Effects of nitrogen limitation on species replacement dynamics during early secondary succession on a semiarid sagebrush site

Summary A soil nitrogen (N) availability gradient was induced on a disturbed sagebrush site in northwestern Colorado by fertilizing with nitrogen (high available N), applying sucrose (low available N), and applying neither nitrogen nor sucrose (control). Species composition was studied for 3 years. At the end of the study, N concentration of aboveground tissue of 3 major species was determined. The rate of species replacement was most rapid on plots receiving the sucrose treatment and was slowest on plots receiving the N treatment. Early-seral dominats had greater tissue N concentrations when availability of the resource was high but lower tissue N concentrations when available soil N became limited. Midseral dominants displayed the opposite pattern. These results suggest that the supply of available soil N, and therefore the dynamics of N incorporation in perennial plant tissue, is a primary mechanism in controlling the rate of secondary succession within this semiarid ecosystem.     

Restoration,succession and climatic change

Two outstanding papers on restoration and succession are briefly discussed as model papers for the type of research papers Appl. Veg.Sci. should publish. The paper on restoration concentrates on the introduction of hay to a site in order to speed up the introduction of target species. The paper on succession discusses the importance of plant colonization ‘windows’ opened by extreme weather events for succession and for offering optimum periods for intervention in restoration practice. Some remarks are also made on the electronic availability of ecology papers.     

Oscillating vegetation dynamics in a wet heathland

Abstract. Question: The significance of disturbances caused by periodical inundation was investigated with respect to its effects on vegetation dynamics, species richness and fluctuations, and to the relevance of certain plant properties. Location and Method: At a sod‐cut stand within nutrient‐poor inland sand dunes, permanent plots along a transect were surveyed over a period of up to nine years after sod cutting. Results: In contrast to never inundated plots, periodically inundated plots were characterized by low vegetation cover and by high numbers of species belonging to many different communities, each of them with a low cover. Periodical inundations favoured the presence of pioneers, species tolerant of disturbances, species adapted to wet conditions and stoloni‐ferous species. Furthermore, annual fluctuations of species within each plot were higher and most species occurred only sporadically. Discussion: A comprehensive model is presented describing the relevant processes identified in the littoral zone. Changing water tables result in the creation of gaps. The re‐colonization of these gaps follows mainly from vegetative regeneration and less to the dispersion of diaspores. Highest species numbers in the zone of moderate disturbances result from a high rate of re‐colonization in spite of local extinctions following each disturbance event. It is suggested that colonization abilities are among the most important features for species occurrence at a site rich in disturbances (more important than competitive abilities and more important than a slow rate of displacement). For nature conservation such sites are very important, because they allow (rare) pioneer species to survive for longer periods of time.     

Effects of temporal variability of disturbance on the succession in marine fouling communities in northern-central Chile

We investigated the effects of temporal variability in a disturbance regime on fouling communities at two study sites in a northern-central Chilean bay. Fouling assemblages grown on artificial settlement substrata were disturbed by mechanical removal of biomass at different time intervals. Using one single disturbance frequency (10 disturbance events over 5 months) we applied 7 different temporal disturbance treatments: a constant disturbance regime (identical intervals between disturbance events), and 6 variable treatments where both variableness and sequences of intervals between disturbance events were manipulated. Two levels of temporal variableness (low and high, i.e. disturbance events were either dispersed or highly clumped in time) in the disturbance regime were applied by modifying the time intervals between subsequent disturbance events. To investigate the temporal coupling between disturbance events and other ecological processes (e.g. larval supply and recruitment intensity), three different sequences of disturbance intervals were nested in each of the two levels of temporal variableness. Species richness, evenness, total abundance, and structure of communities that experienced the various disturbance regimes were compared at the end of the experiment (15 days after the last disturbance event). Disturbance strongly influenced the community structure and led to a decrease in evenness and total abundance but not species richness. In undisturbed reference communities, the dominant competitor Pyura chilensis (Tunicata) occupied most available space while this species was suppressed in all disturbed treatments. Surprisingly, neither temporal variableness in the disturbance regime nor the sequence of intervals between disturbance events had an effect on community structure. Temporal variability in high disturbance regimes may be of minor importance for fouling communities, because they are dominated by opportunistic species that are adapted to rapidly exploit available space.     

Comparison of the response of meio- and macrobenthos to disturbance and organic enrichment

The effects of the interaction between physical disturbance and organic enrichment with respect to various aspects of subtidal meiobenthic nematode community structure have been examined in a large-scale mesocosm experiment. The relationship between meiobenthic and macrobenthic community structure within this experiment has been explored. Meiofauna and macrofauna showed similar and strongly related responses. Highest diversity was observed in treatment combinations of low levels of disturbance and enrichment and this supports the “Dynamic Equilibrium Model” of Huston [Huston, M.A., 1979. A general hypothesis of species diversity. Am. Nat. 113, 81-101.]. Faunal community structure was more variable at treatment combinations of low levels of disturbance and high levels of organic enrichment. Physical disturbance subdued the effects of high levels of nutrient enrichment. The meio-macrobenthic comparison has been extended to other studies where both meiofaunal and macrofaunal responses have been determined. The response of meiobenthos is often, but not always, the same as that of macrobenthos. These inconsistencies can probably be explained by the constraints of the experimental design of the mesocosm, where macrobenthic recruitment is not possible, as well as the different ecology of these two faunal groups.     

The role of the permanent soil seed bank in early stages of a post-agricultural succession in the Inland Pampa,Argentina

We studied the soil seed bank composition in four old fields of different ages, after abandonment from agriculture. Complete seed bank composition was assessed by direct seed separation from soil samples and identification to species. Most species found in the seed bank were not important in the present seral communities. Seed of the species that dominated the early succession were generally not found. Additionally, there were very few propagules rather than on the germination of in situ propagules. We suggest that pampean grasses evolved under that the course of post-agricultural succession will depend strongly on the pattern of arrival of exogenous propagules rather than the germination of in situ propagules. We suggest that pampean grasses evolved under disturbances of low intensity and/or a disturbance regime dominated by small gaps, in which open areas could be rapidly colonized from the edges and/or by remnant vegetative propagules. The changes produced by the introduction of agriculture triggered the invasion by exotic species adapted to the new disturbance regime.     

A note on the dynamics of montane virginPicea yezoensis forest in the Greater Khingan Mts,Northeastern China

The vertical growth ofLarix gmelini in the north of the Greater Khingan Mts., north-eastern China is very fast during the first 50 yr. ThoughPinus sylvestris var.mongolica shows a higher growth rate, it will be replaced by the larch population.Picea jezoensis is the most tolerant tree species and will replace all other trees.The fire cycle is 110–120 yr. Most of the montane areas from 800 to 1000 m a.s.l. are not occupied by spruce forest.There is a sapling bank with large numbers of saplings at ages below 20 yr in the understory in the spruce forest. The sapling bank is the key to maintain the steady state of the spruce forest. The persistent sapling bank in the life history of spruce governs the dynamics of the spruce forest controlled by the intensity and frequency of disturbances.The author is H. Q. Wu     

Short-term effects of a catastrophic beaver dam collapse on a stream fish community

Synopsis We examined the short-term effects of the natural catastrophic collapse of a beaver dam on downstream benthic insect density and fish community structure in a headwater tributary of the Mississippi River. The catastrophic collapse of the dam and ensuing flash flood resulted in a dramatic (< 90%) decrease in benthic insect density in riffle and pool habitats. Sixty days after collapse of the dam, insect densities in riffles were 62% of pre-collapse densities. Insect recolonization of pools was slower than for riffles; 60 days after collapse of the dam insect densities in pools were 8% of pre-collapse levels. Collapse of the beaver dam altered the structure of the downstream fish community by causing a short-term (2–4 days) influx of pond species, resulting in a brief increase in species richness and abundance. Fish species richness and abundance then decreased for 4–60 days to levels below those prior to the collapse.     

Spontaneous vegetation dynamics and restoration prospects for limestone quarries in Lebanon

Abstract. In terms of restoration planning, the analysis of natural regeneration processes represents a valuable starting point for the selection of suitable species to be used. This paper aims to identify colonizing key species among the pioneer vegetation of limestone quarries in Lebanon, to identify potential restoration strategies in terms of reconstitution of the pre‐mining vegetation cover. Characteristics of the major ground colonizers after disturbance were identified in a representative quarry in the thermo‐Mediterranean biozone. The floristic inventories resulted in a matrix of cover values of 107 species × 14 plots. Correspondence analyses were used to pinpoint similarities in the distribution of key species among the different environmental characteristics of sites. The main results reveal a heterogeneous floristic composition along the regeneration gradient, where annual R‐strategy taxa such as Inula viscosa and Ainsworthia cordata dominate on very perturbed and degraded sites. Less degraded areas within the quarry were rich in herbaceous perennial or shrub species such as Geranium dissectum, Stachys distans, Salvia triloba and Ptilostemon chamaepeuce. On relatively non‐degraded areas woody and shrub perennials such as Pinus brutia, Pistacia palaestina and Quercus calliprinos dominate, along with less stress tolerant taxa such as Arbutus andrachne and Cistus creticus. Species to be used in restoration projects should be chosen from among the local vegetation, according to frequency of occurrence during the whole succession process which will reflect their adaptability to local conditions and their relevance to restoration objectives.     

Trap-nesting bees and wasps colonizing set-aside fields: succession and body size,management by cutting and sowing

Trap-nesting bees and wasps (Hymenoptera Aculeata) colonizing crop and fallow fields in an agricultural landscape were studied using 20 sown fields (pea, barley, rye, clover-grass mixtures, Phacelia tanacetifolia) and 20 fields with naturally developed vegetation (1- and 2-year old fields, both mown and unmown, and old meadows). Fourteen species of Apoidea, 4 of Sphecidae, 1 of Eumenidae and 4 of parasitoids were reared from reed nests exposed in these 40 fields of 10 field-types. Fields with naturally developed vegetation had twice as many species as sown fields, due to the distribution pattern of the 14 bee species, whereas the 9 predatory species (wasps and parasitoids) showed a rather uniform distribution. None of the trap-nesting bees were found in Phacelia fields, despite contrasting expectations of beekeepers. Old meadows showed a particularly high abundance and species richness, since only 10% of all traps were exposed, but 32% of all bee nests were sampled in old meadows, including 4 bee species that were not found elsewhere. Accordingly, species richness of fields with naturally developed vegetation showed a significant increase with age. Variability in Hymenoptera species numbers could be explained by corresponding differences in plant species numbers. The alternative hypothesis that field size or field connectivity influenced species richness was not supported. Habitats with great floral diversity appeared to offer better and richer food resources for the flower-visiting bees, whereas food availability apparently did not influence predatory wasps. The bees Osmia caerulescens and Megachile versicolor that had colonized early-successional fields took twice as long to provision cells as those that colonized late-successional meadows characterized by a greater plant species richness. In contrast, the eumenid wasp Ancistrocerus gazella took a similar period of time to provision cells in both field types. In addition, bee and wasp species of plant-species-poor fields were on average significantly larger than those of plant-species-rich fields. Thus, body size appeared to be a good predictor of colonization ability. Management by cutting greatly increased plant species richness in early-successional set-aside fields and thus doubled species richness of bees. Cutting of early-successional habitats can be expected to benefit insects and plants in general, whereas older grassland should show the greatest insect diversity when both mown and unmown parts are present.     

Stand dynamics of a yellow-poplar (Liriodendron tulipifera L.) forest in the Appalachian Mountains, Virginia, USA

Yellow-poplar (Liriodendron tulipifera L.) is a large, rapidly growing, shade-intolerant tree species common after disturbances on moist sites in the Appalachian Mountains. The species is typically scattered throughout old-growth mesophytic forests, where periodic gap formation creates conditions favorable for yellow-poplar establishment and growth. On abandoned agricultural fields, however, it is common for nearly monospecific forests of yellow-poplar to develop.This study examines stand dynamics of a yellow-poplar forest in western Virginia, USA that was established on agricultural fields abandoned in the late 1940s. Increment cores were collected from yellow-poplar trees growing on exposed ridgetops and in a more sheltered hollow. Tree-ring data show that the forest is even-aged. Tree establishment began about 5 years earlier on the ridgetops than in the hollow. Major ice storms disturbed the forest in 1978 and 1994, with two separate events in 1994. Ice storms disturb forests by depositing heavy loads of freezing rain on trees, breaking or uprooting them. The dendroecological data collected for this study provide little support for the hypothesis that ice storm disturbance promotes the establishment of new yellow-poplar cohorts. However, the data show that radial growth of some trees increased after ice storm disturbance, a pattern that reflects the increased availability of light following disturbance. Radial growth declined in some other trees as a consequence of severe injury during the storms.Radial growth responses following the 1978 ice storm were stronger on the ridgetops than in the hollow, suggesting that tree damage was more severe on the higher, more exposed sites. Growth responses were relatively mild following the storms of 1994, and did not exhibit pronounced topographic variations.     

Osmolytes in salinity-stressed Iris hexagona

We characterize the salinity stress response of Iris hexagona, a freshwater species, by measuring three putative osmolytes, betaine, proline, and dimethylsulphonio-propionate (DMSP) in plants after short (3 days) and long-term (4–5 months) exposure to NaCl. HPLC analyses show that untreated control leaves contained 2.1, 0.2 and 3.2 mg g⁻¹ DW of betaine, proline, and DMSP, respectively, and establish the presence of these compounds in the Iridaceae. Within 4 days of salinity stress (200 mM NaCl) betaine, proline and DMSP increased significantly. Among vegetative tissues, the highest level of proline occurred in roots; betaine and DMSP were highest in leaves. Analyses of generative tissue (flowers and flower stalks) after long-term exposure showed the highest levels (>10 mg g⁻¹ DW) of all examined compounds in petals but only proline and betaine increased with salinity. All three substances showed a basipetal gradient in flower stalks. Although the examined compounds responded to salinity, the osmometry of the sap indicated that they comprise less than 10% of the osmotically active solutes. The temporal and spatial changes in the distribution of the analyzed compounds indicate complex responses to salinity.     

Two different effects of calcium on aquaporins in salinity-stressed pepper plants

Two different effects of calcium were studied, respectively, in plasma membrane vesicles and in protoplasts isolated from roots of control pepper plants (Capsicum annuum L cv. California) or of plants treated with 50 mM NaCl, 10 mM CaCl(2) or 10 mM CaCl(2) + 50 mM NaCl. Under saline conditions, osmotic water permeability (P ( f )) values decreased in protoplasts and plasma membrane vesicles, and the same reduction was observed in the PIP1 aquaporin abundance, indicating inhibitory effects of NaCl on aquaporin functionality and protein abundance. The cytosolic Ca(2+) concentration, [Ca(2+)](cyt), was reduced by salinity, as observed by confocal microscope analysis. Two different actions of Ca(2+) were observed. On the one hand, increase in free cytosolic calcium concentrations associated with stress perception may lead to aquaporin closure. On the other hand, when critical requirements of Ca(2+) were reduced (by salinity), and extra-calcium would lead to an upregulation of aquaporins, indicating that a positive role of calcium at whole plant level combined with an inhibitory mechanism at aquaporin level may work in the regulation of pepper root water transport under salt stress. However, a link between these observations and other cell signalling in relation to water channel gating remains to be established.     

The significance of waterfowl grazing in the primary vegetation succession on embanked sandflats

In the annual stage of a primary plant succession on desalinating sandflats of the Lauwerszeepolder, N Netherlands, grazing by waterfowl is a dominant factor, as is illustrated by exclosures. It is concluded from population dynamical data, that both the removal of biomass and the predation of seeds sustain the annual species and retard succession to wards the next stage of a perennial grassland by several years. Yearly mowing in September slightly enhances succession by selectively affecting the dominant annual Salicornia species. The recurrence of high salinities in the upper soil layer of bare, c.q. grazed surfaces is expected to limit the invasion of grass species and other perennials.Nomenclature of plant taxa follows Heukels/van der Meijden (1983), Flora van Nederland, 20th ed. Wolters-Noordhoff, Groningen.This article would not have been written without the contributions of many students and colleagues, equally aware of the rare opportunity we had to study a genuine primary succession. I wish to thank them all. The help in the moving experiment of J. Franke, S. Nijdam, the late M. Robertus, H. Vrieling and J. D. D. Hofman is gratefully acknowledged. Thanks are due to the State Authority of the IJssellake Polders, for permitting access to and use of the experimental areas.     

Exogenous spermidine affects polyamine metabolism in salinity-stressed Cucumis sativus roots and enhances short-term salinity tolerance

We investigated the effects of short-term salinity stress and spermidine application to salinized nutrient solution on polyamine metabolism and various stress defense reactions in the roots of two cucumber (Cucumis sativus L.) cultivars, Changchun mici and Jinchun No. 2. Seedlings grown in nutrient solution salinized with 50 mM NaCl for 8 d displayed reduced relative water content, net photosynthetic rates and plant growth, together with increased lipid peroxidation and electrolyte leakage in the roots. These changes were more marked in cv. Jinchun No. 2 than in cv. Changchun mici, confirming that the latter cultivar is more salinity-tolerant than the former. Salinity stress caused an increase in superoxide and hydrogen peroxide production, particularly in cv. Jinchun No. 2 roots, while the salinity-induced increase in antioxidant enzyme activities and proline contents in the roots was much larger in cv. Changchun mici than in cv. Jinchun No. 2. In comparison to cv. Jinchun No. 2, cv. Changchun mici showed a marked increase in arginine decarboxylase, ornithine decarboxylase, S-adenosylmethionine decarboxylase and diamine oxidase activities, as well as free spermidine and spermine, soluble conjugated and insoluble bound putrescine, spermidine and spermine contents in the roots during exposure to salinity. On the other hand, spermidine application to salinized nutrient solution resulted in alleviation of the salinity-induced membrane damage in the roots and plant growth and photosynthesis inhibition, together with an increase in polyamine and proline contents and antioxidant enzyme activities in the roots of cv. Jinchun No. 2 but not of cv. Changchun mici. These results suggest that spermidine confers short-term salinity tolerance on cucumber probably through inducing antioxidant enzymes and osmoticants.     

Temporal patterns of benthic community development in an Arctic fjord (Kongsfjorden, Svalbard): results of a 24-year manipulation study

We investigated temporal patterns of recolonisation and disturbance in a benthic hard bottom community in high-arctic Kongsfjorden from 1980 to 2003 through annual photographic surveys. A manipulative sampling design was applied, where half of the study area (treatment areas) was cleared at the beginning of the study. Twenty-three different taxa and groups of benthic epifauna were found in the photographs. The benthic community structures of treatments and controls converged within the first decade, but significant differences prevailed until ≤13 years after the start of the study. We could distinguish between three different time intervals with increased inter-annual changes. While the observed differences during the first two intervals could be attributed to recolonisation and succession, the changes in interval 3 were mostly due to increased external forcing and characterised by low inter-group and high inter-annual differences. During this interval, brown algae (mainly Desmarestia) and the sea urchin Strongylocentrotus droebachiensis emerged in high densities, while sea anemone populations declined. Different recolonisation patterns for individual species were related to life span, rate of maturity, predators and larval settlement. We could not find a climax stage in the succession for the benthic community at Kvadehuken, presumably due to the constant level of disturbances at the site.     

高岚河大型底栖动物时空分布及影响因子研究

为了解大型底栖动物沿河流分布的物种丰富度变化及环境因子对其种群的影响,于2016年12月对高岚河的大型底栖动物进行了调查,并对理化指标进行了测定。研究结果发现,物种丰富度沿河流纵向梯度变化差异性不显著,生物多样性指数沿河流变化差异不显著,说明高岚河大型底栖动物群落结构稳定。高岚河上游的生态指示种为高翔蜉和四节蜉,中游的生态指示种为四节蜉和萝卜螺,下游的指示种为萝卜螺。模糊聚类分析发现高岚河大型底栖动物类群可划分为3个生境类群,且在梯度分布下群落生境较相似,大型底栖动物群落间关系接近,分布均匀。典范对应分析发现影响大型底栖动物群落结构沿梯度分布的主要因子是海拔、pH、浊度和深度。     

Seed dynamics during forest succession in Costa Rica

Soil seed banks and current seed inputs each play a role in tropical succession. We compared the abundance and floristic composition of seeds from these two sources at a Costa Rican site by germinating seeds from the soil, measuring seed inputs for 3 yr, and monitoring the earliest colonists in a forest clearing.There were an estimated 6800 viable seeds/m² in the soil of 3.3-yr-old vegetation, 9500 seeds/m² in 11-yr-old vegetation, and 7000 seeds/m² in a 75-yr-old forest. An estimated 10100 seeds/m² fell on the soil surface of the young successional vegetation during 3 yr and 3700 seeds/m² fell during that same time in the forest.Locally produced seeds accounted for about 75% of the seed input to the soil surface early in succession. Seeds dispersed out of young successional vegetation increased the quantity and species richness of the seed input and storage in an adjacent forest. Much of the species richness of the young successional vegetation resulted from seeds dispersed there from other communities by animals.Deforestation stimulated germination of most seeds in the surface soil of the old forest, including seeds of the dominant canopy tree. The recruitment of seedlings from the soil seed bank numerically overwhelmed that from post-disturbance seed rain and sprouts.We evaluated patterns of soil seed storage during succession and predicted the ability of vegetation of differing ages to respond to disturbance. Immediately after disturbance the number of seeds in the soil plummeted due to mortality, low inputs, and germination. As the vegetation regrew, the soil seed bank increased to a peak after 4 to 7 yr, then gradually decreased to its pre-disturbance size. High-frequency pulses of disturbance should result in reduced species richness, dominance by species with long-lived seeds, and fast recovery by seedling recruitment from the soil seed bank.Journal series number 6459 from the Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida 32611, USA.Reprint requests to J. J. E. at Florida.