Spatio‐temporal variation in the effect of herbaceous layer on woody seedling survival in a Chilean mediterranean ecosystem

Questions: What is the effect of herbaceous layer on seedling establishment of three woody pioneer species in open areas of central Chile under a semi‐arid mediterranean climate? How do inter‐annual and habitat conditions (slope aspect) modulate this effect? Under high stress conditions such as the drier year and habitat (north‐facing slope) do herbs reach low abundance and have neutral effects on woody seedlings? Under medium stress conditions for these woody species, such as the wetter year and south‐facing slope, does the herbaceous layer reach greater abundance and have positive effects on woody seedlings due to increasing soil water content? Location: A watershed on the outskirts of Santiago, Chile, subjected to clearing of woody vegetation through firewood extraction and human‐set fires. Methods: In spring 2007, we set up 20 plots (3 m × 2 m). Half of each plot had herbs removed manually and by application of herbicide. In both halves of each plot, one seedling (8 months old) of each of the three native woody species (Colliguaya odorifera, Schinus polygamus and Quillaja saponaria) was planted and survival monitored subsequently. The experiment was repeated in two consecutive growing seasons (2007–2008 and 2008–2009) that differed significantly in total precipitation (152 and 256.5 mm, respectively), and replicated in two sites that differed in aspect and abiotic conditions: a moister south‐ and a drier north‐facing slope. Results: In the first and drier year, the herbaceous layer had low cover and no significant effect on seedling survival of woody species. During the second year, herbs had greater cover and a significant positive effect on spring survival of C. odorifera in the north‐facing slope, which was lost after summer. During this wetter year on the south‐facing slope, herb cover had a positive effect on survival of S. polygamus (mainly during summer). Conclusions: The role of mostly ruderal herbs on woody seedling establishment depended on the species, rainfall of the current year and slope aspect, and may be explained by soil moisture patterns. This suggests that the effect of ruderal herbs on woody seedlings shifts from neutral under high stress conditions produced by drought to positive under moderate stress conditions. Our results contribute to understand interactions between ruderal herbs and woody species under contrasting abiotic conditions. Therefore, control of the herbaceous layer may not be needed in restoration programmes for this region. Moreover, herbs may benefit restoration of woody cover in mesic habitats.     

Effects of leaf litter and precipitation on germination and seedling survival of the endangered tree Beilschmiedia miersii

Question: What effects do leaf litter and rainfall regime have on seed germination (time and probability) and seedling survival of the endangered tree species Beilschmiedia miersii (Lauraceae)? Location: The species is a native tree from the mediterranean climate region of Chile. Seeds were collected from La Campana National Park (Chile). The study was carried out under controlled conditions at the Laboratory of Ecology, University of Chile, Santiago, Chile. Methods: During April 2001,200 seeds were assigned to four experimental treatments: high precipitation, with and without litter and low precipitation, with and without litter. Each treatment had 50 individual seeds, each seed in an individual pot. For statistical purposes, we considered each seed as one replicate. High and low values of artificial rainfall corresponded to mean dry and wet years, respectively, for the period 1958–1993 in the central zone of Chile. Results: Seeds germinated earlier, and in higher proportion, in the presence of leaf litter, but only under low rainfall. Seedling survival was insensitive to both litter and precipitation. Conclusions: We conclude that the presence of litter in native populations facilitates seed germination and recruitment of B. miersii, particularly during dry years. We suggest that the reduction of leaf litter due to extraction for gardens and horticultural activities might preclude regeneration of this endangered species.     

Impacts of simulated livestock grazing on Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem

Allometric foraging theory suggests that herbivores of greatly differing size should co-exist through niche segregation, but a few studies of large–small herbivore foraging relationships have reported competitive interactions. This study addresses the potential roles of habitat productivity and large herbivore grazing intensities on large–small herbivore foraging interactions. We examined effects of different intensity simulated grazing treatments on forage abundance and quality for Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem, and consequent effects on prairie dog individual growth rates, foraging preferences, and activity budgets. We hypothesized that simulated grazing would have predominantly facilitative impacts on Utah prairie dogs, as was found for black-tailed prairie dogs in higher productivity ecosystems. To test this hypothesis, we measured the effects of simulated grazing on forage nitrogen, digestibility, and biomass. Simulated grazing increased average forage nitrogen and digestibility while decreasing forage biomass. These effects were associated with reduced individual growth rates, increased juvenile foraging time, and reduced juvenile vigilance. Results suggest that the negative effects of reduced vegetation biomass greatly outweighed positive treatment effects in this study. However, prairie dogs in the moderate intensity defoliation treatment showed some preference for “grazed” plots over “ungrazed” plots, and this preference increased with time. Our study lends support to the idea that habitat productivity and herbivore densities may mediate shifts between facilitative and competitive interactions between different-sized herbivores.     

Facilitative effect of Lotustenuis on Paspalumdilatatum in a lowland grassland of Argentina

We studied the responses in growth and N content of the perennial grass Paspalum dilatatum to the substitution of Lotus tenuis for a whole group of species, the dicotyledons of a natural grassland community, in the Salado lowland Pampas of Argentina. Two kinds of manipulations were performed in the field: removal of alien dicots with herbicide application, and introduction of L. tenuis, resulting in a combination of four treatments, arranged in a 2 × 2 factorial randomized block design. Leaf area per tiller of P. dilatatum was higher when it was growing near L. tenuis; this increase was the result of a greater leaf elongation rate and slower leaf senescence. In the vicinity of L. tenuis, P. dilatatum exhibited an increase in tiller production and a decrease in tiller death. More tillers were functional at the end of the growing season and their aboveground biomass was 5␣times higher than for plants growing in plots where the community dicots were removed. This increment was accompanied by a higher N content. Growth enhancement of P. dilatatum plants when L. tenuis was the␣immediate neighbour is interpreted as the result of facilitation mediated by higher N availability, and not as a consequence of a release from competition exerted by the community dicots. Competition and facilitation did not interact to produce an increase in the vegetative output of Paspalum dilatatum plants growing under these field conditions. It is on these grounds that Lotus tenuis might be considered as a keystone species in the managed grassland. Received: 5 July 1997 / Accepted: 12 December 1997     

Influence of tree cover on the diversity of herbaceous communities in subalpine wooded pastures

Abstract. The relationships between several diversity indices and tree cover in subalpine wooded pastures with Larix de-cidua were analysed at two spatial scales: phytocoenoses (2500-m² plots) and herbaceous synusia (1-m² quadrats). Diversity indices have a non-linear relationship to Larix cover; the best fit was obtained with Gaussian regressions. Species richness was influenced by Larix cover more than evenness. At the phytocoenosis level, the optimum for species and synusial richness in the herb layer was near 25 % tree cover. At the synusia level, species richness of the herb layer showed an optimum when mean distance to the four nearest Larix trees was ca. 30 m, or when the potential number of sunshine hours between April and September was close to 1250 h. Canonical Correspondence Analysis confirmed the influence of Larix cover on the composition of phytocoenoses and herbaceous synusiae, but included also the role of altitude, slope and aspect.     

用光合-蒸散耦合模型模拟冬小麦水热通量的日变化

从SPAC理论出发,建立了一个冬小麦光合和蒸散的耦合模型．感热通量和潜热通量采用Shuttleworth-Wallace的双层模型计算,并通过冠层阻力的参数化,将光合作用与蒸腾作用耦合起来．用涡度相关方法,观测了感热通量和潜热通量,对模型进行了验证．结果表明,模拟值与观测值比较一致,模型可以很好地模拟感热通量和潜热通量的日变化过程．对模型的敏感性分析发现,冬小麦蒸腾比较敏感的参数有凋萎点、气孔导度参数、叶对红外辐射的反射率和光响应曲线凸度;土壤蒸发只对土壤阻力参数的敏感性较强．本模型对水热通量与环境因子作用过程的理论研究和指导农田的灌溉制度等有一定的意义．     

Influence of an overstorey tree (Prosopis glandulosa) on associated shrubs in a savanna parkland: implications for patch dynamics

The arborescent legume, honey mesquite (Prosopis glandulosa), appears to play a central role in patch dynamics of southern Texas savannas by modifying soils and microclimate and by facilitating the ingress, establishment and/or growth of shrubs in its understorey. As an indirect test for the occurrence and persistence of facilitation in mature shrub clusters (patches), we examined the gas exchange, water relations and production of associated shrubs growing in patches where a Prosopis overstorey was present and in patches where Prosopis had succumbed to natural mortality. Surface (0–10 cm) soils associated with shrub patches were enriched in total [N] and [C] compared to soils of neighboring herbaceous zones. However, there were no detectable differences in soil [N] or [C] in patches with and without Prosopis. Foliar [N] and biomass of various shrub species were also statistically comparable for patches with and without Prosopis. These results are in accordance with other studies that indicate the nutrient legacy associated with Prosopis occupation of a patch may persist for decades after its demise. In comparison to plants growing in the absence of Prosopis, leaf water potentials (predawn and midday), and net photosynthesis and water vapor conductance (morning and midday) of outer-canopy sunlit leaves over an annual growth cycle were comparable for two common evergreen shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing in patches with a live Prosopis. These findings indicate that the presence of Prosopis was not enhancing the growth or activity of mature understorey shrubs; facilitation may, therefore, be important only during early stages of cluster development. In addition, we found no indication that the loss of Prosopis has initiated a downward phase in a cyclic succession of patch initiation, growth and death. Rather, the understorey shrubs appear to be able to maintain growth and productivity in the absence of a Prosopis overstorey, and may, therefore, represent persistent components of woody patches on these savanna landscapes.     

Development of an herbaceous cover in a Sudano-Sahelian savanna in North Cameroon in relation to available soil water

Dynamics of annual communities are studied in Sudano-Sahelian savannas of the north of the Cameroon. The vegetation composition of three types of soil (a vertisol, a degraded vertisol and a ferruginous soil) under two rainfall conditions is compared. The study supports the idea that intense fluctuations in water availability at the beginning of the rainy season determine which species germinate, grow and survive under conditions of water stress. However soil conditions can accentuate or reduce the influence of the rainfall fluctuations while the land use history of each station tends to buffer the variations of the overall pattern determining potential vegetation.     

Convergence patterns and multiple species interactions in a designed plant mixture of five species

It is known that convergence and divergence can occur in complex plant communities, but the relative importance of biotic and abiotic factors driving these processes is less clear. We addressed this issue in an experiment using a range of mixed stands of five species that are common in Swiss fens (Carex elata, C. flava, Lycopus europaeus, Lysimachia vulgaris and Mentha aquatica) and two levels of water and nutrients. One hundred and seventy-six experimental mixtures were maintained in large pots (75 l) for two consecutive growing seasons in an experimental garden. The stands varied systematically in the initial relative abundance of each of the five species and in overall initial stand abundance. The changes in biomass over 2 years were modelled as linear functions of treatments and the initial biomass of each species. The dynamics of the system were mainly driven by differences in the identity of species and by a negative feedback mechanism but also by different abiotic conditions. In all mixtures, C. elata became more dominant over time, which caused an overall convergence of community composition. In addition, the rate of change of each species’ biomass was negatively related to its own initial abundance. Thus, a negative feedback further contributed to the convergence of communities. Species responded differently to water level and nutrient supply, causing community dynamics to differ among treatments. However, the different abiotic conditions only slightly modified the overall convergence pattern. Competitive interactions between more than two species were weaker than the negative feedback but still significantly influenced the species’ final relative abundance. The negative feedback suggests that there is niche partitioning between the species, which permits their coexistence. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Influence of management and precipitation on carbon fluxes in great plains grasslands

Suitable management and sufficient precipitation on grasslands can provide carbon sinks. The net carbon accumulation of a site from the atmosphere, modeled as the Net Ecosystem Productivity (NEP), is a useful means to gauge carbon balance. Previous research has developed methods to integrate flux tower data with satellite biophysical datasets to estimate NEP across large regions. A related method uses the Ecosystem Performance Anomaly (EPA) as a satellite-derived indicator of disturbance intensity (e.g., livestock stocking rate, fire, and insect damage). To better understand the interactions among management, climate, and carbon dynamics, we evaluated the relationship between EPA and NEP data at the 250 m scale for grasslands in the Central Great Plains, USA (ranging from semi-arid to mesic). We also used weekly estimates of NEP to evaluate the phenology of carbon dynamics, classified by EPA (i.e., by level of disturbance impact). Results show that the cumulative carbon balance over these grasslands from 2000 to 2008 was a weak net sink of 13.7 g C m⁻² yr⁻¹. Overall, NEP increased with precipitation (R² = 0.39, P < 0.05) from west to east. Disturbance influenced NEP phenology; however, climate and biophysical conditions were usually more important. The NEP response to disturbance varies by ecoregion, and more generally by grassland type, where the shortgrass prairie NEP is most sensitive to disturbance, the mixed-grass prairie displays a moderate response, and tallgrass prairie is the least impacted by disturbance (as measured by EPA). Sustainable management practices in the tallgrass and mixed-grass prairie may potentially induce a period of average net carbon sink until a new equilibrium soil organic carbon is achieved. In the shortgrass prairie, management should be considered sustainable if carbon stocks are simply maintained. The consideration of site carbon balance adds to the already difficult task of managing grasslands appropriately to site conditions. Results clarify the seasonal and interannual dynamics of NEP, specifically the influence of disturbance and moisture availability.     

The influence of water level and salinity on plant assemblages of a seasonally flooded Mediterranean wetland

We studied the key environmental variables shaping plant assemblages in Mediterranean abandoned ricefields with contrasting freshwater inputs over saline sediments. Plant species cover, water levels and soil variables were studied following a stratified random sampling design. Multivariate analysis identified water regime, particularly summer and autumn irrigation, as the most important environmental variable associated with vegetation composition. Distribution of annual and emergent macrophytes was not associated to salinity as found at the study site (0.57–4.1 mS/cm). Increased soil salinity, caused by summer irrigation near the soil surface did affect shallow-marsh assemblage distribution. These key environmental characteristics allowed us to identify six main assemblages. Annual macrophytes (such as Zannichellia palustris) were defined by high (over 10 cm) annual mean water level (MWL) and early successional conditions; emergent macrophytes (such as Typha spp., Scirpus lacustris) by annual MWL of 10 to − 25 cm and continuous shallow flooding in summer and autumn (MWL of 0–10 cm). The shallow-marsh group, correlated with annual MWL − 25 to − 100 cm, separated into two subgroups by salinity: grassland (including Paspalum distichum) with summer and autumn MWL below − 25 cm and brackish (with Juncus subulatus or Agrostis stolonifera) with summer and autumn MWL just below the soil surface (0 to − 25 cm). Water levels for the grassland subgroup may equate with a salinity ‘refuge’ for P. distichum. Time was a further determinant of variation in the full data set. Abundance of a large group of agricultural annuals (such as Sonchus tenerrimus) and damp ground annuals (including ricefield weeds such as Ammania robusta) decreased with time as bare ground disappeared. Maintenance of spatial vegetation heterogeneity in abandoned ricefields is contingent on continued water regime management.     

Drought susceptibility in emergent wetland angiosperms: a comparison of water deficit growth in five herbaceous perennials

Freshwater wetlands often exist as transitional areas between terrestrial uplands and deep open water. Thus they are fundamentally sensitive to changes in hydrology. Some of the more dramatic changes in wetland water supply occur during extensive droughts, where both precipitation and soil water table markedly decline. While it is generally understood that herbaceous wetland macrophytes are more sensitive to decreased water availability than wetland trees, the degree of susceptibility among wetland herbs remains relatively unexplored. Therefore, the purpose of this study was to evaluate plant growth responses of five herbaceous wetland species (monocots Carex alata, Juncus effusus, and Peltandra virginica, and dicots Saururus cernuus, and Justicia americana) to simulated drought conditions (up to 6 weeks in a 1-in-25-year precipitation low with receding soil water tables). Of the five species studied, three (J. americana, S. cernuus, and J. effusus) had no survivors after 6 weeks of simulated drought. J. americana, appeared to be the most sensitive to water deprivation with a 67% decrease in plant phytomass and an 85% decrease in leaf area with only 2 weeks of drought, and complete mortality after 3 weeks. While P. virginica also had significant decreases in biomass, leaf area, relative growth rate (RGR) and unit leaf rate (ULR), in as little as 2 weeks of drought, no noticeable decreases in survival were observed. In contrast, when J. effusus experienced between 2- and 4-weeks of water deprivation, there were significant increases in RGR, ULR, phytomass, leaf area, and shoot:root ratios. S. cernuus and C. alata remained relatively unaffected following 4 weeks of drought; however by the fifth week, there were significant declines in leaf area for both species. In general, this study provides experimental evidence on how herbaceous macrophytes grow under drought conditions. This basic understanding is fundamental if we are to develop better working models on how wetlands will respond to changing environmental conditions that lead to decreased water supply.     

Phenological interactions between tree and understory herbaceous vegetation of a sahelian semi-arid savanna

This study was conducted during the 1990 planting cycle and focused on interactions between trees and the herbaceous layer caused by their phenological behaviour in a Sahelian semi-arid savanna.The results show that (1) the plant phenological development (tree and grassy group) was synchronous; (2) the tree phenological behaviours were most clearly differentiated because reproduction, flowering and fructification phases were spread out throughout the year and (3) the herbaceous species cycle was then essentially determined by the short biologically damp period, 60 days in 1990. The foliage stage of the ligneous species spread on the considered period, with a reduced realization rate (30 to 40%) due to an invasion of locusts.At the grassy group level, the considerable growth rate in every development stage indicated that there was apparently no competition for soil water. Both trees and grass used the available soil water with grass only exploiting the upper soil layer.Mots-clés: Arbre, Compétition, Couvert, Sahel, Stades phénologiques, Strate herbacéeRésumé Cette étude se propose d'établir les interactions Herbe/Arbre à travers les stades phénologiques des principales espèces ligneuses et de la strate herbacée établie sous l'arbre à partir d'observations menées au cours du cycle 1990.Les résultats montrent une superposition des cycles phénologiques du peuplement ligneux et du groupement herbeux. La phase de feuillaison des espèces ligneuses étudiées s'étale sur toute la période considérée. Le taux d'accomplissement des différentes phénophases est généralement faible (30 à 40%) en raison de l'invasion par des criquets.Le taux d'accomplissement des phénophases au niveau du groupement herbeux indique qu'il n'y a apparemment pas de compétition pour l'eau; le partage de l'eau du sol entre l'herbe et l'arbre se faisant principalement dans l'espace.     

Influence of seed size on performance of non-native annual plant species in a novel community at two planting densities

Climate warming enables plant species to migrate to higher latitudes and altitudes. Within Europe, the Mediterranean harbours many species that might expand their ranges towards Western Europe. Small seed size may facilitate dispersal, however, it may impair establishment of the range-expanding plant species in the novel vegetation. In a greenhouse experiment, we examined effects of average seed size of Mediterranean plant species on their establishment in a mixed community of Western European plant species. Applying two levels of densities of the natives and a herbivory treatment, we tested how seed size is linked to response in plant growth and fitness in novel vegetation. While all non-native plant species showed a negative response to increased planting density, species with small seeds showed a less negative response. This effect persisted under herbivory. Our data suggest that small-seeded non-native plant species may tolerate competitive pressure from novel plant communities better than large-seeded species, so that small seed size may confer a higher probability of establishment of non-native species in novel communities.     

Effects of intraspecific and community density on the lifetime fecundity of long-lived shrubs

Intra- and interspecific density dependence has profound consequences for plant population and community dynamics. In long-lived plants, however, lifetime patterns and mechanisms of density dependence are difficult to study. Here, we examine effects of intraspecific and community density on the lifetime fecundity of two long-lived shrub species from South African Fynbos: Protea repens (animal-pollinated, hermaphroditic) and Leucadendron rubrum (wind-pollinated, dioecious). Both species are serotinous, retaining seeds in cones until fire kills the mother plant. We measured lifetime fecundity as the product of cone number, proportion of cones that are not damaged by predation and seed set (fertile seeds per intact cone). Intraspecific and community densities were quantified by counting individuals of target species and all Proteaceae in small- and large-scale neighbourhoods (10 m and 50 m radius) around each focal individual. Additionally, we determined the age and size of focal individuals. We found that lifetime fecundity of the wind-pollinated L. rubrum is density independent. In contrast, the lifetime fecundity of the animal-pollinated P. repens increases with large-scale intraspecific density and shows a hump-shaped relationship to large-scale community density. Community density has a hump-shaped effect on seed set (probably through partial absence of generalized pollinators at low and competition for pollinators at high densities) and negatively affects cone number per individual. For both species, plant age decreases seed set while increasing lifetime fecundity. The qualitative differences in the density dependence of lifetime fecundity may arise from differences between animal and wind pollination. In particular, interactions with generalized animal pollinators may cause community-level Allee effects with profound consequences for the future dynamics of long-lived plant populations and communities.     

Spatial heterogeneity in the herbaceous layer of a semi-arid savanna ecosystem

Despite increasing recognition of the role spatial pattern can play in ecosystem function, few studies have quantified spatial heterogeneity in savanna ecosystems. The spatial distribution of herbaceous biomass and species composition was measured across three scales in a semi-arid savanna in central Kenya, and patterns were related to environmental variables at different scales. Herbaceous biomass declined across a rainfall gradient and from upper to lower topographic positions, but variation within a site (across 5–50 m) was similar in magnitude to among-site variation associated with rainfall and topography. Geostatistical analyses showed that patchiness at scales of 5–25 m explained 20% of total variation in herbaceous biomass. This pattern arose from the presence of both 5–10-m diameter patches containing high herbaceous biomass (> 170 g m^–2) and 5–10-m diameter patches characterized by nearly bare soil surfaces (< 40 g m^–2). Patch structure was contingent on topography, with larger bare patches at ridgeline and upper hillslope positions. Grass species distributions showed the greatest degree of patch structure and species turnover across distances of 5–45 m. Additional community variation was associated with topography, with minimal variation in species composition across the rainfall gradient. Pattern diversity significantly exceeded levels reported for four other grassland ecosystems, suggesting fundamental differences in local processes generating spatial pattern. It is hypothesized that heterogeneously distributed grazing pressure, interacting with the distribution of shrub canopies, is an important factor generating such high levels of small-scale patch structure in this savanna.     

Sources and sinks of aquatic carbon in a peatland stream continuum

Streams draining peatland systems contain a number of different C-species, all of which are linked either directly or indirectly to the cycling of C in the terrestrial environment. Concentrations and fluxes of dissolved, particulate and gaseous forms of carbon were measured along a network of streams draining an acidic peatland catchment (46.3km2) in NE Scotland. The main aim was to identify sources and sinks of all the major forms of C in the drainage network and use this to develop a conceptual understanding of the evolution of streamwater chemistry along a peatland stream continuum. The investigation included a small-scale intra-catchment study of three contiguous sites in a 1.3km2 headwater catchment (Brocky Burn) and a larger scale integrated study of seven sites. Mean annual fluxes of the main carbon species varied from 115–215 (DOC), 8.15–97.0 (POC), 0.32–6.90 (HCO3--C) and 2.62–10.4 (free CO2-C)kgCha–1year–1; all contributed to the overall carbon flux to varying degrees. Methane-C was only measurable at sites within areas of deep peat (<0.01–0.09kgCha–1year–1). Downstream spatial changes in the intra-catchment study (Brocky Burn) were characterised by a decrease in DOC, CO2-C and CH4-C and an increase in POC fluxes over a distance of 1.1km from the Upper to the Lower sites. In the context of the integrated catchment study estimated losses and gains of carbon from the water column showed no net change in DOC, a large decrease in POC (–55%) and a slight increase in (HCO3--C) (+7.7%) and CO2-C (+4.5%). A significant decrease in the CO2-C flux: HCO3-C flux ratio with distance downstream from the stream source, illustrates the importance of outgassing of CO2 from streams draining peatland C reservoirs. These data are interpreted in the context of losses and gains of the various components of the aquatic C flux along the peatland stream continuum.     

Comparing and evaluating process-based ecosystem model predictions of carbon and water fluxes in major European forest biomes

Process‐based models can be classified into: (a) terrestrial biogeochemical models (TBMs), which simulate fluxes of carbon, water and nitrogen coupled within terrestrial ecosystems, and (b) dynamic global vegetation models (DGVMs), which further couple these processes interactively with changes in slow ecosystem processes depending on resource competition, establishment, growth and mortality of different vegetation types. In this study, four models – RHESSys, GOTILWA+, LPJ‐GUESS and ORCHIDEE – representing both modelling approaches were compared and evaluated against benchmarks provided by eddy‐covariance measurements of carbon and water fluxes at 15 forest sites within the EUROFLUX project. Overall, model‐measurement agreement varied greatly among sites. Both modelling approaches have somewhat different strengths, but there was no model among those tested that universally performed well on the two variables evaluated. Small biases and errors suggest that ORCHIDEE and GOTILWA+ performed better in simulating carbon fluxes while LPJ‐GUESS and RHESSys did a better job in simulating water fluxes. In general, the models can be considered as useful tools for studies of climate change impacts on carbon and water cycling in forests. However, the various sources of variation among models simulations and between models simulations and observed data described in this study place some constraints on the results and to some extent reduce their reliability. For example, at most sites in the Mediterranean region all models generally performed poorly most likely because of problems in the representation of water stress effects on both carbon uptake by photosynthesis and carbon release by heterotrophic respiration (R_h). The use of flux data as a means of assessing key processes in models of this type is an important approach to improving model performance. Our results show that the models have value but that further model development is necessary with regard to the representation of the some of the key ecosystem processes.