Warming the tundra: reciprocal responses of invertebrate herbivores and plants

Rapid warming in northern ecosystems is simultaneously influencing plants, herbivores and the interactions among them. Recent studies suggest that herbivory could buffer plant responses to environmental change, but this has only been shown for vertebrate herbivores so far. The role of invertebrate herbivory in tundra ecosystems is often overlooked, but can be relevant in determining the structure and dynamics of tundra plant communities and may also affect how plants respond to warming. Invertebrate herbivores are also likely to respond more rapidly to warming than vertebrates because their behaviour and life cycles strongly depend on temperature. We investigated the effects of current season warming on Arctic moth caterpillars, their herbivory rates, and the subsequent responses of two common tundra plants, Salix arctica and Dryas octopetala. We manipulated both herbivore presence and temperature in a full‐factorial field experiment at two elevations, using enclosures and passive warming chambers. Changes in temperature achieved through elevation and/or experimental warming directly affected caterpillars, herbivory and the responses of plants. Caterpillars performed worse (higher respiration rates and lower growth rates) in warmer, lower elevation plots and shifted their diets towards more nutritious foods, such that the relative intensity of herbivory changed for the two studied plants. Within‐season responses of both forage plant species were weak, but invertebrate herbivores affected the responses of plants to elevation or experimental warming. Our results suggest that increased temperatures can reduce the performance of cold‐adapted invertebrate herbivores, with potential consequences to the longer term responses of tundra plants to warming due to changes in herbivory rates and selective foraging.     

Impact of grasshoppers and an invasive grass on establishment and initial growth of restoration plant species

Exotic plant invasion can have dramatic impacts on native plants making restoration of native vegetation at invaded sites challenging. Though invasives may be superior competitors, it is possible their dominance could be enhanced by insect herbivores if native plants are preferred food sources. Insect herbivory can regulate plant populations, but little is known of its effects in restoration settings. There is a need to better understand relationships between insect herbivores and invasive plants with regard to their combined potential for impacting native plant establishment and restoration success. The objective of this study was to assess impacts of grasshopper herbivory and the invasive grass Bromus tectorum (cheatgrass) on mortality and growth of 17 native plant species used in restoration of critical sagebrush steppe ecosystems. Field and greenhouse experiments were conducted using moderate densities of a common, generalist pest grasshopper (Melanoplus bivittatus). Grasshoppers had stronger and more consistent impacts on native restoration plants in field and greenhouse studies than cheatgrass. After 6 weeks in the greenhouse, grasshoppers were associated with 36% mortality over all native restoration species compared to 2% when grasshoppers were absent. Herbivory was also associated with an approximately 50% decrease in native plant biomass. However, effects varied among species. Artemisia tridentata, Chrysothamnus viscidiflorus, and Coreopsis tinctoria were among the most negatively impacted, while Oenothera pallida, Pascopyrum smithii, and Leymus cinerus were unaffected. These findings suggest restoration species could be selected to more effectively establish and persist within cheatgrass infestations, particularly when grasshopper populations are forecasted to be high.     

Temperature response of CO<Subscript>2</Subscript> exchange and dissolved organic carbon release in a maritime Antarctic tundra ecosystem

We examined the temperature response of CO₂ exchange and soil biogeochemical processes in an Antarctic tundra ecosystem using laboratory incubations of intact tundra cores. The cores were collected from tundra near Anvers Island along the west coast of the Antarctic Peninsula that was dominated by the vascular plants Colobanthus quitensis and Deschampsia antarctica. After the initial 8-week incubation at moderate growth temperatures (12/7°C, day/night), the tundra cores were incubated for another 8 weeks at either a higher (17/12°C) or lower (7/4°C) temperature regime. Temperature responses of CO₂ exchange were measured at five temperatures (4, 7, 12, 17, and 27°C) following each incubation and soil leachates were collected biweekly over the second incubation. Daytime net ecosystem CO₂ exchange (NEE) per unit core surface area was higher across the five measurement temperatures after the warmer incubation (17/12°C > 7/4°C). Responses of ecosystem respiration (ER) were similar at each measurement temperature irrespective of incubation temperature regimes. ER, expressed on a leaf-area basis, however, was significantly lower following the warmer incubation, suggesting a downregulation of ER. Warmer incubation resulted in a greater specific leaf area and N concentration, and a lower δ¹³C in live aboveground C. quitensis, but a higher δ¹³C in D. antarctica, implying species-specific responses to warming. Concentrations of dissolved organic C and N and inorganic N in soil leachates showed that short-term temperature changes had no noticeable effect on soil biogeochemical processes. The results suggest that downregulation of ER, together with plant species differences in leaf-area production and N use, can play a crucial role in constraining the C-cycle response of Antarctic tundra ecosystems to warming.     

Response of plants and the dominant microarthropod, Cryptopygus antarcticus, to warming and contrasting precipitation regimes in Antarctic tundra

We examined the influence of warming and supplemental precipitation on plant production and abundance of the dominant microarthropod, the springtail Cryptopygus antarcticus (Collembola), in tundra dominated by the vascular plants Colobanthus quitensis and Deschampsia antarctica along the Antarctic Peninsula. Tundra cores were placed in plots near Palmer Station where they were warmed with infrared heaters in combination with receiving supplemental precipitation. Diel canopy air and soil temperatures and air vapor pressure deficits in warmed plots were elevated 0.8 °C, 2.2 °C and 0.13 kPa, respectively. After two growing seasons, total aboveground plant production was greater under warming as a result of enhanced production by C. quitensis, which more than offset declines in moss biomass. Total aboveground plant production was also greater under supplemental precipitation primarily as a result of enhanced moss production. Total aboveground plant production was greatest under the combination of warming and supplemental precipitation, primarily as a result of enhanced C. quitensis production. C. antarcticus were more abundant in cores receiving supplemental precipitation and there was a strong treatment interaction; these springtails were most abundant in warmed cores receiving supplemental precipitation. Over 50% of the variability in the abundance of C. antarcticus could be explained by differences in aboveground plant biomass. However, plant production did not appear directly responsible for differences in C. antarcticus abundance; when we examined C. antarcticus abundance per unit of aboveground plant biomass, differences in its abundance among treatments were still apparent implying these differences were not the direct result of plant biomass. The responses of C. antarcticus were consistent with its known moisture and thermal preferences, suggesting that abiotic factors played a dominant role in controlling its abundance. Precipitation regime had large impacts on warming responses and these were species specific, illustrating the importance of future precipitation regimes in predicting system responses to warming.     

Linking responses of native and invasive plants to hurricane disturbances: implications for coastal plant community structure

Hurricane disturbances produce significant changes in forest microclimates, creating opportunities for seedling regeneration of native and invasive plant species alike. However, there is limited information on how changes in microclimates and pre-existing forest conditions affect native and invasive plants responses to hurricane disturbances. In this manipulative study, we examined the responses of three common shrub/small stature tree species, two of which are native to the coastal region of the southeastern USA (Baccharis halimifolia and Morella cerifera) and one that is invasive (Triadica sebifera), to two key components of hurricane disturbance (canopy damage and saline storm surge). In a greenhouse, we grew seedlings of these species under a range of shade levels that mimicked pre-and post-hurricane canopy conditions for wet pine forest and mixed hardwood forest, two forest communities common in coastal areas of the southeastern USA. Seedlings were subjected to saline storm surges equivalent to full strength sea water for 3 days. Seedling responses (mortality and growth) to the treatments were monitored for 16 months. All species benefitted from higher canopy openness. Storm surge effects were short-lived and seedlings readily recovered under high light conditions. The storm surge had stronger negative effects on survival and growth of all species when coupled with high shade, suggesting storm surge has greater negative impacts on seedlings where hurricane winds cause minimal or no canopy damage. The invasive T. sebifera was by far more shade tolerant than the natives. Survival of T. sebifera seedlings under highly shaded conditions may provide it a competitive edge over native species during community reassembly following tropical storms. Differential responses of native and invasive species to hurricane disturbances will have profound consequences on community structure across coastal forest stands, and may be regulated by legacies of prior disturbances, community structure, extent of canopy damage, and species’ tolerance to specific microclimates.

     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.     

Sensitivity of US Gulf of Mexico coastal marsh vegetation to crude oil: Comparison of greenhouse and field responses

Greenhouse and field studies were conducted to evaluate the effect of crude oil on selected US Gulf of Mexico coastal marsh species. Species showed different levels of sensitivity to oiling between greenhouse and field conditions. In greenhouse studies, two crude oils were used: South Louisiana crude oil (SLC) and Arabian Medium crude oil (AMC). The majority of Spartina patens plants died within one month following oiling with little or no recovery after three months. Panicum hemitomon and Spartina alterniflora were also adversely affected by oiling under greenhouse conditions but to a lesser extent than S. patens. The SLC or AMC oiling led to biomass reductions in S. alterniflora and S. patens. The dry biomass was not affected by oiling in P. hemitomon, Sagittaria lancifolia, Typha latifolia, and Scirpus olneyi. Results showed that S. patens plants were more sensitive to SLC as compared to AMC oil. Gross CO₂-C fixation data collected in the greenhouse indicated no differences in recovery among species across oiling treatments for S. lancifolia, S. olneyi, and T. latifolia. Field studies with S. alterniflora, S. patens and S. lancifolia demonstrated initial sensitivity of these species to oiling, and recovery following oiling with SLC. Our data also showed that caution must be employed whenever results from greenhouse studies are extrapolated to predict oil impact on vegetation under field conditions. Development of any sensitivity index of plant responses to oiling should not be based on greenhouse experiments only. Field evaluations should be included which best depict plant responses to oiling. Thus, restoration measures of US Gulf of Mexico coastal marshes following oiling should rely primarily on field studies. The field research suggests that the US Gulf of Mexico coastal marsh vegetation are likely to recover from oil spills naturally without the need for remediation procedures.     

Shrub canopies influence soil temperatures but not nutrient dynamics: An experimental test of tundra snow–shrub interactions

Shrubs are the largest plant life form in tundra ecosystems; therefore, any changes in the abundance of shrubs will feedback to influence biodiversity, ecosystem function, and climate. The snow–shrub hypothesis asserts that shrub canopies trap snow and insulate soils in winter, increasing the rates of nutrient cycling to create a positive feedback to shrub expansion. However, previous work has not been able to separate the abiotic from the biotic influences of shrub canopies. We conducted a 3‐year factorial experiment to determine the influences of canopies on soil temperatures and nutrient cycling parameters by removing ~0.5 m high willow (Salix spp.) and birch (Betula glandulosa) shrubs, creating artificial shrub canopies and comparing these manipulations to nearby open tundra and shrub patches. Soil temperatures were 4–5°C warmer in January, and 2°C cooler in July under shrub cover. Natural shrub plots had 14–33 cm more snow in January than adjacent open tundra plots. Snow cover and soil temperatures were similar in the manipulated plots when compared with the respective unmanipulated treatments, indicating that shrub canopy cover was a dominant factor influencing the soil thermal regime. Conversely, we found no strong evidence of increased soil decomposition, CO₂ fluxes, or nitrate or ammonia adsorbtion under artificial shrub canopy treatments when compared with unmanipulated open tundra. Our results suggest that the abiotic influences of shrub canopy cover alone on nutrient dynamics are weaker than previously asserted.     

Increasing substrate heterogeneity as a bet‐hedging strategy for restoring wetland vegetation

Mimicking the natural heterogeneity of wetland substrates, e.g. by roughening surface soil or constructing hummocks, has been shown to facilitate wetland plant establishment. We asked if incorporating substrate heterogeneity could also help plants withstand variation in moisture levels. In a wetland with Carex stricta (tussock sedge) as the main restoration target, we manipulated substrates to create different soil moisture environments for planted C. stricta plugs. Our artificial mounds mimicked tussocks formed by C. stricta in natural meadows (circa 10–40 cm in height); we also varied mound compositions and created shallow depressions. Monitoring demonstrated variation in soil moisture among our treatments and natural differences in soil moisture between experimental blocks. Additionally, rainfall varied from severe drought in year 1 to extreme rainfall in year 2. Plug survival, flowering, cover, biomass, leaf length, and growth rate all varied with treatment, block, and/or year. Interactions among those factors were common. Planting plugs in shallow depressions exacerbated stress in a wet block during a wet year, causing low survival. Planting plugs in moisture‐retaining peat pots allowed them to survive and sustain growth even in a dry block during a dry year. We conclude that heterogeneous substrates can be used to hedge against environmental variability by widening the range of microsites available within a restoration site and thereby moderating stressful conditions in some areas.     

Mycorrhizal colonization does not affect tolerance to defoliation of an annual herb in different light availability and soil fertility treatments but increases flower size in light-rich environments

Heterogeneous distribution of resources in most plant populations results in a mosaic of plant physiological responses tending to maximize plant fitness. This includes plant responses to trophic interactions such as herbivory and mycorrhizal symbiosis which are concurrent in most plants. We explored fitness costs of 50% manual defoliation and mycorrhizal inoculation in Datura stramonium at different light availability and soil fertility environments in a greenhouse experiment. Overall, we showed that non-inoculated and mycorrhiza-inoculated plants did not suffer from 50% manual defoliation in all the tested combinations of light availability and soil fertility treatments, while soil nutrients and light availability predominately affected plant responses to the mycorrhizal inoculation. Fifty percent defoliation had a direct negative effect on reproductive traits whereas mycorrhiza-inoculated plants produced larger flowers than non-inoculated plants when light was not a limiting factor. Although D. stramonium is a facultative selfing species, other investigations had shown clear advantages of cross-pollination in this species; therefore, the effects of mycorrhizal inoculation on flower size observed in this study open new lines of inquiry for our understanding of plant responses to trophic interactions. Also in this study, we detected shifts in the limiting resources affecting plant responses to trophic interactions.     

Facilitation and herbivory during restoration of California coastal sage scrub

Using nurse plants to facilitate native plant recruitment in degraded habitats is a common restoration practice across various arid and semiarid environments. Living trees or shrubs are typically considered nurse plants, whereas dead shrubs left in the landscape from prolonged drought are understudied prospective facilitators for native plant recruitment. The interaction between nurse plants and biotic pressures, such as herbivory, on native recruitment is also not well understood in semiarid plant communities. We investigated the effects of facilitation and herbivory on native seedling germination, growth, and survival in the restoration of degraded coastal sage scrub (CSS) habitat. Native shrub seedlings (Artemisia californica and Salvia mellifera) were planted, and native annual species (Amsinckia intermedia, Deinandra fasciculata, Phacelia distans, and Pseudognaphalium californicum) were sown in three Shrub Type treatments (live shrub, dead shrub, and exposed areas), with a nested Cage treatment (no cage and cage) in each Shrub Type treatment. Annual species grew equally well in all Shrub Type treatments; shrub seedlings grew largest in exposed areas. While there was little evidence of facilitation for all species tested, there were strong positive effects of caging on growth and establishment of all species. Caging palatable native species or planting species with anti‐herbivory traits around target plants may be more strategic approaches compared to using nurse plants in restoring degraded CSS after extended drought.     

Local soil,but not commercial AMF inoculum,increases native and non‐native grass growth at a mine restoration site

Soil communities are often degraded in mined sites, and facilitating the recovery of soil mutualists such as arbuscular mycorrhizal fungi (AMF) may assist with the restoration of native plants. At a grassland mine restoration site, I compared a commercial AMF inoculum with soil collected from beneath native grasses as a source of inoculum, as well as a control treatment. Field plots were broadcast‐inoculated and seeded with native grasses, and biomass of native and non‐native species was measured in three consecutive years. In addition, greenhouse‐grown seedlings of a native bunchgrass (Stipa pulchra) were inoculated with similar treatments, transplanted into the field, and assessed after 18 months. When broadcast inoculation was used, the local soil inoculum tended to increase non‐native grass biomass, and marginally decreased non‐native forb biomass in the second year of study, but did not significantly affect native grass biomass. Broadcast commercial inoculum had no detectable effects on biomass of any plant group. Stipa pulchra transplants had greater N content and mycorrhizal colonization, and marginally higher shoot mass and K content, when pre‐inoculated with local soil (relative to controls). Pre‐inoculation with commercial AMF increased AMF colonization of the S. pulchra transplants, but did not significantly affect biomass or nutrient content. The findings indicate that at this site, the use of local soil as an inoculum had greater effects on native and non‐native plants than the commercial product used. In order to substantially increase native grass performance, inoculation of transplanted plugs may be one potential strategy.     

Ecosystem disturbance reduces the allelopathic effects of Empetrum hermaphroditum humus on tundra plants

Question: Do naturally occurring ecosystem disturbances interfere with allelopathic effects and alleviate growing conditions for plants typical of more productive ecosystem states? Location: An unproductive tundra ecosystem in Northern Fennoscandia. Methods: We designed a factorial phytotron experiment to monitor the growth of seedlings of two herbaceous species in undisturbed and disturbed humus of the dominant dwarf shrub Empetrum hermaphroditum, which releases allelopathic substances that accumulate in soil over time. Herbivore defecation and wildfire are two major ecosystem disturbances likely to alleviate effects of these allelopathic substances. Humus and seeds of herbaceous species were sampled from Empetrum‐dominated heaths. Disturbance effects on humus were simulated by adding reindeer faeces and by low‐severity burning. Results: The concentration of the allelopathic substance batatasin‐III in Empetrum humus was 0.32±0.16 (SE) μg g^?1, which is similar to previous reports from boreal areas. Reindeer faeces addition and burning treatments to Empetrum humus caused three‐ and five‐fold increases in growth of the herbaceous species Avenella flexuosa and Solidago virgaurea respectively. Changes in soil chemistry caused by the disturbance, i.e. increased pH and nutrient availability, likely interfered with allelopathic effects and improved conditions for growth. Conclusions: Humus from Empetrum‐dominated tundra areas is infertile for seedlings of local herbaceous plants. Simulated herbivore defecation and wildfire can alleviate this infertility. Based on our results, we discuss the potential role of these naturally occurring disturbances in switching vegetation composition from Empetrum to herbaceous dominance, i.e. from an unproductive to a more productive tundra ecosystem.     

Restoration impacts on fuels and fire potential in a dryland tropical ecosystem dominated by the invasive grass Megathyrsus maximus

Ecological restoration often attempts to promote native species while managing for disturbances such as fire and non‐native invasions. The goal of this research was to investigate whether restoration of a non‐native, invasive Megathyrsus maximus (guinea grass) tropical grassland could simultaneously promote native species and reduce fire potential. Megathyrsus maximus was suppressed with herbicide, and three suites of native species—each including the same groundcover and shrub, and one of three tree species—were outplanted in a randomized, complete block design that also included herbicide control (herbicide with no outplantings) and untreated control treatments. Fuels were quantified 27 months after outplanting, and potential fire behavior (rate of spread and flame length) was modeled with BehavePlus. Compared with untreated controls, native outplant treatments reduced M. maximus cover by 76–91% and M. maximus live and dead fuel loads by greater than 92 and 68%, respectively. Despite reductions in M. maximus fuels, neither treatment‐level (grass + native) total fuel loads and fuel moistures, nor modeled fire behavior differed between outplant treatments and controls. The best performing native woody species (Dodonaea viscosa) had significantly lower average individual plant live fuel moisture (84%) than M. maximus (156%) or other native woody outplant species (201–328%), highlighting the need for careful species selection. These results demonstrate that restoring native species to degraded tropical dry forests is possible, but that ecological restoration will not necessarily alter the potential for fire, at least in the short term, making selection of species with beneficial fuel properties and active fire management critical components of ongoing restoration.     

A GROWTH COMPARISON OF SPARTINA ALTERNIFLORA LOISEL. ECOPHENES UNDER AEROBIC AND ANAEROBIC CONDITIONS

Spartina alterniflora Loisel. culms were collected from tall (creekbank), short (highmarsh) and dieback sites in a North Carolina salt marsh and grown in aerobic and anaerobic simulated marsh systems in the greenhouse. There were no significant differences between density, aerial live biomass, height, leaf width or root biomass and sprig source. All variables were significantly different between aeration treatments. Aerated systems had an average of 6.3 times more biomass than the unaerated treatments. There was a significant interaction between sprig source and aeration treatments based on a multivariate analysis of variance representing overall plant performance. These results demonstrated that pioneer sprigs from the dieback sites had an advantage over the other sprigs when exposed to unaerated systems. Tall plants performed better than plants from short or dieback zones in the aerobic systems.     

Comparisons of Mycorrhizal Responsiveness with Field Soil and Commercial Inoculum for Six Native Montane Species and Bromus tectorum

Reestablishing native perennial plants and reducing invasive species are pivotal for many ecological restoration projects. The interactions among plant species, arbuscular mycorrhizal fungi, and soil P availability may be critical determinants of the success of native and non‐native plants in restoration and species invasions. Here we assessed mycorrhizal responsiveness for three late‐successional and three early‐successional plant species native to Rocky Mountain National Park and for the non‐native Downy brome, cheatgrass (Bromus tectorum L.) using field soil and commercial inoculum. Factorial greenhouse experiments were conducted to compare biomass of plant species with and without field soil and commercial inoculum treatments along a phosphorus (P) gradient, which ranged from ambient field levels to 12% of field levels, using dilutions of native soils. The two field soil inoculum treatments resulted in significant biomass differences for all species studied. Late‐successional species responded positively to field inoculum, whereas early‐successional species responded negatively. The two commercial inocula had low colonization rates (14 of 166 inoculated plants). The commercial inocula substrates had significant treatment effects on five of seven species included in the study in the apparent absence of mycorrhizal symbiosis. Soil P levels influenced mycorrhizal responsiveness in only one species, Smooth blue aster (Aster laevis L.). Our results show that, at least for the species studied here, locally collected field inoculum is the best choice for reestablishment of late‐successional native plant species.     

Can commercial soil microbial treatments remediate plant–soil feedbacks to improve restoration seedling performance?

Seedling performance is often a limiting factor in ecological restoration. Changes in the soil microbial community generated by invasive plants contribute to seedling failure. A method to remediate invasive species‐induced changes to the soil microbial community that results in increased native species seedling performance and decreased invasive species seedling performance could have a large impact on the success of many restoration efforts. In a greenhouse experiment, we first examined the changes in the soil microbial community created by invasive compared to native grasses. Then, we investigated four microbial treatments (bacterial inoculant, fungal inoculant, fungicide, and bactericide/fungicide) to remediate microbial plant–soil feedbacks (PSFs) created by invasive species Bromus inermis and Poa pratensis and increase the performance of natives Andropogon gerardii, Elymus canadensis, Pascopyrum smithii, and Schizachyrium scoparium. We found that the PSF mitigation treatments had some context‐dependent utility for restoration. For example, all of the treatments decreased the performance of B. inermis and fungal inoculant decreased the performance of P. pratensis. However, no single treatment increased the performance of all natives. Fungicide increased the performance of A. gerardii and E. canadensis in soil previously occupied by B. inermis and the performance of S. scoparium in soil previously occupied by P. pratensis. If validated in the field, PSF mitigation treatments may have utility for restoration practitioners.     

Thermal acclimation of shoot respiration in an Arctic woody plant species subjected to 22 years of warming and altered nutrient supply

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long‐term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short‐term temperature (T) response and sensitivity of leaf respiration (R), the high‐T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q₁₀); overall, Q₁₀ values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high‐resolution measurements of R across a range of measuring Ts (25–70 °C) yielded insights into the T at which maximal rates of R occurred (T_max). Although growth temperature did not affect T_max, N+P fertilization increased T_max values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms.     

Induced responses in Ipomoea hederacea: simulated mammalian herbivory induces resistance and susceptibility to insect herbivores

Multispecies interactions between plants and natural enemies are ubiquitous, and often lead to diffuse interactions between plants and their herbivores. Non-specific induced responses, where responses induced by one species affect other species, are one potential mechanism generating diffuse interactions. Using 57 inbred lines of the Ivyleaf morning glory, Ipomoea hederacea, in a greenhouse experiment, we examined whether simulated mammalian herbivory induced responses that could affect plant resistance to the generalist insect herbivore, Spodoptera exigua. Inbred lines were highly variable for induced responses, ranging from induced resistance to induced susceptibility, with the rank-order for resistance in inbred lines changing between clipping and control treatments. We failed to detect significant genetic correlations between induced responses and trichome density, or that clipping modified the negative relationship between trichome density and Spodoptera exigua consumption and biomass. Our results suggest that non-specific induced responses can mediate the diffuse evolutionary relationship between I. hederacea and its herbivores, and that genetic variation in induced responses are an important component of this interaction. Handling Error: Heikki Hokkanen     

Biogeochemical pools and fluxes of carbon and nitrogen in a maritime tundra near penguin colonies along the Antarctic Peninsula

There is limited information regarding biogeochemical pools and fluxes in maritime tundra ecosystems along the Antarctic Peninsula. To collect baseline information on biogeochemical processes in a tundra ecosystem dominated by two vascular plant species (Colobanthus quitensis and Deschampsia antarctica) at Biscoe Point off the coast of Anvers Island, we measured pools and fluxes of C and N in transplanted tundra microcosm cores, complemented with sampling of precipitation and surface runoff. Snow and snowmelt from the tundra collection site and soil leachates from the cores were enriched with N and dissolved organic carbon compared to precipitation and snowmelt samples collected at Palmer Station, indicating high loading of N and organic matter from the penguin colonies adjacent to the tundra site. Relatively high values of δ¹⁵N in the live and dead biomass of D. antarctica and C. quitensis (5.6–25.1‰) indicated an enrichment of N in this tundra ecosystem, possibly through N inputs from adjacent penguin colonies. Stepwise multiple linear regressions found that ecosystem respiration and gross primary production were best predicted by live biomass of D. antarctica, suggesting a disproportionately high contribution of D. antarctica to CO₂ fluxes. The cores with higher δ¹⁵N and lower δ¹³C in the soil organic horizon exhibited higher CO₂ fluxes. The results suggest that abundant N inputs from penguin colonies and the competitive balance between plant species might play a critical role in the response of tundra ecosystems along the Antarctic Peninsula to projected climate change.