Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO2 enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon

Fine root chemistry and decomposition in model communities of north-temperate tree species show little response to elevated atmospheric CO 2 and varying soil resource availability

Rising atmospheric [CO 2 ] has the potential to alter soil carbon (C) cycling by increasing the content of recalcitrant constituents in plant litter, thereby decreasing rates of decomposition. Because fine root turnover constitutes a large fraction of annual NPP, changes in fine root decomposition are especially important. These responses will likely be affected by soil resource availability and the life history characteristics of the dominant tree species. We evaluated the effects of elevated atmospheric [CO 2 ] and soil resource availability on the production and chemistry, mycorrhizal colonization, and decomposition of fine roots in an early- and late-successional tree species that are economically and ecologically important in north temperate forests. Open-top chambers were used to expose young trembling aspen ( Populus tremuloides ) and sugar maple ( Acer saccharum ) trees to ambient (36 Pa) and elevated (56 Pa) atmospheric CO 2 . Soil resource availability was composed of two treatments that bracketed the range found in the Upper Lake States, USA. After 2.5 years of growth, sugar maple had greater fine root standing crop due to relatively greater allocation to fine roots (30% of total root biomass) relative to aspen (7% total root biomass). Relative to the low soil resources treatment, aspen fine root biomass increased 76% with increased soil resource availability, but only under elevated [CO 2 ]. Sugar maple fine root biomass increased 26% with increased soil resource availability (relative to the low soil resources treatment), and showed little response to elevated [CO 2 ]. Concentrations of N and soluble phenolics, and C/N ratio in roots were similar for the two species, but aspen had slightly higher lignin and lower condensed tannins contents compared to sugar maple. As predicted by source-sink models of carbon allocation, pooled constituents (C/N ratio, soluble phenolics) increased in response to increased relative carbon availability (elevated [CO 2 ]/low soil resource availability), however, biosynthetically distinct compounds (lignin, starch, condensed tannins) did not always respond as predicted. We found that mycorrhizal colonization of fine roots was not strongly affected by atmospheric [CO 2 ] or soil resource availability, as indicated by root ergosterol contents. Overall, absolute changes in root chemical composition in response to increases in C and soil resource availability were small and had no effect on soil fungal biomass or specific rates of fine root decomposition. We conclude that root contributions to soil carbon cycling will mainly be influenced by fine root production and turnover responses to rising atmospheric [CO 2 ], rather than changes in substrate chemistry.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/47710/1/442_2005_Article_191.pd

Relationships among nutrient enrichment, detritus quality and quantity, and large-bodied shredding insect community structure

This is a post-peer-review, pre-copyedit version of an article published in Hydrobiologia. The final authenticated version is available online at: https://doi.org/10.1007/s10750-015-2208-2Anthropogenic nutrient enrichment of forested headwater streams can enhance detrital quality, decrease standing stocks, and alter the community structure of detrivorous insects, reducing nutrient retention and decreasing ecosystem functioning. Our objective was to determine if stoichiometric principles could be used to predict genus-specific shifts in shredding insect abundance and biomass across a dissolved nutrient and detritus food quality/quantity gradient. Detritus, insect, and water samples were collected from 12 Ozark Highland headwater streams. Significant correlations were found between stream nutrients and detrital quality but not quantity. Abundance and biomass responses of four out of five tested genera were accurately predicted by consumerresource stoichiometric theory. Low carbon:phosphorus (C:P) shredders responded positively to increased total phosphorus and/or food quality, and high C:P shredders exhibited neutral or negative responses to these variables. Genus-specific declines were correlated with decreased overall biomass in shredder assemblages, potentially causing disruptions in nutrient flows to higher level consumers with nutrient enrichment. This work provides further evidence that elevated nutrients may negatively impact shredding insect communities by altering the stoichiometry of detritus–detritivore interactions. A better understanding of stoichiometric mechanisms altering macroinvertebrate populations is needed to help inform water quality criteria for the management of headwater streams

Liana seedling growth in response to fertilisation in a neotropical forest understorey

I tested the hypothesis that lianas grow taller and/or produce more biomass with higher soil nutrient availability despite severe light limitation in the forest understorey. Young seedlings of Callichlamys latifolia (Bignoniaceae), Doliocarpus major, and D. olivaceus (Dilleniaceae) were grown along a light gradient (0.8-2.2% of full sun) in the undisturbed soil of a lowland tropical rain forest on Barro Colorado Island (Panama) for 19 months. An equal number of seedlings was either supplied with NPK fertiliser equivalent to 220 kg N ha(-1) year(-1) or remained without fertilisation. Seedling survival was not significantly different among species and was not affected by either light availability or fertilisation. Biomass and height increased linearly with increasing light availability in seedlings of all species. Under fertilisation, biomass production was 50% higher in D. olivaceus irrespective of light availability, increased in C. latifolia only in relatively bright microsites, and was not affected in D. major across the entire light gradient. Seedling height was generally less affected by fertilisation with only C. latifolia seedlings growing taller in fertilised plots of a relatively high light availability. Biomass allocation changed little across the light gradient in any species. The strong biomass response to fertilisation in D. olivaceus was correlated with an increased biomass allocation to leaves at the expense of stems with only small changes in root mass fraction. The other two species showed no fertilisation effects on biomass allocation. I conclude that extreme light limitation does not preclude growth responses to nutrient addition in tropical lianas, but species differ in this respect

Soil animals alter plant litter diversity effects on decomposition

Most of the terrestrial net primary production enters the decomposer system as dead organic matter, and the subsequent recycling of C and nutrients are key processes for the functioning of ecosystems and the delivery of ecosystem goods and services. Although climatic and substrate quality controls are reasonably well understood, the functional role of biodiversity for biogeochemical cycles remains elusive. Here we ask how altering litter species diversity affects species-specific decomposition rates and whether large litter-feeding soil animals control the litter diversity–function relationship in a temperate forest ecosystem. We found that decomposition of a given litter species changed greatly in the presence of litters from other cooccurring species despite unaltered climatic conditions and litter chemistry. Most importantly, soil fauna determined the magnitude and direction of litter diversity effects. Our data show that litter species richness and soil fauna interactively determine rates of decomposition in a temperate forest, suggesting a combination of bottom-up and top-down controls of litter diversity effects on ecosystem C and nutrient cycling. These results provide evidence that, in ecosystems supporting a well developed soil macrofauna community, animal activity plays a fundamental role for altered decomposition in response to changing litter diversity, which in turn has important implications for biogeochemical cycles and the long-term functioning of ecosystems with ongoing biodiversity loss

Forest ageing : an unexpected driver of beech leaf litter quality variability in European forests with strong consequences on soil processes

The role of forest age as a potential driver of intraspecific variation in leaf litter quality, that is a key plant trait determining ecosystem functioning, has largely been neglected. Using a set of fully replicated pure beech (Fagus sylvatica) forest stands differing in age (15, 65, 95 and 130 years), we quantified the forest stand age related variability of twelve leaf litter quality traits. Litter Mg, N and K showed significantly higher concentrations in litter from 15-yrs-old stands and decreased with increasing stand age. Mn was the only nutrient analyzed that was highest in the oldest stands. Hemicellulose and cellulose were lowest, and lignin and lignin/N ratio were highest in stands of intermediate age. The amount of N within the litter lignin fraction was highest in the 95-yrs-old stands (51% of total N) and lowest in the oldest stands (34% of total N). The amount of N associated within the hemicellulose fraction (<3%) showed the opposite pattern along the forest stand age gradient compared to lignin. Using Partial Least Squares regressions, we showed that litter N, C/N, lignin/N, K, Mn and Mg were the most important predictors of litter decomposition along the chronosequence. In contrast the proportions of C fractions and the amount of N within these C fractions were the most significant variables explaining the variation in final litter N content after one year of decomposition. N mineralization in ground litter was highly related to the proportion of total N within lignin and humus N mineralization was mostly explained by Mn and the lignin/N ratio. We showed that forest age is an important driver of litter quality variation and contributed considerably to the overall variation of F. sylvatica leaf litter quality traits observed from a reviewed data of published studies conducted at the continental scale. Furthermore, intraspecific litter quality variation greatly impacted belowground processes. Accounting for forest age related litter trait variation, and for the crucial role of the distribution of N within different litter C fractions, may improve the mechanistic understanding of ecosystem functioning

Plant herbivory responses through changes in leaf quality have no effect on subsequent leaf-litter decomposition in a neotropical rain forest tree community

It is commonly accepted that plant responses to foliar herbivory (e.g. plant defenses) can influence subsequent leaf-litter decomposability in soil. While several studies have assessed the herbivory-decomposability relationship among different plant species, experimental tests at the intra-specific level are rare, although critical for a mechanistic understanding of how herbivores affect decomposition and its consequences at the ecosystem scale. Using 17 tree species from the Yasuni National Park, Ecuadorian Amazonia, and applying three different herbivore damage treatments, we experimentally tested whether the plant intra-specific responses to herbivory, through changes in leaf quality, affect subsequent leaf-litter decomposition in soil. We found no effects of herbivore damage on the subsequent decomposition of leaf litter within any of the species tested. Our results suggest that leaf traits affecting herbivory are different from those influencing decomposition. Herbivore damage showed much higher intra-specific than inter-specific variability, while we observed the opposite for decomposition. Our findings support the idea that interactions between consumers and their resources are controlled by different factors for the green and the brown food-webs in tropical forests, where herbivory may not necessarily generate any direct positive or negative feedbacks for nutrient cycling

Consequences of biodiversity loss for litter decomposition across biomes

The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere1, 2, 3. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs2, 4. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical4, 5, 6 given the rapid loss of species worldwide and the effects of this loss on human well-being7, 8, 9. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition4, 5, 6, 10, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism4, 9, 10, 11, 12. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales