It is proposed that throughfall has an important `priming' function in the rapid decomposition of litter and mineralization of P at the start of the wet season. The contribution of P inputted from the atmosphere appears to be significant when compared to the rates of P mineralization from leaf litter.

Drought avoidance and the effect of local topography on trees in the understorey of Bornean lowland rain forest

The water relations of two tree species in the Euphorbiaceae were compared to test in part a hypothesis that the forest understorey plays an integral role in drought response. At Danum, Sabah, the relatively common species Dimorphocalyx muricatus is associated with ridges whilst another species, Mallotus wrayi, occurs widely both on ridges and lower slopes. Sets of subplots within two 4 -ha permanent plots in this lowland dipterocarp rain forest, were positioned on ridges and lower slopes. Soil water potentials were recorded in 1995-1997, and leaf water potentials were measured on six occasions. Soil water potentials on the ridges (-0.047 MPa) were significantly lower than on the lower slopes (-0.012 MPa), but during the driest period in May 1997 they fell to similarly low levels on both sites (-0.53 MPa). A weighted 40-day accumulated rainfall index was developed to model the soil water potentials. At dry times, D. muricatus (ridge) had significantly higher pre-dawn (-0.21 v. -0.57 MPa) and mid-day (-0.59 v. -1.77 MPa) leaf water potentials than M. wrayi (mean of ridge and lower slope). Leaf osmotic potentials of M. wrayi on the ridges were lower (-1.63 MPa) than on lower slopes (-1.09 MPa), with those for D. muricatus being intermediate (-1.29 MPa): both species adjusted osmotically between wet and dry times. D. muricatus trees were more deeply rooted than M. wrayi trees (97 v. 70 cm). M. wrayi trees had greater lateral root cross-sectional areas than D. muricatus trees although a greater proportion of this sectional area for D. muricatus was further down the soil profile. D. muricatus appeared to maintain relatively high water potentials during dry periods because of its access to deeper water supplies and thus it largely avoided drought effects, but M. wrayi seemed to be more affected yet tolerant of drought and was more plastic in its response. The interaction between water availability and topography determines these species' distributions and provides insights into how rain forests can withstand occasional strong droughts.

Reproductive investment and seedling survival of the mast-fruiting rain forest tree, Microberlinia bisulcata A. chev

In the southern part of Korup National Park, Cameroon, the mast fruiting tree Microberlinia bisulcata occurs as a codominant in groves of ectomycorrhizal Caesalpiniaceae within a mosaic of otherwise species-rich lowland rain forest. To estimate the amount of carbon and nutrients invested in reproduction during a mast fruiting event, and the consequential seed and seedling survival, three related field studies were made in 1995. These provided a complete seed and seedling budget for the cohort. Seed production was estimated by counting woody pods on the forest floor. Trees produced on average 26,000 (range 0-92,000) seeds/tree, with a dry mass of 16.6 kg/tree. Seeds were contained in woody pods of mass 307 kg/tree. Dry mass production of pods and seeds was 1034 kg ha(-1), equivalent to over half (55%) of annual leaf litterfall for this species, and contained 13% of the nitrogen and 21% of the phosphorus in annual leaf litterfall. Seed and young-seedling mortality was investigated with open quadrats and cages to exclude vertebrate predators, at two distances from the parent tree. The proportion of seeds on the forest floor which disappeared in the first 6 wk after dispersal was 84%, of which 26.5% was due to likely vertebrate removal, 36% to rotting, and 21.5% to other causes. Vertebrate predation was greater close to the stem than 5 m beyond the crown (41 vs 12% of seeds disappearing) where the seed shadow was less dense. Previous studies have demonstrated an association between mast years at Korup and high dry-season radiation before flowering, and have shown lower leaf-litterfall phosphorus concentrations following mast fruiting. The emerging hypothesis is that mast fruiting is primarily imposed by energy limitation for fruit production, but phosphorus supply and vertebrate predation are regulating factors. Recording the survival of naturally-regenerating M. bisulcata seedlings (6-wk stage) showed that 21% of seedlings survived to 31 mo. A simple three-stage recruitment model was constructed. Mortality rates were initially high and peaked again in each of the next two dry seasons, with smaller peaks in the two intervening wet seasons, these latter coinciding with annual troughs in radiation. The very poor recruitment of M. bisulcata trees in Korup, demonstrated in previous investigations, appears not to be due to a limitation in seed or young-seedling supply, but rather by factors operating at the established-seedling stage.

Contrasting long-term records of biomass burning in wet and dry savannas of equatorial East Africa

Rainfall controls fire in tropical savanna ecosystems through impacting both the amount and flammability of plant biomass, and consequently, predicted changes in tropical precipitation over the next century are likely to have contrasting effects on the fire regimes of wet and dry savannas. We reconstructed the long-term dynamics of biomass burning in equatorial East Africa, using fossil charcoal particles from two well-dated lake-sediment records in western Uganda and central Kenya. We compared these high-resolution (5 years/sample) time series of biomass burning, spanning the last 3800 and 1200 years, with independent data on past hydroclimatic variability and vegetation dynamics. In western Uganda, a rapid (<100 years) and permanent increase in burning occurred around 2170 years ago, when climatic drying replaced semideciduous forest by wooded grassland. At the century time scale, biomass burning was inversely related to moisture balance for much of the next two millennia until ca. 1750 ad, when burning increased strongly despite regional climate becoming wetter. A sustained decrease in burning since the mid20th century reflects the intensified modern-day landscape conversion into cropland and plantations. In contrast, in semiarid central Kenya, biomass burning peaked at intermediate moisture-balance levels, whereas it was lower both during the wettest and driest multidecadal periods of the last 1200 years. Here, burning steadily increased since the mid20th century, presumably due to more frequent deliberate ignitions for bush clearing and cattle ranching. Both the observed historical trends and regional contrasts in biomass burning are consistent with spatial variability in fire regimes across the African savanna biome today. They demonstrate the strong dependence of East African fire regimes on both climatic moisture balance and vegetation, and the extent to which this dependence is now being overridden by anthropogenic activity.