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The stability of soil organic matter (SOM) pools exposed to elevated CO and warming has not been evaluated adequately in long-term experiments and represents a substantial source of uncertainty in predicting ecosystem feedbacks to climate change. We conducted a 6-year experiment combining free-air CO enrichment (FACE, 550 ppm) and warming (+2 °C) to evaluate changes in SOM accumulation in native Australian grassland. In this system, competitive interactions appear to favor C over C species under FACE and warming. We therefore investigated the role of plant functional type (FT) on biomass and SOM responses to the long-term treatments by carefully sampling soil under patches of C- and C-dominated vegetation. We used physical fractionation to quantify particulate organic matter (POM) and long-term incubation to assess potential decomposition rates. Aboveground production of C grasses increased in response to FACE, but total root biomass declined. Across treatments, C : N ratios were higher in leaves, roots and POM of C vegetation. CO and temperature treatments interacted with FT to influence SOM, and especially POM, such that soil carbon was increased by warming under C vegetation, but not in combination with elevated CO. Potential decomposition rates increased in response to FACE and decreased with warming, possibly owing to treatment effects on soil moisture and microbial community composition. Decomposition was also inversely correlated with root N concentration, suggesting increased microbial demand for older, N-rich SOM in treatments with low root N inputs. This research suggests that C-C vegetation responses to future climate conditions will strongly influence SOM storage in temperate grasslands. [ABSTRACT FROM AUTHOR]

The increasing atmospheric concentration of carbon dioxide ([CO2]) contributes to global warming and the accompanying shifts in climate. However, [CO2] itself has the potential to impact on Australia's terrestrial biodiversity, due to its importance in the photosynthetic process, which underlies all terrestrial food webs. Here, we review our knowledge regarding the impacts of elevated [CO2] on native terrestrial species and ecosystems, and suggest key areas in which we have little information on this topic. Experimental information exists for 70 (or less than 0.05%) of Australia's native terrestrial plant and animal species. Of these, 68 are vascular plants. The growth of Australian woody species is more reliably increased by elevated [CO2] than it is in grasses. At the species level, the most overwhelming responses to increased [CO2] are a reduction in plant nitrogen concentration and an increase in the production of secondary metabolites. This is of particular concern for Australia's unique herbivorous and granivorous marsupials, for which no information is available. While many plant species also displayed increased growth rates at higher [CO2], this was far from universal, indicating that changes in community structure and function are likely, leading to alterations of habitat quality. Future research should be directed to key knowledge gaps including the relationship between [CO2], fire frequency and fire tolerance and the impacts of increasing [CO2] for Australia's iconic browsing mammals. We also know virtually nothing of the impacts of the increasing [CO2] on Australia's unique shrublands and semi-arid/arid rangelands. In conclusion, there is sufficient information available to be certain that the increasing [CO2] will affect Australia's native biodiversity. However, the information required to formulate predictions concerning the long-term future of almost all organisms is far in excess of that currently available. [ABSTRACT FROM AUTHOR]

While the influence of elevated CO2 on the production, mass and quality of plant seeds has been well studied, the effect of warming on these characters is largely unknown; and there is practically no information on possible interactions between warming and elevated CO2, despite the importance of these characters in population maintenance and recovery. Here, we present the impacts of elevated CO2 and warming, both in isolation and combination, on seed production, mass, quality, germination success and subsequent seedling growth of Austrodanthonia caespitosa, a dominant temperate C3 grass from Australia, using seeds collected from the TasFACE experiment. Mean seed production and mass were not significantly affected by either elevated CO2 or warming, but elevated CO2 more than doubled the proportion of very light, inviable seeds ( P < 0.05) and halved mean seed N concentration ( P < 0.04) and N content ( P < 0.03). The dependence of seed germination success on seed mass was affected by an elevated CO2× warming interaction ( P < 0.004), such that maternal exposure to elevated CO2 or warming reduced germination if applied in isolation, but not when applied in combination. Maternal effects were retained when seedlings were grown in a common environment for 6 weeks, with seedlings descended from warmed plants 20% smaller ( P < 0.008) with a higher root : shoot ratio ( P < 0.001) than those from unwarmed plants. Given that both elevated CO2 and warming reduced seed mass, quality, germinability or seedling growth, it is likely that global change will reduce population growth or distribution of this dominant species. [ABSTRACT FROM AUTHOR]

• Flowering is a critical stage in plant life cycles, and changes might alter processes at the species, community and ecosystem levels. Therefore, likely flowering-time responses to global change drivers are needed for predictions of global change impacts on natural and managed ecosystems. • Here, the impact of elevated atmospheric CO2 concentration ([CO2]) (550 µmol mol−1) and warming (+2ºC) is reported on flowering times in a native, species-rich, temperate grassland in Tasmania, Australia in both 2004 and 2005. • Elevated [CO2] did not affect average time of first flowering in either year, only affecting three out of 23 species. Warming reduced time to first flowering by an average of 19.1 d in 2004, acting on most species, but did not significantly alter flowering time in 2005, which might be related to the timing of rainfall. Elevated [CO2] and warming treatments did not interact on flowering time. • These results show elevated [CO2] did not alter average flowering time or duration in this grassland; neither did it alter the response to warming. Therefore, flowering phenology appears insensitive to increasing [CO2] in this ecosystem, although the response to warming varies between years but can be strong. [ABSTRACT FROM AUTHOR]

• Flowering is a critical stage in plant life cycles, and changes might alter processes at the species, community and ecosystem levels. Therefore, likely flowering-time responses to global change drivers are needed for predictions of global change impacts on natural and managed ecosystems. • Here, the impact of elevated atmospheric CO2 concentration ([CO2]) (550 µmol mol−1) and warming (+2ºC) is reported on flowering times in a native, species-rich, temperate grassland in Tasmania, Australia in both 2004 and 2005. • Elevated [CO2] did not affect average time of first flowering in either year, only affecting three out of 23 species. Warming reduced time to first flowering by an average of 19.1 d in 2004, acting on most species, but did not significantly alter flowering time in 2005, which might be related to the timing of rainfall. Elevated [CO2] and warming treatments did not interact on flowering time. • These results show elevated [CO2] did not alter average flowering time or duration in this grassland; neither did it alter the response to warming. Therefore, flowering phenology appears insensitive to increasing [CO2] in this ecosystem, although the response to warming varies between years but can be strong. [ABSTRACT FROM AUTHOR]

Rising atmospheric carbon dioxide concentration ([CO2]) has the potential to stimulate ecosystem productivity and sink strength, reducing the effects of carbon (C) emissions on climate. In terrestrial ecosystems, increasing [CO2] can reduce soil nitrogen (N) availability to plants, preventing the stimulation of ecosystem C assimilation; a process known as progressive N limitation. Using ion exchange membranes to assess the availability of dissolved organic N, ammonium and nitrate, we found that CO2 enrichment in an Australian, temperate, perennial grassland did not increase plant productivity, but did reduce soil N availability, mostly by reducing nitrate availability. Importantly, the addition of 2 °C warming prevented this effect while warming without CO2 enrichment did not significantly affect N availability. These findings indicate that warming could play an important role in the impact of [CO2] on ecosystem N cycling, potentially overturning CO2-induced effects in some ecosystems. [ABSTRACT FROM AUTHOR]

• Species differ in their responses to global changes such as rising CO2 and temperature, meaning that global changes are likely to change the structure of plant communities. Such alterations in community composition must be underlain by changes in the population dynamics of component species. • Here, the impact of elevated CO2 (550 µmol mol−1) and warming (+2°C) on the population growth of four plant species important in Australian temperate grasslands is reported. Data collected from the Tasmanian free-air CO2 enrichment (TasFACE) experiment between 2003 and 2006 were analysed using population matrix models. • Population growth of Themeda triandra, a perennial C4 grass, was largely unaffected by either factor but population growth of Austrodanthonia caespitosa, a perennial C3 grass, was reduced substantially in elevated CO2 plots. Warming and elevated CO2 had antagonistic effects on population growth of two invasive weeds, Hypochaeris radicata and Leontodon taraxacoides, with warming causing population decline. Analysis of life cycle stages showed that seed production, seedling emergence and establishment were important factors in the responses of the species to global changes. • These results show that the demographic approach is very useful in understanding the variable responses of plants to global changes and in elucidating the life cycle stages that are most responsive. [ABSTRACT FROM AUTHOR]

Increasing concentrations of atmospheric carbon dioxide are expected to affect carbon assimilation and evapotranspiration (ET), ultimately driving changes in plant growth, hydrology, and the global carbon balance. Direct leaf biochemical effects have been widely investigated, whereas indirect effects, although documented, elude explicit quantification in experiments. Here, we used a mechanistic model to investigate the relative contributions of direct (through carbon assimilation) and indirect (via soil moisture savings due to stomatal closure, and changes in leaf area index) effects of elevated CO2 across a variety of ecosystems. We specifically determined which ecosystems and climatic conditions maximize the indirect effects of elevated CO2. The simulations suggest that the indirect effects of elevated CO2 on net primary productivity are large and variable, ranging from less than 10% to more than 100% of the size of direct effects. For ET, indirect effects were, on average, 65% of the size of direct effects. Indirect effects tended to be considerably larger in water-limited ecosystems. As a consequence, the total CO2 effect had a significant, inverse relationship with the wetness index and was directly related to vapor pressure deficit. These results have major implications for our understanding of the CO2 response of ecosystems and for global projections of CO2 fertilization, because, although direct effects are typically understood and easily reproducible in models, simulations of indirect effects are far more challenging and difficult to constrain. Our findings also provide an explanation for the discrepancies between experiments in the total CO2 effect on net primary productivity. [ABSTRACT FROM AUTHOR]

The net annual exchange of carbon between the atmosphere and terrestrial ecosystems is of prime importance in determining the concentration of CO2 ([CO2]) in the atmosphere and consequently future climate. Carbon loss occurs primarily through soil respiration; it is known that respiration is sensitive to the global changes in [CO2] and temperature, suggesting that the net carbon balance may change in the future. However, field manipulations of temperature and [CO2] alter many important environmental factors so it is unclear how much of the observed alterations in soil respiration is due to changes of microbial function itself instead of changes to the physical and chemical environment. Here we focus on resolving the importance of changes in the microbial community in response to warming and elevated [CO2] on carbon mineralisation, something not possible in field measurements. We took plant material and soil inocula from a long running experiment where native grassland had been exposed to both warming and elevated CO2 and constructed a reciprocal transplant experiment. We found that the rate of decomposition (heterotrophic respiration) was strongly determined by the origin of the microbial community. The combined warming + elevated CO2 treatment produced a soil community that gave respiration rates 30% higher when provided with shoot litter and 70% for root litter than elevated CO2 treatment alone, with the treatment source of the litter being unimportant. Warming, especially in the presence of elevated CO2, increased the size of the apparent labile carbon pool when either C3 or C4 litter was added. Thus, the metabolic activity of the soil community was affected by the combination of warming and elevated CO2 such that it had an increased ability to mineralise added organic matter, regardless of its source. Therefore, soil C efflux may be substantially increased in a warmer, high CO2 world. Current ecosystem models mostly drive heterotrophic respiration from plant litter quality, soil moisture and temperature but our findings suggest equal attention will need to be paid to capturing microbial processes if we are to accurately project the future C balance of terrestrial ecosystems and quantify the feedback effect on atmospheric concentrations of CO2. [ABSTRACT FROM AUTHOR]

When a plant is introduced to a new ecosystem it may escape from some of its coevolved herbivores. Reduced herbivore damage, and the ability of introduced plants to allocate resources from defence to growth and reproduction can increase the success of introduced species. This mechanism is known as enemy release and is known to occur in some species and situations, but not in others. Understanding the conditions under which enemy release is most likely to occur is important, as this will help us to identify which species and habitats may be most at risk of invasion. We compared in situ measurements of herbivory on 16 plant species at 12 locations within their native European and introduced Australian ranges to quantify their level of enemy release and understand the relationship between enemy release and time, space and climate. Overall, plants experienced approximately seven times more herbivore damage in their native range than in their introduced range. We found no evidence that enemy release was related to time since introduction, introduced range size, temperature, precipitation, humidity or elevation. From here, we can explore whether traits, such as leaf defences or phylogenetic relatedness to neighbouring plants, are stronger indicators of enemy release across species.

Our limited understanding of terrestrial ecosystem responses to elevated CO2 is a major constraint on predicting the impacts of climate change. A change in botanical composition has been identified as a key factor in the CO2 response with profound implications for ecosystem services such as plant production and soil carbon storage. In temperate grasslands, there is a strong consensus that elevated CO2 will result in a greater physiological stimulus to growth in legumes and to a lesser extent forbs, compared with C3 grasses, and the presumption this will lead in turn to a greater proportion of these functional groups in the plant community. However, this view is based on data mainly collected in experiments of three or less years in duration and not in experiments where defoliation has been by grazing animals. Grazing is, however, the most common management of grasslands and known in itself to influence botanical composition. In a long-term Free Air Carbon Dioxide Enrichment (FACE) experiment in a temperate grassland managed with grazing animals (sheep), we found the response to elevated CO2 in plant community composition in the first 5 years was consistent with the expectation of increased proportions of legumes and forbs. However, in the longer term, these differences diminished so that the proportions of grasses, legumes and forbs were the same under both ambient and elevated CO2 . Analysis of vegetation before and after each grazing event showed there was a sustained disproportionately greater removal ('apparent selection') of legumes and forbs by the grazing animals. This bias in removal was greater under elevated CO2 than ambient CO2 . This is consistent with sustained faster growth rates of legumes and forbs under elevated CO2 being countered by selective defoliation, and so leading to little difference in community composition., (© 2013 John Wiley & Sons Ltd.)

Leaf morphology varies reliably with increasing altitude in many species, and this is generally considered to be related to temperature. Changes in irradiance with elevation may confound any relationships between a morphological character and altitude, particularly if altitude of origin affects the response to irradiance. Here we describe the interaction between irradiance and altitude of origin on leaf morphology of Southern beech, Nothofagus cunninghamii. Cuttings from each of four altitudes were grown in a glasshouse under full sunlight or 50% shade, and leaf morphology was related to irradiance, altitude of origin and accession. There was a significant interaction between irradiance and altitude of origin for leaf length, width, thickness, area, weight, specific leaf area and stomatal density. There was no effect of altitude on leaf length to width ratio or stomatal index, nor was there an interaction between irradiance and altitude of origin for these variables. These results show that the altitude of origin of a plant has an overriding impact on the leaf morphological response to irradiance. This must be considered in climatic reconstructions.

•  Leaf morphology varies predictably with altitude, and leaf morphological features have been used to estimate average temperatures from fossil leaves. The altitude-leaf morphology relationship is confounded by the two processes of acclimation and adaptation, which reflect environmental and genetic influences, respectively. •  Here we describe the relationship between altitude and leaf morphology for Southern beech, Nothofagus cunninghamii (Hook.) Oerst.. Cuttings from several trees from each of four altitudes were grown in a common glasshouse experiment, and leaf morphology related to both genotype and altitude of origin. •  Genotype had a significant impact on leaf morphology, but in the field there was also a significant, overriding effect of altitude. This altitude effect disappeared in glasshouse-grown plants for all morphological variables other than leaf thickness and specific leaf area. •  These results show that, while leaf length, width and area are partially controlled by genetic factors, these variables are plastic and respond to environmental influences associated with a particular altitude. Thus altitudinal trends in leaf size in N. cunninghamii are unlikely to be the result of adaptation.

Eucalyptus nitens is a species that is adapted to low temperature. This study examines xanthophyll-cycle engagement in E. nitens seedlings exposed to cold-induced photoinhibitory conditions under different levels of irradiance and nutrient status. Xanthophyll-cycle pool size indicated an increased requirement for light energy dissipation under high irradiance and low nutrient status. Greater sensitivity to photoinhibition of non-shaded seedlings indicated that sustained xanthophyll-cycle engagement may occur in response to damaged chlorophyll. Within irradiance treatments, fertilised seedlings had higher photochemical efficiency and faster recovery from photoinhibition than unfertilised seedlings. These results demonstrate that fertilised compared to unfertilised seedlings can utilise a greater proportion of incident light under cold temperature conditions

Alpine environments are characterised by low temperatures and high light intensities. This combination leads to high light stress owing to the imbalance between light energy harvesting and its use in photochemistry. In extreme cases, high light stress can lead to the level of photo-oxidative damage exceeding the rate of repair to the photosynthetic apparatus. Plant species may vary in the mechanisms they use to prevent photodamage, but most comparisons are of geographically and ecologically distinct species. Differences in leaf colouration suggested that photoprotective strategies might differ among Tasmanian evergreen alpine shrub species. We compared chlorophyll fluorescence and leaf pigment composition in six co-occurring alpine shrub species on the summit of Mt Wellington, southern Tasmania, Australia, during spring and autumn. We found marked differences among species in light energy utilisation, attenuation and dissipation. Ozothamnus ledifolius maintained a large capacity for photosynthetic light utilisation and thus, had a low requirement for light dissipation. All five of the other species relied on xanthophyll-cycle-dependent thermal energy dissipation. In addition Epacris serpyllifolia, Richea sprengelioides and Leptospermum rupestre had foliar anthocyanins that would attenuate photosynthetically active light in the leaf. During spring, all species retained de-epoxidised xanthophylls through the night and the pre-dawn concentration of antheraxanthin and zeaxanthin was significantly correlated with reductions in pre-dawn F v / F m . We propose that these species use three photoprotective strategies to cope with the combination of high light and low temperature.

Photoinhibition of photosynthesis at low temperatures was investigated in two species of subalpine eucalypt, Eucalypts nitens (Deane and Maiden) Maiden and E. pauciflora Sieb. ex Spreng. Imposition of an artificial cold-hardening treatment increased the frost tolerance of leaf tissue and increased tolerance to excess light. Cold-hardened seedlings of both species had a higher photosynthetic capacity than non-hardened seedlings at 6 and 16°C and lower levels of non-photochemical quenching (NPQ) at 20 and 5°C. Furthermore, hardened seedlings had faster rates of NPQ development at 5 and -3.5°C. An increase in minimal fluorescence, which indicates slowly reversible photoinhibition, was evident in all seedlings at -1.5 and -3.5°C but was less pronounced in hardened seedlings, with a threefold faster rate of development of NPQ, at -3.5°C than non-hardened seedlings. Hardened seedlings also recovered faster from photoinhibition at -3.5°C. Thus cold hardening increased tolerance to high light in these species. Differences between E. nitens and E. pauciflora in their response to excess light were small and significant only at -3.5°C. Faster recovery from photoinhibition of E. pauciflora was consistent with its occurrence in colder habitats than E. nitens.