Primary Succession Gradient Research Articles

Shift from facilitative to neutral interactions by the cushion plant Silene acaulis along a primary succession gradient

AbstractBackgroundThe stress‐gradient hypothesis predicts a shift from facilitative to competitive plant interactions with decreasing abiotic stress. This has been supported by studies along elevation and temperature gradients, but also challenged by the hypothesis of a facilitation collapse at extremely harsh sites. Although facilitation is known to be important in primary succession, few studies have examined these hypotheses along primary succession gradients.AimTo examine whether there is a relationship between the presence of the circumpolar cushion plant Silene acaulis and other species, and if so, whether there is a shift between positive and negative interactions along a primary succession gradient in a glacier foreland.LocationFinse, southern Norway.MethodsWe examined the performance of the common alpine forb Bistorta vivipara, species richness of vascular plants, bryophytes and lichens, and the number of seedlings and fertile vascular plants in S. acaulis cushions, and control plots without S. acaulis, along a succession gradient with increasing distance from a glacier front, and thus decreasing abiotic stress. To examine if S. acaulis cushions modify the abiotic environment, we recorded soil temperature, moisture, organic content and pH in cushions and control plots.ResultsBistorta vivipara performed better, as shown by bigger leaves in S. acaulis cushions compared to control plots in the harshest part of the gradient close to the glacier. There were few differences in B. vivipara performance between cushion and control plots in the more benign environment further away from the glacier. This suggests a shift from facilitative to mainly neutral interactions by S. acaulis on the performance of B. vivipara with decreasing abiotic stress. A trend, although not significant, of higher vascular species richness and fertility inside S. acaulis cushions along the whole gradient, suggests that S. acaulis also facilitates community‐level species richness. The causal mechanism of this facilitation is likely that the cushions buffer extreme temperatures.ConclusionsOur results support the stress‐gradient hypothesis for the relationship between the cushion plant S. acaulis and the performance of a single species along a primary succession gradient in a glacier foreland. S. acaulis also tended to increase vascular plant species richness and fertility regardless of stress level along the gradient, suggesting facilitation at the community level. We found no collapse of facilitation at the most stressful end of the gradient in this alpine glacier foreland.

Feedback effects between plant and flower-visiting insect communities along a primary succession gradient

Primary successions of glacier forelands are unique model systems to investigate community dynamics and assembly processes. However, successional changes of plant and insect communities have been mainly analysed separately. Therefore, changes in plant–insect interactions along successional gradients on glacier forelands remain unknown, despite their relevance to ecosystem functioning. This study assessed how successional changes of the vegetation influenced the composition of the flower-visiting insect assemblages of two plant species, Leucanthemopsis alpina (L.) Heyw. and Saxifraga bryoides L., selected as the only two insect-pollinated species occurring along the whole succession. In addition, we investigated the links between reproductive output of these plants and pollinator abundance through experimental exclusion of pollinators. Plant community structure changed along the succession, affecting the distribution and the abundance of insects via idiosyncratic responses of different insect functional groups. L. alpina interacted with ubiquitously distributed pollinators, while S. bryoides pollinators were positively associated with insect-pollinated plant species density and S. bryoides abundance. With succession proceeding, insect assemblages became more functionally diverse, with the abundance of parasitoids, predators and opportunists positively related to an increase in plant cover and diversity. The reproductive output of both plant species varied among successional stages. Contrary to our expectation, the obligate insect-pollinated L. alpina showed a reproductive output rather independent from pollinator abundance, while the reproductive output of the self-fertile S. bryoides seemed linked to pollinator abundance. Observing ecological interactions and using functional traits, we provided a mechanistic understanding of community assembly processes along a successional gradient. Plant community diversity and cover likely influenced insect community assembly through bottom-up effects. In turn, pollinators regulate plant reproductive output through top-down control. We emphasise that dynamics of alpine plant and insect communities may be structured by biotic interactions and feedback processes, rather than only be influenced by harsh abiotic conditions and stochastic events.

Structure-dynamic relationship of plant–insect networks along a primary succession gradient on a glacier foreland

There is a growing interest in understanding the structure–dynamic relationship of ecological networks. Ecological network changes along primary successions are poorly known: to address such topic, gradient of primary succession on glacier forelands is an ideal model, as sites of different age since deglaciation stand for different ecosystem developmental stages. We aimed to investigate the assembly processes of plant–insect networks and to elucidate its functional implications for ecosystem stability along this time sequence succession. We collected data on the functional role of anthophilous insect groups and performed network analysis to evaluate their relative importance in the structure of plant–insect interaction networks with increasing time since deglaciation along the primary succession of a debris-covered glacier foreland. We sampled anthophilous insects visiting the flowers of two models plant species, Leucanthemopsis alpina and Saxifraga bryoides. Insects were identified and trophic roles were attributed to each taxon (detritivores, parasitoids, phytophagous, pollinators, predators, and opportunists) at five sites representing the primary succession gradient. Plant–insect interactions were visually represented by a bipartite network for each successional stage. For each plant species and insect group, centrality indices were computed quantifying their community importance. For the whole network, centralization and link density were calculated. Pollinators dominated pioneer communities on the debris-covered glacier and in recently deglaciated areas, while parasitoids, predators and opportunists characterised late-succession stages. Plant species centrality varied along the succession. Pollinators showed initially higher but then decreasing centrality, while the centrality of predators and parasitoids increased with time since deglaciation. Along the same gradient link density showed an increasing trend while network centralization tended to decrease. The present study provides new insight into the initial steps of plant–insect network assembly and sheds light on the relationship between structure and dynamic in ecological networks. In particular, during the succession process, more links are formed and plant–anthophilous insect interactions change from a network dominated by pollinators to a functionally more diversified food web. We conclude that applying network theory to the study of primary succession provides a useful framework to investigate the relationship between community structure and ecosystem stability.

Taxonomic and functional successional patterns in macroinvertebrates related to flying dispersal abilities: a case study from isolated manmade ponds at reclaimed opencast coal mines

It has been proposed that communities change from r to K strategies during primary succession. However, because strong-flying organisms are expected to arrive first in newly created habitats and they show trait characteristics associated more often with K strategies, we hypothesized that the r to K trajectories would be more closely followed by flightless and poorly-flying organisms. Moreover, we expected that macroinvertebrate communities would converge in their functional composition due to deterministic forces while diverging as taxonomic assemblages due to stochastic drift and biotic interactions. However, we also expected that dispersal abilities of the organisms would affect these tendencies. To address these issues, macroinvertebrates were sampled from isolated manmade ponds of different ages (1–22 years old) constructed at reclaimed opencast coal mines. In accordance with our expectations, only flightless and poorly-flying organisms exhibited a slight shift from r to K strategies, the community taxonomically diverged along the primary succession gradient, and stochastic drift showed greater effects on strong-flying organisms. In contrast, the community did not converge in its functional composition. The weak differences observed among the macroinvertebrates from ponds of different ages suggested that limiting environmental conditions prevented the organisms from evolving to a more structured community.

Changes in the root‐associated fungal communities along a primary succession gradient analysed by 454 pyrosequencing

We investigated changes in the root-associated fungal communities associated with the ectomycorrhizal herb Bistorta vivipara along a primary succession gradient using 454 amplicon sequencing. Our main objective was to assess the degree of variation in fungal richness and community composition as vegetation cover increases along the chronosequence. Sixty root systems of B. vivipara were sampled in vegetation zones delimited by dated moraines in front of a retreating glacier in Norway. We extracted DNA from rinsed root systems, amplified the ITS1 region using fungal-specific primers and analysed the amplicons using 454 sequencing. Between 437 and 5063 sequences were obtained from each root system. Clustering analyses using a 98.5% sequence similarity cut-off yielded a total of 470 operational taxonomic units (OTUs), excluding singletons. Between eight and 41 fungal OTUs were detected within each root system. Already in the first stage of succession, a high fungal diversity was present in the B. vivipara root systems. Total number of OTUs increased significantly along the gradient towards climax vegetation, but the average number of OTUs per root system stayed unchanged. There was a high patchiness in distribution of fungal OTUs across root systems, indicating that stochastic processes to a large extent structure the fungal communities. However, time since deglaciation had impact on the fungal community structure, as a systematic shift in the community composition was observed along the chronosequence. Ectomycorrhizal basidiomycetes were the dominant fungi in the roots of B. vivipara, when it comes to both number of OTUs and number of sequences.

Colonization of experimentally created gaps along an alpine successional gradient

The colonization of artificially created gaps was analyzed along an alpine successional gradient from pioneer to early, late, and old successional stages. The presence/absence of species and the abundances of seedlings and adults in the gaps were recorded and compared with those of the surrounding areas. We hypothesized that in the older successional stages, the gaps were likely to be colonized by clonal ingrowth of the surrounding species. In the younger stages, we expected to find a high presence of seedlings and adults recruited by seeds. Micro-succession in the gaps occurred at each successional stage, with all life forms among the colonizers. The abundance of seedlings was significantly higher in the gaps compared with the surrounding area. At the early and late successional stages, the surrounding areas provided safe sites for seedling establishment, with the abundance of adults recruited by seeds higher at the gap edges than in the gap centers. We can confirm the first hypothesis of a higher clonal ingrowth in the old successional stage. Clonal ingrowth also occurred in the younger successional stages. Despite the lower species richness in the gaps, a positive correlation was found between gap and surrounding species frequencies, which were the highest in the pioneer and the lowest in the old successional stage. We conclude that gaps are relevant for seedling recruitment along the entire primary succession gradient. New species invasions from greater distances were not observed in the gaps. The dominant species on each site were identified to be successful gap colonizers.

Estimating phosphorus availability for microbial growth in an emerging landscape

Estimating phosphorus (P) availability is difficult—particularly in infertile soils such as those exposed after glacial recession—because standard P extraction methods may not mimic biological acquisition pathways. We developed an approach, based on microbial CO 2 production kinetics and conserved carbon:phosphorus (C:P) ratios, to estimate the amount of P available for microbial growth in soils and compared this method to traditional, operationally-defined indicators of P availability. Along a primary succession gradient in the High Andes of Perú, P additions stimulated the growth-related (logistic) kinetics of glutamate mineralization in soils that had been deglaciated from 0 to 5 years suggesting that microbial growth was limited by soil P availability. We then used a logistic model to estimate the amount of C incorporated into biomass in P-limited soils, allowing us to estimate total microbial P uptake based on a conservative C:P ratio of 28:1 (mass:mass). Using this approach, we estimated that there was < 1 μg/g of microbial-available P in recently de-glaciated soils in both years of this study. These estimates fell well below estimates of available soil P obtained using traditional extraction procedures. Our results give both theoretical and practical insights into the kinetics of C and P utilization in young soils, as well as show changes in microbial P availability during early stages of soil development.

Differences in CO2 dynamics between successional mire plant communities during wet and dry summers

Questions: What impact do a wet and a dry growing season have on CO2 dynamics of mire plant communities along a primary succession gradient from the initiation stage to the bog stage? Location: Mires on a land uplift coast, Finland. Methods: We measured CO2 dynamics and vascular plant green area development on five mires that form a sequence of mire succession. TWINSPAN was used to define successional mire plant communities and regression analyses were used to explore the temporal variation in CO2 dynamics of the communities. Results: CO2 dynamics of successional plant communities reacted differently to a wet and a dry growing season. The net CO2 uptake rate of the earlier successional communities decreased in the dry growing season due to a decrease in photosynthesizing leaf area. Concurrently, CO2 uptake of the later successional communities moderately increased or did not change. Generally, the difference in net ecosystem exchange (NEE) between a dry and a wet year resulted from the altered rate of gross photosynthesis (PG) rather than ecosystem respiration (RE). Conclusions: Critical factors for the more stable carbon (C) gas dynamics in the later stages of mire succession were (1) higher autogenic control of the physical environment and (2) an increase in the number of factors regulating the PG rate. These factors may buffer mire ecosystems (in terms of the C sink function) from extreme and unfavourable variations in environmental conditions.

Demographic Consequences of the Two Reproductive Modes in Poa alpina L. along a Primary Succession Gradient in the Central Alps

Poa alpina grass plays a predominant role across the entire range of primary succession on alpine glacier forelands. One demographic factor that reacts clearly to changing environmental conditions is reproduction. Using permanent plot data, the complete life cycle of Poa alpina was studied along a successional gradient of the Rotmoos glacier foreland (2300–2400 m a.s.l., Central Alps, Austria) over a period of three years. We used matrix modeling to study the importance of the generation of plantlets and seedlings along the successional gradient and their ability to form adult individuals, and we hypothesized that plantlets develop faster to adults than seedlings because they start already with 3–4 developed leaves.The study showed that plantlet and seedling fecundities of Poa alpina changed differently in the course of succession: seedling establishment was observed over the entire range of the successional stages, whereas plantlet establishment almost vanished with ongoing succession. In the pioneer stage, plantlets were more important than reproduction by seedlings. But we found neither a higher survival rate nor a significant advantage in development to adults for plantlets compared to seedlings. Opportunistic reproduction—plantlets under harsh abiotic conditions, seeds under increasing density—may therefore explain the fact that the species is ubiquitous along the whole glacier foreland.

Root endophytes along a primary succession gradient in northern Finland

Primary successional vegetation gradients are characterized by changes in the soil microbial communities. However, information on possible shifts of the root endophytes along these gradients is scarce. The objective of the current study was to identify root endophytic fungi from a primary successional gradient on land uplift seashore of a geographically isolated island area. We applied a sequencing approach by amplifying the ITS region with fungal specific primers. We used mainly an isolate-based method, and to compare the abundance of culturable and unculturable endophytes, direct sequencing of one representative plant specimen Deschampsia flexuosa was also carried out. A total of 38 cultured endophytic strains were sequenced from Empetrum nigrum (Empetraceae), Vaccinium vitis-idaea (Ericaceae) and Deschampsia flexuosa (Poaceae). Out of these, 27 were identified as Phialocephala fortinii, three as Mollisia minutella, four as Phialophora sp., one as Ascomycetes sp. and three remained unidentified. The strains clustered into five clades in the phylogram, mostly irrespective of the successional stages and hosts from which they had been isolated. The early successional seashore dune ridge plants however, seemed to host a distinct fungal taxon, Phialophora sp. Culture-independent methods were applied on a root sample of a mid-successional Deschampsia flexuosa specimen and a total of 16 clones were randomly selected and sequenced. Out of 16 sequences, 13 were identified as unculturable strains and three showed closest similarity to a basidiomycete Cortinarius callisteus. The unculturable sequences were grouped into two main clades and were different from any culturable isolate in this study. Our results suggest that (i) P. fortinii dominates the isolate data at mid to late successional stages, (ii) roots of the ericaceous plants and the grass Deschampsia flexuosa are colonized by the same endophytic fungi in this ecosystem, and (iii) unculturable endophytes are common and potentially more abundant than the culturables. To our knowledge, this is the first report of the molecular phylogenies of the DSE in the mid-boreal zone and also the first report of the unculturable root endophytes of D. flexuosa.

Population dynamics along a primary succession gradient: do alpine species fit into demographic succession theory?

Understanding processes and mechanisms governing changes in plant species along primary successions has been of major importance in ecology. However, to date hardly any studies have focused on the complete life cycle of species along a successional gradient, comparing pioneer, early and late-successional species. In this study it is hypothesized that pioneer species should initially have a population growth rate, lambda, greater than one with high fecundity rates, and declining growth rates when they are replaced by late-successional species. Populations of late-successional species should also start, at the mid-successional stage (when pioneer species are declining), with growth rates greater than one and arrive at rates equal to one at the late successional stage, mainly due to higher survival rates that allow these species to persist for a long time. The demography of pioneer- (Saxifraga aizoides), early (Artemisia genipi) and late-successional species (Anthyllis vulneraria ssp. alpicola) was investigated together with that of a ubiquitous species (Poa alpina) along the Rotmoos glacier foreland (2300-2400 m a.s.l., Central Alps, Austria) over 3 years. A matrix modelling approach was used to compare the main demographic parameters. Elasticity values were plotted in a demographic triangle using fecundity, individual growth and survival as vital rates contributing to the population growth rates. The results largely confirmed the predictions for population growth rates during succession. However, high survival rates of larger adults characterized all species, regardless of where they were growing along the succession. At the pioneer site, high mortality rates of seedlings, plantlets and young individuals were recorded. Fecundity was found to be of minor relevance everywhere, but it was nevertheless sufficient to increase or maintain the population sizes. Demographically, all the species over all sites behaved like late-successional or climax species in secondary successions, mainly relying on survival of adult individuals. Survival serves as a buffer against temporal variation right from the beginning of the primary succession, indicating a major difference between primary and secondary succession.

Patch dynamics and local succession in a sandstone area with frequent disturbance

Abstract. A system of sand talus cones in a small forested rocky sandstone area was investigated to determine the importance of disturbance‐related dynamics and mesoclimate to vegetation differentiation. These cones (usually 3–12 m long) are formed by the accumulation of sand at the foot of sandstone rocks and are subject to frequent disturbance by the transport of sand. Vegetation was recorded both at the whole‐cone level and at the within‐cone level; the latter was approached by means of a transect of small squares along each cone. Soil profiles were recorded at the upper, centre and bottom parts of the cones. To express mesoclimatic differentiation among the cones within the rocky area, the potential insolation was calculated from the horizon angular height and the likelihood of thermal inversions was estimated by the height of the cones above the valley bottom.The major environmental factor correlated with variation in cone vegetation is the disturbance dynamics as determined by the structure of the soil profile; active cones (with sand at the surface) are colonized by different species as compared with non‐active cones (with surface covered by humus sediments). There is a clear primary succession gradient from plants able to cope with continuous sand transport to stands of tall bryophytes, small shrubs and herbs. Quite often the same gradient is found within cones, with late successional vegetation confined to the lower, stabilized parts of the cones, while the upper part is still being affected by sand transport. The differentiation along the gradient of disturbance is much stronger than the differentiation due to climatic or other gradients.Indirect data indicate that the long‐term average rate of sand accumulation is ca. 1 mm/yr. If sand transport ceases, Sphagnum peat accumulates on some of the cones; the depth of accumulated Sphagnum remains may reach 50 cm. The development of the peat layer is but weakly correlated with the measured variables suggesting that random processes at the beginning of Sphagnum establishment may be a driving force. By measuring Sphagnum growth and decomposition, and the peat density, we estimated the time needed for their development to be several hundreds of years.

Microbial and Plant Community Structure Across a Primary Succession Gradient

The formation of the organic layer within Scandinavian forest soil started about 10000 yr ago, following the retreat of the continental ice sheet. Since then the land has been slowly rising in northern Europe and uplift still occurs on the coast of the Bothnian Bay at a rate of about 0.6-0.9 m per 100 yr. Four, 300 m long, successional gradients were studied from the shoreline to a Scots pine (Pinus sylvestris) stand with a fully developed humus layer of a few centimetres' thickness. The plantless shoreline was followed by small foredunes and dunes, characterized by Agrostis stolonifera and Leymus arenarius, respectively, and the deflation basin characterized by lichens and sparse Festuca ovina and Deschampsia flexuosa. The study sites situated in Scots pine stands of about 25 and 40 yr age were characterized by sparse dwarf shrubs, lichens and bryophytes. The amount of organic matter in soil increased along the gradient. When the microbial biomass, estimated as indicative phospholipid fatty acids (PLFAs), was calculated on the organic matter basis, the total microbial biomass as well the amount of bacterial PLFAs decreased from the earlier stages of succession towards the pine forest. The ratio of fungal to bacterial PLFAs increased along the succession. The bacterial community structure in the shore soil was different to that in the dune soil or in forested zones. On the plantless shore the microbial community was almost completely described by PLFAs indicative of Gram-negative bacteria. In addition to these bacterial PLFAs, dunes were characterized by PLFAs indicative of actinomycetes. Thus, the fungal part of the microbial community seemed to respond most to the accumulation of organic matter and increasing C/N ratio, while the bacterial biomass and bacterial community structure seemed also to reflect the composition of the vegetation and the varying quality of the organic matter.