Jon Lloyd

Centre for Ecology & Hydrology, Wallingford, ENG, United Kingdom

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Publications (38)217.92 Total impact

  • Article: Photosynthetically relevant foliar traits correlating better on a mass vs an area basis: of ecophysiological relevance or just a case of mathematical imperatives and statistical quicksand?
    New Phytologist 04/2013; · 6.64 Impact Factor
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    Article: On the delineation of tropical vegetation types with an emphasis on forest/savanna transitions
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    ABSTRACT: Background: There is no generally agreed classification scheme for the many different vegetation formation types occurring in the tropics. This hinders cross-continental comparisons and causes confusion as words such as ‘forest’ and ‘savanna’ have different meanings to different people. Tropical vegetation formations are therefore usually imprecisely and/or ambiguously defined in modelling, remote sensing and ecological studies. Aims: To integrate observed variations in tropical vegetation structure and floristic composition into a single classification scheme. Methods: Using structural and floristic measurements made on three continents, discrete tropical vegetation groupings were defined on the basis of overstorey and understorey structure and species compositions by using clustering techniques. Results: Twelve structural groupings were identified based on height and canopy cover of the dominant upper stratum and the extent of lower-strata woody shrub cover and grass cover. Structural classifications did not, however, always agree with those based on floristic composition, especially for plots located in the forest–savanna transition zone. This duality is incorporated into a new tropical vegetation classification scheme. Conclusions: Both floristics and stand structure are important criteria for the meaningful delineation of tropical vegetation formations, especially in the forest/savanna transition zone. A new tropical vegetation classification scheme incorporating this information has been developed. Keywords: canopy cover, cluster analysis forest, savanna, tropics, vegetation categorisation
    Plant Ecology & Diversity 01/2013; 6:101-137. · 1.04 Impact Factor
  • Article: Quantifying the abundance and stable isotope composition of pyrogenic carbon using hydrogen pyrolysis.
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    ABSTRACT: Pyrogenic carbon (C(P) ) is an important component of the global carbon budget. Accurate determination of the abundance and stable isotope composition of C(P) in soils and sediments is crucial for understanding the dynamics of the C(P) cycle and interpreting records of biomass burning, climate and vegetation change in the past. Here we test hydrogen pyrolysis (hypy) as a new technique potentially capable of eliminating labile organic carbon (C(L) ) from total organic carbon (C(T) ) in a range of matrices in order to enable reliable quantification of both the C(P) component of C(T) and the stable carbon isotope composition of C(P) (δ(13) C(P) ). We mixed C(P) at a range of concentrations with common C(P) -free matrices (C(L)  = cellulose, chitin, keratin, decomposed wood, leaf litter, grass and algae) and determined the amount of residual carbon not removed by hydrogen pyrolysis (C(R) ) as a ratio of C(T) (C(R) /C(T) ). Mixing C(P) with a unique δ(13) C value provided a natural abundance isotope label from which to precisely determine the ratio of C(P) to residual C(L) remaining after hypy. All C(P) -free matrices contained trace carbon after hypy, indicating that hypy does not remove all the C(L) . However, there was a strong correlation between C(R) /C(T) and C(P) /C(T) , viz. C(R) /C(T)  = 1.02(C(P) /C(T) ) + 4.0 × 10(-3) , r(2)  = 0.99, p <0.001, suggesting that only a small and reasonably constant fraction of C(L) remains after hypy. Uncertainties associated with the correction for contamination of C(R) by residual C(L) are minimal allowing for reliable determinations of both C(P) and δ(13) C(P) in many cases. Hydrogen pyrolysis appears to be a robust technique for estimating C(P) abundance and δ(13) C(P) across a range of materials. Nevertheless, caution is required in interpreting δ(13) C(P) values when C(P) /C(T) is low, with C(P) /C(T) >4% being required for the determination of the δ(13) C(P) values within an interpretable error under our experimental conditions. Copyright © 2012 John Wiley & Sons, Ltd.
    Rapid Communications in Mass Spectrometry 12/2012; 26(23):2690-6. · 2.79 Impact Factor
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    Article: Variation in soil carbon stocks and their determinants across a precipitation gradient in West Africa
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    ABSTRACT: We examine the influence of climate, soil properties and vegetation characteristics on soil organic carbon (SOC) along a transect of West African ecosystems sampled across a precipitation gradient on contrasting soil types stretching from Ghana (15°N) to Mali (7°N). Our findings derive from a total of 1108 soil cores sampled over 14 permanent plots. The observed pattern in SOC stocks reflects the very different climatic conditions and contrasting soil properties existing along the latitudinal transect. The combined effects of these factors strongly influence vegetation structure. SOC stocks in the first 2 m of soil ranged from 20 Mg C ha−1 for a Sahelian savanna in Mali to over 120 Mg C ha−1 for a transitional forest in Ghana. The degree of interdependence between soil bulk density (SBD) and soil properties is highlighted by the strong negative relationships observed between SBD and SOC (r2 > 0.84). A simple predictive function capable of encompassing the effect of climate, soil properties and vegetation type on SOC stocks showed that available water and sand content taken together could explain 0.84 and 0.86 of the total variability in SOC stocks observed to 0.3 and 1.0 m depth respectively. Used in combination with a suitable climatic parameter, sand content is a good predictor of SOC stored in highly weathered dry tropical ecosystems with arguably less confounding effects than provided by clay content. There was an increased contribution of resistant SOC to the total SOC pool for lower rainfall soils, this likely being the result of more frequent fire events in the grassier savannas of the more arid regions. This work provides new insights into the mechanisms determining the distribution of carbon storage in tropical soils and should contribute significantly to the development of robust predictive models of biogeochemical cycling and vegetation dynamics in tropical regions.
    Global Change Biology 01/2012; · 6.86 Impact Factor
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    Article: Variations in Amazon forest productivity correlated with foliar nutrients and modelled rates of photosynthetic carbon supply.
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    ABSTRACT: The rate of above-ground woody biomass production, W(P), in some western Amazon forests exceeds those in the east by a factor of 2 or more. Underlying causes may include climate, soil nutrient limitations and species composition. In this modelling paper, we explore the implications of allowing key nutrients such as N and P to constrain the photosynthesis of Amazon forests, and also we examine the relationship between modelled rates of photosynthesis and the observed gradients in W(P). We use a model with current understanding of the underpinning biochemical processes as affected by nutrient availability to assess: (i) the degree to which observed spatial variations in foliar [N] and [P] across Amazonia affect stand-level photosynthesis; and (ii) how these variations in forest photosynthetic carbon acquisition relate to the observed geographical patterns of stem growth across the Amazon Basin. We find nutrient availability to exert a strong effect on photosynthetic carbon gain across the Basin and to be a likely important contributor to the observed gradient in W(P). Phosphorus emerges as more important than nitrogen in accounting for the observed variations in productivity. Implications of these findings are discussed in the context of future tropical forests under a changing climate.
    Philosophical Transactions of The Royal Society B Biological Sciences 11/2011; 366(1582):3316-29. · 6.40 Impact Factor
  • Article: Investigating diversity dependence of tropical forest litter decomposition: experiments and observations from Central Africa
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    ABSTRACT: QuestionsMixed litter may decompose at different rates to single-species litter, leading to differences in ecosystem functioning and decomposition. Studies of the effects of different litter species and combinations are rare in tropical forests and absent from African forests. Therefore we investigated: (1) Are there differences in litter decomposition in two forest types differing in tree diversity; and (2) is litter decomposition diversity-dependent?LocationOld-growth moist evergreen tropical forest Dja Faunal Reserve, southeast Cameroon.Methods We calculate decomposition rates (leaf litter fall/leaf litter standing crop) along a tree diversity gradient in two forest types (naturally occurring low-diversity monodominant and adjacent higher-diversity mixed forest). Both forests experience the same climate on the same soil type; the former is dominated by a single species, Gilbertiodendron dewevrei (De Wild.) J. Léonard, probably due to lack of a long-term disturbance and has similar edaphic factors. Decomposition experiments were conducted in both forest types using single and mixed species litter bags of standard high-quality (bay leaves; Laurus nobilis L.) and low-quality (G. dewevrei) litter over 9 months.ResultsThe estimated decomposition rate in mixed forest was four times faster than in monodominant forest, and not significantly correlated with local quadrat-scale tree species diversity. The litter bag experiment showed that decomposition of high-quality leaves was faster than low-quality leaves (k values: 2.0 yr−1 vs 0.6 yr−1). Decay rates for each single species litter type were not significantly different in both forest types. However, G. dewevrei litter in mixed bags decomposed faster than in single-species bags in mixed forest, suggesting an impact of litter mixing on decomposition. In addition, bay litter in mixed bags decomposed faster in mixed than in monodominant forests across the three study sites.Conclusion The observed difference in litter decomposition rate between low-diversity monodominant and adjacent high-diversity forest is more likely due to dominance of low-quality G. dewevrei litter, rather than low-diversity of the litter itself.
    Journal of Vegetation Science 10/2011; 23(2):223 - 235. · 2.77 Impact Factor
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    Article: Mechanisms of monodominance in diverse tropical tree‐dominated systems
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    ABSTRACT: Summary1. The existence of many types of monodominant forests is readily explainable by ecological theory (e.g. early successional forests). Nevertheless, monodominant stands sometimes occur in areas where a much higher diversity typically occurs. Such ‘classical monodominance’ is not currently readily explained by ecological theory.2. We briefly review the published mechanisms suggested to cause classical monodominance and then combine them into a new probabilistic conceptual framework to better understand why these systems occur. We build on two theories proposed to explain monodominance: a lack of exogenous disturbance over long periods and species-specific life-history traits. We suggest that certain traits under certain conditions may generate positive feedbacks leading to a greater probability of monodominance being achieved. Such positive feedbacks have the potential to drive a typically diverse system towards a monodominant one.3. Synthesis. Classical monodominance in tropical forests is hypothesized to be attained when a group of traits occur together under low exogenous disturbance conditions, this giving rise to a series of positive feedbacks. The presented framework links the differing mechanisms proposed in the literature to explain classical monodominance and shows there are potentially alternative routes to monodominance, thus reconciling apparently contradictory observational and experimental results.
    Journal of Ecology 03/2011; 99(4):891 - 898. · 4.69 Impact Factor
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    Article: Soil does not explain monodominance in a Central African tropical forest.
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    ABSTRACT: Soil characteristics have been hypothesised as one of the possible mechanisms leading to monodominance of Gilbertiodendron dewerei in some areas of Central Africa where higher-diversity forest would be expected. However, the differences in soil characteristics between the G. dewevrei-dominated forest and its adjacent mixed forest are still poorly understood. Here we present the soil characteristics of the G. dewevrei forest and quantify whether soil physical and chemical properties in this monodominant forest are significantly different from the adjacent mixed forest. We sampled top soil (0-5, 5-10, 10-20, 20-30 cm) and subsoil (150-200 cm) using an augur in 6 × 1 ha areas of intact central Africa forest in SE Cameroon, three independent patches of G. dewevrei-dominated forest and three adjacent areas (450-800 m apart), all chosen to be topographically homogeneous. Analysis--subjected to Bonferroni correction procedure--revealed no significant differences between the monodominant and mixed forests in terms of soil texture, median particle size, bulk density, pH, carbon (C) content, nitrogen (N) content, C:N ratio, C:total NaOH-extractable P ratio and concentrations of labile phosphorous (P), inorganic NaOH-extractable P, total NaOH-extractable P, aluminium, barium, calcium, copper, iron, magnesium, manganese, molybdenum, nickel, potassium, selenium, silicon, sodium and zinc. Prior to Bonferroni correction procedure, there was a significant lower level of silicon concentration found in the monodominant than mixed forest deep soil; and a significant lower level of nickel concentration in the monodominant than mixed forest top soil. Nevertheless, these were likely to be the results of multiple tests of significance. Our results do not provide clear evidence of soil mediation for the location of monodominant forests in relation to adjacent mixed forests. It is also likely that G. dewevrei does not influence soil chemistry in the monodominant forests.
    PLoS ONE 01/2011; 6(2):e16996. · 4.09 Impact Factor
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    Article: Drought-mortality relationships for tropical forests.
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    ABSTRACT: *The rich ecology of tropical forests is intimately tied to their moisture status. Multi-site syntheses can provide a macro-scale view of these linkages and their susceptibility to changing climates. Here, we report pan-tropical and regional-scale analyses of tree vulnerability to drought. *We assembled available data on tropical forest tree stem mortality before, during, and after recent drought events, from 119 monitoring plots in 10 countries concentrated in Amazonia and Borneo. *In most sites, larger trees are disproportionately at risk. At least within Amazonia, low wood density trees are also at greater risk of drought-associated mortality, independent of size. For comparable drought intensities, trees in Borneo are more vulnerable than trees in the Amazon. There is some evidence for lagged impacts of drought, with mortality rates remaining elevated 2 yr after the meteorological event is over. *These findings indicate that repeated droughts would shift the functional composition of tropical forests toward smaller, denser-wooded trees. At very high drought intensities, the linear relationship between tree mortality and moisture stress apparently breaks down, suggesting the existence of moisture stress thresholds beyond which some tropical forests would suffer catastrophic tree mortality.
    New Phytologist 08/2010; 187(3):631-46. · 6.64 Impact Factor
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    Article: Co-limitation of photosynthetic capacity by nitrogen and phosphorus in West Africa woodlands.
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    ABSTRACT: Photosynthetic leaf traits were determined for savanna and forest ecosystems in West Africa, spanning a large range in precipitation. Standardized major axis fits revealed important differences between our data and reported global relationships. Especially for sites in the drier areas, plants showed higher photosynthetic rates for a given N or P when compared with relationships from the global data set. The best multiple regression for the pooled data set estimated V(cmax) and J(max) from N(DW) and S. However, the best regression for different vegetation types varied, suggesting that the scaling of photosynthesis with leaf traits changed with vegetation types. A new model is presented representing independent constraints by N and P on photosynthesis, which can be evaluated with or without interactions with S. It assumes that limitation of photosynthesis will result from the least abundant nutrient, thereby being less sensitive to the allocation of the non-limiting nutrient to non-photosynthetic pools. The model predicts an optimum proportionality for N and P, which is distinct for V(cmax) and J(max) and inversely proportional to S. Initial tests showed the model to predict V(cmax) and J(max) successfully for other tropical forests characterized by a range of different foliar N and P concentrations.
    Plant Cell and Environment 06/2010; 33(6):959-80. · 5.22 Impact Factor
  • Article: Modelling basin-wide variations in Amazon forest photosynthesis
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    ABSTRACT: Given the importance of Amazon rainforest in the global carbon and hydrological cycles, there is a need to use parameterized and validated ecosystem gas exchange and vegetation models for this region in order to adequately simulate present and future carbon and water balances. Recent research has found major differences in above-ground net primary productivity (ANPP), above ground biomass and tree dynamics across Amazonia. West Amazonia is more dynamic, with younger trees, higher stem growth rates and lower biomass than central and eastern Amazon (Baker et al. 2004; Malhi et al. 2004; Phillips et al. 2004). A factor of three variation in above-ground net primary productivity has been estimated across Amazonia by Malhi et al. (2004). Different hypotheses have been proposed to explain the observed spatial variability in ANPP (Malhi et al. 2004). First, due to the proximity to the Andes, sites from western Amazonia tend to have richer soils than central and eastern Amazon and therefore soil fertility could possibly be highly related to the high wood productivity found in western sites. Second, if GPP does not vary across the Amazon basin then different patterns of carbon allocation to respiration could also explain the observed ANPP gradient. However since plant growth depends on the interaction between photosynthesis, transport of assimilates, plant respiration, water relations and mineral nutrition, variations in plant gross photosynthesis (GPP) could also explain the observed variations in ANPP. In this study we investigate whether Amazon GPP can explain variations of observed ANPP. We use a sun and shade canopy gas exchange model that has been calibrated and evaluated at five rainforest sites (Mercado et al. 2009) to simulate gross primary productivity of 50 sites across the Amazon basin during the period 1980-2001. Such simulation differs from the ones performed with global vegetation models (Cox et al. 1998; Sitch et al. 2003) where i) single plant functional type parameter values are assigned and assumed invariant with environmental condition but also ii) these models use leaf N as a factor that limit photosynthesis. Instead, since leaf P may also limit photosynthesis of the tropical forest (Reich et al. 2009), we use a more specific description of photosynthetic capacity across the basin based on the model evaluation done in Mercado et al. (2009) in which canopy photosynthetic capacity is related to foliar P but also using the relationships derived between canopy photosynthesis and leaf nutrients (N and P) from measurements in tropical trees (Domingues et al.In review). A study of this kind can inform the global vegetation/climate community as to the need for variability in key model parameters in order to accurately simulate carbon fluxes across the Amazon basin. Baker, T. R., et al. 2004. Increasing biomass in Amazonian forest plots. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences 359 (1443):353-365. Phillips, O. L. et al. 2004. Pattern and process in Amazon tree turnover, 1976-2001. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences 359 (1443):381-407. Malhi, Y. et al. 2004. The above-ground coarse wood productivity of 104 Neotropical forest plots. Global Change Biology 10 (5):563-591. Mercado, L.M. et al. 2009. Impact of changes in diffuse radiation on the global land carbon sink. Nature 458 (7241), 1014. Cox, P. M. et al. 1998. A canopy conductance and photosynthesis model for use in a GCM land surface scheme. Journal of Hydrology 213 (1-4):79-9 Sitch, S. et al. 2003. Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model. Global Change Biology 9 (2):161-185. Reich B. R. et al. 2009. Leaf phosphorus influences the photosynhtesis-nitrogen relation: a cross-biome analysis of 314 species. Oecologia, doi 10.1007/s00442-009-1291-3. Domingues, T. et al. In review. Co-limitation of photosynthetic capacity by nitrogen and phosphorus along a precipitation gradient in West Africa. Plant Cell and Environment.
    04/2010; 12:13607.
  • Article: Are the dynamics of tropical forests dominated by large and rare disturbance events?
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    ABSTRACT: A recent Ecology Letters paper of Fisher et al. (2008) utilized a modelling framework to investigate disturbance effects on forest biomass dynamics. But it contains serious methodological and conceptual errors. Associated conclusions are unlikely to be correct.
    Ecology Letters 12/2009; 12(12):E19-21; discussion E22-5. · 17.56 Impact Factor
  • Article: Drought sensitivity of the Amazon rainforest.
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    ABSTRACT: Amazon forests are a key but poorly understood component of the global carbon cycle. If, as anticipated, they dry this century, they might accelerate climate change through carbon losses and changed surface energy balances. We used records from multiple long-term monitoring plots across Amazonia to assess forest responses to the intense 2005 drought, a possible analog of future events. Affected forest lost biomass, reversing a large long-term carbon sink, with the greatest impacts observed where the dry season was unusually intense. Relative to pre-2005 conditions, forest subjected to a 100-millimeter increase in water deficit lost 5.3 megagrams of aboveground biomass of carbon per hectare. The drought had a total biomass carbon impact of 1.2 to 1.6 petagrams (1.2 x 10(15) to 1.6 x 10(15) grams). Amazon forests therefore appear vulnerable to increasing moisture stress, with the potential for large carbon losses to exert feedback on climate change.
    Science 04/2009; 323(5919):1344-7. · 31.20 Impact Factor
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    Article: Increasing carbon storage in intact African tropical forests.
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    ABSTRACT: The response of terrestrial vegetation to a globally changing environment is central to predictions of future levels of atmospheric carbon dioxide. The role of tropical forests is critical because they are carbon-dense and highly productive. Inventory plots across Amazonia show that old-growth forests have increased in carbon storage over recent decades, but the response of one-third of the world's tropical forests in Africa is largely unknown owing to an absence of spatially extensive observation networks. Here we report data from a ten-country network of long-term monitoring plots in African tropical forests. We find that across 79 plots (163 ha) above-ground carbon storage in live trees increased by 0.63 Mg C ha(-1) yr(-1) between 1968 and 2007 (95% confidence interval (CI), 0.22-0.94; mean interval, 1987-96). Extrapolation to unmeasured forest components (live roots, small trees, necromass) and scaling to the continent implies a total increase in carbon storage in African tropical forest trees of 0.34 Pg C yr(-1) (CI, 0.15-0.43). These reported changes in carbon storage are similar to those reported for Amazonian forests per unit area, providing evidence that increasing carbon storage in old-growth forests is a pan-tropical phenomenon. Indeed, combining all standardized inventory data from this study and from tropical America and Asia together yields a comparable figure of 0.49 Mg C ha(-1) yr(-1) (n = 156; 562 ha; CI, 0.29-0.66; mean interval, 1987-97). This indicates a carbon sink of 1.3 Pg C yr(-1) (CI, 0.8-1.6) across all tropical forests during recent decades. Taxon-specific analyses of African inventory and other data suggest that widespread changes in resource availability, such as increasing atmospheric carbon dioxide concentrations, may be the cause of the increase in carbon stocks, as some theory and models predict.
    Nature 03/2009; 457(7232):1003-6. · 36.28 Impact Factor
  • Article: Changing Ecology of Tropical Forests: Evidence and Drivers
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    ABSTRACT: Global environmental changes may be altering the ecology of tropical forests. Long-term monitoring plots have provided much of the evidence for large-scale, directional changes in tropical forests, but the results have been controversial. Here we review evidence from six complementary approaches to understanding possible changes: plant physiology experiments, long-term monitoring plots, ecosystem flux techniques, atmospheric measurements, Earth observations, and global-scale vegetation models. Evidence from four of these approaches suggests that large-scale, directional changes are occurring in the ecology of tropical forests, with the other two approaches providing inconclusive results. Collectively, the evidence indicates that both gross and net primary productivity has likely increased over recent decades, as have tree growth, recruitment, and mortality rates, and forest biomass. These results suggest a profound reorganization of tropical forest ecosystems. We evaluate the most likely drivers of the su...
    02/2009; 40:529-549.
  • Article: Do species traits determine patterns of wood production in Amazonian forests?
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    ABSTRACT: Understanding the relationships between plant traits and ecosystem properties at large spatial scales is important for predicting how compositional change will affect carbon cycling in tropical forests. In this study, we examine the relationships between species wood density, maximum height and above-ground, coarse wood production of trees ≥10 cm diameter (CWP) for 60 Amazonian forest plots. Average species maximum height and wood density are lower in Western than Eastern Amazonia and are negatively correlated with CWP. To test the hypothesis that variation in these traits causes the variation in CWP, we generate plot-level estimates of CWP by resampling the full distribution of tree biomass growth rates whilst maintaining the appropriate tree-diameter and functional-trait distributions for each plot. These estimates are then compared with the observed values. Overall, the estimates do not predict the observed, regional-scale pattern of CWP, suggesting that the variation in community-level trait values does not determine variation in coarse wood productivity in Amazonian forests. Instead, the regional gradient in CWP is caused by higher biomass growth rates across all tree types in Western Amazonia. Therefore, the regional gradient in CWP is driven primarily by environmental factors, rather than the particular functional composition of each stand. These results contrast with previous findings for forest biomass, where variation in wood density, associated with variation in species composition, is an important driver of regional-scale patterns in above-ground biomass. Therefore, in tropical forests, above-ground wood productivity may be less sensitive than biomass to compositional change that alters community-level averages of these plant traits.
    01/2009;
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    Article: Effects of rising temperatures and [CO2] on the physiology of tropical forest trees.
    Jon Lloyd, Graham D Farquhar
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    ABSTRACT: Using a mixture of observations and climate model outputs and a simple parametrization of leaf-level photosynthesis incorporating known temperature sensitivities, we find no evidence for tropical forests currently existing "dangerously close" to their optimum temperature range. Our model suggests that although reductions in photosynthetic rate at leaf temperatures (TL) above 30 degrees C may occur, these are almost entirely accountable for in terms of reductions in stomatal conductance in response to higher leaf-to-air vapour pressure deficits D. This is as opposed to direct effects of TL on photosynthetic metabolism. We also find that increases in photosynthetic rates associated with increases in ambient [CO2] over forthcoming decades should more than offset any decline in photosynthetic productivity due to higher D or TL or increased autotrophic respiration rates as a consequence of higher tissue temperatures. We also find little direct evidence that tropical forests should not be able to respond to increases in [CO2] and argue that the magnitude and pattern of increases in forest dynamics across Amazonia observed over the last few decades are consistent with a [CO2]-induced stimulation of tree growth.
    Philosophical Transactions of The Royal Society B Biological Sciences 06/2008; 363(1498):1811-7. · 6.40 Impact Factor
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    Article: Contributions of woody and herbaceous vegetation to tropical savanna ecosystem productivity: a quasi-global estimate.
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    ABSTRACT: To estimate the relative contributions of woody and herbaceous vegetation to savanna productivity, we measured the 13C/12C isotopic ratios of leaves from trees, shrubs, grasses and the surface soil carbon pool for 22 savannas in Australia, Brazil and Ghana covering the full savanna spectrum ranging from almost pure grassland to closed woodlands on all three continents. All trees and shrubs sampled were of the C3 pathway and all grasses of the C4 pathway with the exception of Echinolaena inflexa (Poir.) Chase, a common C3 grass of the Brazilian cerrado. By comparing the carbon isotopic compositions of the plant and carbon pools, a simple model relating soil delta 13C to the relative abundances of trees + shrubs (woody plants) and grasses was developed. The model suggests that the relative proportions of a savanna ecosystem's total foliar projected cover attributable to grasses versus woody plants is a simple and reliable index of the relative contributions of grasses and woody plants to savanna net productivity. Model calibrations against woody tree canopy cover made it possible to estimate the proportion of savanna productivity in the major regions of the world attributable to trees + shrubs and grasses from ground-based observational maps of savanna woodiness. Overall, it was estimated that 59% of the net primary productivity (Np) of tropical savannas is attributable to C4 grasses, but that this proportion varies significantly within and between regions. The C4 grasses make their greatest relative contribution to savanna Np in the Neotropics, whereas in African regions, a greater proportion of savanna Np is attributable to woody plants. The relative contribution of C4 grasses in Australian savannas is intermediate between those in the Neotropics and Africa. These differences can be broadly ascribed to large scale differences in soil fertility and rainfall.
    Tree Physiology 04/2008; 28(3):451-68. · 2.88 Impact Factor
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    Article: Growth form and seasonal variation in leaf gas exchange of Colophospermum mopane savanna trees in northwest Botswana.
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    ABSTRACT: We investigated differences in physiological and morphological traits between the tall and short forms of mopane (Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard) trees growing near Maun, Botswana on a Kalahari sandveld overlying an impermeable calcrete duricrust. We sought to determine if differences between the two physiognomic types are attributable to the way they exploit available soil water. The tall form, which was located on deeper soil than the short form (5.5 versus 1.6 m), had a lower leaf:fine root biomass ratio (1:20 versus 1:6), but a similar leaf area index (0.9-1.0). Leaf nitrogen concentrations varied between 18 and 27 mg g(-1) and were about 20% higher in the tall form than in the short form. Maximum net assimilation rates (A sat) occurred during the rainy seasons (March-April 2000 and January-February 2001) and were similar in the tall and short forms (15-22 micromol m(-2) s(-1)) before declining to less than 10 micromol m(-2) s(-1) at the end of the rainy season in late April. As the dry season progressed, A sat, soil water content, predawn leaf water potential (Psi pd) and leaf nitrogen concentration declined rapidly. Before leaf abscission, Psi pd was more negative in the short form (-3.4 MPa) than in the tall form (-2.7 MPa) despite the greater availability of soil water beneath the short form trees. This difference appeared attributable to differences in root depth and density between the physiognomic types. Stomatal regulation of water use and carbon assimilation differed between years, with the tall form having a consistently more conservative water-use strategy as the dry season progressed than the short form.
    Tree Physiology 04/2008; 28(3):417-24. · 2.88 Impact Factor
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    Article: Seasonal variations in soil water in two woodland savannas of central Brazil with different fire history.
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    ABSTRACT: Changes in soil water content were determined in two cerrado (sensu stricto) areas with contrasting fire history and woody vegetation density. The study was undertaken near Brasília, Brazil, from 1999 to 2001. Soil water content was measured with a neutron probe in three access tubes per site to a depth of 4.7 m. One site has been protected from fire for more than 30 years and, as a consequence, has a high density of woody plants. The other site had been frequently burned, and has a high herbaceous vegetation density and less woody vegetation. Soil water uptake patterns were strongly seasonal, and despite similarities in hydrological processes, the protected area systematically used more water than the burned area. Three temporarily contiguous patterns of water absorption were differentiated, characterized by variation in the soil depth from which water was extracted. In the early dry season, vegetation used water from throughout the soil profile but with a slight preference for water in the upper soil layers. Toward the peak of the dry season, vegetation had used most or all available water from the surface to a depth of 1.7 m, but continued to extract water from greater depths. Following the first rains, all water used was from the recently wetted upper soil layers only. Evaporation rates were a linear function of soil water availability, indicating a strong coupling of atmospheric water demand and the physiological response of the vegetation.
    Tree Physiology 04/2008; 28(3):405-15. · 2.88 Impact Factor

Institutions

  • 2011
    • Centre for Ecology & Hydrology
      Wallingford, ENG, United Kingdom
    • University of Cambridge
      Cambridge, ENG, United Kingdom
  • 2008–2011
    • University of Leeds
      • School of Geography
      Leeds, ENG, United Kingdom
    • University of Botswana
      Gaborone, South East District, Botswana
  • 2010
    • The University of Edinburgh
      • School of GeoSciences
      Edinburgh, SCT, United Kingdom
  • 2001–2008
    • Max-Planck-Institut für Biogeochemie Jena
      Jena, Thuringia, Germany
  • 2006
    • Australian National University
      Canberra, Australian Capital Territory, Australia