Drought-induced forest decline: causes, scope and implications

A large number of episodes of forest mortality associated with drought and heat stress have been detected worldwide in recent decades, suggesting that some of the world's forested ecosystems may be already responding to climate change. Here, we summarize a special session titled 'Drought-induced forest decline: causes, scope and implications' within the 12th European Ecological Federation Congress, held in ávila (Spain) from 25 to 29 September 2011. The session focused on the interacting causes and impacts of die-off episodes at the community and ecosystem levels, and highlighted recent events of drought- and heat-related tree decline, advances in understanding mechanisms and in predicting mortality events, and diverse consequences of forest decline. Talks and subsequent discussion noted a potentially important role of carbon that may be interrelated with plant hydraulics in the multi-faceted process leading to drought-induced mortality; a substantial and yet understudied capacity of many forests to cope with extreme climatic events; and the difficulty of separating climate effects from other anthropogenic changes currently shaping forest dynamics in many regions of the Earth. The need for standard protocols and multi-level monitoring programmes to track the spatio-temporal scope of forest decline globally was emphasized as critical for addressing this emerging environmental issue.     

Patterns and trends in tropical forest cover

Abstract

In the 2005 edition of the Global Forest Resources Assessment of the Food and Agriculture Organization of the United Nations, a moderate negative trend was reported regarding the change of tropical forests: the net annual change was estimated at ?11.8 million ha for the period 2000–2005, while the rate was ?11.65 for the previous decade. Tropical Asia showed the highest rate and most negative trend, passing from ?0.8% to ?0.96% per year. The remote sensing survey done for previous Forest Resource Assessment editions covering the period 1980–2000 revealed distinct change processes in the three tropical regions. Survey results indicated that socio‐economic and cultural aspects that characterise and differentiate the geographic regions determine the nature of the change processes and underlying cause–effect mechanisms, while the ecological setting determines the intensity of change and reveals its environmental implications. A comparison of deforestation processes of the two decades indicated an on‐going process of “radicalisation” of the dynamics determined by an increasing frequency of high‐gradient changes (e.g. total clearing rather than fragmentation and degradation) and by a shift of deforestation fronts towards wetter zones, with a consequent higher per‐hectare carbon emission associated with deforested areas.     

Where can managers effectively resist climate-driven ecological transformation in pinyon–juniper woodlands of the US Southwest?

Pinyon–juniper (PJ) woodlands are an important component of dryland ecosystems across the US West and are potentially susceptible to ecological transformation. However, predicting woodland futures is complicated by species-specific strategies for persisting and reproducing under drought conditions, uncertainty in future climate, and limitations to inferring demographic rates from forest inventory data. Here, we leverage new demographic models to quantify how climate change is expected to alter population demographics in five PJ tree species in the US West and place our results in the context of a climate adaptation framework to resist, accept, or direct ecological transformation. Two of five study species, Pinus edulis and Juniperus monosperma, are projected to experience population declines, driven by both rising mortality and decreasing recruitment rates. These declines are reasonably consistent across various climate futures, and the magnitude of uncertainty in population growth due to future climate is less than uncertainty due to how demographic rates will respond to changing climate. We assess the effectiveness of management to reduce tree density and mitigate competition, and use the results to classify southwest woodlands into areas where transformation is (a) unlikely and can be passively resisted, (b) likely but may be resisted by active management, and (c) likely unavoidable, requiring managers to accept or direct the trajectory. Population declines are projected to promote ecological transformation in the warmer and drier PJ communities of the southwest, encompassing 37.1%–81.1% of our sites, depending on future climate scenarios. Less than 20% of sites expected to transform away from PJ have potential to retain existing tree composition by density reduction. Our results inform where this adaptation strategy could successfully resist ecological transformation in coming decades and allow for a portfolio design approach across the geographic range of PJ woodlands.     

Amazonian peatlands: an ignored C sink and potential source

In tropical lowlands, peatlands are commonly reported from Southeast Asia, and especially Indonesian tropical peatlands are known as considerable C sinks and sources. In contrast, Amazonia has been clearly understudied in this context. In this study, based on field observations from 17 wetland sites in Peruvian lowland Amazonia, we report 0–5.9 m thick peat deposits from 16 sites. Only one of the studied sites did not contain any kind of peat deposit (considering pure peat and clayey peat). Historic yearly peat and C accumulation rates, based on radiocarbon dating of peat samples from five sites, varied from 0.94 ± 0.99 to 4.88 ± 1.65 mm, and from 26 ± 3 to 195 ± 70 g C m⁻², respectively. The long-term apparent peat and C accumulation rates varied from 1.69 ± 0.03 to 2.56 ± 0.12 mm yr⁻¹, and from 39 ± 10 to 85 ± 30 g C m⁻² yr⁻¹, respectively. These accumulation rates are comparable to those determined in the Indonesian tropical peatlands. Under altered conditions, Indonesian peatlands can release globally relevant amounts of C to the atmosphere. Considering the estimated total area of Amazonian peatlands (150 000 km²) close to that of the Indonesian ones (200 728 km²) as well as several factors threatening the Amazonian peatlands, we suggest that the total C stocks and fluxes associated with Amazonian peatlands may be of global significance.     

Uptake of water by lateral roots of small trees in an Amazonian Tropical Forest

A pulse chase technique was used to determine depth and breath of plant water uptake in an Amazonian evergreen forest. Two 2×2 m² plots were irrigated with deuterated water. The deuterium pulse, measured as D values of soil and plant sap water, was followed in the soil water profile and in stem water of small trees inside and up to 12 m away from the irrigated plots. The deuterium pulse percolation rate was measured to be approximately 0.25 m/month and similar to a previous study in central Amazon. There was little horizontal movement of label through the soil profile; allowing us to conclude that any evidence of label in plants away from the irrigation plots implies the presence of their roots inside the irrigation plots. The bulk of label uptake occurred in plants inside the irrigation plots. However, there were a few individuals as far as 10 m away picking up the label from the irrigation plots. This labeling pattern leads to the conclusion that small trees may have a core of water absorbing roots close to their main trunk, with some roots meandering far from their main trunk.     

Canopy recovery after drought dieback in holm-oak Mediterranean forests of Catalonia (NE Spain)

Climate change is likely to produce more frequent and longer droughts in the Mediterranean region, like that of 1994, which produced important changes in the Quercus ilex forests, with up to 76% of the trees showing complete canopy dieback. At the landscape level, a mosaic of responses to the drought was observed, linked to the distribution of lithological substrates. Damage to the dominant tree species (Q. ilex) and the most common understorey shrub (Erica arborea) was more noticeable on the compact substrates (breccia) than on the fissured ones (schist). This result was consistent with observations documenting deeper root penetration in schist than in breccia materials, allowing the plants growing on fissured substrates to use water from deeper soil levels. Smaller plants were more vulnerable to drought than larger plants in the trees, but not in the shrubs. Overall, Q. ilex was more affected than E. arborea. The resilience of the system was evaluated from the canopy recovery 1 year after the episode. Stump and crown resprouting was fairly extensive, but the damage pattern in relation to substrate, plant size, and species remained similar. The effect of recurrent drought episodes was studied on vegetation patches of Q. ilex located on mountain slopes and surrounded by bare rock. We observed that plants that resprouted weakly after a previous drought in 1985 were more likely to die or to produce poor regeneration in 1995 than plants that had resprouted vigorously. Vegetation patches located on the lower part of the slope were also less damaged than patches situated uphill. The study provides evidence of relevant changes in forest canopy as a consequence of extreme climate events. The distribution of this effect across the landscape is mediated by lithological substrate, causing patchy patterns. The results also support the hypothesis that recurrent droughts can produce a progressive loss of resilience, by depleting the ability of surviving plants to regenerate.     

Entomogenous fungi in tropical forest ecosystems: an appraisal

Abstract. 1. Species of the genus Cordyceps (Ascomycotina; Clavicipitales) are the commonest fungi encountered on arthropods in tropical forests.
2. The asexual states of Cordyceps may occur in conjunction with or separately from the perfect state and are classified in the genera Hirsutella, Hymenostilbe, Nomuraea, Paecilomyces and Verticillium of the imperfect fungi (Deuteromycotina; Moniliales).
3. Most Cordyceps species have a restricted host range and this rigid host specificity, at the generic or tribal level, is especially evident in ant-fungal associations.
4. Pathogenicity was not tested but circumstantial evidence is presented which supports the view that Cordyceps species are primary pathogens of arthropods.
5. Observations of living, infected ants suggest that behavioural patterns are radically altered, possibly altruistically orientated.
6. It is thought that entomogenous fungi are involved in the regulation of arthropod populations and may help to maintain stability in tropical forest ecosystems.
7. The value of these organisms to man may be in the metabolites they produce rather than in their direct usage as biological control agents of insect pests.     

Tree Mortality and Collecting Botanical Vouchers in Tropical Forests1

There is growing concern about the potential impact of reseatchers on tropical forest ecology, but few data. The aim of this paper is to examine the effects of collecting botanical specimens from tropical forest trees on their subsequent survivorship, using mortality data from plots in Amazonian Peru that were established in 1989 and reinventoried in 1994. In total, 2017 trees were originally tagged and collections were made from 948 trees. Making voucher collections always involved using unsterilized telescopic plant collecting poles to cut representative small branches, and sometimes also involved using iron-spiked tree-climbing gear to gain access to the canopy. Annual mortality in the four plots averaged 1.99 percent. Among the whole population of dicotyledenous trees, there was no detectable difference between the mortality rate of collected trees (1.96%) and noncollected trees (2.29%). We conclude that in spite of the physical damage caused to collected trees, collecting voucher specimens from tropical moist forest trees may not affect their survivorship, at least in the short-term. Further studies are needed to fully evaluate the potential impacts of research activities on petmanent forest plots in the tropics.     

Annual wood production in a tropical rain forest in NE Costa Rica linked to climatic variation but not to increasing CO2

Increased atmospheric [CO₂] could theoretically lead to increased forest productivity (‘CO₂ fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long‐term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO₂] on annual wood production in a neotropical rain forest. Our study area was a meso‐scale section (600 ha) of old‐growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO₂] on wood production. A remarkably simple two‐factor model explained 91% of the interannual variance in stand‐level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r²=0.85) and night‐time temperature (negative effect, r²=0.42). Stand‐level tree mortality increased significantly with night‐time temperature. After accounting for dry season rainfall and night‐time temperature, there was no effect of annual [CO₂] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night‐time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer.     

Potential Biomass Accumulation in Amazonian Regrowth Forests

Biomass accumulation in the secondary forests of abandoned pastures and slash-and-burn agricultural fallows is an important but poorly constrained component of the regional carbon budget for the Brazilian Amazon. Using empirical relationships derived from a global analysis, we predicted potential aboveground biomass accumulation (ABA) for the region's regrowth forests based on soil texture and climate data. For regrowth forests on nonsandy soils, the globally derived relationship provided a nearly unbiased linear predictor of Amazonian validation data consisting of 66 stands at seven sites; there was no significant difference between stands that regrew following use as pasture land and those that regrew following slash-and-burn agriculture. For regrowth forests on nonsandy soil, the 1 sigma error range of our ABA model was 58%–171% for the Amazonian validation data. For regrowth forests on sandy soils, the validation data were limited to 19 stands at one site, and the globally derived relationship was substantially biased multiplicatively and nonlinearly. Hence we developed a regional refinement by adding to our validation data ABA values from the two Amazonian sites with sandy soil that had previously been included in the global analysis. Based on a conservative jackknife goodness-of-fit assessment (leaving out one site at a time), we calculated a 1 sigma error range of 42%–158% for our sandy soil Amazonian regrowth forest ABA model. We present our predictions of potential regrowth forest ABA as a set of 0.5° resolution maps for the region at 5, 10, and 20 years following abandonment. Received 6 September 2000; accepted 19 April 2001.     

Resource distribution and soil moisture content can regulate bait control in an ant assemblage in Central Amazonian forest

Resources influence population growth, interspecific interactions, territoriality and, in combination with moisture content, affect terrestrial arthropod distribution and abundance. Ants are usually described as interactive and compete in transitive hierarchies, where the dominants behaviourally exclude subordinate species from food resources. In this study, we evaluated the effects of (i) dominant ants, soil moisture and an artificial resource gradient on the number of ant species attracted to baits; and (ii) how soil moisture and an artificial resource gradient change the number of controlled baits in a Central Amazonian rain forest. We sampled 30 100‐m‐long transects, located at least 200 m apart. The transects were established with six different bait densities varying between six and 41 baits and the soil moisture content was measured at 10 points for each transect. Six ant species were considered dominant, and had negative correlations with the number of species at baits (r² = 0.186; F_1,28 = 6.419; P = 0.017). However, almost half of the transects showed low abundance of dominant species (<30%), and relatively high number of species (mean of 20.1 ± 8.75). Resource availability and soil moisture had negative and positive correlations, with number of controlled baits. These results suggest that, even though the dominance is relatively poorly developed on the floor of this tropical forest, both resource availability and soil moisture affect resource control, and thus, the number of species that use baits.     

Relationships between branch spacing, growth rate and light in tropical forest saplings

1. The spacing of branches along central stems was related to growth rate and light level in forest saplings and trees in tropical moist forest on Barro Colorado Island (BCI), Panama. The study included 14 species with tiers of plagiotropic branches (having planar leaf arrangements) and four species with continuous distributions of plagiotropic branches.
2. All species showed increases in branch spacing with increasing light and growth rate of diameter, similar to the patterns in leaf spacing noted previously in species which initially bear large leaves on unbranched stems.
3. Non-tiered species had shorter internodes than tiered species but because the latter bear more branches per node, both groups had similar numbers of branches per unit stem length, when compared at similar growth rates.
4. Differences in the relationship between internode length and growth rate among tiered species were related to demographic characteristics, suggesting that tree architecture may influence forest composition.
5. The strong correlation observed between branch spacing and growth rate suggests that branch spacing may be used to estimate past growth histories of forest tree species with plagiotropic branches.     

The regional variation of aboveground live biomass in old-growth Amazonian forests

The biomass of tropical forests plays an important role in the global carbon cycle, both as a dynamic reservoir of carbon, and as a source of carbon dioxide to the atmosphere in areas undergoing deforestation. However, the absolute magnitude and environmental determinants of tropical forest biomass are still poorly understood. Here, we present a new synthesis and interpolation of the basal area and aboveground live biomass of old‐growth lowland tropical forests across South America, based on data from 227 forest plots, many previously unpublished. Forest biomass was analyzed in terms of two uncorrelated factors: basal area and mean wood density. Basal area is strongly affected by local landscape factors, but is relatively invariant at regional scale in moist tropical forests, and declines significantly at the dry periphery of the forest zone. Mean wood density is inversely correlated with forest dynamics, being lower in the dynamic forests of western Amazonia and high in the slow‐growing forests of eastern Amazonia. The combination of these two factors results in biomass being highest in the moderately seasonal, slow growing forests of central Amazonia and the Guyanas (up to 350 Mg dry weight ha^?1) and declining to 200–250 Mg dry weight ha^?1 at the western, southern and eastern margins. Overall, we estimate the total aboveground live biomass of intact Amazonian rainforests (area 5.76 × 10⁶ km² in 2000) to be 93±23 Pg C, taking into account lianas and small trees. Including dead biomass and belowground biomass would increase this value by approximately 10% and 21%, respectively, but the spatial variation of these additional terms still needs to be quantified.     

Carabidae diversity along an altitudinal gradient in a Peruvian cloud forest (Coleoptera)

Carabid beetles were sampled at five sites, ranging from 1500 m to 3400 m, along a 15 km transect in the cloud forest of Manu National Park, Perú. Seasonal collections during a one year period yielded 77 morphospecies, of which 60% are projected to be undescribed species. There was a significant negative correlation between species richness and altitude, with the number of carabid species declining at the rate of one species for each 100 m increase in altitude. The majority of species (70.1 %) were restricted to only one altitudinal site and no species was found at more than three of the five altitudinal sites. Only one genus, Pelmatellus (Tribe Harpalini), was found at all five sites. Active (hand) collections yielded approximately twice as many species per individuals collected than passive (pitfall trap) collections. This study is the first systematic sampling ofcarabid beetles of a high altitude gradient in the cloud forests of southeastern Perú and supports the need to conserve the zone of extremely high biodiversity present on the eastern slopes of the Peruvian Andes.