Modelling the effect of El Niño on the persistence of small populations: The Galápagos penguin as a case study

Small populations are vulnerable to long-term declines, even where short-term censuses indicate increasing trends in numbers. Census data for the Galápagos penguin (Spheniscus mendiculus) collected between 1970 and 2004 provide evidence that despite year-to-year population increases detected in most of the annual censuses, the strong El Niño events of 1982-83 and 1997-98 were followed by population declines of more than 60% from which the species has yet to recover. Such large declines raise concerns about the future viability of the species because the frequency and severity of El Niño events are predicted to increase. We used the simulation software VORTEX to evaluate the potential effects of El Niño on the risk of extinction of the Galápagos penguin population and its four constituent subpopulations. Weak and strong El Niño events were treated as catastrophes, with varying frequencies, which simulated past, current and future effects on the penguin population. The “Current El Niño” scenario, based on the frequency of El Niño events recorded in the Galápagos between 1965 and 2004, indicated an approximately 30% probability of extinction within the next 100 years for the penguin population. More ominously, the species may be at a greater risk if the frequency of strong El Niño episodes increases only marginally. A probability of extinction greater than 80% was predicted when the current frequency (5%) of strong El Niño events was doubled (to 10%). The probabilities of extinctions were higher for each subpopulation treated individually, ranging from 34% for Isabela and Fernandina, 64% for Bartolomé-Santiago to 78% for the smallest subpopulation on Floreana. Sensitivity analyses identified survival of penguins during El Niño events and sex ratio as influential parameters. The estimates of extinction risk may be conservative as other threats associated with increased human activities on the islands may further compromise species persistence.     

Evaluating effects of deforestation, hunting, and El Niño events on a threatened lemur

Madagascar ranks as one of the world’s top extinction hotspots because of its high endemism and high rate of habitat degradation. Global climate phenomena such as El Niño Southern Oscillations may have confounding impacts on the island’s threatened biota but these effects are less well known. We performed a demographic study of Propithecus edwardsi, a lemur inhabiting the eastern rainforest of Madagascar, to evaluate the impact of deforestation, hunting, and El Niño on its population and to re-evaluate present endangerment categorization under the IUCN. Over 18 years of demographic data, including survival and fecundity rates were used to parameterize a stochastic population model structured with three stage classes (yearlings, juveniles, and adults). Results demonstrate that hunting and deforestation are the most significant threats to the population. Analysis of several plausible scenarios and combinations of threat revealed that a 50% population decline within three generations was very likely, supporting current IUCN classification. However, the analysis also suggested that changing global cycles may pose further threat. The average fecundity of lemurs was over 65% lower during El Niño years. While not as severe as deforestation or hunting, if El Niño events remain at the current high frequency there may be negative consequences for the population. We suggest that it is most critical for this species continued survival to create more protected areas, not only to thwart hunting and deforestation, but also to give this endangered lemur a better chance to recover from and adapt to altered climate cycles in the future.     

Incidental and intentional catch threatens Galápagos waved albatross

As large, long-lived seabirds with delayed and slow reproduction, albatrosses have low intrinsic mortality rates and are especially vulnerable to extinction from extrinsic sources of mortality such as fishery bycatch. Leg-band recovery information for waved albatrosses revealed mortality from both incidental catch and intentional catch for human consumption. Annual adult survival in 1999-2005, estimated from capture-mark-recapture data, was lower than historical estimates. This recent increase in adult mortality probably contributed to recent and dramatic shrinkage of the breeding population; periodic matrix models confirm that population growth rate is most sensitive to changes in adult survival. Banding data and recovery information also suggest that capture by fisheries is male-biased, which should reduce fecundity in this species with obligate bi-parental care. This new documentation of bycatch, harvesting, and associated demographic consequences provides reason for serious concern about the persistence of the single breeding population of the waved albatross.     

Assessing the risks of introduced chickens and their pathogens to native birds in the Galápagos Archipelago

Poultry production is an important economic activity on inhabited islands of the Galápagos archipelago. There has been a recent surge in both small-scale backyard chickens and larger scale broiler production associated with growth in the human population and the tourist industry. With increased poultry production, concerns have been expressed about the increasing risk of transfer of disease from chickens to native Galápagos bird species that may have little resistance to introduced pathogens [Wikelski, M., Foufopoulos, J., Vargas, H., Snell, H., 2004. Galápagos birds and diseases: invasive pathogens as threats for island species. Ecology and Society 9(5). Available from: URL:http://www.ecologyandsociety.org/vol9/iss1/art5]. This study evaluates risks posed by chicken disease to endemic and native Galápagos bird species, based on empirical evidence of pathogens present in chickens on the islands and a literature review of effects of these pathogens in wild species. Pathogens identified in domestic chicken populations of immediate avian conservation concern are Newcastle disease, Mycoplasma gallisepticum, and the proventricular parasite Dispharynx sp. Newcastle disease (avian paramyxovirus-1) poses an imminent threat to Galápagos penguins (Spheniscus mendiculus), flightless cormorants (Phalacrocorax harrisi), and lava gulls (Larus fuliginosus), species with very small population sizes (less than 1500 animals each). Additionally, litter from broiler farms could affect ecological processes in local ecosystems. Improved poultry biosecurity measures are urgently needed on the Galápagos Islands for avian disease management, yet developing these strategies presents political, social, and economic challenges.     

Spatial and temporal variation in the Argentine ant edge effect: Implications for the mechanism of edge limitation

One consequence of human land-use is the exposure of native communities to invasive human commensal species along edges. Argentine ants (Linepithema humile) invade a variety of habitats in California with consequent dramatic declines in native ants. In coastal southern California, USA Argentine ants appear to be an edge effect in small habitat fragments [Suarez, A.V., Bolger, D.T., Case, T.J., 1998. Effects of fragmentation and invasion on native ant communities in coastal southern California. Ecology 79, 2041-2056]. They invade fragments from the urban edge, but only penetrate coastal sage scrub (CSS) habitat to a distance of approximately 200 m. Using pitfall sampling in edge (<250 m from the urban edge) and interior (>600 m from the edge) habitat I tested whether there is also an Argentine ant edge effect in the largest blocks of habitat in the landscape and investigated patterns of spatial and temporal variation in native and Argentine ants. Argentine ants were common in coastal sage scrub habitat within 250 m of urban edges, but rare in interior areas. Correspondingly, native ants were significantly less abundant and diverse in edge areas as compared to interior. Over the period 1997-2000 Argentine ants did not become more abundant in interior habitat suggesting it will remain a refuge for native ants. Argentine ant abundance in edge plots varied greatly among years and sites. Annual variation in abundance was positively related to annual rainfall. Increased soil moisture near edges due to urban runoff has often been suggested as the mechanism that allows the invasion of edge but not interior habitat. This hypothesis predicts that edge habitat downslope of the urban edge should support invasions of higher abundance and greater spatial penetration than habitat upslope. However, I found that edge slope did not predict the extent of invasion, whereas, soil type did. Coarse, well-drained soils supported an Argentine ant invasion of lower abundance and lesser spatial penetration than soils that should retain more moisture. These patterns of spatial and annual variation are more consistent with a biotic flow mechanism where ants move from urban habitat into CSS sites that are temporarily favorable, rather than an abiotic flow where urban runoff causes a physical change to CSS habitat near edges. This invasion affects a substantial area of habitat, however, the inability of Argentine ants to invade interior habitat suggests that refugia for native ants will persist if large unfragmented blocks of habitat are maintained.     

Hazard,risk and agrarian adaptations in a hyperarid watershed: El Niño floods,streambank erosion,and the cultural bounds of vulnerability in the Andean Middle Horizon

Peru's Osmore drainage, also known as the Moquegua Valley, is one of the driest regions on Earth, yet agricultural development has supported complex societies in the basin for almost 4000 yrs because of canal construction and irrigation. We compare the distinct agrarian and settlement systems of three coeval archaeological cultures in this arid region, Huaracane, Wari and Tiwanaku, and how each adapted—or failed to adapt—to geomorphic and climatic hazards. Systematic settlement pattern survey and radiocarbon dating along with geomorphological analysis of flood history and riverine processes permit detailed discussion of agrarian strategies during the Formative (1800 BC–AD 500) and Middle Horizon (AD 500–1000) periods, with distinct settlement “niches” in terms of agricultural practices and longitudinal position in the drainage, and lateral location relative to the floodplain. These adaptive strategies each manifest distinct “bounds of adaptability” to natural hazard, climate events and social stressors, and thus varying risk profiles. Besides the continual risk associated with sustained droughts, the Atacama and Peruvian Coastal Desert and culturally connected highland Altiplano are also vulnerable to the vagaries of the El Niño-Southern Oscillation (ENSO) leading to catastrophic floods in the mid-valley during warm phase ENSOs, in tandem with simultaneous hazards in other regions inhabited by these transregional cultures. By comparing the archaeological record of Wari, Huaracane, and Tiwanaku culture settlements with geomorphic signatures of catastrophic El Niños, we show that the viability of each cultural sequence depended on specific relationships to floodplain streambank erosion, construction and reworking over multiple time scales.     

Effective population size in a bottlenecked fur seal population

The census population size (N) is usually the only information available for most threatened species. For evolutionary matters, the effective population size (N_e), not the census number, is a prime concern. Factors such as variation in the sex ratio of breeding individuals, variation of population size in different generations and mating system are important. The South American fur seal, Arctocephalus australis, has been exploited in Peru by humans since ca. 2000 BC and now the original population declined 72%, as a result of low food availability during the severe El Niño in 1997-1998. In this sense A. australis is now classified as in danger of extinction in Peru. We present the first estimate of N_e of the Peruvian population of A. australis that takes into account the effects of mating system and variation in population size caused by the 1997-1998 El Niño. The resulting N_e was 2153 specimens. We believe that the estimated N_e for the Peruvian population is a critical value, because it is significantly lower than the mean minimum viable population for vertebrates (7000 breeding age adults). This estimated N_e is of critical importance because combined with the current El Niño events are reasons of great concern for the survival of the species and should be taken into account in future management plans to ensure the conservation and protection of the species in the Peruvian coast.     

An experimental study on the effects of an introduced parasite in Darwin’s finches

It is widely recognised that the main threats to the biota of the Galápagos Islands stem from newly introduced diseases, plants and animals. Introduced organisms may lead to the rapid extinction of naïve insular populations. Previous research on Galápagos showed that the parasitic larvae of the fly Philornis downsi occurred in virtually all nests of land bird species examined, with high nestling mortality (27% of all broods) in relation to high parasite infestation. In this study, we used an experimental approach to measure the fitness impacts of the fly larvae in two species of Darwin’s finch (Geospizinae). We created parasite-reduced nests by applying a 1% pyrethrin solution to the nest during the early nesting phase, and measured: (1) nestling growth, (2) nestling haemoglobin (Hb) concentration and (3) fledging success. A single insecticide treatment was sufficient to reduce parasite intensity to nearly zero, and revealed reduced mass gain and reduced fledging success in parasitized nests compared to parasite-reduced nests. This is the first experimental study of fitness impacts by Philornis on its host species. The results are discussed in the light of short-term protection measures for nests of rare finch species in the Galápagos archipelago.     

Extinction behind our backs: the possible fate of one of the Darwin’s finch species on Isla Floreana, Galápagos

Without regular monitoring a rare species may slip into extinction unnoticed. We report a possible case from the Galápagos archipelago. The warbler finch (Certhidea fusca) has not been recorded by scientists on Isla Floreana in recent years, and we have been concerned about its status. On a two-day visit to prime habitat in the breeding season of 2004 we used playback of warbler finch song and calls recorded on another island to stimulate an approach of local birds. We failed to find a single warbler finch, whereas we encountered numerous small tree finches (Camarhynchus parvulus), medium tree finches (C. pauper) and yellow warblers (Dendroica petechia). Searches in 1979, 1983, 1997 and 1999 were also unsuccessful. Our continuing failure to find any warbler finches indicates the species must be extremely rare on the island, if not actually extinct. We discuss possible reasons for their demise and draw attention to other species that could be heading in the same direction.     

The effects of cattle grazing on tall-herb fen vegetation and molluscs

The effects of light year-round cattle grazing on tall-herb fen vegetation and wetland molluscs were compared to the effects of non-intervention over a period of four years using grazing exclosures. The distribution of cattle within the area of fen was investigated by plotting the position of the herd at 3-4 day intervals throughout the year. Cattle distributed themselves randomly throughout the fen in spring, autumn and winter, but showed a more aggregated distribution in summer. Grazing reduced the biomass of Phragmites australis and increased stem densities of Glyceria maxima, resulting in a shift of dominance from Phragmites to Glyceria. Plant species-richness was also significantly higher in areas open to grazing.Grazing decreased total densities of molluscs and substantially reduced densities of the rare snail Vertigo moulinsiana. V. moulinsiana was particularly associated with areas of fen that had a high water table and high biomass of ungrazed Carex riparia. However, because of the patchy nature of the grazing, V. moulinsiana survived at reasonably high densities in patches of ungrazed vegetation within the grazing unit.     

Small-scale habitat fragmentation effects on pollinator behaviour: experimental evidence from the bumblebee Bombus veteranus on calcareous grasslands

We examined visiting patterns of pollinators of Betonica officinalis L. (Lamiaceae) in experimentally fragmented calcareous grasslands and corresponding control plots at two study sites (Movelier and Nenzlingen) in the north-western Swiss Jura mountains. Fragments (1.5×1.5 m) were isolated by a 5-m wide strip of frequently mown vegetation while the control plots were situated in the adjacent undisturbed vegetation. The most common pollinator, the bumblebee Bombus veteranus (Apidae), visited fragments 53.7% less frequently than control plots. Furthermore, a change in foraging behaviour of Bombus veteranus was observed. In fragments the bumblebees visited more inflorescences, flew longer total visiting distances and the visiting time per patch tended to be higher than in control plots. The distribution of angles between arrival and departure direction (turning angles) differed from a uniform distribution in fragments but not in control plots. The increased directionality of bumblebee flight might be due to a decrease in floral rewards. Our results show that small-scale habitat fragmentation can affect plant pollination at two levels both relevant for plant fitness. First, lower visitation rates indicate a limitation of pollinators which might result in reduced seed set of the pollinated plant. Second, changes in pollinator behaviour might reduce pollen dispersal among flowers, increase inbreeding and hence reduce genetic variability in populations of this bumblebee pollinated plant.     

Species-specific barrier effects of a motorway on the habitat use of two threatened forest-living bat species

It is generally recognized that roads can adversely affect local animal populations but little is known how roads affect bats. In particular, no study compared the response of bats that differ in foraging ecology to motorways that cut through the breeding habitat. As bats are key species in conservation, such data are urgently needed for designing management plans. Using radio-telemetry, mist netting, and mark-recapture data we investigated the effects of a motorway with heavy traffic on the habitat use of two threatened forest-living bats. We compared barbastelle bats (Barbastella barbastellus), which forage in open space, to Bechstein’s bats (Myotis bechsteinii), which glean prey from the vegetation. Five of six radio-tracked barbastelle bats crossed the motorway during foraging and roost switching, flying through underpasses and directly over the motorway. In contrast, only three of 34 radio-tracked Bechstein’s bats crossed the motorway during foraging, all three using an underpass. Bechstein’s bats, unlike barbastelle bats, never crossed the motorway during roost switching. Moreover, only in Bechstein’s bats individuals foraging close to the motorway had smaller foraging areas than individuals foraging further away, whereas other forest edges had no such effect. Our data show that motorways can restrict habitat accessibility for bats but the effect seems to depend on the species’ foraging ecology and wing morphology. We suggest that motorways have stronger barrier effects on bats that forage close to surfaces than on bats that forage in open space, and discuss the implications of our findings for bat conservation during road construction.     

Coppice management effects on experimentally established populations of three herbaceous layer woodland species

Traditional coppice management creates a temporal release of resources. We determined how this affected three herbaceous species (Cardamine pratensis, Primula elatior and Geum urbanum) and if it was suitable for their conservation within woodland given their differing phenologies and habitat affinities for woodland.Reproductive adults were transplanted and their fate, i.e. survival and counts of leafs and flowers, plus the fate of their progeny, were monitored for three years following cutting of coppice shoots (three light levels) or yearly autumn mowing.Cardamine pratensis and P. elatior produced more flowers with increasing light availability. Mowing increased flower and leaf production with time for P. elatior. Seedling numbers followed a similar trend. Geum urbanum initially produced more flowers with increasing light and when mown, but the effect disappeared and did not increase seedling numbers. Its basal leaves showed the opposite pattern. Population growth rates (λ), calculated for P. elatior and G. urbanum, confirmed the strong treatment effects on the former and the absence of effects on the latter. Yet, decomposition of treatment effects, showed considerable flexibility in life history of G. urbanum, except for contributions of fecundity. The latter, however, contributed most to positive effects on λ for P. elatior.Early flowering species with an affinity for open habitats (C. pratensis > P. elatior) benefited more from temporal resource release than the later flowering, typical woodland species. Coppice management thereby maintains both typical forest herbs and herbs with affinity for more open habitats. This is an important conservation tool especially in intensively managed landscapes.     

Beneficial effects of earthworms and arbuscular mycorrhizal fungi on establishment of leguminous trees on Pb/Zn mine tailings

Planting trees to stabilize metalliferous mine tailings is a widely used form of land reclamation although substantial soil amendment is invariably required, both to improve the physico-chemical status of the tailings and to ameliorate toxicity prior to planting. Here, we report a glasshouse study of the combined effects of burrowing earthworms (Pheretima guillelmi) and arbuscular mycorrhizal fungi (Glomus spp., AMF) on establishment of a naturally invasive, woody, nitrogen-fixing legume, Leucaena leucocephala, on topsoil-amended Pb/Zn mine tailings. AMF provided the most effective preliminary inoculant, improving N, P and K uptake, but earthworms had more influence improving N nutrition. In most cases, the combined effects of AMF and earthworms were additive and proved to be beneficial to plant growth, plant nutrition and for protection against uptake of toxic metals. AMF influenced metal uptake more than earthworms, but together they reduced mobility of Pb and Zn in soil by as much as 25%. Some minor but significant negative interactions were also evident; for example, earthworms enhanced soil microbial activity but inhibited the beneficial effects of AMF on N₂-fixation. We argue that increased attention to ecological interactions in soil could reduce costs and improve the efficacy of restoring a vegetation cover to land impacted by contaminated spoils.     

Impact of human dwellings on the distribution of the exotic Argentine ant: a case study in the Doñana National Park, Spain

The invasive Argentine ant Linepithema humile was found in close perimeters to inhabited houses situated in various habitat types in Doñana National Park (Southern Spain). We suggest that the Argentine ant is found in these sites largely due to passive importation by man, but from there may have the potential to spread into surrounding favourable natural habitats. Ant species richness and diversity around houses was similar to that in natural habitats, except in the cases where the Argentine ant dominated, where a considerable lower ant species diversity and richness was observed. The species composition of ants in the immediate vicinity of houses was very different from that of the surrounding natural habitats, although the species composition between houses was very similar, regardless of the type of surrounding natural habitat. Ant species around houses were typically generalists, opportunists or open habitat specialists. Many of these species are considered dominant in that they are characterised by having large nests, aggressive behaviour and mass recruitment in response to attack. However, these species are displaced by the Argentine ant when introduced. Not all ant species show the same responses to invasions by the Argentine ant, and some species, such as Cardiocondyla batesii, Oxyopomyrmex saulcyi or Cataglyphis floricola, may be able to persist for a period following an invasion of the Argentine ant. These species are characterised by small nests and submissive behaviour, and may survive by avoiding conflict with the invasive species.     

The effects of earthworms on the demography of annual plant assemblages in a long-term mesocosm experiment

Earthworms have been shown to influence plant growth, survival and fecundity. They can therefore affect plant demography in plant communities changing their composition. A long term mesocosm experiment was set-up to test the effects of an endogeic (Aporrectodea caliginosa) and an anecic (Lumbricus terrestris) earthworm species on assemblages of four species of annuals: one grass (Poa annua), two forbs (Veronica persica and Cerastium glomeratum) and one legume (Trifolium dubium). The number of individuals and the biomass of each species were investigated. A. caliginosa and L. terrestris affected the density of T. dubium at each of the three monitored census dates. The other plant species responded to A. caliginosa and L. terrestris at the second and third generations. The presences of A. caliginosa and L. terrestris reduced the total number of plant individuals from the second to the third generation. At harvest (3rd generation), T. dubium and V. persica had more and larger individuals in the presence of A. caliginosa. When both earthworm species were present, T. dubium had few but larger individuals. Our study confirms that earthworms affect plant demography and plant community structure. Our results also show that accurate prediction of long-term effects of earthworms on plant communities cannot be achieved using results on their short-term effects on plant growth. This is due to the poor understanding of the effects of earthworms on plant resource allocation and demography, and also the possibility that earthworms may exert the opposite effect on the short and long-term availability of nutrients.     

Cause and duration of mustard incorporation effects on soil-borne plant pathogenic fungi

Two fungal plant pathogens, Rhizoctonia solani AG 2-2 and Fusarium oxysporum f.sp. lini, were studied in relation to general responses of soil fungi and bacteria following incorporation of Brassica juncea. Our aim was to understand to what extent the changes in the biological and physicochemical characteristics of the soil could explain the effects on the studied pathogens and diseases, and to determine the temporal nature of the responses. Short-term effects of mustard incorporation (up to 4 months) were investigated in a microcosm experiment, and compared with a treatment where composted plant material was incorporated. In a field experiment, the responses were followed up to 11 months after removal or incorporation of a mustard crop. In general, responses in the variables measured changed more after incorporation of fresh mustard material than after addition of similar amounts of composted plant material (microcosms) or after removal of the mustard crop (field). The soil inoculum potential of R. solani AG 2-2 decreased directly after incorporation of mustard, but increased later to disease levels above those in the untreated soil. Neither of these effects could be explained by changes in the population density of R. solani AG 2-2. Fusarium spp. were less influenced, although an increase in the suppressiveness to Fusarium wilt was observed after mustard incorporation as compared with the treatment where mustard was removed. The microbial responses to mustard incorporation were more pronounced for bacteria than for fungi. After an initial substantial increase, the bacterial density decreased but remained above the levels in the control treatment throughout the experimental periods. The bacterial community structure was modified up to 8 months after mustard incorporation. We conclude that incorporation of fresh mustard influences soil microbial communities, especially the bacteria, and has a potential to control the pathogenic activity of R. solani 2-2 on a short-term perspective. The time dependency in microbial responses is important and should be taken into consideration for the evaluation of the potential of Brassicas to control plant disease on a field scale.     

Sampling effects on the assessment of genetic diversity of rhizobia associated with soybean and common bean

Biological nitrogen fixation plays a key role in agriculture sustainability, and assessment of rhizobial diversity contributes to worldwide knowledge of biodiversity of soil microorganisms, to the usefulness of rhizobial collections and to the establishment of long-term strategies aimed at increasing contributions of legume-fixed N to agriculture. Although in recent decades the use of molecular techniques has contributed greatly to enhancing knowledge of rhizobial diversity, concerns remain over simple issues such as the effects of sampling on estimates of diversity. In this study, rhizobia were isolated from nodules of plants grown under field conditions, in pots containing soil, or in Leonard jars receiving a 10⁻² or a 10⁻⁴ serially-diluted soil inoculum, using one exotic (soybean, Glycine max) and one indigenous (common bean, Phaseolus vulgaris) legume species. The experiments were performed using an oxisol with a high population (10⁵ cells g⁻¹ soil) of both soybean rhizobia, composed of naturalized strains introduced in inoculants and of indigenous common-bean rhizobia. BOX-PCR was used to evaluate strain diversity, while RFLP-PCR of the ITS (internally transcribed spacer) region with five restriction enzymes aimed at discriminating rhizobial species. In both analyses the genetic diversity of common-bean rhizobia was greater than that of soybean. For the common bean, diversity was greatly enhanced at the 10⁻⁴ dilution, while for the soybean dilution decreased diversity. Qualitative differences were also observed, as the DNA profiles differed for each treatment in both host plants. Differences obtained can be attributed to dissimilarity in the history of the introduction of both the host plant and the rhizobia (exotic vs. indigenous), to host-plant specificity, rhizobial competitiveness, and population structure, including ease with which some types are released from microcolonies in soil. Therefore, sampling method should be considered both in the interpretation and comparison of the results obtained in different studies, and in the setting of the goals of any study, e.g. selection of competitive strains, or collection of a larger spectrum of rhizobia. Furthermore, effects of sampling should be investigated for each symbiosis.     

Five years of simulated atmospheric nitrogen deposition have only subtle effects on the fate of newly synthesized carbon in Calluna vulgaris and Eriophorum vaginatum

To understand the implications of atmospheric nitrogen deposition on carbon turnover in peatlands, we conducted a ¹³C pulse labeling experiment on Calluna vulgaris and Eriophorum vaginatum already receiving long-term (5 years) amendments of 56 kg N ha⁻¹ y⁻¹ as ammonium or nitrate. We examined shoot tissue retention, net ecosystem respiration returns of the ¹³C pulse, and soil porewater DOC content under the two species. ¹³C fixation in Eriophorum leaves was enhanced with nitrogen addition and doubled with nitrate supply. This newly fixed C appeared to be relocated below-ground faster with nitrogen fertilization as respiration returns were unaffected by N inputs. By contrast, increases in ¹³C fixation were not observed in Calluna. Instead, net ecosystem respiration rates over Calluna increased with N fertilization. There was no significant label incorporation into DOC, suggesting a conservative strategy of peatland vegetation regarding allocation of C through root exudation. Greater concentrations of total DOC were identified with nitrate addition in Calluna. Given the long-term nature of the experiment and the high N inputs, the overall impacts of nitrogen amendments on the fate of recently synthesized C in Eriophorum and Calluna in this ombrotrophic peatland were surprisingly more moderate than originally hypothesized. This may be due to N being effectively retained within the bryophyte layer, thus limiting, and delaying the onset of, below-ground effects.     

Rhizosphere activity, grass species and N availability effects on the soil C and N cycles

We examined whether grass species and soil nitrogen (N) availability could enhance Carbon (C) and N turnover during root litter decay in grassland. Three species with increasing competitiveness (Festuca ovina, Dactylis glomerata and Lolium perenne) were grown at two N fertiliser levels in an undisturbed grassland soil, in which soil organic fractions derived for the last 9 years from Lolium root litter which was ¹³C-depleted. During the subsequent experimental year, the C turnover was calculated using the respective δ¹³C values of the old and new C in the root phytomass, in two Particulate Organic Matter (POM) fractions above 200 μm and in the lightest part of the aggregated soil fraction between 50 and 200 μm. Soil N availability was monitored during the regrowth periods with ion exchange resins (IER). The C decay rates of each particle size fraction were calculated with a simple mechanistic model of C dynamics. The N mineralisation immobilisation turnover (MIT) was characterised by dilution of ¹⁵N-labelled fertiliser in the N harvestThe C:N ratio and the residence time of C in the fractions decreased with particle size. The presence of a grass rhizosphere increased the decay rate of old C. Accumulation of new C in particle size fractions increased with species competitiveness and with N supply. Species competitiveness increased C turnover in the aggregated fraction, as a result of greater accumulation of new C and faster decay of old C. Fertiliser N increased N turnover and C mineralisation in the SOM. Species competitiveness decreased soil -N exchanged with the IER and increased dissolved organic C (DOC) content. The nature of the current rhizosphere is thus an important factor driving C and N transformations of the old root litter, in relation with grass species strategy. Plant competitiveness may stimulate the C and N turnover in the more evolved SOM fractions in a similar way to the mineral N supply.