Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change

Global environmental change, related to climate change and the deposition of airborne N‐containing contaminants, has already resulted in shifts in plant community composition among plant functional types in Arctic and temperate alpine regions. In this paper, we review how key ecosystem processes will be altered by these transformations, the complex biological cascades and feedbacks that might result, and some of the potential broader consequences for the earth system. Firstly, we consider how patterns of growth and allocation, and nutrient uptake, will be altered by the shifts in plant dominance. The ways in which these changes may disproportionately affect the consumer communities, and rates of decomposition, are then discussed. We show that the occurrence of a broad spectrum of plant growth forms in these regions (from cryptogams to deciduous and evergreen dwarf shrubs, graminoids and forbs), together with hypothesized low functional redundancy, will mean that shifts in plant dominance result in a complex series of biotic cascades, couplings and feedbacks which are supplemental to the direct responses of ecosystem components to the primary global change drivers. The nature of these complex interactions is highlighted using the example of the climate‐driven increase in shrub cover in low‐Arctic tundra, and the contrasting transformations in plant functional composition in mid‐latitude alpine systems. Finally, the potential effects of the transformations on ecosystem properties and processes that link with the earth system are reviewed. We conclude that the effects of global change on these ecosystems, and potential climate‐change feedbacks, cannot be predicted from simple empirical relationships between processes and driving variables. Rather, the effects of changes in species distributions and dominances on key ecosystem processes and properties must also be considered, based upon best estimates of the trajectories of key transformations, their magnitude and rates of change.     

Nitrogen‐dependent recovery of subarctic tundra vegetation after simulation of extreme winter warming damage to Empetrum hermaphroditum

Vast areas of (sub)arctic tundra are dominated by the ericoid dwarf shrub Empetrum hermaphroditum. Recent experimental and observational data have shown that Empetrum can be damaged heavily by recurrent extreme winter warming. In addition, summer warming leads to increased soil N availability in tundra ecosystems. In a 7‐year experiment, I investigated the recovery of subarctic Empetrum‐dominated tundra vegetation using a factorial combination of various degrees of aboveground Empetrum removal (simulating the damaging effects of extreme winter warming) and N addition (simulating one of the effects of summer warming). After 7 years no new species had established in the plots. The growth of planted Betula nana seedlings was stimulated by Empetrum removal and reduced by N addition. This Empetrum‐dominated tundra ecosystem was resilient against severe disturbances. Only when Empetrum was 100% removed did it fail to recover, and only in combination with high N supply the subordinate species (notably Eriophorum vaginatum and Rubus chamaemorus, a graminoid and a forb) could benefit. In the 50% removal treatment Empetrum recovered in 7 years when no N was supplied and the cover of the subordinate species did not change. However, when N was added Empetrum recovered faster (in 4 years) and the subordinates decreased. When Empetrum was not removed and N was added, Empetrum even increased in abundance at the expense of the subordinate species. Thus, profound changes in tundra ecosystems can only be expected when Empetrum is very heavily damaged as a result of recurrent extreme winter warming and when soil N availability is increased as a result of summer warming. These changes in species composition upon extreme disturbance events may lead to a wide variety of ecosystem feedbacks and cascade processes as this tundra system is relatively species‐poor, and can be hypothesized to have low functional redundancy.     

Global mapping of Jatropha curcas yield based on response of fitness to present and future climate

Although acclaimed as a biofuel crop with high potential to sustainably replace fossil fuels, Jatropha curcas L. remains a poorly studied plant. Reliable yield assessments with conventional methods require agroclimatic and physiological knowledge, which is not yet available for Jatropha. To fill this gap, we tested a novel two‐step approach integrating knowledge from biogeography and population biology with available Jatropha field data. In the first step, using MaxEnt, a widely implemented model in biogeography, we predicted Jatropha fitness in response to climate by relating natural occurrence recorded in herbaria with bioclimatic geodatasets. In the second step, we relied on population biology principles supported by seed mass addition experiments to relate fitness to reproductive potential, hence seed yield. Jatropha seed yield in response to climate was mapped worldwide for actual (1950–2000 average) and future (2020) climate conditions. The modelled Jatropha seed yield was validated against a set of on‐field yield assessments (R²=0.67, P<0.001). The discrepancies between estimated and measured yields were partially explained by model uncertainties, as quantified by the sensitivity analysis of our modelling (R²=0.57, P=0.001). Jatropha has a pan‐tropical distribution, plus specific adaptability to hot temperate areas. Climate variables most significantly affecting modelled yield response were annual average temperature, minimum temperature, annual precipitation and precipitation seasonality.     

A Race for Space? How Sphagnum fuscum stabilizes vegetation composition during long‐term climate manipulations

Strong climate warming is predicted at higher latitudes this century, with potentially major consequences for productivity and carbon sequestration. Although northern peatlands contain one‐third of the world's soil organic carbon, little is known about the long‐term responses to experimental climate change of vascular plant communities in these Sphagnum‐dominated ecosystems. We aimed to see how long‐term experimental climate manipulations, relevant to different predicted future climate scenarios, affect total vascular plant abundance and species composition when the community is dominated by mosses. During 8 years, we investigated how the vascular plant community of a Sphagnum fuscum‐dominated subarctic peat bog responded to six experimental climate regimes, including factorial combinations of summer as well as spring warming and a thicker snow cover. Vascular plant species composition in our peat bog was more stable than is typically observed in (sub)arctic experiments: neither changes in total vascular plant abundance, nor in individual species abundances, Shannon's diversity or evenness were found in response to the climate manipulations. For three key species (Empetrum hermaphroditum, Betula nana and S. fuscum) we also measured whether the treatments had a sustained effect on plant length growth responses and how these responses interacted. Contrasting with the stability at the community level, both key shrubs and the peatmoss showed sustained positive growth responses at the plant level to the climate treatments. However, a higher percentage of moss‐encroached E. hermaphroditum shoots and a lack of change in B. nana net shrub height indicated encroachment by S. fuscum, resulting in long‐term stability of the vascular community composition: in a warmer world, vascular species of subarctic peat bogs appear to just keep pace with growing Sphagnum in their race for space. Our findings contribute to general ecological theory by demonstrating that community resistance to environmental changes does not necessarily mean inertia in vegetation response.     

On the sex-specific mechanisms of butterfly flight: flight performance relative to flight morphology, wing kinematics, and sex in Pararge aegeria

Many evolutionary ecological studies have documented sexual dimorphism in morphology or behaviour. However, to what extent a sex-specific morphology is used differently to realize a certain level of behavioural performance is only rarely tested. We experimentally quantified flight performance and wing kinematics (wing beat frequency and wing stroke amplitude) and flight morphology (thorax mass, body mass, forewing aspect ratio, and distance to centre of forewing area) in the butterfly Pararge aegeria (L.) using a tethered tarsal reflex induced flight set-up under laboratory conditions. On average, females showed higher flight performance than males, but frequency and amplitude did not differ. In both sexes, higher flight performance was partly determined by wing beat frequency but not by wing stroke amplitude. Dry body mass, thorax mass, and distance to centre of forewing area were negatively related to wing beat frequency. The relationship between aspect ratio and wing stroke amplitude was sex-specific: females with narrower wings produced higher amplitude whereas males show the opposite pattern. The results are discussed in relation to sexual differences in flight behaviour.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 675–687.     

A functional analysis of how frogs jump out of water

Euphlyctis cyanophlyctis and E. hexadactylus are two common frog species from south-east Asia. Both species have the remarkable ability to leap from a floating position out of the water. Rana esculenta is a European species from the same family that barely manages to do so. The species' morphology, however, looks roughly the same. We studied the unique abilities of the Euphlyctis species by videotaping the three species mentioned above whilst they jumped out of the water. These recordings enabled us to study the hind leg movements and to test several hypotheses. Differences between the species only became apparent when details of the behaviour were analysed. The three species mainly differed in their slip factor at the start of the propulsive phase, resulting in a difference in slip distance. This difference could be traced back to the difference in web shape by means of a physical model. The Euphlyctis species can prevent their feet from slipping because of their rounded and convex webbing, resulting in a larger take-off velocity and thereby a successful emergence out of the water.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 413–420.