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Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta-analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non-native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non-native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm-invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long-lasting effects on ecosystem functioning. [ABSTRACT FROM AUTHOR]

Dissolved organic matter (DOM), the mobile form of soil organic matter (SOM), plays an important role in soil C cycling and in nutrient transport. We investigated the effects of 5 years of CO2 enrichment (370 versus 570 µmol CO2 mol-1) on DOM dynamics at the alpine treeline, including the analysis of fast-cycling components such as low molecular weight organic acids (LMWOAs), dissolved organic carbon (DOC) biodegradability, and the decomposition of 14C-labeled oxalate. Concentrations of DOC in canopy throughfall were 20% higher at elevated CO2, probably driven by higher carbohydrate concentrations in leaves. In the organic soil layer, 5 years of CO2 enrichment increased water-extractable organic C by 17% and soil solution DOC at 5 cm depth by 20%. The 13C tracing of recently assimilated CO2 revealed that the input of recent plant-derived C (<15% of total DOC) was smaller than the CO2-induced increase in DOC. This strongly suggests that CO2 enrichment enhanced the mobilization of native DOC, which is supported by significant increases in dissolved organic nitrogen (DON). We mainly attribute these increases to a stimulated microbial activity as indicated by higher basal and soil respiration rates (+27%). The 14C-labeled oxalate was more rapidly mineralized from high CO2 soils. The concentrations of LMWOAs, but also those of "hydrophilic" DOC and biodegradable DOC (6% of total DOC), were, however, not affected by elevated CO2, suggesting that production and consumption of "labile" DOC were in balance. In summary, our data suggest that 5 years of CO2 enrichment speeded up the cycling of "labile" DOM and SOM in a late successional treeline ecosystem and increased the mobilization of older DOM through a stimulated microbial activity. Such a "priming effect" implies that elevated CO2 can accelerate the turnover of native SOM, and thus, it may induce increasing losses of old C from thick organic layers. [ABSTRACT FROM AUTHOR]

1. Plants are frequently observed to increase carbon allocation to below-ground sinks and particularly, to accelerate fine root turnover in response to rising atmospheric CO2 concentration. While these strong below-ground responses have predominantly been observed in rapidly expanding systems, late successional plant communities have rarely been studied. 2. In an ongoing free air CO2 enrichment (FACE) experiment, we assessed below-ground responses to elevated CO2 after 4 years, in a treeline ecosystem in the Swiss Central Alps (2180 m a.s.l.) dominated by a late successional ericaceous dwarf shrub community ( Vaccinium myrtillus, V. uliginosum, Empetrum hermaphroditum), and a sparse overstorey of 30-year-old Larix decidua and Pinus uncinata trees. Measurements included quantification of fine root growth using ingrowth root cores and parallel standing crop harvests and decomposition of roots using litter bags. 3. Elevated CO2 did not stimulate root growth of the treated vegetation (although some significant above-ground growth responses were observed), nor did altered root decomposition occur. Root quality measurements indicated that elevated CO2 resulted in significantly higher starch concentrations, but no change in N concentration, or root dehydrogenase activity. 4. The use of the stable isotope δ13C permitted us to trace the new carbon entering the system through our CO2 enrichment treatment. We observed that only c. 30% of new root biomass (< 2 mm) was formed by new carbon indicating a rather slow root turnover in this system. 5. Our data show that fine root growth may be much less stimulated by elevated CO2 in systems with late successional elements than has been reported in ecosystems with a rapidly expanding plant community biomass. [ABSTRACT FROM AUTHOR]

Northern latitude and upper altitude climatic treelines have received increasing attention given their potential sensitivity to atmospheric and climate change. While greater radial stem growth at treeline sites in recent decades has been attributed largely to increasing temperature, rising atmospheric CO2 concentration may also be contributing to this growth stimulation. Tree ring increments of mature Larix decidua and Pinus uncinata were measured over 4 years in a free air CO2 enrichment experiment at treeline in the Swiss Central Alps (2180 m a.s.l.). In addition, a one-time defoliation treatment in the second year (2002) of the experiment was used to simulate one of the common natural insect outbreak events. In response to elevated atmospheric CO2, Larix showed a cumulative 4-year growth response of+41%, with particularly strong responses in the third and fourth year. This increase in radial stem wood growth was the result of more latewood production, in particular, the formation of larger tracheids, rather than a greater number of cells. In contrast, Pinus showed no change in ring width to elevated [CO2], neither in each of the treatment years, nor in the cumulative response over 4 years, although an increase in tracheid size was observed in the third year. Defoliation led to a pronounced decrease in annual ring width of both species, marked in particular by less latewood production, in the treatment, as well as subsequent years. There was no significant interaction between defoliation and CO2 enrichment. Although Pinus showed no growth response to CO2, the positive growth response observed in Larix after 4 years of CO2 enrichment implies that the sensitivity of treeline trees to global change may not be purely temperature driven. We conclude that the open sparse canopy in the treeline ecotone favours the indeterminate growth strategy of the early successional Larix when neither weather nor carbon are limiting, whereas the later successional Pinus does not show any indication of more vigorous growth under future higher atmospheric CO2 concentrations. [ABSTRACT FROM AUTHOR]

Historically, carbon limitation, through a shortage of photoassimilates has been argued to limit the growth of trees at the upper altitudinal treeline. In a three-year free-air CO2 enrichment (FACE) experiment, two species of 30-year-old alpine conifers (Larix decidua and Pinus uncinata) were studied to test this hypothesis in situ in the Swiss Central Alps (2180 m above sea level). CO2 enrichment was combined with foliage removal to test the effect of altered source-sink relationships on tree growth and leaf level responses. Elevated CO2 enhanced photosynthesis and increased nonstructural carbohydrate (NSC) concentrations in the needles of both species. While the deciduous larch trees showed longer needles and a stimulation of shoot growth over all three seasons when grown in situ under elevated CO2, pine trees showed no such responses. Irrespective of CO2 concentration, defoliation in both species stimulated photosynthesis and increased stomatal conductance in remaining current-year needles in the treatment year and reduced leaf nitrogen concentration in the year following defoliation. Defoliated larch trees had fewer and shorter needles with reduced NSC concentrations in the year following defoliation and showed no stimulation in shoot elongation when exposed to elevated CO2. In contrast, defoliation of evergreen pine trees had no effect on needle NSC concentrations, but stimulated shoot elongation when defoliated trees were exposed to elevated CO2. After three years, our results suggest that deciduous larch is carbon limited at treeline, while evergreen pine is not. However, as indicated by the defoliation treatment, the carbon economy of these trees can clearly be modified by extreme events. The expected changes in growth of these treeline trees with improving carbon availability as atmospheric CO2 continues to increase will thus depend on both the interplay between biotic and abiotic processes, and the species or tree functional types involved. [ABSTRACT FROM AUTHOR]

Summary • Experimental CO2 enrichment of mature Larix decidua and Pinus uncinata trees and their understory vegetation was used to test the carbon limitation hypothesis of treeline formation at the alpine treeline in Switzerland. • Forty plots (each 1.1 m2 ) were established; half of them were exposed to elevated (566 ppm) atmospheric CO2 using a free air CO2 enrichment (FACE) system releasing pure CO2 , and the other half were treated as controls at current ambient [CO2 ]. • Reliable and adequate CO2 control was achieved, with 63% and 90% of 1-min averages having a [CO2 ] within ±10% and ±20% of the target value, respectively, which is comparable to previous FACE systems. Both tree species showed higher net photosynthesis, lower stomatal conductance, and increased accumulation of nonstructural carbohydrates in response to CO2 in the first year of treatment. Quite unexpectedly, shoot length increment increased significantly at elevated CO2 (up to 23%) compared with controls in both species. • The pure CO2 release technology proved suitable for CO2 enrichment of native trees on this remote mountain slope. Our results suggest an improved C balance and growth of treeline trees in response to elevated CO2 . However, it is unclear whether this initial growth stimulation will persist in the longer term. [ABSTRACT FROM AUTHOR]