Species richness and cover along a 60-year chronosequence in old-fields of southeastern Spain

We analyse changes in plant cover and species richness along a 60-year chronosequence in semi-arid Mediterranean old-fields of southeastern Spain. The objectives were: (i) to study patterns of species richness along the abandonment gradient in semi-arid conditions (e.g., to test the “humped-back model” in our system); (ii) to test whether different broad life forms (annuals, forbs, grasses and woody species) showed different patterns along the abandonment gradient, and (iii) to examine to what extent plants with different dispersal strategies dominate at different stages of succession. The explained variance of the regression relating species richness to years since abandonment is improved when considering different life forms. The results suggest that cover and richness of different functional groups show a non-linear unimodal (often positive-skewed) pattern along the gradient (age since abandonment). Maximum total richness is found at young stages of abandonment (<20 years), when most life forms and dispersal strategies coexist. Annuals and perennial forbs reached their maximum richness during the first 10 years of abandonment. About 45% of total woody species richness is reached at this time as a consequence of early colonization of zoochorous shrubs. While the results showed a tendency towards a life-form replacement sequence, the pattern is not so clear when looking at the different dispersal strategies. The results complement previous results in Mediterranean conditions and emphasise the importance of considering different functional types when studying successional patterns. This revised version was published online in June 2006 with corrections to the Cover Date.     

Drivers of secondary succession rates across temperate latitudes of the Eastern USA: climate,soils, and species pools

Climate change is widely expected to induce large shifts in the geographic distribution of plant communities, but early successional ecosystems may be less sensitive to broad-scale climatic trends because they are driven by interactions between species that are only indirectly related to temperature and rainfall. Building on a biogeographic analysis of secondary succession rates across the Eastern Deciduous Forest (EDF) of North America, we describe an experimental study designed to quantify the relative extent to which climate, soil properties, and geographic species pools drive variation in woody colonization rates of old fields across the EDF. Using a network of five sites of varying soil fertility spanning a latitudinal gradient from central New York to northern Florida, we added seeds of nine woody pioneer species to recently tilled old fields and monitored first-year growth and survivorship. Results suggest seedlings of southern woody pioneer species are better able to quickly establish in fields after abandonment, regardless of climate regime. Sites of lower soil fertility also exhibited faster rates of seedling growth, likely due to the slower development of the successional herbaceous community. We suggest that climate plays a relatively minor role in community dynamics at the onset of secondary succession, and that site edaphic conditions are a stronger determinant of the rate at which ecosystems develop to a woody-dominated state. More experimental research is necessary to determine the nature of the herbaceous–woody competitive interface and its sensitivity to environmental conditions.     

黄土高原自然植被演替过程中的植物特征与土壤元素动态

在3～149年的时间尺度上,对黄土高原植被次生演替过程中植物特征和土壤养分元素动态进行了研究.结果表明:1) 随着演替时间尺度的延伸,土壤全C、全N含量呈增加趋势,而土壤全K、全Na和土壤pH值呈下降趋势,土壤全P变化趋势不明显;此外,表层(0～10 cm)土壤CaO 含量在演替系列上呈下降趋势,深层(20～30 cm,40～50 cm)则呈增加趋势.演替过程对几种土壤化学元素含量影响的程度随着土壤深度的增加而减弱.2)植物群落物种丰富度在演替的中间阶段最高.3) 在演替的早期阶段,植物群落优势种往往具有稳定的土壤种子库、CR-生活对策和S-繁殖对策,在贫瘠的土壤上具有较强的竞争能力,且具有较强的水平扩展能力和克隆繁殖能力,C-生活对策、在土壤全N含量较高的生境中具有较强竞争能力的多年生植物,在演替中后期占据群落的优势地位.此外,在所涉及的植物特征中,多年生生活史,C-、CR-、SC-、SR-、S-生活对策,以及R-、W-、Bs-、VBs- 和V-繁殖对策等特征在非优势物种中出现的频率较高.4) C-、 SC-生活对策, 克隆能力, 多年生生活史, 水平扩展能力, 种子的动物传播方式,秋季开花,荚果、坚果等特征出现的比例在一定程度上与土壤全C、全N 和全K含量正相关; 而 S-、SR-、R-、CR-生活对策,一、二年生生活史, 种子繁殖, S-繁殖对策,以及胞果、蒴果等特征的比例与土壤全Na、CaO含量和土壤pH正相关.5)在演替过程中出现的植物均属草本植物生活型,因此,草原可能是黄土高原上受制于大尺度环境条件(显域生境)下的优势植被类型(特别是降雨量不超过550 mm的地区).     

Composition,size and dynamics of the seed bank in a mediterranean shrubland of Chile

Abstract Analysis was performed of the richness and abundance of woody species, forbs, and annual grasses in the easily germinating soil seed bank (henceforth seed bank) in a mediterranean shrubland of central Chile. The effects of successional development after fire and by microsite type (underneath or outside shrubs) on the density of seeds in the soil, and the relationship of species abundance in the seed bank with its abundance in the above‐ground vegetation was examined. A total of 64 plant species were recorded in the seed bank, of which 44 were annual or biannual. Eight species were woody and another eight were perennial herbs. Four could not be identified to species level. The highest richness of established herbaceous species was recorded in late spring, with 31 species. The regeneration of the herbaceous vegetation was driven by the annual production of seeds and by a reserve of short‐lived propagules in the soil. Density of all germinating seeds was significantly higher during late spring and late summer. Density of grass seeds was greater during late spring, while that of all other species was greater during late summer. Annual grass seeds accumulated in higher proportion at exposed microsites rather than under woody canopy, and in young (< 5 years old) and intermediate‐age patches (10–20 years old) rather than in mature vegetation (30–50 years old). The abundance of established woody and herb species was uncorrelated with that of the seed bank.     

Restoration of a Landfill Site in Berlin, Germany by Spontaneous and Directed Succession

We report on spontaneous and directed succession on a dry sandy landfill site of low fertility at Berlin‐Malchow, Germany. Changes in species composition and cover were followed on unmown and mown permanent plots of 2 × 2 m size through 5 years of vegetation development. Species richness on unmown plots was relatively constant during the time of observation, with 20 to 25 species per 4 m². Total cover of unmown plots continuously increased from approximately 10% in the first year to 80% in the fifth year. There are no clearly discernible sequential successional stages until present. The species composition includes species of all life forms, which colonized the site immediately after the initiation of the succession process representing the initial floristic composition type of vegetation development. However, perennial grasses and herbs gradually increased in cover up to approximately 40%. Woody plants were also present from the first year of succession and increased up to more than 20% cover in the fifth year, forming a shrub layer (>0.5 m) after the second year. Mowing significantly increased species richness, which was evident from the third year onward. This effect was mainly due to the reduction of the tall perennial grass Calamagrostis epigejos. Solidago canadensis and woody species were also significantly affected (lower cover and height), whereas short perennial herbs like Plantago lanceolata and Trifolium repens benefited from mowing.     

Colonization and early succession on anthropogenic soils

Abstract. This paper reports on vegetation development on permanent experimental plots during five years of succession. Nine (1 m²) plots were filled with three typical substrates from man-made habitats of urban and industrial areas in the region of Berlin. The three substrates (a commercial ‘topsoil’, a ruderal ‘landfill’ soil and a sandy soil), differ in organic matter and nutrient contents. Relevés of species composition and percent cover of each species present were made monthly during the growing season from the start of vegetation development. This paper describes the different successional pathways on topsoil and ruderal soil and the colonization process on sandy soil. On topsoil, ruderal annuals are dominant in the first year and are replaced by short-lived perennials from the second year. Those species were replaced by long-lived perennial herbs (Ballota nigra or Urtica dioica) from the third year of succession onwards. On the ruderal land-fill soil the early successional stages are less sharp and the perennial Solidago canadensis is able to dominate within one year after the succession was initiated. On sandy soil there is still an ongoing colonization process, where pioneer tree species like Betula pendula and Populus nigra play a main role. The importance of ‘initial floristic composition’, the role of substrate for community structure and the peculiarities of successional sequences on anthropogenic soils in the context of primary and secondary successions are discussed.     

Divergent Pathways of Successional Recovery for In Situ Oil Sands Exploration Drilling Pads on Wooded Moderate‐Rich Fens in Alberta,Canada

Peatlands in northern Alberta, Canada, are being rapidly impacted by oil sands activities, with potentially long‐term consequences for their recovery. In situ oil sands extraction requires exploration of oil resources on a dense network of drilling pads across the landscape. This study examined the recovery of wooded moderate‐rich (WMR) fens 10 years after abandonment of these sites with minimal restorative measures. Bryophyte and vascular plant diversity, site microtopography, and water chemistry were assessed on drilling pads and in adjacent areas of undisturbed reference habitat. WMR fens affected by drilling activities were divided a priori into two groups based on strongly divergent trends in their successional development. One group represented the majority of WMR fens observed on the land base; at these sites hummock‐forming mosses including minerotrophic Sphagnum species were infrequent and tree recruitment was almost absent. The other group was dominated by Sphagnum species, had Picea mariana and Larix laricina recruitment, and appeared to recover more quickly. Both groups had high abundance of wetland sedges, notably Carex aquatilis. Further, drilling pads belonging to the first group had a high water table, limited elevated microsites, and had surface flooding over a portion of the growing season, in contrast to Sphagnum‐dominated sites. Development of the aquatic, bryophyte‐poor wetland type is comparable to early stages of wetland succession and these systems will recover relatively slowly, likely from decades to more than a century. Restoring part of the vertical distribution of microhabitats before abandonment of these pads could stimulate the successional recovery of vegetation.     

Seed dispersal by wind,birds, and bats between Philippine montane rainforest and successional vegetation

In the moist Neotropics, vertebrate frugivores have a much greater role in the dispersal of forest and successional woody plants than wind, and bats rather than birds play the dominant role in dispersing early successional species. I investigated whether these patterns also occurred in a Philippine montane rainforest and adjacent successional vegetation. I also asked whether seed mass was related to probability of dispersal between habitats. A greater number of woody species and stems in the forest produced vertebrate-dispersed seeds than wind-dispersed seeds. Although input of forest seeds into the successional area was dominated by vertebrate-dispersed seeds in terms of species richness, wind-dispersed seeds landed in densities 15 times higher. Frugivorous birds dispersed more forest seeds and species into the successional area than bats, and more successional seeds and species into the forest. As expected, seed input declined with distance from source habitat. Low input of forest seeds into the successional area at the farthest distance sampled, 40 m from forest edge, particularly for vertebrate-dispersed seeds, suggests very limited dispersal out of forest even into a habitat in which woody successional vegetation provides perches and fruit resources. For species of vertebrate-dispersed successional seeds, probability of dispersal into forest declined significantly with seed mass.     

Woodland regeneration on grazed former arable land: A question of tolerance,defence or protection?

Ecological restoration of native woodlands and wooded pastures on former agricultural land is an important topic in modern conservation practice. The introduction of large herbivores is increasingly used to achieve these aims. We investigated how grazing, resistance traits of plants (concerning herbivory) and associational resistance interact and affect the establishment pattern of woody species on abandoned arable land (N-Belgium, W-Europe). In these early successional tree assemblages, we tested whether grazing increased or decreased spatial heterogeneity, which is supposed to be a crucial factor for biodiversity.With repeated measurements, 2-3 and 5-7 years after the cessation of agricultural use, we sampled 87 grazed and 56 ungrazed plots (314 m²) in 14 sampling areas (former arable land) on nutrient rich, (sandy) loam soils. We recorded established tree frequencies, related to grazing, time, resistance traits and unpalatable/spiny vegetation cover in the herb and low shrub layer. We investigated horizontal and vertical heterogeneity using variances in establishment frequencies and variances in frequencies of trees that were able to grow beyond the browse line, respectively.We found massive colonisation of grazing tolerant and resistant woody species in early successional stages. Grazing decreased frequencies and height of the most abundant tolerant species (mainly Salix caprea L.). After 5-7 years, frequencies of defensive and tolerant species were equal, but the former (mainly Betula pendula Roth) were able to grow beyond the browse line. When the cover of unpalatable/spiny vegetation was high enough (>60% of plot size), it also provided suitable nurse sites for tolerant species to grow out. In early assemblages, grazing increased horizontal and vertical heterogeneity, resulting in intermediate successional stages. In the long-term, the mechanism of associational resistance will also allow non-resistant and tolerant species to grow beyond the browse line and promote forest succession and the order of species establishment and replacement.     

Effects of habitat fragmentation on the movement patterns and dispersal ability of the brown spiny rat (<Emphasis Type="Italic">Maxomys rajah</Emphasis>) in the Planted Forest Zone of Sarawak,Eastern Malaysia

Brown spiny rats (Maxomys rajah) were translocated from continuous secondary forest to small isolated patches of remnant native forest embedded within Acacia mangium plantation in the Planted Forest Zone of Sarawak, East Malaysia, and fitted with tracking spools to monitor behaviours in novel environments and to record responses to a range of habitat edge features. Forest roads, large clearings and acacia plantation compartments were found to pose barriers to dispersal of brown spiny rats over short temporal scales, whereas old regenerating haul trails were readily crossed on 50% of the encounters. Downed woody debris accounted for a greater proportion of the travel route compared with brown spiny rats tracked in secondary and primary forest in Sabah, which may represent heightened predator avoidance in new environments. Provision of downed woody debris within plantation compartments may improve the dispersal ability of brown spiny rats in this modified landscape, and thus promote metapopulation dynamics and colonisation of vacant habitat patches.     

Spontaneous Establishment of Woody Plants in Central European Derelict Sites and their Potential for Reclamation

The performance of woody plants was analyzed in 15 successional seres starting at bare ground in central European manmade habitats. The total cover of woody species after 10 years of succession was significantly related neither to initial soil moisture nor to nitrogen (expressed using Ellenberg indicator values). But the comparison of seres indicates that establishment of woody plants was easier under moderate environmental conditions and retarded in extreme habitats (dry, nutrient-poor, or acid). The arrival of the first woody plants was delayed in dry sites. No significant differences were found between primary and secondary seres, either with respect to the total cover of woody plants reached after 10 years of succession or considering the time of their arrival. In total, 24 woody species (10 shrubs and 14 trees) appeared in the series investigated. Their successional performance (in terms of the number of seres in which the species occurred and maximum cover reached in any sere) was not related to species traits (life strategy, type of mycorrhizae, mode of dispersal, diaspore weight), except for the regeneration strategy, species with seasonal regeneration by seeds were capable of creating higher cover. Betula pendula (European birch) was the most successful species in spontaneous succession, especially on moist sites. Practical suggestions for the management of particular habitats (sites disturbed by mining, sites reclaimed after acid rain deforestation, urban sites, abandoned fields) are provided regarding the establishment of woody plants.     

Secondary Succession and Natural Habitat Restoration in Abandoned Rice Fields of Central Korea

Floristic composition and soil characteristics (moisture, pH, nutrient contents) in abandoned upland rice paddies of different ages were analyzed to clarify the regenerative aspects of succession as a tool for habitat restoration. The study sites represented five seral stages: newly abandoned paddy fields; successional paddy fields abandoned for 3, 7, and 10 years; and a 50‐year‐old Alnus japonica forest. A vegetation sere was apparent in changes of dominant plant species in the order Alopecurus aequalis var. amurensis (annual grass), Aneilema keisak (annual forb), Juncus effusus var. decipiens (rush), Salix koriyanagi (willow), and Alnus japonica (alder) communities. These temporal stages resemble the spatial zonation of vegetation in local riparian floodplain ecosystems, indicating a hydrosere, with soil moisture decreasing over time. Age distributions and life forms of the dominant plant species support a “tolerance” model of secondary succession, in which the established species persist into later successional stages. Persistence of earlier colonizers led to a net cumulative increase in species richness and a more even distribution of species cover with increasing field age. Between 10 and 50 years, vegetation stabilizes as an alder community. Soil moisture content decreased steadily with paddy field age after an initial rise immediately after their abandonment, whereas pools of organic matter, N, P, K, Ca, and Mg, increased with field age. The pace and direction of recovery of native vegetation and natural soil properties in these abandoned rice paddies resembled classic old field succession, a form of secondary succession that often serves as a template for guiding restoration efforts. Active intervention, in particular dismantling artificial levees, could accelerate the recovery process, but natural habitat recovery generally appears sufficiently robust to achieve “passive” restoration of this rare community without intervention.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

Consequences of abandonment for a regional fen flora and mechanisms of successional change

Abstract. Until the 1960s, species-rich vegetation on minero-trophic peaty soüs (fen sites) were characteristic of the alluvial plains in Schleswig-Holstein (Northwest Germany). Today, many of these habitats undergo successional changes due to abandonment. Vegetation development after abandonment can be characterized as a sequence of different successional stages and described in terms of a successional model. Successional stage I includes grazed, mown and recently abandoned sites without dominants. Stages II and III are characterized by the dominance of highly competitive herbaceous species whüe stage IV consists of woody vegetation. Ca. 3000 phytosociological relevés were assigned to the respective successional stages. Mean cover values were calculated for 250 species of the regional fen flora and assigned to successional categories according to their changes in cover in the successional series. According to our results 141 species decrease during succession, while 100 species were restricted to early successional stages and 85 species increased. Abandonment of all fen sites in Schleswig-Holstein will probably lead to the regional loss of 23 species of the fen flora. To identify mechanisms underlying successional change, the successional categories were correlated with life history traits and ecological requirements of the species. Results indicate that both light competition and limitation of sexual reproduction of small-seeded species might play a major role in the decrease and extinction of species during succession. Finally, conservation strategies for endangered species in a cultural landscape are discussed.     

Forest recovery in abandoned agricultural lands in a karst region of the Dominican Republic

This study documents the status of forest vegetation in the karst region of Los Haitises National Park, Dominican Republic, following the abandonment of pastures (5 years), young (5 years) `conucos' (mixed plantings), old (7–30 years) conucos, and cacao plantations (>25 years). We compared these sites to vegetation characteristics of patches of forest in karst valleys (`old forest'–too old to know their exact land use) and on mogote tops with no recent history of human disturbance. The youngest sites date to when squatters were removed from Los Haitises National Park. Forest structure (density, basal area, and species richness of woody plants 1 cm DBH) were all significantly affected by land use. Density was highest in intermediate-aged valley sites (old conucos) and mogote tops, while both basal area and species richness tended to increase with age of abandonment. Although cacao plantations had been abandoned for more than 25 years the species diversity was low, due to continued regeneration of this persistent crop. Abandoned pastures had the greatest nonwoody biomass and were dominated by the fern Nephrolepis multiflora which had completely replaced pasture grasses. An ordination of the woody plant communities separated the mogote tops from valleys, emphasizing the strong control that topography has on the forest community in moist and wet tropical forests on karst substrates. Valley sites were arranged in the ordination in order of their age, suggesting a successional sequence converging on the composition of the `old forest' sites.     

Spontaneous development of urban woody vegetation on differing soils

The general aim of the study was to follow the development of spontaneous woody vegetation on man-made urban habitats in Berlin, Germany. In spring 1968, the vegetation was removed completely in five groups of plots, which had been filled with five different transferred soil material in 1928. The spontaneous vegetation was monitored annually from 1968 to 2006. Especially, the development of shrubs and trees was followed. In addition the growth of the most important herbaceous species, Solidago canadensis, was investigated.

The soils varied from moderately rich sandy soils to rich sandy or loamy soils. On the two sandy soils S. canadensis developed poorly. Tree species dominated (>50% of total cover) on these plots after 14–18 years. On the soils rich in nutrients, S. canadensis developed strongly. Tree species did not dominate there before 21–25 years. The diversity of woody species increased over time and reached 33 species altogether. Indigenous Acer, Quercus and Betula species prevailed. Alien species, typical for urban ruderal sites, were rare.

The results support the idea of establishing urban woodlands by spontaneous succession as a cheap way to develop near-natural plant communities rich in species.     

Secondary succession on sandy old‐fields in Hungary

Abstract. Development of semi‐natural vegetation has recently been a primary concern of restoration efforts. A primary management question is whether active intervention is required or spontaneous secondary succession could suffice. We studied 54 old‐fields in central Hungary, which differed in time since abandonment but which had similar environmental conditions and management histories. The sites were grouped into four age groups according to the time elapsed since cultivation abandonment: 1–5, 6–10, 11–23 and 24–33 yr. In each old‐field we recorded the species and estimated their abundances. We grouped species in two ways: according to life form (annuals, biennials, perennials, woody plants) and according to coenological behaviour (weeds, sand and steppe generalists, specialists). We analysed the changes in species number and abundance in these categories as a function of site age. Contrary to other successional studies, the total number of species did not change significantly among the four age groups. A significant change was detected between the first two age groups as to life‐form composition. Species number and abundance of annuals decreased, while the perennials and woody plants increased. As to coenological behaviour, species number changed only in the first two age groups, while abundance changed in the first three. Weeds quickly disappeared and specialists established and spread, while the species number and abundance of generalists did not change significantly. We concluded that the basic shifts in species composition are almost completed within 10 yr. Most of the late successional species colonized and weeds disappeared. We conclude that there was no need for active intervention in this system: the spontaneous secondary succession leads to semi‐natural vegetation.     

Spatio-temporal patterns along a primary succession on alluvial sediments

Dynamic alluvial landscapes offer many possibilities to study primary succession processes on newly developed habitats. However, within the Central European environmental conditions, where watercourses and their riparian spaces are under severe anthropogenic pressures — water regulation, deforestation, lowering of groundwater — natural processes are limited. We studied primary succession on alluvial stream deposits in an artificial lake, where we aimed to follow the terrestrialisation rate and habitat turnover, along with plant species richness and composition across successional stages. In 30 years, a pristine white-willow riparian forest developed. One half of the initially aquatic habitat became terrestrial. The frequency of change, studied on 11250 quadrats 10×10 m each (on a scale from “no change” to 8 changes) and the mean of change per habitat type (most of the habitats changed 2 to 3 times) revealed only one successional trajectory. The percentage flow chart showed a deterministic pathway of succession. The “time since formation” of a terrestrial habitat shows that more than 20% of the lake was terrestrialised within in the first ten years. We studied species richness and composition along three composed transects, following successional stages. We found that the newly assembled riparian white willow woodland has a surprisingly low colonisation rate of plant species.     

Ecological relationships between seedling emergence and litter cover in the earliest stage of plant succession on sandy soil

The relationships between seedling emergence and litter cover were studied in the earliest successional stage of a plant community. During a period of 3 years, changes in vegetation cover and species composition were assessed on three permanent quadrats with virgin sandy soil substrates A successional pattern from summer annual grasses to winter annual herbacoeus plants to biennial and perennial plants was confirmed, and invasion and replacement of the component species were conspicuous. The spatial distributions of seedlings and litter were heterogeneous in winter, and some patches consisting of both seedlings and litter were observed. There was a strong positive correlation between distribution patterns of seedlings and litter; seedling density increased with increasing litter cover. The results suggest that the litter cover of the previous stage plays an imporrant role in promoting successional changes in the early stage of plant succession.