Improved evaporative flux partitioning and carbon flux in the land surface model JULES: impact on the simulation of land surface processes in temperate Europe

The primary role of land surface models embedded in climate models is to partition surface available energy into upwards, radiative, sensible and latent heat fluxes. Partitioning of evapotranspiration, ET, is of fundamental importance: as a major component of the total surface latent heat flux, ET affects the simulated surface water balance, and related energy balance, and consequently the feedbacks with the atmosphere. In this context it is also crucial to credibly represent the CO2 exchange between ecosystems and their environment. In this study, JULES, the land surface model used in UK weather and climate models, has been evaluated for temperate Europe. Compared to eddy covariance flux measurements, the CO2 uptake by the ecosystem is underestimated and the ET overestimated. In addition, the contribution to ET from soil and intercepted water evaporation far outweighs the contribution of plant transpiration. To alleviate these biases, adaptations have been implemented in JULES, based on key literature references. These adaptations have improved the simulation of the spatio-temporal variability of the fluxes and the accuracy of the simulated GPP and ET, including its partitioning. This resulted in a shift of the seasonal soil moisture cycle. These adaptations are expected to increase the fidelity of climate simulations over Europe. Finally, the extreme summer of 2003 was used as evaluation benchmark for the use of the model in climate change studies. The improved model captures the impact of the 2003 drought on the carbon assimilation and the water use efficiency of the plants. It, however, underestimates the 2003 GPP anomalies. The simulations showed that a reduction of evaporation from the interception and soil reservoirs, albeit not of transpiration, largely explained the good correlation between the carbon and the water fluxes anomalies that was observed during 2003. This demonstrates the importance of being able to discriminate the response of individual component of the ET flux to environmental forcing.

Palaeovegetation (Communications arising): diversity of temperate plants in east Asia

The exceptionally broad species diversity of vascular plant genera in east Asian temperate forests, compared with their sister taxa in North America, has been attributed to the greater climatic diversity of east Asia, combined with opportunities for allopatric speciation afforded by repeated fragmentation and coalescence of populations through Late Cenozoic ice-age cycles1. According to Qian and Ricklefs1, these opportunities occurred in east Asia because temperate forests extended across the continental shelf to link populations in China, Korea and Japan during glacial periods, whereas higher sea levels during interglacial periods isolated these regions and warmer temperatures restricted temperate taxa to disjunct refuges. However, palaeovegetation data from east Asia2, 3, 4, 5, 6 show that temperate forests were considerably less extensive than today during the Last Glacial Maximum, calling into question the coalescence of tree populations required by the hypothesis of Qian and Ricklefs1.

Spatial structuring of arbuscular mycorrhizal communities in benchmark and modified temperate eucalypt woodlands

Arbuscular mycorrhizal fungi (AMF) are crucial to the functioning of the plant–soil system, but little is known about the spatial structuring of AMF communities across landscapes modified by agriculture. AMF community composition was characterized across four sites in the highly cleared south-western Australian wheatbelt that were originally dominated by forb-rich eucalypt woodlands. Environmentally induced spatial structuring in AMF composition was examined at four scales: the regional scale associated with location, the site scale associated with past management (benchmark woodlands with no agricultural management history, livestock grazing, recent revegetation), the patch scale associated with trees and canopy gaps, and the fine scale associated with the herbaceous plant species beneath which soils were sourced. Field-collected soils were cultured in trap pots; then, AMF composition was determined by identifying spores and through ITS1 sequencing. Structuring was strongest at site scales, where composition was strongly related to prior management and associated changes in soil phosphorus. The two fields were dominated by the genera Funneliformis and Paraglomus, with little convergence back to woodland composition after revegetation. The two benchmark woodlands were characterized by Ambispora gerdemannii and taxa from Gigasporaceae. Their AMF communities were strongly structured at patch scales associated with trees and gaps, in turn most strongly related to soil N. By contrast, there were few patterns at fine scales related to different herbaceous plant species, or at regional scales associated with the 175 km distance between benchmark woodlands. Important areas for future investigation are to identify the circumstances in which recolonization by woodland AMF may be limited by fungal propagule availability, reduced plant diversity and/or altered chemistry in agricultural soils.

The effect of temperature and inorganic phosphorus supply on growth and acid phosphatase production in arctic and temperate strains of ectomycorrhizal Hebeloma spp. in axenic culture

Acid phosphatase production by 12 Hebeloma strains was usually derepressed when inorganic phosphorus in the growth medium was limited, but appeared to be constitutive in some strains. At low temperatures (≤ 12°) arctic strains produced more extracellular and wall-bound acid phosphatase, yet grew more slowly than the temperate strains. We suggest that low growth rates in arctic strains may be a physiological response to cold whereby resources are diverted into carbohydrate accumulation for cryoprotection. At near freezing temperatures, increased extracellular phosphatase production may compensate for a loss of enzyme activity at low temperature and serve to hydrolyse organic phosphorus in frozen soil over winter.

Utilization of organic nitrogen by ectomycorrhizal fungi (Hebeloma spp.) of arctic and temperate origin

Arctic and temperate strains of Hebeloma spp. were grown in axenic culture on glutamic acid, alanine, lysine and NH4+ as sole sources of nitrogen (N), with excess carbon (C) or deficient C (supplied as glucose). Their ability to utilize seed protein as a natural N source was also assessed. All strains tested had the capacity to assimilate amino acids and generally utilized alanine and glutamic acid more readily than NH4+. Some strains were able to utilize amino C when starved of glucose C, and could mineralize amino-N to NH3-N. Arctic strains, in particular, appeared to be pre-adapted to the utilization of seed protein N and glutamic acid N, which is often liberated in high concentrations after soil freezing. The results are discussed in relation to their possible ecological importance.

Plastic and evolutionary responses to heat stress in a temperate dung fly: negative correlation between basal and induced heat tolerance?

Extreme weather events such as heat waves are becoming more frequent and intense. Populations can cope with elevated heat stress by evolving higher basal heat tolerance (evolutionary response) and/or stronger induced heat tolerance (plastic response). However, there is ongoing debate about whether basal and induced heat tolerance are negatively correlated and whether adaptive potential in heat tolerance is sufficient under ongoing climate warming. To evaluate the evolutionary potential of basal and induced heat tolerance, we performed experimental evolution on a temperate source 4 population of the dung fly Sepsis punctum. Offspring of flies adapted to three thermal selection regimes (Hot, Cold and Reference) were subjected to acute heat stress after having been exposed to either a hot-acclimation or non-acclimation pretreatment. As different traits may respond differently to temperature stress, several physiological and life history traits were assessed. Condition dependence of the response was evaluated by exposing juveniles to different levels of developmental (food restriction/rearing density) stress. Heat knockdown times were highest, whereas acclimation effects were lowest in the Hot selection regime, indicating a negative association between basal and induced heat tolerance. However, survival, adult longevity, fecundity and fertility did not show such a pattern. Acclimation had positive effects in heat-shocked flies, but in the absence of heat stress hot-acclimated flies had reduced life spans relative to nonacclimated ones, thereby revealing a potential cost of acclimation. Moreover, body size positively affected heat tolerance and unstressed individuals were less prone to heat stress than stressed flies, offering support for energetic costs associated with heat tolerance. Overall, our results indicate that heat tolerance of temperate insects can evolve under rising temperatures, but this response could be limited by a negative relationship between basal and induced thermotolerance, and may involve some but not other fitness-related traits.

The role of CO2 and dynamic vegetation on the impact of temperate land use change in the HadCM3 coupled climate model

Human induced land-use change (LUC) alters the biogeophysical characteristics of the land surface influencing the surface energy balance. The level of atmospheric CO2 is expected to increase in the coming century and beyond, modifying temperature and precipitation patterns and altering the distribution and physiology of natural vegetation. It is important to constrain how CO2-induced climate and vegetation change may influence the regional extent to which LUC alters climate. This sensitivity study uses the HadCM3 coupled climate model under a range of equilibrium forcings to show that the impact of LUC declines under increasing atmospheric CO2, specifically in temperate and boreal regions. A surface energy balance analysis is used to diagnose how these changes occur. In Northern Hemisphere winter this pattern is attributed in part to the decline in winter snow cover and in the summer due to a reduction in latent cooling with higher levels of CO2. The CO2-induced change in natural vegetation distribution is also shown to play a significant role. Simulations run at elevated CO2 yet present day vegetation show a significantly increased sensitivity to LUC, driven in part by an increase in latent cooling. This study shows that modelling the impact of LUC needs to accurately simulate CO2 driven changes in precipitation and snowfall, and incorporate accurate, dynamic vegetation distribution.

Net Energy Analysis of Double Glazing for Residential Buildings in Temperate Climates

Energy used in buildings is a major contributor to Australia’s energy consumption and associated environmental impacts. The advent of complex glazing systems such as double glazing, particularly in northern America and Europe, has partially closed a weak thermal link in the building envelope. In milder climates, however, building envelope features may not be as effective in life cycle energy terms, i.e. including the embodied energy of their manufacture. A net energy analysis compares the savings in operational energy to the additional requirements for embodied energy, in terms of the energy payback period and energy return on investment. The effectiveness of double glazing is determined for an Australian residential building. A wide range of building operation regimes was simulated. These results support the principle of installing double glazing in residential buildings in Melbourne, Australia, at least in terms of net primary energy savings.

Colonisation by epibionts and meiofauna of real and mimic pneumatophores in a cool temperate mangrove habitat

The size and pace of change in meiofaunal assemblages suggest that meiofauna make excellent subjects for testing theories about how ecological communities change. A field experiment was performed in which the  abundance and composition of epibionts and meiofauna on natural,  transplanted and mimic pneumatophores were monitored over a 47 wk period. Meiofaunal density increased with growth of algal epibionts, both reaching maximum values after 24 wk, at the end of winter. At this time the assemblages from the 3 substrata were similar, although the combined abundances of meiofauna on transplants and mimics were only 28% of the average on natural pneumatophores. Meiofaunal abundance on all substrata decreased rapidly during spring as algal cover declined due to desiccation. Twenty-three species of nematode were recorded from mimics compared with 8 and 7 from transplants and pneumatophores, respectively. A temporal sequence of feeding groups occurred in the order of epigrowth feeders, deposit feeders, and omnivore/predators, with the latter 2 adding to rather than replacing earlier trophic groups. Scavengers were found only on natural pneumatophores. The turnover rates of nematode species between all census times were similar, peaking at 63%, but there was no trend in the turnover rates with time. We conclude that mimics are more suitable than transplanted pneumatophores for colonisation studies because of their greater persistence and more easily standardised surface area. Moreover, the composition of colonising assemblages on them closely resembled assemblages on natural pneumatophores at the time of peak meiofaunal abundance.

Nematode assemblages from Avicennia marina leaf litter in a temperate mangrove forest in south-eastern Australia

Meiofauna from Avicennia marina leaf litter in a temperate mangrove forest was enumerated, and the nematode assemblages compared on the bases of leaf colour (used as a guide to leaf age) and shore horizon where samples were collected. Twenty-one putative nematode species were collected from 48 leaf litter samples. Univariate analyses indicated that neither the colour of the leaf nor the shore horizon significantly affected abundance of nematodes. However, of the four (222) treatment groups, rarefaction curves revealed highest diversity on brown leaves from under the shade of the tree canopy (H'=0.751-0.126 SE, n=17). Species diversity of leaf litter nematodes was lower in this temperate mangrove system than reported from tropical mangrove studies. ANOSIM tests confirmed a significant effect of shore horizon on nematode assemblages. The dominant feeding group among nematodes was non-selective deposit feeders (7/21 species, but 77% of all nematodes). Epigrowth grazers were represented by 8/21 species of nematodes, but only 19% of the total number. Excised leaves became skeletonised by about 15 weeks. Shorter temporal scales of life cycles of nematodes compared with leaf degradation, and the dynamic nature of epibiontic assemblages, probably explain the similar assemblage structure on yellow and brown leaves.

The effects of land use on leaf-litter processing by macro-invertebrates in an Australian temperate coastal stream

Replacement of riparian vegetation by pasture has occurred worldwide and is predicted to have strong effects on macroinvertebrate community structure and function in streams, but this has rarely been examined. In this study, leaf processing and macroinvertebrate community structure were examined in a single stream using experimental leaf-packs and surveys of natural leaf-packs. Two sites in each of three land use categories were selected to represent reaches in forest, pasture and forest-pasture boundary regions. In two experiments using tethered leaf-packs, no differences were found in mean leaf breakdown between land use types. However, shredding invertebrates were absent from the pasture sites, so leaf breakdown in pasture resulted from chemical, physical and microbial processes only. Amounts of fine particulate organic matter in experimental leaf-packs were higher in pasture reaches than the forest and boundary reaches but did not influence leaf breakdown. Macroinvertebrate species richness did not differ between land uses. A predictive model developed for species richness and total abundance enabled direct comparison of assemblages on experimental packs to natural leaf-packs. In the forest reach and at the forest-pasture boundary, macroinvertebrate species richness and total abundance increased proportionally with the number of leaves within a pack, but this relationship was not observed in the pasture reach. Pasture land use on Skenes Creek was therefore associated with weakened relationships between allochthonous inputs and macroinvertebrate communities, but this did not alter leaf breakdown.

Meiofaunal recruitment to mimic pneumatophores in a cool-temperate mangrove forest: spatial context and biofilm effects

A field experiment was devised to test whether meiofauna that colonised mimic pneumatophores (artificial substrates) resembled the assemblage on adjacent live pneumatophores in three randomly chosen intertidal, estuarine sites. The experiment showed that the close proximity of particular biota on living pneumatophores did not reliably influence subsequent development of assemblages upon mimic pneumatophores within a scale of 10 m during a colonisation period of less than 20 weeks. There was some convergence of the composition of the colonising assemblage of meiofauna on mimic pneumatophores with the local assemblages in sites dominated by barnacles, or where the natural pneumatophores were free from macroscopic epibionts. However, tychopelagic meiofauna from algal epiphytes did not significantly colonise mimic pneumatophores during the 20-week trial, probably due a lack of growing algae. During the conditioning phase suspended in water at a marine site 20 km from the mangroves, mimic pneumatophores acquired an assemblage of meiofauna different from the estuarine assemblage that colonised mimics following implantation in the estuarine mudflat. Enhanced colonisation rates of mimics in suspended bags at the conditioning site may be explained by the absence of benthic macroinvertebrates, and the lack of intertidal exposure. Biofilms aged 2, 7, and 11 weeks had no consistent, different effect on the subsequent colonisation of meiofauna. We conclude that divergence of phytal-based assemblages of meiofauna depends upon the amount of coverage, as well as the type, of fouling macro-epibionts on the pneumatophores. Meiofaunal assemblages on artificial substrates after 20 weeks colonisation displayed less intrinsic patchiness than mature phytal assemblages on natural pneumatophores, and so present a potentially useful way of improving the power of biomonitoring applications using meiofauna.

How important are different types of temperate woodlands for ground-foraging birds?

There is widespread concern about population decline in a number of woodland-dependent birds in southern Australia. Of all declining species, approximately half forage on the ground. This study examined the avifaunal assemblages of temperate woodlands of the Northern Plains, Victoria, to investigate the importance of woodland habitats for ground-foraging species. Four main types of woodland were surveyed (white cypress-pine, black box, grey box and river red gum) and, in total, 89 bird species were detected. All four woodland types differed in habitat structure and, in turn, supported significantly different avifaunal assemblages. Forty of the 89 species (45%) foraged, at least in part, on the ground. Species richness and abundance of ground-foragers differed significantly between woodland types, being highest in white cypress-pine and black box. There was a greater richness of ground-foragers during the breeding than non-breeding season, but abundance did not vary seasonally. Overall, ground-foraging birds comprised a greater proportion of species (>55%) and individuals (>60%) in white cypress-pine and black box woodland than in grey box and river red gum (42–48% of species, <50% individuals). Those ground-foragers regarded as declining also occurred in greatest richness in white cypress-pine woodlands, one of the most depleted habitats in the region. The lowest richness of ‘declining’ ground-foraging species was in river red gum woodland, the most widespread woodland type. Throughout Australia, the proportion of ground-foraging species in bird assemblages tends to be greater in temperate, semi-arid or arid woodlands than in moist forests and rainforests. However, in many regions woodland habitats are severely depleted and their open ground layer is particularly vulnerable to degradation. The extent of suitable habitat for ground-foraging birds in temperate woodlands may be much less than is apparent from current measures of tree cover. Sustainable management of drier (non-riverine) temperate woodlands is required to conserve this important element of the Australian avifauna.