Experimental manipulation of succession in heathland ecosystems

An experiment was carried out in two heathland ecosystems, one dominated by Calluna vulgaris and the other by Molinia caerulea, to analyse the effects of soil organic matter accumulation and nutrient mineralization on plant species dynamics during succession. The experiment included one treatment that received nutrient solution and two treatments where the rate of soil organic matter accumulation was reduced by removing litter or accelerated by adding litter. In a fourth treatment the C. vulgaris litter produced in the C. vulgaris-dominated plots was replaced by litter of M. caerulea and vice versa. Treatments were applied over 8 years. Addition of nutrient solution caused C. vulgaris to decline, and grass species to increase sharply, compared to the control plots. Addition of litter enhanced both N mineralization and the biomass of M. caerulea and Deschampsia flexuosa but reduced the biomass of C. vulgaris. The effects of replacing C. vulgaris litter by M. caerulea litter, or vice versa, on N mineralization and species dynamics could not be attributed to differences between the decomposability of the different litter materials that were transferred. The results confirm the hypothesis that increased litter inputs accelerate the rate of species replacement during succession.     

Rapid response of Orthoptera to restoration of montane heathland

Understanding how to restore threatened ecosystems is of special relevance for nature conservation. The aim of this study was to use Orthoptera as ecological indicators for the effects of montane heathland restoration in Central Europe. We analysed the three following treatments: (i) montane heathlands (MONHEATH) (N = 7), (ii) restoration sites (RESSITE) (N = 3) and (iii) clear-cuts of spruce forests as unprocessed and ungrazed control sites (CONTROL) (N = 3). Vegetation structure and microclimate differed considerably between MONHEATH on the one hand and RESSITE and CONTROL on the other hand. Orthoptera species richness and density did so too. MONHEATH was characterised by a high-growing dense dwarf-shrub and moss layer having a cool microclimate and high soil moisture. In contrast, RESSITE and CONTROL had sparse vegetation and a warm microclimate; Orthoptera species richness and density was highest on these sites. Our study clearly showed that heathland Orthoptera responded rapidly to restoration measures, while Ericaceae dwarf shrubs slowly established. The vast majority of Orthoptera species found on the restoration sites are early and mid-successional species. The colonization of the sites by late-successional Orthoptera species in the future will depend on the further development of the heathland vegetation; that is, if Ericaceae will expand to the sites. We conclude that the realised restoration measures are suitable to promote heathland Orthoptera of early and mid-successional stages. However, the current management of montane heathlands is insufficient and needs to be intensified in order to provide structurally diverse habitats with their characteristic orthopteran assemblages.     

Phylogenetic and functional dissimilarity does not increase during temporal heathland succession

Succession has been a focal point of ecological research for over a century, but thus far has been poorly explored through the lens of modern phylogenetic and trait-based approaches to community assembly. The vast majority of studies conducted to date have comprised static analyses where communities are observed at a single snapshot in time. Long-term datasets present a vantage point to compare established and emerging theoretical predictions on the phylogenetic and functional trajectory of communities through succession. We investigated within, and between, community measures of phylogenetic and functional diversity in a fire-prone heathland along a 21 year time series. Contrary to widely held expectations that increased competition through succession should inhibit the coexistence of species with high niche overlap, plots became more phylogenetically and functionally clustered with time since fire. There were significant directional shifts in individual traits through time indicating deterministic successional processes associated with changing abiotic and/or biotic conditions. However, relative to the observed temporal rate of taxonomic turnover, both phylogenetic and functional turnover were comparatively low, suggesting a degree of functional redundancy among close relatives. These results contribute to an emerging body of evidence indicating that limits to the similarity of coexisting species are rarely observed at fine spatial scales.     

Non-native grass invasion suppresses forest succession

Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

Markov models in the study of post-fire succession in heathland communities

Data on the presence of a number of vegetation states (defined in terms of species dominance in areas of 10×10 cm) and transition probabilities were derived from permanent quadrats in a number of recently burned heath stands. Data were taken from a species-rich community, a species-poor type and a high-level Calluna-Eriophorum bog. Simple Markovian models were constructed using these data, and the model predictions were compared with known or expected trends. Models for species-rich heath yielded poor simulations of expected trends since matrices derived from data for the first years after fire did not contain sufficient information on transitions to states important later in the developmental sequence. Model results for the simpler species-poor and bog communities were more satisfactory and simulated expected trends. In these types all species recovered quickly after fire and less rearrangement of species abundances took place. Maximum likelihood statistics carried out on the transition matrices produced inconclusive results for the species-rich and species-poor types, but indicated that the data from the Calluna-Eriophorum bog approximated a first-order time-homogeneous Markov chain. It was concluded that Markov models lack predictive ability except in relatively simple systems, but that they may be useful in illustrating variations in short-term community dynamics.Nomenclature follows Tutin et al. (1964–80) for vascular plants and Smith (1978) for mosses.I am grateful to Prof. C. H. Gimingham for help and advice during this study, which was carried out during tenure of a Natural Environment Research Council studentship. Thanks are also due to Dr M. B. Usher and an anonymous referee for their comments on the draft.     

Leaf-nutrient accumulation and turnover at three stages of succession from heathland to forest

Abstract. Accumulation of nutrients in leaves of the dominating species of three ecosystems, characterizing the secondary succession from Genisto-Callunetum heathland through Leucobryo-Pinetum birch-pine woodland to mature Querco-Fagetum oak-beech forest, as well as nutrient turnover within these ecosystems was studied. The objective of the study was to establish potential variations in quantity and quality of nutrient supply to the plants with respect to succession dynamics. The results show very low leaf nutrient concentrations of all species investigated, coinciding with low nutrient availability in the soil. However, the nutrient content of leaves and leaf litter of Quercus petraea and Fagus sylvatica, which dominate the late succession stages, and in Betulapéndula are higher than in the photosynthetic organs (leaves and young shoots) of Calluna vulgaris and Pinus sylvestris. The combination of the higher nutrient content of the leaves and an increasing leaf-litter production during succession results in an increased nutrient turnover via leaf-litter fall. However, due to the high leaf biomass, the storage of nutrients in the leaf biomass is highest within the birch-pine woodland. From this, it may be assumed that the low demand and the low loss of nutrients via leaf-litter fall are favourable for Pinus at the early stages of forest succession on poor sandy soils. In contrast, Quercus and Fagus are provided with better growth conditions at the later stages of succession resulting from the accumulation of plant-available nutrients in the ecosystem by Pinus sylvestris, combined with a higher nutrient turnover as compared with the heathland.     

Soil seed bank dynamics in post-fire heathland succession in south-eastern Australia

Soil seed banks can exert a strong influence on the path of vegetation succession following fire, with species varying in their capacity to persist in the seed bank over time, leading to changes in seed bank composition and propagules available for post-fire colonisation. This study examined the effect of time since fire on soil seed bank dynamics in a chronosequence of seven sites spanning 26 years in a south-eastern Australian sand heathland. No significant change was evident in the species richness and density of the germinable soil seed bank, but species composition differed significantly among young (0–6 years since fire), intermediate (10–17 years since fire) and old-aged (24–26 years since fire) sites (using presence/absence data). No significant trend was observed in the similarity between the extant vegetation and the soil seed bank with time since fire. A total of 32% of the species recorded in the soil seed bank were not present in the above-ground vegetation at the same site, which suggests that species requiring fire for germination may be present in the seed bank. Most species present in the extant vegetation were not recorded (63%) or were in very low abundances in the soil seed bank (29%). The mode of regeneration appears to be the major determinant of species absence in the soil seed bank, as 66% of species occurring in the extant vegetation but not in the seed bank have the capacity to regenerate by resprouting. This study shows that a major shift in the successional pathway after fire due to altered seed bank composition is unlikely in this vegetation; most species not recorded in the seed bank are either resprouters (obligate or facultative) or serotinous, suggesting that they will readily regenerate following fire. Unless fire frequencies are high and kill fire-sensitive obligate seeders before they reach maturity, the chance that the soil seed bank could substantially alter vegetation composition within the study area after fire is low. However, it is unclear how successional pathways may alter in response to severe fires with the potential to kill both seeders and resprouters.     

Interspecific competition: A mechanism for rodent succession after fire in wet heathland

Abstract A competitive interaction between two species of sympatric native rodents was demonstrated experimentally in Myall Lakes National Park (NSW, Australia). The previously documented replacement of Pseudomys gracilicaudatus by Rattus lutreolus in areas of wet heath in this region implied that competition may be the mechanism facilitating this succession. We present experimental evidence to support this suggestion. Removal of the larger R. lutreolus caused a significant increase in abundance of P. gracilicaudatus on five experimental sites in comparison to five unmanipulated control sites. The sites used in this experiment were chosen from three ages of wet heath regenerating after fire. These three ages represent key stages in the replacement of P. gracilicaudatus by R. lutreolus and the occurrence of competition in each of these ages indicates that competition plays an important role in this succession. In the initial stages of the experiment when R. lutreolus were removed they were replaced by new R. lutreolus individuals. This has been interpreted as evidence of strong intraspecific competition. After removal of R. lutreolus, P. gracilicaudatus expanded its habitat range into microhabitats that had formerly been occupied by R. lutreolus and reduced its range in microhabitats previously occupied by P. gracilicaudatus. This leads us to believe that P. gracilicaudatus was occupying inferior microhabitat before the removal of R. lutreolus, which had excluded it from more preferable microhabitat.     

The effects of fire on woody plant encroachment are exacerbated by succession of trees of decreased palatability

The ingression of woody plants into the grassy layer of savannas and grasslands has become a global concern. The increase of woody plants has been primarily attributed over grazing, fire and more recently to the increase of atmospheric CO₂. We used long-term observations and analyses to assess changes in woody vegetation in Ithala Game Reserve (IGR), South Africa. Textural analysis of aerial photographs was used to detect changes in woody vegetation, from 1943 to 2007 in Ithala Game Reserve (IGR), South Africa. Daily rainfall data from 1905 to 2009 were used in a time-series analysis to determine if rainfall patterns have changed. The time-series analysis showed that the low magnitude (0–10 mm) rainfall events decreased from 1916 to 2009 and high magnitude rainfall events increased (10–20 and >20 mm). The mean annual rainfall increased from ～700 to ～850 mm from the 1930s to the 2000s. This change in rainfall was a key factor in the increase in woody vegetation from 1943 to 2009. We also used field data from the same reserve collected over 30 years to assess the increases in tree cover. Tree cover and density increased significantly by 32.5% and 657.9 indiv ha^?1 respectively, over 64 years. Before the proclamation of IGR in 1972, increases in woody vegetation from 1943 were non-significant. After the proclamation of IGR, herbivore population numbers and spatial distribution influenced the accumulation of grassy biomass required to fuel fires. In areas with reduced fuel loads, the consequential suppression of fire accelerated the rate of woody plant invasion into savannas. The increase in woody vegetation coincided with a decrease in palatable (e.g. Acacia gerrardii and Acacia davyi) and an increase in unpalatable woody plants. The avoidance of the unpalatable trees (e.g. Euclea and Searsia species) by large mammalian herbivores has allowed these trees to increase in density relatively unhindered.     

Impacts of long-term enhanced UV-B radiation on bryophytes in two sub-Arctic heathland sites of contrasting water availability

Background and Aims

Anthropogenic depletion of stratospheric ozone in Arctic latitudes has resulted in an increase of ultraviolet-B radiation (UV-B) reaching the biosphere. UV-B exposure is known to reduce above-ground biomass and plant height, to increase DNA damage and cause accumulation of UV-absorbing compounds in polar plants. However, many studies on Arctic mosses tended to be inconclusive. The importance of different water availability in influencing UV-B impacts on lower plants in the Arctic has been poorly explored and might partially explain the observed wide variation of responses, given the importance of water in controlling bryophyte physiology. This study aimed to assess the long-term responses of three common sub-Arctic bryophytes to enhanced UV-B radiation (+UV-B) and to elucidate the influence of water supply on those responses.

Methods

Responses of three sub-Arctic bryophytes (the mosses Hylocomium splendens and Polytrichum commune and the liverwort Barbilophozia lycopodioides) to +UV-B for 15 and 13 years were studied in two field experiments using lamps for UV-B enhancement with identical design and located in neighbouring areas with contrasting water availability (naturally mesic and drier sites). Responses evaluated included bryophyte abundance, growth, sporophyte production and sclerophylly; cellular protection by accumulation of UV-absorbing compounds, β-carotene, xanthophylls and development of non-photochemical quenching (NPQ); and impacts on photosynthesis performance by maximum quantum yield (F_v /F_m) and electron transport rate (ETR) through photosystem II (PSII) and chlorophyll concentrations.

Results

Responses were species specific: H. splendens responded most to +UV-B, with reduction in both annual growth (–22 %) and sporophyte production (–44 %), together with increased β-carotene, violaxanthin, total chlorophyll and NPQ, and decreased zeaxanthin and de-epoxidation of the xanthophyll cycle pool (DES). Barbilophozia lycopodioides responded less to +UV-B, showing increased β-carotene and sclerophylly and decreased UV-absorbing compounds. Polytrichum commune only showed small morphogenetic changes. No effect of UV-B on bryophyte cover was observed. Water availability had profound effects on bryophyte ecophysiology, and plants showed, in general, lower growth and ETR, together with a higher photoprotection in the drier site. Water availability also influenced bryophyte responses to +UV-B and, in particular, responses were less detectable in the drier site.

Conclusions

Impacts of UV-B exposure on Arctic bryophytes were significant, in contrast to modest or absent UV-B effects measured in previous studies. The impacts were more easily detectable in species with high plasticity such as H. splendens and less obvious, or more subtle, under drier conditions. Species biology and water supply greatly influences the impact of UV-B on at least some Arctic bryophytes and could contribute to the wide variation of responses observed previously.     

Impacts of drought and native grass competition on buffelgrass (Pennisetum ciliare)

Biological Invasions - Invasive species are detrimental to ecosystems worldwide, and long-term invasive treatment outcomes are generally poor. Integrating active restoration into an invasive...     

Impacts of anthropogenic disturbances on forest succession in the mid-montane forests of Central Himalaya

We studied the influence of anthropogenic drivers on the distribution and regeneration of tree species in vegetation at different stages of succession from grasslands to oak forests in mid-montane Central Himalaya. We found fire, grazing, and lopping as the main factors hindering a progressive successional regime towards a late-successional oak community. Succession was studied in five vegetation formations (grasslands, pine, pine–oak, open oak, and dense oak), with similar site conditions, representing a theoretical successional sequence from early- to late-successional stages. A structured survey with uniform distribution of sampling plots in the five selected vegetation formations was conducted to gather information abut the vegetation communities. Early-successional grasslands and pine forests were found to harbour high densities of pine and oak seedling and sapling regeneration. However, recurring fires and chronic unsustainable levels of grazing in these vegetation formations obstructed progressive succession by eliminating regenerating seedling and saplings from the forest understorey. Similarly, in intermediate- and late-successional stages (including pine–oak, open oak, and dense oak), overexploitation of existing oaks trees via lopping and grazing of regenerating oak seedlings and saplings hampered oak regeneration and development. The possibility to convert pine forests into oak as well as the conservation of existing oak forests through controlled grazing and lopping are management options that can contribute to an enhanced functioning of forest ecosystems in the study area. We conclude that with strategic management that restricts the current anthropogenic disturbances, the extent of oak forest in the study area can be increased.     

Mixtures with grass litter may hasten shrub litter decomposition after shrub encroachment into mountain grasslands

Aims

Shrub encroachment in mesic grasslands alters the identity and quality of litters entering the system. As litter from shrubs and grasses can differ in their quality, this can lead to differences in litter decomposition by the direct effect of quality, but also to litter interaction during decomposition. The objective of this study was to examine the occurrence of non-additive effects of litter mixtures on the decomposition rates of legume shrub litter (poor in P) or conifer shrub litter (poor in N) and grass litter.

Methods

In addition to single litter type litterbags for the three species, we mixed litters of each pair of possible combinations to determine the influence of each species on mass loss. Litterbags were placed in the field and collected after 1, 6, 8, 12 and 24 months. In each collection, litter of each species remaining in mixed bags was separated, dry weighed and analyzed for C, N and P.

Results

With respect to shrub litter decomposing alone, mass loss of shrub litter when mixed with grass showed a 9–10 % increase in decomposition rate for conifer and a 3 % increase for legume litter. These litter mixture effects varied with time and they were detected after a decomposition period of 1 year in legume litter and of 2 years in conifer litter.

Conclusions

Grass litter hastened conifer and legume litter decomposition in leaf litter mixtures, at least during the first stages of the process. The potential consequences of this result to alter litter accumulation patterns and thus carbon sequestration rates after shrub encroachment into grasslands will depend on whether the observed trends are maintained in the advanced decomposition stages.     

Head size and shape in relation to grass feeding in Acridoidea (Orthoptera)

Among grasshoppers, grass-feeding species had relatively very large heads and species feeding on other herbaceous plants had small ones. Those feeding on both food types were intermediate in size. These results are consistent with the hypothesis that head size is determined by the need for large mandibles and large mandibular adductor muscles, which are in turn associated with hard or tough diets.     

The invertebrates of heather and heathland

WEBB, N. R., 1989. The invertebrates of heather and heathland. The southern heathlands of Great Britain have long been renowned amongst entomologists. The invertebrates fall into two groups: species which feed on heathland plants–about 40 in the case of Calluna–and a second group which depends on the particular physical conditions provided by heathland, such as hot dry sandy areas or the micro-topography of Sphagnum bogs. The ecology of representative examples will be described, and the effects of fragmentation and isolation of the heathlands on the distribution and abundance of invertebrates discussed.     

Impacts of extreme winter warming in the sub‐Arctic: growing season responses of dwarf shrub heathland

Climate change scenarios predict an increased frequency of extreme climatic events. In Arctic regions, one of the most profound of these are extreme and sudden winter warming events in which temperatures increase rapidly to above freezing, often causing snow melt across whole landscapes and exposure of ecosystems to warm temperatures. Following warming, vegetation and soils no longer insulated below snow are then exposed to rapidly returning extreme cold. Using a new experimental facility established in sub‐Arctic dwarf shrub heathland in northern Sweden, we simulated an extreme winter warming event in the field and report findings on growth, phenology and reproduction during the subsequent growing season. A 1‐week long extreme winter warming event was simulated in early March using infrared heating lamps run with or without soil warming cables. Both single short events delayed bud development of Vaccinium myrtillus by up to 3 weeks in the following spring (June) and reduced flower production by more than 80%: this also led to a near‐complete elimination of berry production in mid‐summer. Empetrum hermaphroditum also showed delayed bud development. In contrast, Vaccinium vitis‐idaea showed no delay in bud development, but instead appeared to produce a greater number of actively growing vegetative buds within plots warmed by heating lamps only. Again, there was evidence of reduced flowering and berry production in this species. While bud break was delayed, growing season measurements of growth and photosynthesis did not reveal a differential response in the warmed plants for any of the species. These results demonstrate that a single, short, extreme winter warming event can have considerable impact on bud production, phenology and reproductive effort of dominant plant species within sub‐Arctic dwarf shrub heathland. Furthermore, large interspecific differences in sensitivity are seen. These findings are of considerable concern, because they suggest that repeated events may potentially impact on the biodiversity and productivity of these systems should these extreme events increase in frequency as a result of global change. Although climate change may lengthen the growing season by earlier spring snow melt, there is a profound danger for these high‐latitude ecosystems if extreme, short‐lived warming in winter exposes plants to initial warm temperatures, but then extreme cold for the rest of the winter. Work is ongoing to determine the longer term and wider impacts of these events.