Nutrient content and dynamics in north Swedish shrub tundra areas

Forbs and leaves of deciduous shrubs had high concentrations of Ca, Mg, K, P, and N. Deciduous dwarf shrubs had intermediate concentrations but higher than evergreen dwarf shrubs. Monocots, cryptogams, woody and belowground tissues had low concentrations.
Plant nutrient concentrations and nutrient content in soil organic matter increased from dry towards moist tundra areas.
The residence time of nutrients was considerably less than ten years in surface litter, but several decades or centuries in total organic matter. The longest residence time was found in the moist part of the tundra.
N, K, and P in Betula nana leaves were translocated to a great extent prior to leaf fall, whereas Mg and particularly Ca were only slightly translocated. As on other tundra areas shortage of nutrients probably limits plant growth. In that case short supply of N and F seems most probable due to retention in litter and soil organic matter.     

Role of litter decomposition for the increased primary production in areas heavily grazed by reindeer: a litterbag experiment

Heavy grazing and trampling by reindeer increase nutrient cycling and primary production in areas where grasslands have replaced shrub and moss tundra. One way in which herbivores can affect nutrient cycling is through changing the litter decomposition processes. We studied the effect of herbivory on litter decomposition rate by reciprocal transplantation of litter between lightly grazed and heavily grazed areas, using the litterbag technique. We used litter from two of the most common species on the lightly grazed side, Betula nana and Empetrum nigrum , and two of the most common species on the heavily grazed side, Carex bigelowii and Deschampsia flexuosa . We found that herbivory improved litter quality by favouring species with easily decomposed litter. However, herbivory also improved litter quality by increasing the nitrogen content and lowering the C/N ratio of each species. Decomposition rates even correlated with the abundance of the plant category in question. Shrub litter decomposed faster in the lightly grazed area where shrubs were common, and graminoid litter decomposed faster in the heavily grazed area where graminoids were common. These results indicate that the decomposer micro-organisms are adapted to the most common litter types. This study shows that detailed information about the effect of herbivory on litter quality is important to understand differences between the short-term and long-term effects of herbivory on nutrient cycling and primary production.     

Plant–herbivore interactions in Alaskan arctic tundra change with soil nutrient availability

Herbivores in nutrient‐limited systems such as arctic tundra have been suggested to play a minor role in controlling plant growth simply because they are relatively few in number. However, theory predicts that as net primary productivity (NPP) increases because of greater inputs of nutrients or energy, herbivores may have greater effects on plant growth. This prediction has not been tested in the context of climate warming in arctic tundra, which may increase soil nutrient availability and thus NPP. We examined a long‐term experiment that excluded small and large mammalian herbivores and increased soil nutrients in two arctic Alaskan tundra communities: dry heath (DH) and moist acidic tussock (MAT). In the ninth year of manipulations, we measured weekly growth of three plant species of three growth forms: tussock‐forming graminoid, rhizomatous graminoid, and dwarf deciduous shrub, in each community. All species grew better when fertilized. In DH, this increase in growth was exaggerated when plants were protected from herbivores, confirming that herbivory had a negative effect on plant growth under increased nutrient conditions, but was unimportant under ambient soil conditions. However, in MAT, the importance of herbivory differed among species with fertilization. The tussock‐forming sedge at MAT, Eriophorum vaginatum, grew better and flowered more when fenced under both ambient and amended nutrients compared to plants exposed to herbivores. This species decreases in abundance in long‐term fertilized plots when mammals are present, and our results suggest that herbivory may be accounting for at least some of that loss, in addition to shifts in competitive relationships. Although we only focused on individual plants here rather than the entire community, our results suggest that under the increased soil nutrient conditions expected with continued climate warming in the Arctic, herbivores may become more important in affecting several abundant tundra plant populations, and should not be ignored.     

Dwarf shrubs are stronger competitors than graminoid species at high nutrient supply in peat bogs

Climate warming is likely to increase nutrient mineralization rates in bog ecosystems which may change the plant species composition. We examined the competitive relationships between two graminoid species, Eriophorum vaginatum and Rhynchospora alba, and two ericoid species, Calluna vulgaris and Vaccinium oxycoccus, at different nutrient supply rates. In a greenhouse, the plants were grown in monocultures and mixtures at four nutrient treatments: control, high N, high P, and high N + P. The results show that the ericoids responded more strongly to the nutrient treatments than the graminoids. The dwarf shrubs showed higher growth rates and reduced root:shoot ratio at high N + P supply. When grown in mixture the ericoids increased their growth, while graminoids decreased in biomass or showed signs of nutrient limitation compared to their monoculture plants. This suggests that under increased nutrient availability, bogs are more likely to turn into dwarf shrub dominated ecosystems and not grassland.     

Increased ectomycorrhizal fungal abundance after long-term fertilization and warming of two arctic tundra ecosystems

Shrub abundance is expected to increase with enhanced temperature and nutrient availability in the Arctic, and associated changes in abundance of ectomycorrhizal (EM) fungi could be a key link between plant responses and longer-term changes in soil organic matter storage. This study quantifies the response in EM fungal abundance to long-term warming and fertilization in two arctic ecosystems with contrasting responses of the EM shrub Betula nana. Ergosterol was used as a biomarker for living fungal biomass in roots and organic soil and ingrowth bags were used to estimate EM mycelial production. We measured 15N and 13C natural abundance to identify the EM-saprotrophic divide in fungal sporocarps and to validate the EM origin of mycelia in the ingrowth bags. Fungal biomass in soil and EM mycelial production increased with fertilization at both tundra sites, and with warming at one site. This was caused partly by increased dominance of EM plants and partly by stimulation of EM mycelial growth. We conclude that cycling of carbon and nitrogen through EM fungi will increase when strongly nutrient-limited arctic ecosystems are exposed to a warmer and more nutrient-rich environment. This has potential consequences for below-ground litter quality and quantity, and for accumulation of organic matter in arctic soils.     

Inhibition of growth,and effects on nutrient uptake of arctic graminoids by leaf extracts — allelopathy or resource competition between plants and microbes?

Previous research has shown that plant extracts, e.g. from boreal dwarf shrubs and trees, can cause reduced growth of neighbouring plants: an effect known as allelopathy. To examine whether arctic and subarctic plants could also be affected by leaching of phytochemicals, we added extracts from the commonly occurring arctic dwarf shrubs Cassiope tetragona and Empetrum hermaphroditum, and from mountain birch, Betula pubescens ssp. tortuosa to three graminoid species, Carex bigelowii, Festuca vivipara and Luzula arcuata, grown in previously sterilized or non-sterilized arctic soils. The graminoids in non-sterilized soil grew more slowly than those in sterilized soil. Excised roots of the plants in non-sterilized soil had higher uptake rate of labelled P than those in sterilized soil, demonstrating larger nutrient deficiency. The difference in growth rate was probably caused by higher nutrient availability for plants in soils in which the microbial biomass was killed after soil sterilization. The dwarf shrub extracts contained low amounts of inorganic N and P and medium high amounts of carbohydrates. Betula extracts contained somewhat higher levels of N and much higher levels of P and carbohydrates. Addition of leaf extracts to the strongly nutrient limited graminoids in non-sterilized soil tended to reduce growth, whereas in the less nutrient limited sterilized soil it caused strong growth decline. Furthermore, the N and P uptake by excised roots of plants grown in both types of soil was high if extracts from the dwarf shrubs (with low P and N concentrations) had been added, whereas the P uptake declined but the N uptake increased after addition of the P-rich Betula extract. In contrast to the adverse extract effects on plants, soil microbial respiration and soil fungal biomass (ergosterol) was generally stimulated, most strongly after addition of the Betula extract. Although we cannot exclude the possibility that the reduced plant growth and the concomitant stimulation of microbial activity were caused by phytochemicals, we believe that this was more likely due to labile carbon in the extracts which stimulated microbial biomass and activity. As a result microbial uptake increased, thereby depleting the plant available pool of N and P, or, for the P-rich Betula extract, depleting soil inorganic N alone, to the extent of reducing plant growth. This chain of events is supported by the negative correlation between plant growth and sugar content in the three added extracts, and the positive correlation between microbial activity, fungal biomass production and sugar content, and are known reactions when labile carbon is added to nutrient deficient soils.     

Testing the Stress‐Gradient Hypothesis During the Restoration of Tropical Degraded Land Using the Shrub Rhodomyrtus tomentosa as a Nurse Plant

The relative importance of facilitation and competition between pairwise plants across abiotic stress gradients as predicted by the stress‐gradient hypothesis has been confirmed in arid and temperate ecosystems, but the hypothesis has rarely been tested in tropical systems, particularly across nutrient gradients. The current research examines the interactions between a pioneer shrub Rhodomyrtus tomentosa (the nurse plant) and seedlings of a transplanted native woody Schima superba (the target species) in a tropical system in which position on a slope corresponds with a nutrient gradient; high soil nutrients at the slope bottom and relatively low soil nutrients at the slope top. In contrast, soil physical traits were more favorable for seedling growth under the shrub than in open spaces. The effect of R. tomentosa on S. superba survival was positive (facilitation) at the top of the slope, as indicated by the relative interaction index (RII), but negative in the bottom (competition). RII indicated a positive effect on seedling height at the top of the slope but was not at the bottom. Seedling survival was positively related to soil nutrient level and negatively related to soil acidity, but seedling growth of S. superba seemed to be enhanced by the shrub canopy. Thus, the results seem to support stress‐gradient hypothesis in terms of target species survival but not growth. We suggest using the shrub as a nurse plant in forest restoration in tropical degraded land with caution because not all of its effects on target species are positive .     

Exclusion of grass roots from soil organic layers by Calluna: the role of ericoid mycorrhizas

The role of ericoid mycorrhizal colonization in competition between the dwarf shrub Calluna vulgaris and coarse grass Nardus stricta was investigated. Nardus was grown alone, or in competition with Calluna, in a layered organic/sand substrate with and without inoculation with the ericoid mycorrhizal endophyte Hymenoscyphus ericae, and with and without the addition of nitrogen. Root length and allocation between different substrate layers was assessed along with plant biomass, nutrient uptake and mycorrhizal colonization. Calluna was the superior competitor for nutrients, probably because of its ability to concentrate root growth in the upper organic layer. In the presence of Calluna both the absolute amount and proportion of Nardus root length in the organic layer were reduced, and this reduction was greatest when Calluna was mycorrhizal. The presence of ericoid mycorrhizal colonization did not reduce Nardus shoot nutrient content or concentration, suggesting that ericoid mycorrhizal suppression of Nardus growth was not due to nutrient competition: alternative mechanisms of interference are discussed.     

Twenty-two years of warming, fertilisation and shading of subarctic heath shrubs promote secondary growth and plasticity but not primary growth

Most manipulation experiments simulating global change in tundra were short-term or did not measure plant growth directly. Here, we assessed the growth of three shrubs (Cassiope tetragona, Empetrum hermaphroditum and Betula nana) at a subarctic heath in Abisko (Northern Sweden) after 22 years of warming (passive greenhouses), fertilisation (nutrients addition) and shading (hessian fabric), and compare this to observations from the first decade of treatment. We assessed the growth rate of current-year leaves and apical stem (primary growth) and cambial growth (secondary growth), and integrated growth rates with morphological measurements and species coverage. Primary- and total growth of Cassiope and Empetrum were unaffected by manipulations, whereas growth was substantially reduced under fertilisation and shading (but not warming) for Betula. Overall, shrub height and length tended to increase under fertilisation and warming, whereas branching increased mostly in shaded Cassiope. Morphological changes were coupled to increased secondary growth under fertilisation. The species coverage showed a remarkable increase in graminoids in fertilised plots. Shrub response to fertilisation was positive in the short-term but changed over time, likely because of an increased competition with graminoids. More erected postures and large, canopies (requiring enhanced secondary growth for stem reinforcement) likely compensated for the increased light competition in Empetrum and Cassiope but did not avoid growth reduction in the shade intolerant Betula. The impact of warming and shading on shrub growth was more conservative. The lack of growth enhancement under warming suggests the absence of long-term acclimation for processes limiting biomass production. The lack of negative effects of shading on Cassiope was linked to morphological changes increasing the photosynthetic surface. Overall, tundra shrubs showed developmental plasticity over the longer term. However, such plasticity was associated clearly with growth rate trends only in fertilised plots.     

Variable sensitivity of plant communities in Iceland to experimental warming

Facing an increased threat of rapid climate change in cold‐climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open‐top chambers in two widespread but contrasting tundra communities in Iceland. In a species‐poor and nutrient‐deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species‐rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land‐use history.     

Post‐fire vegetation dynamics in nutrient‐enriched and non‐enriched sclerophyll woodland

Abstract Exotic plant invasions are a significant problem in urban bushland in Sydney, Australia. In low‐nutrient Hawkesbury Sandstone communities, invasive plants are often associated with urban run‐off and subsequent increases in soil nutrients, particularly phosphorus. Fire is an important aspect of community dynamics in Sydney vegetation, and is sometimes used in bush regeneration projects as a tool for weed control. This study addressed the question: ‘Are there differences in post‐fire resprouting and germination of native and exotic species in nutrient‐enriched communities, compared with communities not disturbed by nutrient enrichment?’ We found that in non‐enriched areas, few exotic species emerged, and those that did were unable to achieve the rapid growth that was seen in exotic plants in the nutrient‐enriched areas. Therefore, fire did not promote the invasion of exotic plants into areas that were not nutrient‐enriched. In nutrient‐enriched areas after fire, the diversity of native species was lower than in the non‐enriched areas. Some native species were able to survive and compete with the exotic species in terms of abundance, per cent cover and plant height. However, these successful species were a different suite of natives to those commonly found in the non‐enriched areas. We suggest that although fire can be a useful tool for short‐term removal of exotic plant biomass from nutrient‐enriched areas, it does not promote establishment of native species that were not already present.     

Changing leaf litter feedbacks on plant production across contrasting sub-arctic peatland species and growth forms

Plant species and growth forms differ widely in litter chemistry, which affects decay and may have important consequences for plant growth via e.g. the release of nutrients and growth-inhibitory compounds. We investigated the overall short-term (9.5 months) and medium-term (21.5 months) feedback effects of leaf litter quality and quantity on plant production, and tested whether growth forms can be used to generalise differences among litter species. Leaf litter effects of 21 sub-arctic vascular peatland species on Poa alpina test plants changed clearly with time. Across all growth forms, litter initially reduced plant biomass compared with untreated plants, particularly litters with a high decomposition rate or low initial lignin/P ratio. In the second year, however, litter effects were neutral or positive, and related to initial litter N concentration (positive), C/N, polyphenol/N and polyphenol/P ratios (all negative), but not to decomposability. Differences in effect size among several litter species were large, while differences in response to increasing litter quantities were not significant or of similar magnitude to differences in response to three contrasting litter species. Growth forms did not differ in initial litter effects, but second-year plant production showed a trend (P < 0.10) for differences in response to litters of different growth forms: evergreen shrubs < graminoids or deciduous shrubs < forbs. While long-persisting negative litter effects were predominant across all growth forms, our data indicate that even within nutrient-constrained ecosystems such as northern peatlands, vascular plant species, and possibly growth forms, differ in litter feedbacks to plant growth. Differences in the composition of undisturbed plant communities or species shifts induced by external disturbance, such as climate change, may therefore feedback strongly to plant biomass production and probably nutrient cycling rates in northern peatlands. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Plants retard litter decay in a nutrient-limited soil: a case of exploitative competition?

Litter was incubated in pasteurized and unpasteurized soils, with and without the presence of prairie grasses (Bouteloua gracilis or Schizachyrium scoparium), to determine if competition between plants and saprophytes altered rates of litter decay. A soil pasteurization treatment was included to ascertain if the presence of vesicular-arbuscular mycorrhizae inocula would affect the competitive interaction. Analyses of variance detected significant effects (P<0.5) of soil pasteurization, plant species and litter presence on root mass, and significant effects of plant species and litter presence on shoot and total plant mass. The degree of mycorrhizal colonization varied little between plant species but mycorrhizae were entirely absent in pasteurized soils; soil pasteurization usually reduced plant growth. Analyses of covariance detected (1) a highly significant negative relationship (P<0.01) between litter mass loss and plant growth (as covariates), when controlling for the effects of time, and (2) a significant positive relationship (P<0.05) between litter nitrogen content and mass loss (as covariates), when controlling for the effects of time and soil pasteurization. These relationships would be expected if exploitative competition for nutrients existed between plants and decomposers such that nutrient uptake by plants reduced concomitant litter decay. Received: 27 December 1996 / Accepted: 11 September 1997     

Photosynthetic and respiratory responses to temperature and light of three Alaskan tundra growth forms

Photosynthetic and respiratory response of four Alaskan tundra species comprising three growth forms were investigated in the laboratory using an infrared gas analysis system. Vaccinium vitis-idaea , a dwarf evergreen shrub, demonstrated a low photosynthetic capacity: P_max= 1 mg CO₂ g dry wt⁻¹ h⁻¹; T_opt < 10°C. Betula nana , a deciduous shrub, had a high relatively photosynthetic capacity: P_max= 14 mg CO₂ g dry wt⁻¹ h⁻¹; T_opt 17°C. Two graminoid (sedge) species, Carex aquatilis and Eriophorum vaginalum , showed different responses. Carex showed a high photosynthetic capacity: P_max= 20 mg CO₂ g dry wt⁻¹ h⁻¹; T_opt 22°C. Eriophorum vaginatum demonstrated an intermediate photosynthetic capacity of 4 mg CO₂ g dry wt⁻¹ h⁻¹ at saturated light levels. Leaf dark respiration, up to 20°C, was approximately the same for all species. The patterns of root respiration among species was opposite to the trend in photosynthesis. Vaccinium vitis-idaea had the highest rate of root respiration and B. nana the lowest ( C aquatilis was not measured). Correlation between leaf nitrogen content (%) and photosynthetic capacity was high. Hypothesized growth form relationships explained differences in photosynthetic capacity between the deciduous shrub and evergreen shrub, but did little to account for differences between the two sedges. Differences in rooting patterns between species may affect tissue nutrient content, carbon flux rates, and carbon balance.     

A dominant dwarf shrub increases diversity of herbaceous plant communities in a Trans-Himalayan rangeland

Plant communities are structured by both competition and facilitation. The interplay between the two interactions can vary depending on environmental factors, nature of stress, and plant traits. However, whether positive or negative interactions dominate in regions of high biotic and abiotic stress remains unclear. We studied herbaceous plant communities associated with a dwarf shrub Caragana versicolor in semi-arid, high altitude Trans-Himalayan rangelands of Spiti, India. We surveyed 120 pairs of plots (within and outside shrub canopies) across four watersheds differing in altitude, aspect, and dominant herbivores. Herbaceous communities within shrub canopies had 25% higher species richness, but similar abundance when compared to communities outside the canopy, with the shrub edge having higher diversity than the centre of the canopy. Grasses and erect forbs showed positive associations with the shrub, while prostrate plants occurred at much lower abundance within the canopy. Rare species showed stronger positive associations with Caragana than abundant species. Experimental removal of herbaceous vegetation from within shrub canopies led to 42% increase in flowering in Caragana, indicating a cost to the host shrubs. Our study indicates a robust pattern of a dwarf shrub facilitating local community diversity across this alpine landscape, increasing diversity at the plot level, facilitating rare species, and yet incurring a cost to hosts from the presence of herbaceous plants. Given these large influences of this shrub on the vegetation of these high altitude rangelands, we suggest that the shrub microhabitat be explicitly considered in any analyses of ecosystem health in such rangelands.     

Increased precipitation modulates the influence of nitrogen and litter inputs on the nutrient resorption proficiency rather than efficiency of <Emphasis Type="Italic">Leymus chinensis</Emphasis>

Resorption of nutrients from senescing organs is an important conservation mechanism that is usually influenced by the supply of soil nutrients and plant growth requirements. Therefore, it is likely that increases in nitrogen (N), precipitation, and litter could lead to changes in nutrient resorption because of changes in nutrients in the soil and accelerated plant growth in response to the alleviation of water limitations in arid and semiarid environments. In the current study, we investigated the effects of water, N, and litter addition on the nutrient resorption efficiency and proficiency of N and phosphorus (P) in leaves and stems of Leymus chinensis in Inner Mongolia, China. Our results showed that N addition significantly decreased the N resorption efficiency in leaves under water addition, and increased P resorption efficiency under ambient precipitation conditions. There was no apparent influence of either litter or water addition on N and P resorption efficiencies. However, N and litter addition significantly altered N and P resorption proficiencies, and these effects were modulated by water availability. Furthermore, changes in resorption proficiencies were mainly associated with alterations in the nutritional status of green organs in response to water, N and litter addition, except for leaf P. Our findings highlight the importance of increased precipitation in modulating the nutrient resorption proficiency of plants under potentially increased nutrient availability in semiarid grasslands. Therefore, global changes in precipitation and N, and corresponding litter changes could result in complex effects on plant nutrient economies and, in turn, could influence the return of nutrients to the soil.     

Lack of compensatory growth under phosphorus deficiency in grazing-adapted grasses from the Serengeti Plains

Summary Two shortgrass species (Sporobolus ioclados and Eustachys paspaloides) and two midgrass species (E. paspaloides and Pennisetum mezianum) from the Serengeti grasslands of Tanzania were grown under conditions of extreme phosphorus (P) deficiency. Production of each of these species is maintained or enhanced by defoliation under adequate nutrient supply (McNaughton et al. 1983). However, under the P-deficient conditions of our experiment, defoliation caused a reduction in biomass of all plant parts of each species. Green leaf biomass was reduced most strongly by defoliation, and crowns were least affected. Yield of biomass and nutrients to grazers (green leaves+clipped material) was enhanced by weekly defoliation in the shortgrass grazing-adapted species, whereas yield to producers (live biomass and nutrients retained by the plant) and yield to decomposers (litter) were strongly reduced by defoliation in all species. Phosphate absorption capacity (V _max) measured on excised roots was enhanced by defoliation in the grazing-adapted Sporobolus, but, due to low affinity (high K _m) of roots of defoliated plants for phosphate, absorption rate was not greatly altered at low solution concentrations. Phosphate absorption capacity was reduced or unaffected by defoliation in other species. We conclude that under conditions of P deficiency, plants are unable to acquire the nutrients necessary to replenish large nutrient losses to grazers. In low-nutrient environments, compensatory growth (stimulation of production by grazing) is not a viable strategy. Therefore, in these environments plants respond evolutionarily to herbivores by developing chemical or morphological defenses.     

Resource competition and fire‐regulated nutrient demand in carnivorous plants of wet pine savannas

Question: What are the mechanisms by which fire reduces competition for both a short‐lived and a long‐lived species in old‐growth ground‐cover plant communities of wet pine savannas (originally Pinus palustris, replaced by P. elliottii)? Location: Outer coastal plain of southeastern Mississippi, USA. Methods: I reviewed previous competition experiments and proposed a new hypothesis to explain the relationship between fire, competition, and species co‐existence in wet longleaf pine savannas. The first study is about growth and seedling emergence responses of a short‐lived carnivorous plant, Drosera capillaris, to reduction in below‐ground competition and above‐ plus below‐ground competition. The second study deals with growth and survival responses of a long‐lived perennial carnivorous plant, Sarracenia alata, to neighbour removal and prey‐exclusion to determine if a reduction in nutrient supply increased the intensity of competition in this nutrient‐poor system. Results: Fire increased seedling emergence of the short‐lived species by reducing above‐ground competition through the destruction of above‐ground parts of plants and the combustion of associated litter. Prey exclusion did not increase competitive effects of neighbours on the long‐lived species. However, because the experiment was conducted in a year without fire, shade reduced nutrient demand, which may have obviated competition for soil nutrients between Sarracenia alata and its neighbours. Conclusion: Repeated fires likely interact with interspecific differences in nutrient uptake to simultaneously reduce both above‐ground competition and competition for nutrients in old‐growth ground cover communities in pine savannas. Restoration practitioners should consider the possibility that the composition of the plant community is just as important as fire in ensuring that frequent fires maintain species diversity.     

Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

Non-native and native shrubs have differing impacts on species diversity and composition of associated plant communities

The spread of non-native plants has been depicted as a serious threat to biodiversity. However, it remains unclear whether the indigenousness of the invading plant plays a marked role for the ecological consequences of an invasion as few studies have compared the ecological impacts of non-native shrubs with structurally or functionally comparable native shrubs. We studied patches of introduced and native shrubs to assess whether there are general differences in plant species composition or biomass between patches formed by non-native versus native shrubs. The indigenousness of the shrub (non-native vs. native) did not explain the variation in soil nutrients, neither the production of shoot biomass or allocation of growth to different parts of the shoot. The amount of light reaching ground level did not differ between patches of a non-native and a native shrub. However, species richness and biomass of herbaceous plants were lower in patches of non-native than native shrubs and the amount of litter was higher below non-native than native shrubs. Our results suggest that the indigenousness of the patch-forming plant may be an important factor for the diversity and composition of associated herbaceous vegetation. Based on our results, resource availability (light and nutrients) is not a sufficient explanation for the negative effects of non-native shrubs on plant communities. Further research is needed to investigate whether alternative explanations, such as the novelty of the toxic compounds produced by non-native plants, can explain the differences we observed.