Do non-native Platanus hybrida riparian plantations affect leaf litter decomposition in streams?

In forest headwater streams where the riparian canopy limits autochthonous primary production, leaf litter decomposition is a key process controlling nutrient and carbon cycling. Any alteration of the riparian vegetation may influence litter decomposition and detrital food webs. We evaluated the effect of non-native Platanus hybrida riparian plantations on leaf litter decomposition in Mediterranean streams. The experiment was conducted in six headwater streams; three lined by native riparian vegetation and three crossing P. hybrida plantations. We have characterized the processing rates of alder leaves and the assemblages of shredder macroinvertebrates and fungi. Litter decomposition was significantly faster in the P. hybrida than in the reference streams. Although the dissolved inorganic nitrogen concentration was higher in P. hybrida, no significant effect was observed in decomposition rates. Differences in decomposition rates reflected the macroinvertebrate and shredder colonization in alder litter, with higher abundance and richness in the P. hybrida streams. However, aquatic hyphomycete sporulation rate was higher in reference streams, suggesting that the variation in decomposition rates is a direct consequence of shredder abundance. Our findings support part of the substrate quality-matrix quality (SMI) hypothesis, which expects that high-quality litter will show increased decomposition rates in a low-quality litter matrix.     

Higher temperature reduces the effects of litter quality on decomposition by aquatic fungi

1. We investigated the effects of riparian plant diversity (species number and identity) and temperature on microbially mediated leaf decomposition by assessing fungal biodiversity, fungal reproduction and leaf mass loss. 2. Leaves of five riparian plant species were first immersed in a stream to allow microbial colonisation and were then exposed, alone or in all possible combinations, at 16 or 24 °C in laboratory microcosms. 3. Fungal biodiversity was reduced by temperature but was not affected by litter diversity. Temperature altered fungal community composition with species of warmer climate, such as Lunulospora curvula, becoming dominant. 4. Fungal reproduction was affected by litter diversity, but not by temperature. Fungal reproduction in leaf mixtures did not differ or was lower than that expected from the weighted sum of fungal sporulation on individual leaf species. At the higher temperature, the negative effect of litter diversity on fungal reproduction decreased with the number of leaf species. 5. Leaf mass loss was affected by the identity of leaf mixtures (i.e. litter quality), but not by leaf species number. This was mainly explained by the negative correlation between leaf decomposition and initial lignin concentration of leaves. 6. At 24 °C, the negative effects of lignin on microbially mediated leaf decomposition diminished, suggesting that higher temperatures may weaken the effects of litter quality on plant litter decomposition in streams. 7. The reduction in the negative effects of lignin at the higher temperature resulted in an increased microbially mediated litter decomposition, which may favour invertebrate‐mediated litter decomposition leading to a depletion of litter stocks in streams.     

Growth of an invertebrate shredder on native (Populus) and non-native (Tamarix, Elaeagnus) leaf litter

1. Large-scale invasions of riparian trees can alter the quantity and quality of allochthonous inputs of leaf litter to streams and thus have the potential to alter stream organic matter dynamics. Non-native saltcedar ( Tamarix sp.) and Russian olive ( Elaeagnus angustifolia ) are now among the most common trees in riparian zones in western North America, yet their impacts on energy flow in streams are virtually unknown.
2. We conducted a laboratory feeding experiment to compare the growth of the aquatic crane fly Tipula (Diptera: Tipulidae) on leaf litter from native cottonwood ( Populus ) and non-native Tamarix and Elaeagnus . Tipula showed positive growth on leaf litter of all three species; however, after 7 weeks, larvae fed Tamarix leaves averaged 1.7 and 2.5 times the mass of those fed Elaeagnus and Populus , respectively. Tipula survival was highest on Populus , intermediate on Tamarix and lowest on Elaeagnus .
3. High Tipula growth on Tamarix probably reflects a combination of leaf chemistry and morphology. Conditioned Tamarix leaf litter had intermediate carbon : nitrogen values (33 : 1) compared to Populus (40 : 1) and Elaeagnus (26 : 1), and it had intermediate proportions of structural carbon (42%) compared to Elaeagnus (57%) and Populus (35%). Tamarix leaves are also relatively small and possibly more easily ingested by Tipula than either Elaeagnus or Populus .
4. Field surveys of streams in the western U.S.A. revealed that Tamarix and Elaeagnus leaf packs were rare compared to native Populus , probably due to the elongate shape and small size of the non-native leaves. Thus we conclude that, in general, the impact of non-native riparian invasion on aquatic shredders will depend not only on leaf decomposition rate and palatability but also on rates of leaf litter input to the stream coupled with streambed retention and subsequent availability to consumers.     

Boreal peat properties link to plant functional traits of ecosystem engineers

Aims

Warming has the potential to alter plant litter mass loss and nutrient release during decomposition. However, a great deal of uncertainty remains concerning how other factors such as litter species or substrate quality might modify the effects of increased temperature on decomposition. Meanwhile, the temperature sensitivity of plant litter decay in tropical and subtropical forest ecosystems remains poorly resolved.

Methods

This study was designed to assess the effects of experimental warming on litter decomposition and nutrient release of two contrasting tree species (Schima superba and Machilus breviflora) by translocating model forest ecosystems from the high-elevation sites to the lower-elevation sites in subtropical China. Translocating model mountain evergreen broad-leaved forest (MEBF) to the altitude of 300 m and 30 m increased the average monthly soil temperature at 5 cm depth by 0.88 and 1.84 °C, respectively during the experimental period. Translocating model coniferous and broad-leaved mixed forest (CBMF) to the altitude of 30 m increased the average monthly soil temperature at 5 cm depth by 0.85 °C.

Results

We found that experimental warming accelerated litter decomposition in both model forest types, and the promoting efficiency was greater when the temperature increased. The litter with high quality (Schima superba) had stronger response to warming than low quality litter (Machilus breviflora). Warming accelerated Na, K, Mg, P, N and Ca release from Schima superba litter, but only simulated Ca release from Machilus breviflora litter. Overall, litter decomposition was controlled by the order: soil temperature > litter quality > soil moisture > litter incubation forest type under experimental warming in the subtropical China.

Conclusion

We conclude that leaf litter decomposition was facilitated by experimental warming in subtropical China. Litter species might modify the effects of increased temperature on litter decomposition; however, forest type has no effect on litter decomposition.

     

Contrasting above‐ and belowground organic matter decomposition and carbon and nitrogen dynamics in response to warming in High Arctic tundra

Tundra regions are projected to warm rapidly during the coming decades. The tundra biome holds the largest terrestrial carbon pool, largely contained in frozen permafrost soils. With warming, these permafrost soils may thaw and become available for microbial decomposition, potentially providing a positive feedback to global warming. Warming may directly stimulate microbial metabolism but may also indirectly stimulate organic matter turnover through increased plant productivity by soil priming from root exudates and accelerated litter turnover rates. Here, we assess the impacts of experimental warming on turnover rates of leaf litter, active layer soil and thawed permafrost sediment in two high‐arctic tundra heath sites in NE‐Greenland, either dominated by evergreen or deciduous shrubs. We incubated shrub leaf litter on the surface of control and warmed plots for 1 and 2 years. Active layer soil was collected from the plots to assess the effects of 8 years of field warming on soil carbon stocks. Finally, we incubated open cores filled with newly thawed permafrost soil for 2 years in the active layer of the same plots. After field incubation, we measured basal respiration rates of recovered thawed permafrost cores in the lab. Warming significantly reduced litter mass loss by 26% after 1 year incubation, but differences in litter mass loss among treatments disappeared after 2 years incubation. Warming also reduced litter nitrogen mineralization and decreased the litter carbon to nitrogen ratio. Active layer soil carbon stocks were reduced 15% by warming, while soil dissolved nitrogen was reduced by half in warmed plots. Warming had a positive legacy effect on carbon turnover rates in thawed permafrost cores, with 10% higher respiration rates measured in cores from warmed plots. These results demonstrate that warming may have contrasting effects on above‐ and belowground tundra carbon turnover, possibly governed by microbial resource availability.     

Organic Matter Dynamics in a Tropical Gallery Forest in a Grassland Landscape

The high biodiversity of tropical forest streams depends on the strong input of organic matter, yet the leaf litter decomposition dynamics in these streams are not well understood. We assessed how seasonal litterfall affects leaf litter breakdown, density and biomass of aquatic invertebrates, and the microbial biomass and sporulation of aquatic hyphomycetes in a South American grassland ‘vereda’ landscape. Although litter production in the riparian area was low, leaf litter breakdown was high compared with other South American systems, with maximum values coinciding with the rainy season. Fungal biomass in decomposing leaves was high, but spore densities in water and sporulation rates were very low. Invertebrates were not abundant in litter bags, suggesting they play a minor role in leaf litter decomposition. Chironomids accounted for ~70 percent of all invertebrates; only 10 percent of non‐Chironomidae invertebrates were shredders. Therefore, fungi appear to be the drivers of leaf litter decomposition. Our results show that despite low productivity and relatively fast litter decomposition, organic matter accumulated in the stream and riparian area. This pattern was attributed to the wet/dry cycles in which leaves falling in the flat riparian zone remain undecomposed (during the dry period) and are massively transported to the riverbed (rainy season).     

Riparian leaf litter decomposition on pond bottom after a retention on floating vegetation

Allochthonous (e.g., riparian) plant litter is among the organic matter resources that are important for wetland ecosystems. A compact canopy of free‐floating vegetation on the water surface may allow for riparian litter to remain on it for a period of time before sinking to the bottom. Thus, we hypothesized that canopy of free‐floating vegetation may slow decomposition processes in wetlands. To test the hypothesis that the retention of riparian leaf litter on the free‐floating vegetation in wetlands affects their subsequent decomposition on the bottom of wetlands, a 50‐day in situ decomposition experiment was performed in a wetland pond in subtropical China, in which litter bags of single species with fine (0.5 mm) or coarse (2.0 mm) mesh sizes were placed on free‐floating vegetation (dominated by Eichhornia crassipes, Lemna minor, and Salvinia molesta) for 25 days and then moved to the pond bottom for another 25 days or remained on the pond bottom for 50 days. The leaf litter was collected from three riparian species, that is, Cinnamomum camphora, Diospyros kaki, and Phyllostachys propinqua. The retention of riparian leaf litter on free‐floating vegetation had significant negative effect on the carbon loss, marginal negative effects on the mass loss, and no effect on the nitrogen loss from leaf litter, partially supporting the hypothesis. Similarly, the mass and carbon losses from leaf litter decomposing on the pond bottom for the first 25 days of the experiment were greater than those from the litter decomposing on free‐floating vegetation. Our results highlight that in wetlands, free‐floating vegetation could play a vital role in litter decomposition, which is linked to the regulation of nutrient cycling in ecosystems.     

Effects of increased temperature and aquatic fungal diversity on litter decomposition

Climate warming and biodiversity loss are two major factors threatening freshwaters. Aquatic hyphomycetes are fungi that play a key role in organic matter turnover in streams. To assess the impacts of temperature increase and aquatic hyphomycete diversity on plant-litter decomposition, we manipulated fungal assemblage composition at two levels of diversity (four and eight species) under ambient temperature of 16 °C and two regimes of temperature increase differing in 8 °C: abrupt versus gradual increase from 16 to 24 °C. The effects were evaluated on leaf-litter decomposition, fungal biomass and reproduction. Results showed faster leaf decomposition under increased temperature, but no differences were found between an abrupt and a gradual increase in temperature. Assemblage composition was the major factor controlling fungal biomass and reproduction, while fungal diversity was only critical to maintain reproduction.     

Implications of water column ammonium uptake and regeneration for the nitrogen budget in temperate, eutrophic Missisquoi Bay, Lake Champlain (Canada/USA)

Invasion by exotic trees into riparian areas has the potential to impact aquatic systems. We examined the effects of the exotic Salix fragilis (crack willow) on the structure and functioning of small streams in northern Patagonian Andes via a field survey of benthic invertebrates and leaf litter and an in situ experiment. We compared leaf decomposition of the native Ochetophila trinervis (chacay) and S. fragilis in reaches dominated by native vegetation versus reaches dominated by crack willow. We hypothesized that S. fragilis affects the quality of leaf litter entering the streams, changing the aquatic biota composition and litter decomposition. Our study showed that crack willow leaves decomposed slower than chacay, likely related to leaf properties (i.e., leaf toughness). Benthic leaf litter mass was similar between the two riparian vegetation types, though in stream reaches dominated by crack willow, leaves of this species represented 82% of the total leaf litter. Benthic invertebrate abundance and diversity were similar between reaches but species composition differed. Our study found little evidence for strong impacts of crack willow on those small streams. Further studies on other aspects of ecosystem functioning, such as primary production, would enhance our understanding of the impacts of crack willow on Patagonian streams.     

Litter decomposition in a temperate and a tropical stream: the effects of species mixing,litter quality and shredders

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Urban riparian systems function as corridors for both native and invasive plant species

Riparian areas are often the only green areas left in urban and suburban landscapes, providing opportunities for conservation and connectivity of both aquatic and terrestrial organisms. While city planners and land managers often tout the importance of riparian networks for these uses, it is not well established if urban riparian plant communities are actually functioning as connected assemblages. Furthermore, urban riparian zones are well known to be highly invaded by non-native plant species and may be functioning to increase the spread of non-native species across the landscape. Here we examine connectivity of plant assemblages in riparian networks within an extensively urbanized landscape. We sampled riparian plant communities at 13 sites along three second-order streams of the Rahway River watershed, New Jersey. We also characterized propagule dispersal at each site by sampling litter packs on the river banks five times between March–October 2011 and identifying germinants from litter packs after cold stratification. Species turnover of both riparian and litter vegetation was more strongly associated with flow distance, particularly for native species, indicating that riverine systems are important for promoting connectivity of native plant assemblages in urban landscapes. However, non-native germinants significantly dominated propagule dispersal along the stream reaches, particularly early in the growing season, suggesting spread utilizing the river system and preemption may be an important mechanism for invasion success in this system. Our data show that management of invasive species should be planned and implemented at the watershed scale to reduce spread via the river system.     

Future increase in temperature may stimulate litter decomposition in temperate mountain streams: evidence from a stream manipulation experiment

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Effects of experimental warming and nitrogen enrichment on leaf and litter chemistry of a wetland grass,Phragmites australis

As global climate is warming and the nitrogen cycle accelerates, plants are likely to respond not only by shifting community composition, but also by adjusting traits such as tissue chemistry. We subjected a widespread wetland plant, Phragmites australis, to increased nitrate supply and elevated temperature in enclosures that were established in a littoral permanently submerged freshwater marsh. The nitrogen (N) and phosphorus (P) concentrations in green leaves ranged from 11.4 to 13.8 mg N and from 1.5 to 2.0 mg P g⁻¹ dry mass. While the N concentration changed little in brown litter, the P concentration decreased to 0.53–0.65 mg P g⁻¹ litter dry mass. Neither experimental warming of the water and sediment surface, nor nitrate enrichment during the growing season affected nitrogen or phosphorus concentrations in green leaves. Concentrations of the two major structural carbon compounds in plant litter, cellulose and lignin, were also unaffected, ranging from 32.1 to 34.2% of dry mass for cellulose and from 16.3 to 17.7% of dry mass for lignin. Warming, however, significantly increased the nitrogen concentration of fully brown leaf litter. Thus, temperature appears to be more important than the supply of dissolved N in the water, especially in affecting leaf litter N concentrations in P. australis, even when only water but not air temperature is increased. This result may have implications for decomposition processes and decomposer food webs, which both depend on the quality of plant litter.     

Over-winter decomposition and associated macroinvertebrate communities of three deciduous leaf species in forest streams on the Canadian Boreal Shield

The decomposition of deciduous leaf material provides a critical source of energy to aquatic food webs. Changes to riparian forests through harvesting practices may alter the species composition of deciduous leaf material entering streams. We compared over-winter decomposition of three different riparian leaf species (speckled alder (Alnus incana ssp. rugosa (Du Roi) J. Clausen), white birch (Betula papyrifera Marsh.), and trembling aspen (Populus tremuloides Michx.)) to determine their importance as a food resource for macroinvertebrate communities within Boreal Shield streams in northeastern Ontario, Canada. Leaf pack decomposition of the three leaf species formed a processing continuum throughout winter, where alder and birch leaf packs decomposed at a medium rate (k = 0.0065/day and 0.0053/day, respectively) and aspen leaf packs decomposed more slowly (k = 0.0035/day). Macroinvertebrate community colonization on leaf packs changed through time regardless of leaf species. Alder leaf packs supported higher abundances of macroinvertebrates in the fall while aspen leaf packs supported greater shredder abundances in the following spring. The study shows that leaf diversity may be important for providing a sustained food resource for aquatic macroinvertebrates throughout the relatively long over-winter period in Canadian Boreal Shield streams. Riparian forest management strategies should ensure that deciduous plant species richness is sustained in riparian areas.     

Interactive effects of global warming and ‘global worming’ on the initial establishment of native and exotic herbaceous plant species

The spread of exotic earthworms (‘worming’) and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.     

Shifts in leaf litter breakdown along a forest–pasture–urban gradient in Andean streams

Tropical montane ecosystems of the Andes are critically threatened by a rapid land‐use change which can potentially affect stream variables, aquatic communities, and ecosystem processes such as leaf litter breakdown. However, these effects have not been sufficiently investigated in the Andean region and at high altitude locations in general. Here, we studied the influence of land use (forest–pasture–urban) on stream physico‐chemical variables (e.g., water temperature, nutrient concentration, and pH), aquatic communities (macroinvertebrates and aquatic fungi) and leaf litter breakdown rates in Andean streams (southern Ecuador), and how variation in those stream physico‐chemical variables affect macroinvertebrates and fungi related to leaf litter breakdown. We found that pH, water temperature, and nutrient concentration increased along the land‐use gradient. Macroinvertebrate communities were significantly different between land uses. Shredder richness and abundance were lower in pasture than forest sites and totally absent in urban sites, and fungal richness and biomass were higher in forest sites than in pasture and urban sites. Leaf litter breakdown rates became slower as riparian land use changed from natural to anthropogenically disturbed conditions and were largely determined by pH, water temperature, phosphate concentration, fungal activity, and single species of leaf‐shredding invertebrates. Our findings provide evidence that leaf litter breakdown in Andean streams is sensitive to riparian land‐use change, with urban streams being the most affected. In addition, this study highlights the role of fungal biomass and shredder species (Phylloicus; Trichoptera and Anchytarsus; Coleoptera) on leaf litter breakdown in Andean streams and the contribution of aquatic fungi in supporting this ecosystem process when shredders are absent or present low abundance in streams affected by urbanization. Finally, we summarize important implications in terms of managing of native vegetation and riparian buffers to promote ecological integrity and functioning of tropical Andean stream ecosystems.     

Assessing the influence of wildfire on leaf decomposition and macroinvertebrate communities in boreal streams using mixed-species leaf packs

1. We investigated how compositional differences in riparian leaf litter derived from burned and undisturbed forests influenced leaf breakdown and macroinvertebrate communities using experimental mixed-species leaf packs in boreal headwater streams. Leaf pack mixtures simulating leaf litter from dominant riparian woody-stem species in burned and undisturbed riparian zones were incubated in two references and two fire-disturbed streams for 5 weeks prior to measuring temperature-corrected breakdown rates and macroinvertebrate community composition, richness, and functional metrics associated with decomposers such as shredder abundance and % shredders.
2. Leaf litter breakdown rates were higher and had greater variability in streams bordered by reference riparian forests than in streams where riparian forests had been burned during a wildfire. Streams bordered by fire disturbance showed significant effects of litter mixture on decomposition rates, observed as significantly higher decomposition rates of a fire-simulated leaf mixture compared to all other mixtures.
3. Variation among sites was higher than variation among litter mixtures, especially for macroinvertebrate community composition. In general, fire-simulated leaf mixtures had greater shredder abundances and proportions, but lower overall macroinvertebrate abundance; however, the shredder abundance trend was not consistent across all leaf mixtures at each stream.
4. These results show that disturbance-driven riparian forest condition and resulting composition of leaf subsidies to streams can influence aquatic invertebrate community composition and their function as decomposers. Therefore, if one of the primary goals of modern forest management is to emulate natural disturbance patterns, boreal forest managers should adapt silvicultural practices to promote leaf litter input that would arise post-fire, thereby supporting stream invertebrate communities and their function.

     

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

Responses of Aquatic Fungal Communities on Leaf Litter to Temperature‐Change Events

Increases of extreme weather events are predicted to occur with ongoing climate change, but impacts to freshwaters have rarely been examined. We assessed the effects of temperature on leaf‐litter associated fungi by exposing leaves colonized in a stream to 18 °C (control), 25 °C, or 18 °C after freezing. Treatments altered fungal dominance on leaves; Lunulospora curvula sporulation was stimulated by increased temperature and stopped by the freeze‐thaw treatment. Fungal biomass and diversity decreased at 18 °C after freezing, but not at 25 °C. Leaf decomposition was retarded by the freeze‐thaw treatment (k = –0.024 day^–1) and stimulated at 25 °C (k = –0.069 day^–1). Results suggest that occasional freezing may constrain fungal diversity and their ecological functions, while warming appears to accelerate plant‐litter decomposition in streams. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Drought and warming induced changes in P and K concentration and accumulation in plant biomass and soil in a Mediterranean shrubland

A field experiment involving drought and warming manipulation was conducted over a 6-year period in a Mediterranean shrubland to simulate the climate conditions projected by IPCC models for the coming decades (20% decreased soil moisture and 1°C warming). We investigated P and K concentration and accumulation in the leaves and stems of the dominant species, and in soil. Drought decreased P concentration in Globularia alypum leaves (21%) and in Erica multiflora stems (30%) and decreased K concentration in the leaves of both species (20% and 29%, respectively). The general decrease of P and K concentration in drought plots was due to the reduction of soil water content, soil and root phosphatase activity and photosynthetic capacity that decreased plant uptake capacity. Warming increased P concentration in Erica multiflora leaves (42%), but decreased it in the stems and leaf litter of Erica multiflora and the leaf litter (33%) of Globularia alypum, thereby demonstrating that warming improved the P retranslocation and allocation from stem to leaves. These results correlate with the increase in photosynthetic capacity and growth of these two dominant shrub species in warming plots. Drought and warming had no significant effects on biomass P accumulation in the period 1999–2005, but drought increased K accumulation in aboveground biomass (10 kg ha⁻¹) in Globularia alypum due to the increase in K concentration in stems. The stoichiometric changes produced by the different responses of the nutrients led to changes in the P/K concentration ratio in Erica multiflora leaves, stems and litter, and in Globularia alypum stems and litter. This may have implications for the nutritional value of these plant species and plant–herbivore relationships. The effects of climate change on P and K concentrations and contents in Mediterranean ecosystems will differ depending on whether the main component of change is drought or warming.