The Inverted Phantom Giant

In his famous children’s book, “Jim Button and Luke the Engine Driver”, Michael Ende describes a curious character: A phantom giant. Clothed in rags and with a long beard, the phantom giant appears enormous from far away, but shrinks to normal size as one gets closer. Most people avoid the poor creature, but the ones that dare approach it encounter a gentle, lonely being called Mr. Tur Tur. Chemical ecology is just the opposite of Mr. Tur Tur: A phantom dwarf. Or, in other words, an inverted phantom giant. From a distance, chemical ecology appears like a slightly odd, marginal section of biology and chemistry. But, as the interested scholar approaches, it starts growing and very quickly reaches gigantic dimensions, because all life is explained by chemistry, and all biological chemistry is guided by ecological principles. Herein lies the difficulty with chemical ecology: As it is not perceived well by biologists and chemists, few approach it to understand its significance, and the ones that do find themselves in front of a giant that defies their attempts to define and contain it. This is where the Journal of Chemical Ecology comes in: It invites us to take a closer look at an underestimated discipline and supports us to explore it and deal with its multidimensionality through the promotion of knowledge and methods. These services are unique and make the journal stand out of the crowd of scientific journals. Writing children’s books has become difficult in the era of information technology. And, so has the job of the Journal of Chemical Ecology. Young scientists gather information through accessible, dynamic websites and social platforms. They want articles that are available through a single mouse click, anywhere, anytime. They prefer advanced interactive hypertext protocols over clumsy pdf files. They care about transparency, non-profit and open access just as much as about traditional journal properties. In my view, reaching “the kids” is the major challenge of the Journal over the next years. Promoting an inverted phantom giant in the 21st century requires a combination of high-quality information and boosted visibility. In Michael Ende’s book, Jim and Luke follow exactly this strategy with Mr. Tur Tur: They become friends and offer him a job as a living lighthouse to protect their small island. They combine a quality relationship with high visibility, et voilà, the story ends well! I am looking forward to seeing if the Journal of Chemical Ecology will follow a similar path to reach the next generation of biologists and chemists. If yes, there is a good chance that in 40 years from now, somebody will write a laudation and refer to another famous book by Michael Ende: “The Neverending Story”.

A nearby M Star with tree Transitting Supter-Earths Discovered by K2

Small, cool planets represent the typical end-products of planetary formation. Studying the architectures of these systems, measuring planet masses and radii, and observing these planets' atmospheres during transit directly informs theories of planet assembly, migration, and evolution. Here we report the discovery of three small planets orbiting a bright (K-s = 8.6 mag) M0 dwarf using data collected as part of K2, the new ecliptic survey using the repurposed Kepler spacecraft. Stellar spectroscopy and K2 photometry indicate that the system hosts three transiting planets with radii 1.5-2.1 R-circle plus, straddling the transition region between rocky and increasingly volatile-dominated compositions. With orbital periods of 10-45 days the planets receive just 1.5-10x the flux incident on Earth, making these some of the coolest small planets known orbiting a nearby star; planet d is located near the inner edge of the system's habitable zone. The bright, low-mass star makes this system an excellent laboratory to determine the planets' masses via Doppler spectroscopy and to constrain their atmospheric compositions via transit spectroscopy. This discovery demonstrates the ability of K2 and future space-based transit searches to find many fascinating objects of interest.

From stellar to planetary composition: Galactic chemical evolution of Mg/Si mineralogical ratio

Aims. The main goal of this work is to study element ratios that are important for the formation of planets of different masses. Methods. We study potential correlations between the existence of planetary companions and the relative elemental abundances of their host stars. We use a large sample of FGK-type dwarf stars for which precise Mg, Si, and Fe abundances have been derived using HARPS high-resolution and high-quality data. Results. A first analysis of the data suggests that low-mass planet host stars show higher [Mg/Si] ratios, while giant planet hosts present [Mg/Si] that is lower than field stars. However, we found that the [Mg/Si] ratio significantly depends on metallicity through Galactic chemical evolution. After removing the Galactic evolution trend only the difference in the [Mg/Si] elemental ratio between low-mass planet hosts and non-hosts was present in a significant way. These results suggest that low-mass planets are more prevalent around stars with high [Mg/Si]. Conclusions. Our results demonstrate the importance of Galactic chemical evolution and indicate that it may play an important role in the planetary internal structure and composition. The results also show that abundance ratios may be a very relevant issue for our understanding of planet formation and evolution.

Gibberellin Deficiency Confers Both Lodging and Drought Tolerance in Small Cereals

Tef [Eragrostis tef (Zucc.) Trotter] and finger millet [Eleusine coracana Gaertn] are staple cereal crops in Africa and Asia with several desirable agronomic and nutritional properties. Tef is becoming a life-style crop as it is gluten-free while finger millet has a low glycemic index which makes it an ideal food for diabetic patients. However, both tef and finger millet have extremely low grain yields mainly due to moisture scarcity and susceptibility of the plants to lodging. In this study, the effects of gibberellic acid (GA) inhibitors particularly paclobutrazol (PBZ) on diverse physiological and yield-related parameters were investigated and compared to GA mutants in rice (Oryza sativa L.). The application of PBZ to tef and finger millet significantly reduced the plant height and increased lodging tolerance. Remarkably, PBZ also enhanced the tolerance of both tef and finger millet to moisture deficit. Under moisture scarcity, tef plants treated with PBZ did not exhibit drought-related symptoms and their stomatal conductance was unaltered, leading to higher shoot biomass and grain yield. Semi-dwarf rice mutants altered in GA biosynthesis, were also shown to have improved tolerance to dehydration. The combination of traits (drought tolerance, lodging tolerance and increased yield) that we found in plants with altered GA pathway is of importance to breeders who would otherwise rely on extensive crossing to introgress each trait individually. The key role played by PBZ in the tolerance to both lodging and drought calls for further studies using mutants in the GA biosynthesis pathway in order to obtain candidate lines which can be incorporated into crop-breeding programs to create lodging tolerant and climate-smart crops.

Conditional expression of Sry, and Wnt4 by gene-targeting: Studies in sexual and skeletal development

Sry and Wnt4 cDNAs were individually introduced into the ubiquitously-expressed Rosa26 ( R26) locus by gene targeting in embryonic stem (ES) cells to create a conditional gene expression system in mice. In the targeted alleles, expression of these cDNAs should be blocked by a neomycin resistance selection cassette that is flanked by loxP sites. Transgene expression should be activated after the blocking cassette is deleted by Cre recombinase. ^ To test this conditional expression system, I have bred R26-stop- Sry and R26-stop-Wnt4 heterozygotes with a MisRII-Cre mouse line that expresses Cre in the gonads of both sexes. Analysis of these two types of bigenic heterozygotes indicated that their gonads developed normally like those of wild types. However, one XX R26-Sry/R26-Sry; MisR2-Cre/+ showed epididymis-like structures resembling those of males. In contrast, only normal phenotypes were observed in XY R26-Wnt4/R26-Wnt4; MisR2-Cre /+ mice. To interpret these results, I have tested for Cre recombinase activity by Southern blot and transcription of the Sry and Wnt4 transgenes by RT-PCR. Results showed that bigenic mutants had insufficient activation of the transgenes in their gonads at E12.5 and E13.5. Therefore, the failure to observe mutant phenotypes may have resulted from low activity of MisR2-Cre recombination at the appropriate time. ^ Col2a1-Cre transgenic mice express Cre in differentiating chondrocytes. R26-Wnt4; Col2a1-Cre bigenic heterozygous mice were found to exhibit a dramatic alteration in growth presumably caused by Wnt4 overexpression during chondrogenesis. R26-Wnt4; Col2a1-Cre mice exhibited dwarfism beginning approximately 10 days after birth. In addition, they also had craniofacial abnormalities, and had delayed ossification of the lumbar vertebrate and pelvic bones. Histological analysis of the growth plates of R26-Wnt4; Col2a1-Cre mice revealed less structural organization and a delay in onset of the primary and secondary ossification centers. Molecular studies confirmed that overexpression of Wnt4 causes decreased proliferation and early maturation of chondrocytes. In addition, R26-Wnt4; Col2a1-Cre mice had decreased expression of vascular endothelial growth factor (VEGF), suggesting that defects in vascularization may contribute to the dwarf phenotype. Finally, 9-month-old R26-Wnt4; Col2a1-Cre mice had significantly more fat cells in the marrow cavities of their metaphysis long bones, implying that long-term overexpression of Wnt4may cause bone marrow pathologies. In conclusion, Wnt4 was activated by Col2a1-Cre recombinase and was overexpressed in the growth plate, resulting in aberrant proliferation and differentiation of chondrocytes, and ultimately leads to dwarfism in mice. ^

A sex difference in seasonal timing of birth in a livebearing fish

Sex differences in seasonal timing include differences in hatch- or birth-date distribution and differences in the timing of migration or maturation such as protandrous arrival timing (PAT), which is early male arrival at breeding sites. I describe a novel form of protandrous arrival timing, as a sex difference in birth-date distribution in a live-bearing fish (Dwarf Perch, Micrometrus minimus). In this species, birth coincides with arrival at breeding sites because newborn males are sexually active. A series of samples of pregnant females and young of year was collected in Tomales Bay, CA. I analyzed the daily age record in otoliths to estimate the conception date of broods and the age that young-of-year individuals were born. Males were born at a younger age than females, as indicated by the daily age record and also by the predominance of females in broods from which some young had already been born, which was a common occurrence in pregnant females with older embryos. Sex ratio of broods varied with conception date such that early-season broods were predominantly male, possibly as a result of temperature-dependent sex determination. The combined effects of the sex difference in age at birth and seasonal shift in sex ratio were to shift the mean birth date of males relative to females by five days. The most likely ultimate explanation for PAT in the Dwarf Perch is that it arises from exploitation (scramble) competition for mating opportunities among recently-born young-of-year males.

Roles of p38 mitogen-activated protein kinase pathway in the regulation of SOX9 activity and chondrogenesis

Sox9 is a master transcription factor in chondrocyte differentiation. Several lines of evidence suggest that the p38 mitogen-activated protein kinase (MAPK) pathway is involved in chondrocyte differentiation. In the present study, we examined the roles of p38 in the regulation of SOX9 activity and chondrogenesis. ^ COS7 cells were transfected with a SOX9 expression vector and 4x48-p89, a luciferase construction harboring four tandem copies of a SOX9-dependent 48-bp enhancer in Col2a1. Coexpression of MKK6EE, a constitutively active mutant of MKK6, a MAPKK that specifically activates p38, further increased the activity of the SOX9-dependent 48-bp enhancer about 5-fold, and SOX9 protein levels were not increased under these conditions. This increase in enhancer activity was not observed in a mutant enhancer construct harboring mutations that abolish SOX9 binding. These data strongly suggested that activation of the p38 pathway results in increased activity of SOX9. In addition, the increase of the activity of the SOX9-dependent 48-bp enhancer by MKK6EE was also observed in primary chondrocytes, and this increase was abolished by coexpression of a p38 phosphatase, MKP5, and p38 specific inhibitors. Furthermore, treatment of primary chondrocytes with p38 inhibitors decreased the expression of Col2a1, a downstream target of Sox9, without affecting Sox9 RNA levels, further supporting the hypothesis that p38 plays a role in regulating Sox9 activity in chondrocytes. ^ To further study the role of the p38 MAPK pathway in chondrogenesis, we generated transgenic mice that express MKK6EE in chondrocytes under the control of the Col2a1 promoter/intron regulatory sequences. These mice showed a dwarf phenotype characterized by reduced chondrocyte proliferation and a delay in the formation of primary and secondary ossification centers. Histological analysis using in situ hybridization showed reduced expression of Indian hedgehog, PTH/PTHrP receptor, cyclin D1 and increased expression of p21. In addition, consistent with the notion that Sox9 activity was increased in these mice, transgenic mice that express MKK6EE in chondrocytes showed phenotypes similar to those of mice that overexpress SOX9 in chondrocytes. Therefore, our study provides in vivo evidence for the role of p38 in chondrocyte differentiation and suggests that Sox9 is a downstream target of the p38 MAPK pathway. ^

Climate and soil characteristics and soil arthropod abundance on Anchorage, Signy and Falkland Islands

Over a 2-year study, we investigated the effect of environmental change on the diversity and abundance of soil arthropod communities (Acari and Collembola) in the Maritime Antarctic and the Falkland Islands. Open Top Chambers (OTCs), as used extensively in the framework of the northern boreal International Tundra Experiment (ITEX), were used to increase the temperature in contrasting communities on three islands along a latitudinal temperature gradient, ranging from the Falkland Islands (51°S, mean annual temperature 7.5 °C) to Signy Island (60°S, -2.3°C) and Anchorage Island (67°S, -3.8°C). At each island an open and a closed plant community were studied: lichen vs. moss at the Antarctic sites, and grass vs. dwarf shrub at the Falkland Islands. The OTCs raised the soil surface temperature during most months of the year. During the summer the level of warming achieved was 1.7 °C at the Falkland Islands, 0.7 °C at Signy Island, and 1.1 °C at Anchorage Island. The native arthropod community diversity decreased with increasing latitude. In contrast with this pattern, Collembola abundance in the closed vegetation (dwarf shrub or moss) communities increased by at least an order of magnitude from the Falkland Islands (9.0 +/- 2 x 10**3 ind./m**2) to Signy (3.3 +/- 8.0 x 10**4 ind./m**2) and Anchorage Island (3.1 +/- 0.82 x 10**5 ind./m**2). The abundance of Acari did not show a latitudinal trend. Abundance and diversity of Acari and Collembola were unaffected by the warming treatment on the Falkland Islands and Anchorage Island. However, after two seasons of experimental warming, the total abundance of Collembola decreased (p < 0.05) in the lichen community on Signy Island as a result of the population decline of the isotomid Cryptopygus antarcticus. In the same lichen community there was also a decline (p < 0.05) of the mesostigmatid predatory mite Gamasellus racovitzai, and a significant increase in the total number of Prostigmata. Overall, our data suggest that the consequences of an experimental temperature increase of 1-2°C, comparable to the magnitude currently seen through recent climate change in the Antarctic Peninsula region, on soil arthropod communities in this region may not be similar for each location but is most likely to be small and initially slow to develop.

(Table 1) Normalized difference vegetation index in control and warming plots, Abisco Research Station

Extreme winter warming events in the sub-Arctic have caused considerable vegetation damage due to rapid changes in temperature and loss of snow cover. The frequency of extreme weather is expected to increase due to climate change thereby increasing the potential for recurring vegetation damage in Arctic regions. Here we present data on vegetation recovery from one such natural event and multiple experimental simulations in the sub-Arctic using remote sensing, handheld passive proximal sensors and ground surveys. Normalized difference vegetation index (NDVI) recovered fast (2 years), from the 26% decline following one natural extreme winter warming event. Recovery was associated with declines in dead Empetrum nigrum (dominant dwarf shrub) from ground surveys. However, E. nigrum healthy leaf NDVI was also reduced (16%) following this winter warming event in experimental plots (both control and treatments), suggesting that non-obvious plant damage (i.e., physiological stress) had occurred in addition to the dead E. nigrum shoots that was considered responsible for the regional 26% NDVI decline. Plot and leaf level NDVI provided useful additional information that could not be obtained from vegetation surveys and regional remote sensing (MODIS) alone. The major damage of an extreme winter warming event appears to be relatively transitory. However, potential knock-on effects on higher trophic levels (e.g., rodents, reindeer, and bear) could be unpredictable and large. Repeated warming events year after year, which can be expected under winter climate warming, could result in damage that may take much longer to recover.

Soil characteristics and Collembola and Acari abundance in control and warming plots at Abisco Research Station

Extreme weather events can have negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (2-10 °C for 2-14 days), but returning to cold winter climate exposes the ecosystem to lower temperatures by the loss of insulating snow. Soil animals, which play an integral part in soil processes, may be very susceptible to such events depending on the intensity of soil warming and low temperatures following these events. We simulated week-long extreme winter warming events - using infrared heating lamps, alone or with soil warming cables - for two consecutive years in a sub-Arctic dwarf shrub heathland. Minimum temperatures were lower and freeze-thaw cycles were 2-11 times more frequent in treatment plots compared with control plots. Following the second event, Acari populations decreased by 39%; primarily driven by declines of Prostigmata (69%) and the Mesostigmatic nymphs (74%). A community-weighted vertical stratification shift occurred from smaller soil dwelling (eu-edaphic) Collembola species dominance to larger litter dwelling (hemi-edaphic) species dominance in the canopy-with-soil warming plots compared with controls. The most susceptible groups to these winter warming events were the smallest individuals (Prostigmata and eu-edaphic Collembola). This was not apparent from abundance data at the Collembola taxon level, indicating that life forms and species traits play a major role in community assembly following extreme events. The observed shift in soil community can cascade down to the micro-flora affecting plant productivity and mineralization rates. Short-term extreme weather events have the potential to shift community composition through trait composition with potentially large consequences for ecosystem development.

Pollen record and age determination of a sediment profile from Frengstadsetra, Norway

Innerdalen was once a mountain valley (ca. 780 m a.s.l.) with birch forests, bogs and several summer farms. Today it is a 6.5 km**2 artifical lake. In 1980 and 1981 archaeological and palynological investigations were carried out due to the hydroelectric power plans. Radiocarbon dated pollen diagrams from 9 different localities in Innerdalen provide information on a mountain environment which has been exploited to varying degrees by human groups for thousands of years. In the Birch Zone, ca. 9500-8500 years B.P., the deglaciated surface is vegetated by the normal sequence of pioneering species, first show-bed communities, then shrub/dwarf-shrub communities, and finally a birch forest community. In the Pine Zone, ca. 8500-7500 years B.P., the mixed Birch-Pine forest which prevailed at the end of the Birch Zone is replaced by a dense pine forest. The tree limit was higher than it is today. In the Alder Zone, ca. 7500-4000 years B.P., the newly arrived alder gradually succeeded pine, particularily on good soils. This alder forest has a modem analog in the pre-alpine gray alder forests in Norway. In the last part of the Alder Zone, ca. 6000-4000 years B.P., elm and hazel are nominally present on particularily rich soils, marking the edaphic and climatic optimum in Innerdalen. During this time the first evidence of human impact on the vegetation is apparent in the pollen diagrams. At both Sætersetra in the south of the valley and Liabekken in the north, forest clearance and the development of grazed grass meadows is documented, and human impact continues until the present. The Herb Zone, ca. 4000 years B.P. to 1600 A.D., is characterized by the rapid decline of alder. The forest is increasingly open, and bog formation is initiated. The sub-alpine belt of birch forest is established, probably due to the shift to a cooler, moister climate. Human activity can also have influenced the vegetational changes, although at 4 of the localities human activity also is first apparent after the alder decline. Some localities show measurably less human impact on the vegetation ca. 2600-2000 years B.P. Grazing intensity increases ca. 2000 years B.P. At the end of the Herb Zone rye and barley pollen is registered at Sætersetra and Flonan, indicating contact between the grazing activities of Innerdal and grain cultivation activities outside the valley. The Spruce Zone, ca. 1600 A.D. to the present, does not begin synchronously since the presence of long-distance transported spruce pollen at a locality is entirely dependent on the density of the vegetation ie. degree of human impact. The youngest spruce rise is ca. 1500 A.D. at Røstvangen, when summerfarming is initiated. Summerfarming activities in Innerdal produce an increasingly open landscape. Rye and barley pollen at several localities may indicate limited local cultivation, but is more likely long-distance transport via humans and domesticated animals from cultivated areas outside Innerdalen.

Strontium isotopic ratios of evaporites of the Tyrrhenian Sea

Strontium isotopic ratios of gypsums recovered from upper Miocene (Messinian) evaporites at ODP Leg 107 Holes 652A, 653B, and 654A (Tyrrhenian Sea) are lower than expected. The values for the Messinian balatino-like gypsum, single gypsum crystals, and anhydrites range from 0.70861 to 0.70886 and are approximately 25 * 10**-5 less than would be expected for evaporites precipitated from Messinian seawater (0.70891-0.70902). Pre-evaporitic planktonic foraminifers from Hole 654A show variable degrees of dolomitization and 87Sr/86Sr values that irregularly decrease upward from normal marine values approximately 81m below the lowest evaporite occurrence. This suggests diagenetic alteration by advecting interstitial water with a low 87Sr/86Sr ratio or that the lower Sr isotopic ratios for the Messinian evaporites could have resulted from a greater influence of fresh water on the Sr isotopic composition of the desiccating Tyrrhenian Sea. Fluctuations of the 87Sr/86Sr-ratio for evaporites in the sedimentary cycles recognized for Holes 653B and 654A, the generally low Sr isotopic ratio of river water entering the Mediterranean Sea, and the presence of dwarf marine microfossils suggest that the 87Sr/86Sr ratio of the evaporites responded to hydrologic variations in a very restricted basin with variable rates of marine and fresh water input. The strontium isotopic ratios of the Messinian anhydrites from the proposed lacustrine sequence at Hole 652A fall in the same range as the marine evaporites from Holes 654A and 653B. This suggests a common or similar origin of the brines at the three locations. The complex depositional and hydrologic conditions in the Mediterranean during the Messinian salinity crisis preclude the use of Sr isotopic values from the evaporites for stratigraphic correlation and dating. They are, however, very useful in the interpretation of the depositional history of the basin. General calculations assuming a closed system suggest that the 87Sr/86Sr ratio of Messinian seawater (-0.7090) could be reduced to that of the evaporites (-0.7087) by mixing with fresh water (e.g., Nile River) in times of 10**4 to 10**5 yr.

Palynology on sediment profile Tso Kar in Ladakh

Palynological investigation of a 410 cm long core section from Tso Kar (33°10'N, 78°E, 4527 m a.s.l.), an alpine lake situated in the arid Ladakh area of NW India at the limit of the present-day Indian summer monsoon, was performed in order to reconstruct post-glacial regional vegetation and climate dynamics. The area was covered with alpine desert vegetation from ca. 15.2 to 14 kyr BP (1 kyr=1000 cal. years), reflecting dry and cold conditions. High influx values of long-distance transported Pinus sylvestris type pollen suggest prevailing air flow from the west and northwest. The spread of alpine meadow communities and local aquatic vegetation is a weak sign of climate amelioration after ca. 14 kyr BP. Pollen data (e.g. influx values of Pinus roxburghii type and Quercus) suggest that this was due to a strengthening of the summer monsoon and the reduced activity of westerly winds. The further spread of Artemisia and species-rich meadows occurred in response to improved moisture conditions between ca. 12.9 and 12.5 kyr BP. The subsequent change towards drier desert-steppe vegetation likely indicates more frequent westerly disturbances and associated snowfalls, which favoured the persistence of alpine meadows on edaphically moist sites. The spread of Chenopodiaceae-dominated vegetation associated with an extremely weak monsoon occurred at ca. 12.2-11.8 kyr BP during the Younger Dryas interstadial. A major increase in humidity is inferred from the development of Artemisia-dominated steppe and wet alpine meadows with Gentianaceae after the late glacial/early Holocene transition in response to the strengthening of the summer monsoon. Monsoonal influence reached maximum activity in the Tso Kar region between ca. 10.9 and 9.2 kyr BP. The subsequent development of the alpine meadow, steppe and desert-steppe vegetation points to a moderate reduction in the moisture supply, which can be linked to the weaker summer monsoon and the accompanying enhancement of the winter westerly flow from ca. 9.2 to 4.8 kyr BP. The highest water levels of Tso Kar around 8 kyr BP probably reflect combined effect of both monsoonal and westerly influence in the region. An abrupt shift towards aridity in the Tso Kar region occurred after ca. 4.8 kyr BP, as evidenced by an expansion of Chenopodiaceae-dominated desert-steppe. Low pollen influx values registered ca. 2.8-1.3 kyr BP suggest scarce vegetation cover and unfavourable growing conditions likely associated with a further weakening of the Indian Monsoon.

Carbon/Nitrogen ratio of plant tissues and biomass of belowground consumers after 11 years of experimental treatment, Toolik lake

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The Exploitation Ecosystem Hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.

Initial biomass and biomass change of tundra and forest plants in three herbivore treatments in Norway and Sweden

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana, has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (> 0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (< 0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.