Soil organic matter in soil physical fractions in adjacent semi-natural and cultivated stands in temperate Atlantic forests

Changes from natural tree species to rapidly growing exotic species as well as intensification of forestry operations with heavy machinery can lead to changes in the quantity and quality of organic matter inputs to soil and to disruption of soil physical structure. These two ecosystem properties are tightly linked to organic matter dynamics. Five adjacent forest stands were selected to study soil organic matter dynamics in soil physical fractions. On one hand, two semi-natural broadleaved forests (Quercus robur, Fagus sylvatica) and an adult radiata pine plantation (40-year-old,) in order to study the effect of species change on these parameters, and on the other, a chronosequence of Pinus radiata plantations (40-year-old; 3-year-old; 16-year-old), to study the effect of mechanization during harvesting and intense site preparation. Samples of intact topsoil (0-5 cm) were collected and aggregate-size distribution, mean weight diameter (MWD), total C and N, particulate organic matter (POM)-C, POM-N and microbial biomass-C were determined in each aggregate size fraction. Microbial respiration and nitrogen mineralization were also assessed in each aggregate size fraction, during a 28 day incubation period.Losses of POM-C and POM-N in the bulk soil due to mechanical site preparation were high relative to total soil C and N, which suggests that POM is a sensitive parameter to the effect of mechanization. The ratio C-POM:SOM was significantly related to MWD (R² = 0.75, P < 0.001) reflecting that POM may play a key role in the topsoil aggregate formation in these stands. Semi-natural stands had a higher proportion of macroaggregates (0.25-2 mm) than the cultivated adult one. Megaaggregates (>2 mm) were the most abundant class in mature stands (82-92%), whereas macro- and microaggregates (<2 mm) were the most abundant ones in the intensely soil prepared P. radiata plantation (49%).Indicators for sustainable forest management related to soil organic matter should not only be assessed in terms of total C stocks but also with respect to sensitive organic matter and its degradability in different size classes.     

Restoring forest herb communities through landscape-level deer herd reductions: Is recovery limited by legacy effects?

White-tailed deer (Odocoileus virginianus) overbrowsing has altered plant species diversity throughout deciduous forest understories in eastern North America. Here we report on a landscape-level (306 km²) project in Pennsylvania, USA that tracked the herbaceous community response to deer herd reductions. From 2001 to 2007, we estimated deer densities, browse impact on woody seedlings, and censused the herbaceous flora in permanent plots throughout the area. We assessed herb layer species richness, abundance, and dominance and measured three known phytoindicators of deer impact: Trillium spp., Maianthemum canadense, and Medeola virginiana. We predicted that browse-sensitive taxa would increase in abundance, size, and flowering as would overall species diversity following deer culls and browse impact that declined by an order of magnitude by 2007. Following intensified deer harvests, we observed a limited recovery of the herbaceous community. Trillium spp. abundance, height, and flowering; M. canadense cover; and M. virginiana height all increased following herd reductions. Similarly, forb and shrub cover increased by 130% and 300%, respectively. Nevertheless, species diversity (i.e., richness and dominance) did not vary. Our work demonstrates that reducing deer densities can provide rapid morphological and population-level benefits to palatable species without a concomitant recovery in diversity. We suggest that decreasing deer populations alone may not promote plant diversity in overbrowsed, depauperate forests without additional restoration strategies to mitigate a browse-legacy layer dominated by browse-resistant species.     

Seeing the forest for the deer: Do reductions in deer-disturbance lead to forest recovery?

High levels of deer browsing can prevent canopy tree regeneration, but little is known about changes to forest size-structure following long-term deer herd reductions. We monitored changes in forest stand structure and composition in southwestern Ontario, Canada, over 28-years using permanent plots. Our study site was the largest remaining tract of Carolinian (deciduous) forest in Canada (11 km²), a habitat type that contains up to a fifth of Canada’s species at risk and is under intense anthropogenic pressures. We recorded declines in all tree size classes between 1981 and 1996, during which densities of white-tailed deer (Odocoileus virginianus) reached a peak of 55 deer km⁻². Despite significant and sustained deer herd reductions between 1996 and 2009, which reduced deer densities to 7 deer km⁻², there was limited recruitment of small trees and declines in basal area of tree species that were sensitive to deer browsing. Our results suggest that recovery from herbivory is a protracted process during which canopy tree regeneration may continue to decline despite a reduction in browsing pressure due to deer culling. Large declines in canopy-tree densities in Carolinian forests may lead to forest size-structures and herbaceous plant communities that resemble rare oak savanna habitat, creating difficult decisions for conservation managers aiming to protect rare and endangered species within native ecosystems. We recommend that managers protect Carolinian forest stands and encourage canopy tree regeneration by increasing seed sources of native trees. While deer control is essential in reducing forest damage, our results highlight the need to explore other forms of active management to expedite otherwise slow increases in tree density.     

Functional diversity of microbial communities in the mixed boreal plain forest of central Canada

This study was designed to examine whether or not specific tree species (Picea glauca, Picea mariana, Pinus banksiana, Populus tremuloides), their post-fire stand age, or their position in a successional pathway had any significant effect on the functional diversity of associated soil microbial communities in a typical mixed boreal forest ecosystem (Duck Mountain Provincial Forest, Manitoba, Canada). Multivariate analyses designed to identify significant biotic and/or abiotic variables associated with patterns of organic substrate utilization (assessed using the BIOLOG™ System) revealed the overall similarity in substrate utilization by the soil microbial communities. The five clusters identified differed mainly by their substrate-utilization value rather than by specific substrate utilization. Variability in community functional diversity was not strongly associated to tree species or post-fire stand age; however, redundancy analysis indicated a stronger association between substrate utilization and successional pathway and soil pH. For example, microbial communities associated with the relatively high pH soils of the P. tremuloides-P. glauca successional pathway, exhibited a greater degree of substrate utilization than those associated with the P. banksiana-P. mariana successional pathway and more acidic soils. Differences in functional diversity specific to tree species were not observed and this may have reflected the mixed nature of the forest stands and of their heterogeneous forest floor. In a densely treed, mixed boreal forest ecosystem, great overlap in tree and understory species occur making it difficult to assign a definitive microbial community to any particular tree species. The presence of P. tremuloides in all stand types and post fire stand ages has probably contributed to the large amount of overlap in utilization profiles among soil samples.     

Predator-prey relationships and responses of ungulates and their predators to the establishment of protected areas: A case study of tigers, leopards and their prey in Bardia National Park, Nepal

Interactions among sympatric large predators and their prey and how they respond to conservation measures are poorly known. This study examines predictions concerning the effects of establishing a protected area in Nepal on tigers (Panthera tigris), leopards (Panthera pardus), and their ungulate prey. Within a part of the park, after 22 years the total density of wild ungulates had increased fourfold, to ca. 200 animals/km², almost exclusively due to a remarkable increase in chital deer (Axis axis). Tiger density also increased markedly to nearly 20 animals/100 km², whereas leopard density did not and was ca. 5 animals/100 km². The prediction that grazers should increase more than browsers was only partially supported. The prediction of positive density dependence in prey selection was not supported. Instead, the most abundant species (chital and hog deer, Axis porcinus) were killed less frequently than expected, whereas the lower-density wild boar (Sus scrofa) was preferred. Predictions that (i) initially rare species suffer highest predation was partially supported, that (ii) predation is highest among the most abundant prey was not supported, and that (iii) predation is highest among the most preferred prey independently of their densities was supported. Clearly, the conservation efforts adopted in Bardia were successful, as both tigers and their natural prey base increased. However, the positive numerical response of tigers limited and depressed the abundance of some prey species. Thus, conservation activities aimed at restoring large predators are likely to change in the composition of the overall mammal community, potentially eliminating rare but preferred prey species.     

Seasonal variation in trehalose contents of roots and nodules of leguminous trees in a tropical deciduous forest in Mexico

Seasonal variations of trehalose contents in roots and root-nodules of five legumes in a tropical deciduous forest in Jalisco, México, were determined. The tree species were: Lonchocarpus eriocarinalis and Erythrina lanata (sub-family Papilionoideae) and Piptadenia constricta, Albizia occidentalis and Lysiloma microphylum (sub-family Mimosoideae). Trehalose accumulation in nodules and roots varied seasonally and among species. For example, the Papilionoid-species retained nodules longer than those in the Mimosoideae (5 and 4 months, respectively), and accumulated the highest amounts of trehalose (average values of 178 vs. 2.88 mg g⁻¹ nodule (d. wt), respectively). Generally, maximum trehalose contents in nodules and roots were observed in November, at the beginning of the dry season.     

Bird species richness in a Bornean exotic tree plantation: A long-term perspective

To provide sustainable income from forestlands, large areas in the tropics are planted with “agricultural” trees, such as oil palm and rubber, and “industrial” trees, such as Acacia mangium and Gmelina arborea. To examine how native forest birds use such plantations, we surveyed in 2005 the avifauna at Sabah Softwoods, a plantation in southeastern Sabah, Malaysian Borneo. We focused on A. mangium, Albizia (Paraserianthes falcataria), oil palm (Elaeis guineensis), and logged native forest, and compared our results to those of a study conducted at the same plantation in 1982. The number of forest species in the industrial groves did not change dramatically between 1982 and 2005, even though the trees had been cropped several times and the plantation was, by 2005, completely surrounded by cleared land and far removed from primary forest. However, as is common in logged or isolated forests throughout the world, certain primary forest groups (e.g., muscicapine flycatchers) have been extirpated from the entire plantation area. The industrial groves also lacked some larger species of kingfisher, woodpeckers, and canopy frugivores. Nevertheless, numerous primary forest taxa (ca. 50% of species) were found in mature industrial tree groves. Albizia attracted the most species of birds, followed closely by Acacia. Both tree types underpinned relatively complex secondary forests that attracted forest birds. In contrast, younger groves of Acacia and Albizia held mainly open country and scrub species. Oil palm, as a remarkably simple and unusual habitat, attracted few bird species. Sustained occurrence of forest birds in all groves of exotic trees at Sabah Softwoods was substantially enhanced by the relatively rich avifauna of the logged native forest remaining in substantial stands throughout the plantation.     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Surface water and ground-water thresholds for maintaining Populus-Salix forests, San Pedro River, Arizona

Ground-water and surface flow depletions are altering riparian ecosystems throughout the southwestern United States, and have contributed to the decline of forests of the pioneer trees Populus fremontii (Fremont cottonwood) and Salix gooddingii (Goodding willow). On some rivers, these forests have been replaced by shrublands of Tamarix ramosissima (tamarisk), a drought-tolerant species from Eurasia. The physiological response of these three riparian plant species to decreases in water availability is well studied, but little attention has been given to shifts in community and population structure in response to declines in surface flow and ground-water levels. Based on study of 17 sites spanning a hydrologic gradient, this research identified hydrologic thresholds above which P. fremontii-S. gooddingii maintain tall dense stands with diverse age classes, and above which they are more abundant than T. ramosissima stands along the San Pedro River in Arizona. Surface flow permanence was the hydrologic variable that explained most of the variance in species abundance and relative importance, with inter-annual ground-water fluctuation and depth to ground water also contributing. P. fremontii and S. gooddingii were dominant over T. ramosissima at sites where surface flow was present more than 76% of the time, inter-annual ground-water fluctuation was less than 0.5 m, and average maximum depth to ground water was less than 2.6 m, during a two-year period of data collection. Because T. ramosissima is a shrub species with smaller leaves and lower canopy heights than P. fremontii and S. gooddingii, these shifts in species composition corresponded to decreases in maximum canopy height and upper stratum (above 8 m) vegetation volume as site water availability declined. As well, sites with deeper water tables and more intermittent flows had greater areal coverage of shrublands and less of woodlands. The altered vegetation structure along dewatered rivers may lower wildlife habitat quality. This study provides river managers with quantitative hydrologic guidelines for conserving tall dense P. fremontii-S. gooddingii stands and emphasizes the importance of maintaining long-term hydrologic conditions that are favorable to these species. The study also has implications for riparian restoration and invasive species management by suggesting that desired outcomes may be achieved through restoration of physical processes.     

Earthworm communities in birch stands with different origin in central Finland

The aim of the study was to compare earthworm communities in anthropogenous birch stands with different origin in Finland. A total of nine forest sites were investigated: three birch stands (Betula pendula) planted ca. 30 years prior to the study after clear-cutting of spruce stands (“Birch after Spruce”, BS), three birch stands planted ca. 30 years earlier on arable soil that had been under normal cultivation until forestation (“Birch after Field”, BF), and three “Natural Deciduous” forests (D). Earthworms were sampled in May and October 1999 using a combination of formaline extraction and modified wet funnels. There were conspicuous differences between replicates of similarly managed forests. Earthworms were totally lacking in one of the D sites, while another had an abundant and diverse community. Only Dendrobaena octaedra was present in one BS site, while the two others harboured also Aporrectodea caliginosa and three Lumbricus species. All these species were also present in the BF sites, where their total biomass (ranging from 70 to 138 g (f.w.)/m²) was 2.6 times the average in BS, and of the same magnitude as the average in natural deciduous stands. A separate experiment revealed that L. terresris and A. caliginosa, which are not found in the surrounding coniferous forest, are able to live and reproduce in the soil of the D site where they were absent. It was concluded that earthworm species survive and reproduce in birch stands established on arable soil, where they have invaded during the long cultivation. On the other hand, their possibilities to disperse from cultural landscapes determine their presence not only in birch stands established in earlier coniferous forests, but also in “natural deciduous” forests where source populations are not present in the surroundings.     

Decomposition of leaves from two indigenous trees of contrasting qualities under shaded-coffee and agricultural land-uses during the dry season at Wondo Genet, Ethiopia

To explore the potential of trees and shrubs on farmlands on traditional systems in southern Ethiopia, mineralization of macronutrients and loss of organics from leaves of Cordia africana and Albizia gummifera were studied under shaded-coffee and agricultural land-uses during the dry season. Leaves in litterbags were incorporated at 15 cm depth in soil under both land uses and residues were recovered after 2, 4, 8, 12 and 16 weeks. Contents of macronutrients and organics in initial and recovered residues were measured. Single- or double-exponential decay or quadratic functions were fitted to describe patterns of decay or release of various leaf constituents. The two species differed significantly (P<0.05) with respect to contents of ash, N, P, K, cellulose (CEL), lignin (LG), total soluble polyphenols (PL), and condensed tannins (CT). Cordia had higher content of ash, K, P, CEL, LG and a higher C-to-N ratio while Albizia had higher contents of N, PL, CT and a higher C-to-P ratio. Albizia had significantly greater mass loss, N loss and release of CT than Cordia. N was immobilized for the first 4 weeks in most treatments. Across land uses and species, mass loss rates varied from −0.023 week⁻¹ in Cordia to −0.034 week⁻¹ in Albizia (R²>0.70). Higher rate of release of CT seems to have facilitated decomposition in Albizia despite higher initial PL and CT in the leaves of this species. There was no significant land-use effect on any of the variables considered. It was concluded that under drier conditions, tree cover might not affect decomposition, and that organic residues with high content of polyphenols, particularly condensed tannins could decompose faster than those with lower content. This suggests that indigenous tree species with high concentrations of tannins, supposedly considered to be of ‘poor quality’, might still be quite useful as an organic input for improving soil fertility and productivity in the tropics.     

Effect of selective logging on the genetic diversity of Scaphium macropodum

We examined the effect of selective logging on the genetic diversity of Scaphium macropodum using RAPD markers via two approaches: (1) to investigate the immediate effect by studying a same population before and after logging, and (2) to determine the long term effect by comparing two regenerated stands with an adjacent unlogged stand, assuming that they were genetically identical before logging. Results showed no negative immediate impact for the first approach, probably due to the high abundance and heterogeneity of S. macropodum in the compartment investigated. However, for the latter approach, substantial genetic erosion (i.e. 31.5% reduction for Shannon diversity, H) was detected in one of the regenerated stands corresponding to its extremely low tree density for S. macropodum. This implies the possible occurrence of genetic drift and increased inbreeding due to population decline as a result of logging. However, the observed genetic differences among the three sub-populations having prevailed before logging cannot be totally discounted in the second approach. This study also demonstrates the use of tree density as a good surrogate measure of genetic diversity. The present harvesting system in Malaysia based on a general cutting limit need to be refined; the basis for determining cutting limit in a forest management unit should consider abundance of commercial species.     

Controlled ectomycorrhization of an exotic legume tree species Acacia holosericea affects the structure of root nodule bacteria community and their symbiotic effectiveness on Faidherbia albida, a native Sahelian Acacia

Many fast growing tree species have been introduced to promote biodiversity rehabilitation on degraded tropical lands. Although it has been shown that plant productivity and stability are dependent on the composition and functionalities of soil microbial communities, more particularly on the abundance and diversity of soil symbiotic micro-organisms (mycorrhizal fungi and rhizobia), the impact of tree introduction on soil microbiota has been scarcely studied. This research has been carried in a field plantation of Acacia holosericea (Australian Acacia species) inoculated or not with an ectomycorrhizal fungus isolate, Pisolithus albus IR100. After 7 year's plantation, the diversity and the symbiotic properties of Bradyrhizobia isolated from the plantation soil or from the surrounding area (Faidherbia albida (Del.) a. Chev. parkland) and able to nodulate F. albida, a native Sahelian Acacia species, have been studied. Results clearly showed that A. holosericea modified the structure of Bradyrhizobia populations and their effectiveness on F. albida growth. This negative effect was counterbalanced by the introduction of an ectomycorrhizal fungus, P. albus, on A. holosericea root systems.In conclusion, this study shows that exotic plant species can drastically affect genotypic and symbiotic effectiveness of native Bradyrhizobia populations that could limit the natural regeneration of endemic plant species such as F. albida. This effect could be counterbalanced by controlled ectomycorrhization with P. albus. These results have to be considered when exotic tree species are used in afforestation programs that target preservation of native plants and soil ecosystem rehabilitation.     

Population responses of oribatids and enchytraeids to ectomycorrhizal and saprotrophic fungi in plant-soil microcosms

The oribatid mites Oppiella nova, Tectocepheus velatus and Nothrus silvestris and the enchytraeid worm Cognettia sphagnetorum are four common animal species in boreal forest soils. According to the literature, they respond differently to clear-cutting of forest stands. O. nova responds with population decreases, T. velatus and N. silvestris with small changes and C. sphagnetorum with population increases. We hypothesised that the presence/absence of ectomycorrhizal (EM) fungi is a major factor in explaining these reactions. The population responses of these soil animals to inoculation of five species of EM fungi growing in symbiosis with their host tree (Pinus sylvestris L.) and one saprotrophic fungus, Hypholoma capnoides, growing on wood were tested in pot microcosms with artificial soil (peat and vermiculite) for 70-84 days. Additionally, plants without inoculation of EM fungi, plants growing in forest soil (FS) and plant-free peat and vermiculite (PV) were included. O. nova increased significantly in abundance in the treatments with the EM fungi Suillus variegatus and Paxillus involutus, but not in the other treatments. T. velatus increased significantly in abundance in FS, but declined in most of the other treatments, and N. silvestris showed a similar, albeit not significant, response. C. sphagnetorum did not increase in abundance in any of the treatments with EM fungi but increased its abundance four times in both FS and PV. The results show that the fungivore O. nova preferentially feeds on certain EM fungi, especially S. variegatus, whereas the EM fungus Piloderma fallax and the saprotrophic H. capnoides are not preferred. In contrast, C. sphagnetorum is restricted in its population growth by the EM fungi studied, and this dominant microbi-detritivore is clearly favoured by the absence of EM fungi.     

The evaluation of key sites for breeding waders in lowland Scotland

This study considers the selection of sites based on the densities of breeding waders that would be considered to be distributed widely across common habitats in Scotland: so-called ‘wider countryside’ species. Five methods to provide a hierarchical classification of sites are assessed. Fifty percent of the populations of redshank (Tringa totanus) and snipe (Gallinago gallinago) occur on just 2 and 4.5% of Scottish lowlands, respectively, yet neither are suitably protected by conservation measures and both would benefit from a carefully targeted approach to their conservation. Three species, lapwing (Vanellus vanellus), curlew (Numenius arquata) and oystercatcher (Haematopus ostralegus) are more appropriately considered to be dispersed species. We propose that, if the density of a breeding wader on a site exceeds the threshold level predicted for the top 1% of Scottish lowland for that species, and if the area of the site exceeds 1 km² then that site be considered to be a key site for farmland waders. The minimum density (in pairs km⁻²) must exceed 16.8 for lapwing, 10.1 for oystercatcher, 6.1 for snipe, 7.5 for curlew or 3.6 for redshank.     

Implications of intraguild predation for sea turtle nest protection

In the United States, raccoons Procyon lotor are often removed from sea turtle nesting beaches to decrease egg mortality. However, raccoons also consume ghost crabs Ocypode quadrata, another common egg predator. Reducing predator populations can benefit secondary predators, inflating total predation pressure and leading to a decline in prey species. We used track and burrow counts to compare raccoon and ghost crab abundance at four beaches in Florida, USA, that differ in management activity and determined predation rates on loggerhead Caretta caretta nests by each predator. Mean raccoon abundance (range 0.12-0.46 tracks plot⁻¹ night⁻¹) and ghost crab density (0.09-0.19 burrows m⁻²) were inversely correlated. Ghost crabs were largest at the site with the fewest raccoons. The stable nitrogen isotope ratios of ghost crabs (mean 9.8‰) were positively correlated with body mass, indicating larger ghost crabs feed at a higher trophic level and suggesting large ghost crabs may consume more loggerhead eggs. The highest rates of egg predation by both predators (31%) occurred where raccoon abundance was lowest and ghost crab abundance was highest, suggesting ghost crab burrows may facilitate predation by raccoons. Our data suggest that predation by raccoons limits ghost crabs and that removing raccoons can increase ghost crab abundance and sea turtle egg mortality. Although predator removal can be effective when nest predation rates are quite high, maintaining moderate raccoon densities may be important for controlling ghost crabs. These results highlight the importance of understanding food web connectivity in developing management strategies to achieve conservation goals, especially when the species of concern are threatened or facing extinction.     

Compound-specific δC and δH analyses of plant and soil organic matter: A preliminary assessment of the effects of vegetation change on ecosystem hydrology

Here we present δ¹³C and δ²H data of long-chained, even-numbered (C₂₇-C₃₁) n-alkanes from C3 (trees) and C4 (grasses) plants and from the corresponding soils from a grassland-woodland vegetation sequence in central Queensland, Australia. Our data show that δ¹³C values of the C4 grassland species were heavier relative to those of C3 tree species from the woodland (Acacia leaves) and woody grassland (Atalaya leaves). However, n-alkanes from the C4 grasses had lighter δ²H values relative to the Acacia leaves, but showed no significant difference in δ²H values when compared with C3 Atalaya leaves. These results differ from those of previous studies, showing that C4 grasses had heavier δ²H values relative to C3 grasses and trees. Those observations have been explained by C4 plants accessing the more evaporation-influenced and isotopically heavier surface water and tree roots sourcing deeper, isotopically lighter soil water (“Two-layered soil-water system”). By comparison, our data suggest that ecosystem changes (vegetation “thickening”) can significantly alter the soil hydrological characteristics. This is shown by the heavier δ²H values in the woodland soil compared with lighter δ²H values in the grassland soil, implying that the recent vegetation change (increased tree biomass) in the woodland had altered soil hydrological conditions. Estimated δ²H values of the source-water for vegetation in the grassland and woodland showed that both trees and grasses in open settings accessed water with lighter δ²H values (avg. −46‰) compared with water accessed by trees in the woodland vegetation (avg. −7‰). These data suggest that in semi-arid environments the “two-layer” soil water concept might not apply. Furthermore, our data indicate that compound-specific δ²H and δ¹³C analyses of n-alkanes from soil organic matter can be used to successfully differentiate between water sources of different vegetation types (grasses versus trees) in natural ecosystems.     

The fate of condensed tannins during litter consumption by soil animals

Condensed tannins (CT) can strongly affect litter decomposition, but their fate during the decomposition process, in particular as influenced by detritivore consumption, is not well understood. We tested the hypothesis that litter CT are reduced by the gut passage of two functionally distinct detritivores of Mediterranean forests, the millipede Glomeris marginata, and the land snail Pomatias elegans, as a fixed proportion of initial litter CT, but more so in Pomatias since snails are known to have a more efficient enzymatic capacity. Contrary to our hypothesis, both detritivore species reduced litter CT to near zero in their faecal pellets irrespective of the wide range in initial leaf litter CT concentrations of 9-188 mg g⁻¹ d m among three Mediterranean tree species (Pistacia terebinthus, Quercus ilex, Alnus glutinosa) and different decomposition stages of their litter. The almost complete disappearance of CT even from some litter types highly concentrated in CT, due to either degradation by gut microorganism or complexation of CT into insoluble high molecular weight structures, suggests a high “de-tanning” efficiency across functionally distinct detritivore species. The transformation of CT-rich litter into virtually CT-free faecal pellets by detritivores might be highly relevant for the subsequent decomposition process in ecosystems with a high macrofauna abundance and CT-rich plant species such as Mediterranean forests.     

Transfer of N fixed by a legume tree to the associated grass in a tropical silvopastoral system

Below-ground transfer of nitrogen (N) fixed by legume trees to associated non-N₂-fixing crops has received little attention in agroforestry, although the importance of below-ground interactions is shown in other ecosystems. We used ¹⁵N natural abundance to estimate N transfer from the legume tree Gliricidia sepium (Jacq.) Kunth ex Walp. to C₄ grass Dichanthium aristatum (Poir.) C.E. Hubb. in a silvopastoral system, where N was recycled exclusively by below-ground processes and N₂ fixation by G. sepium was the sole N input to the system. Finding a suitable reference plant, a grass without contact with tree roots or litter, was problematic because tree roots invaded adjacent grass monocrop plots and soil isotopic signature in soil below distant grass monocrops differed significantly from the agroforestry plots. Thus, we used grass cultivated under greenhouse conditions in pots filled with agroforestry soil as the reference. A model of soil ¹⁵N fractionation during N mineralization was developed for testing the reliability of that estimate. Experimental and theoretical results indicated that 9 months after greenhouse transplanting, the percentage of fixed N in the grass decreased from 35% to <1%, due to N export in cut grass and dilution of fixed N with N taken up from the soil. The effect of soil ¹⁵N fractionation on the estimate of the reference value was negligible. This indicates that potted grass is a suitable reference N transfer studies using ¹⁵N natural abundance. About one third of N in field-grown grass was of atmospheric origin in agroforestry plots and in adjacent D. aristatum grassland invaded by G. sepium roots. The concentration of fixed N was correlated with fine root density of G. sepium but not with soil isotopic signature. This suggests a direct N transfer from trees to grass, e.g. via root exudates or common mycorrhizal networks.     

CO2 and inorganic N supply modify competition for N between co-occurring grass plants, tree seedlings and soil microorganisms

Plant-plant and plant-soil interactions play a key role in determining plant community structure and ecosystem function. However, the effects of global change on the interplay between co-occurring plants and soil microbes in successional communities are poorly understood. In this study, we investigated competition for nitrogen (N) between soil microorganisms, grass plants and establishing tree seedlings under factorial carbon dioxide (CO₂) and N treatments. Fraxinus excelsior seedlings were germinated in the presence or absence of grass competition (Dactylis glomerata) at low (380 μmol mol⁻¹) or high (645 μmol mol⁻¹) CO₂ and at two levels of N nutrition in a mesocosm experiment. Pulse ¹⁵N labelling was used to examine N partitioning among plant and soil compartments. Dactylis exerted a strong negative effect on Fraxinus biomass, N capture and ¹⁵N recovery irrespective of N and CO₂ treatment. In contrast, the presence of Dactylis had a positive effect on the microbial N pool. Plant and soil responses to N treatment were of a greater magnitude compared with responses to elevated CO₂, but the pattern of Fraxinus- and microbial-N pool response to N and CO₂ varied depending on grass competition treatment. Within the Dactylis competition treatment, decreases in Fraxinus biomass in response to N were not mirrored by decreases in tree seedling N content, suggesting a shift from below- to above-ground competition. In the Dactylis-sown pots, ¹⁵N recovery could be ranked Dactylis > microbial pool > Fraxinus in all N and CO₂ treatment combinations. Inequalities between Fraxinus and soil microorganisms in terms of ¹⁵N recovery were exacerbated by N addition. Contrary to expectations, elevated CO₂ did not increase plant-microbe competition. Nevertheless, microbial ¹⁵N recovery showed a small positive increase in the high CO₂ treatment. Overall, elevated CO₂ and N supply did not interact on plant/soil N partitioning. Our data suggest that the competitive balance between establishing tree seedlings and grass plants in an undisturbed sward is relatively insensitive to CO₂ or N-induced modifications in N competition between plant and soil compartments.