Fifty years vegetation development of a xerothermic calcareous grassland in Central Europe after heavy disturbance

Fifty years ago on the south-facing slope of a limestone hill 4 km NE of Göttingen (Germany) a small plot of 4 m² size was chosen in a xerothermic calcareous grassland (Mesobromion alliance) for investigation. The formerly grazed slope was abandoned a few years prior to the study, but in 1987 intermittent grazing started again. The mean annual temperature in Göttingen (1953–2003) was 8.8 °C on the average; the annual mean increased by 1.1 °C during the study period. Mean annual precipitation amounted to 642 mm (393–1093 mm). The mean monthly precipitation during summer (April–August) varied in the range between 29 and 101 mm. In the years 1953–2003 the presence of species was recorded annually in 400 subplots (100 cm² each). In part of this time also the amount of bare soil within the plot was recorded, and standing crop and the nitrogen contents in leaves of Bromus erectus were measured in the immediate surrounding of the plot.In fall 1953 the vegetation was removed completely in half of the plot, in the other half the two dominant grasses Bromus erectus and Brachypodium pinnatum were left. The differences between the two halves vanished already after 2 years (starting phase 1953–1955). Thus, only results concerning the plot as a whole are reported here. During the investigation three periods could be distinguished. After the starting phase in period I (1955–1986) total presence, presence of the most important species, species richness, number of endangered species, diversity and evenness increased. In this way the grassland regenerated after the heavy disturbance, and a dense, species-rich vegetation formed. Only a few species showed maximum values already within the first 10 years and faded out later.Period II (1987–1997) spanned 10 years of grazing. During this time total presence, presence of many important species, species richness, number of endangered species, diversity and evenness decreased, whereas areas with bare soil increased. Only two species showed maximum values during this phase, and just one new species invaded (Lolium perenne). During this period a more open type of grassland was formed. Period III (1998–2003) spanned 5 years without grazing and only one grazing year (2002). Here, a partial regeneration of a more closed grassland took place with traits similar to period I.Several phenomena occurred continuously through all three periods. Bromus erectus always was the dominating species. Nevertheless, proportions of graminoids (in % total presence) were lowered, and the proportion of herbs increased. The ratio Bromus erectus/Brachypodium pinnatum was determined mainly by Brachypodium. Seedlings of woody plants were always present, but in most cases survived only one summer. Shrubs did not settle inside the plot, although small bushes developed in the immediate surrounding. In summary the 50-year process can be described as a slow recovery of a calcareous grassland after very heavy disturbance, modified by intermittent grazing, without formation of shrub vegetation.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

Distribution and conservation status of coastal sage scrub in southwestern California

Abstract. A landscape-based characterization of vegetation has been developed for southwestern California using satellite imagery, air photos, existing vegetation maps, and field data. Distribution maps of nine dominant coastal scrub species and 13 species assemblages that were identified by divisive information analysis have been analyzed to quantify spatial patterns of species co-occurrence. Three general distribution patterns are identified that suggest the Diegan, Ventaran and Riversidian Associations identified by other workers. Vegetation data have also been related to land ownership and management to assess the conservation status of upland plant communities. A large proportion of the mapped distribution of species and vegetation types is on private land, and several taxa show less than 4 % of mapped distribution in nature reserves. The analysis highlights the need to extend current conservation planning efforts into the northern part of the region to encompass areas where Salvia leucophylla is a frequent community dominant.     

Differential responses to nitrogen fertilization in native shrubs and exotic annuals common to mediterranean coastal sage scrub of California

This study examined the growth responses of exotic annuals and native shrubs to elevated N levels to test the hypothesis that increased N availability favors nitrophilous annuals over the slower-growing shrubs. The vegetation structure of the coastal sage scrub ecosystems in southern California is shifting from shrubland to annual grasslands. Over the last 30 years large tracts of wildlands, particularly those adjacent to urban centers, have lost significant native shrub cover, which has been replaced by exotic annuals native to the Mediterranean Basin. During this same time, air pollution has led to increased terrestrial eutrophication by atmospheric deposition. Changes in vegetation are often the result of changes in resource availability. The results of our experiments showed the three native shrubs tested to be more nitrophilous than the three annuals tested, which contrasts with most models of perennial species' adaptation to stressful environments. Under greenhouse conditions the annual grasses exhibited yield depression at the highest N treatments of 80 g g^–1 in soil. The three shrub species evaluated continued to increase shoot biomass at 80 g g^–1 N in soil. The grasses also exhibited increased tissue N concentrations with increased soil N in contrast with the shrubs where there was little difference in tissue N concentrations with increasing availability. Although the differential yield responses to elevated N do not explain the success of the annual vegetation in replacing shrubs, the inability of the shrubs to regulate growth under elevated N levels may explain the poor survival of mature individuals.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Influence of litter removal and mineral soil disturbance on the spread of an invasive grass in a Central Hardwood forest

Soil and litter disturbances within Central Hardwood forests may facilitate exotic plant species invasion of interior forest areas. Microstegium vimineum is an annual exotic grass that has become common throughout the Southeastern United States. Three replicates of three different mineral soil and litter disturbance treatments, plus a control with no disturbance, were established on the leading edge of M. vimineum patches prior to seed fall. All patches were located in areas with similar forest canopy structure and slope in three Central Hardwood forest stands prior to seed fall. At the beginning of the following growing season, each individual M. vimineum seedling was mapped within the treatment plots. The mean number of M. vimineum individuals that established within each treatment did not differ significantly from the control. The distance at which 90% of the individuals had spread, and the overall mean distance spread were significantly farther for the litter removal treatment than the control. The farthest individual seedling from the boundary of existing patches in both the litter removal and the mineral soil disturbance and litter removal treatments were significantly farther than the control. The individuals that spread the farthest are of most concern due to the large number of viable seed that a single M. vimineum plant can produce. These results suggest that disturbance of the litter layer may increase the spread rate of M. vimineum and facilitate its invasion of new habitats, and that leaving litter layers intact may slow the spread of M. vimineum.     

Drought in Southern California coastal sage scrub reduces herbaceous biomass of exotic species more than native species,but exotic growth recovers quickly when drought ends

Plant Ecology - Semi-arid regions with Mediterranean-type climates harbor exceptional biodiversity, but are increasingly threatened by invading exotic annual species and climatic changes, including...     

Carbon and nitrogen turnover in adjacent grassland and cropland ecosystems

The effects of cultivation and soil texture on net and gross N mineralization, CO₂ evolution and C and N turnover were investigated using paired grassland and cropped sites on soils of three textures. Gross N mineralization and immobilization were measured using¹⁵N-isotope dilution. Grassland soils had high CO₂ evolution and gross N mineralization rates, and low net N mineralization rates. Cropland soils had low CO₂ evolution rates but had high net and gross N mineralization rates. Grassland soils thus had high immobilization rates and cropland soils had low immobilization rates. Cultivation increased N turnover but reduced C turnover. The data suggest that the microflora in grassland soils are N limited, while those of cropland soils are limited by C availability. Increasing clay content reduced N turnover. C turnover was less clearly related to texture. Differences in the immobilization potential of substrates help explain why agricultural soils have higher N losses than do grassland soils.     

Response of nematode-trapping fungi to organic substrates in a coastal grassland soil

To understand why Arthrobotrys oligospora and other nematode-trapping fungi are common and sometimes abundant in the coastal grassland soils of the Bodega Marine Reserve (BMR, Sonoma County, CA), we examined how resident trapping fungi responded to the addition of eight organic substrates (lupine leaves, grass leaves, dead isopods, dead moth larvae, isopod faeces, deer faeces, shrimp shells, and powdered chitin). We were especially interested in the effects of dead isopods because isopods are abundant at BMR and because previous studies had documented strong responses of A. oligospora to other arthropods (dead moth larvae). Soil from BMR was packed into vials (40 g dry mass equivalent per vial with water potential at −230 kPa and bulk density at 0.9 g cm⁻³), and one substrate or no substrate was added to the soil surface. After 30 d at 20 °C, trapping fungi were quantified by dilution plating and most probable number procedures. The response of A. oligospora was inversely related to substrate carbon:nitrogen (C:N) ratio: substrates with low C:N ratios (dead isopods, lupine leaves, dead moth larvae) usually caused large increases in A. oligospora whereas those with higher C:N ratios (isopod faeces, deer faeces, grass leaves) did not. An exception was chitin powder, which had a low C:N ratio, but which did not cause A. oligospora to proliferate. Responses of A. oligospora were directly related to the quantity of nitrogen added with each substrate, and those substrates that caused large increases in resident nematodes usually caused large increases in A. oligospora. Other trapping fungi did not respond as strongly as A. oligospora.     

Plant community changes after the reduction of an invasive rangeland weed, diffuse knapweed, Centaurea diffusa

The expected outcome of weed control in natural systems is that the decline of a dominant weed will result in an increase in diversity of the plant community but this has seldom been tested. Here we evaluate the response of the plant community following the decline of diffuse knapweed (Centaurea diffusa) in six different pastures at White Lake, BC, Canada over five years. This period followed the establishment, spread and high levels of attack by the introduced European weevil, Larinus minutus, as part of a biological control program. Knapweed declined immediately before and during the study period, but, contrary to expectations, the species richness and diversity of the rangeland plant community did not increase. The absolute cover of native and introduced forbs and grasses increased following knapweed decline, but only the introduced grasses showed a consistent increase in cover relative to the other life-forms. However, unlike in other studies, the native plants dominated the study site. We conclude that the changes in plant communities following successful biological control are variable among programs and that the impact of replacement species must be evaluated in assessing the success of ecological restoration programs that use biological control to manage an undesirable weed.     

The prevalence of non-indigenous species in southern California embayments and their effects on benthic macroinvertebrate communities

The prevalence of non-indigenous species (NIS) in southern California embayments was assessed from 123 Van Veen grab samples collected in nine bays and harbors during the summer of 1998. NIS occurred in all but two samples. They accounted for only 4.3% of the 633 taxa but contributed 27.5% of the abundance. There was no significant difference in the proportion of NIS abundance among ports harboring large vessels, small boat marinas, and areas where boats were not moored. Three species accounted for 92% of the NIS abundance: a spionid polychaete worm Pseudopolydora paucibranchiata, a mytilid bivalve Musculista senhousia, and a semelid bivalve Theora ubrica. The NIS did not appear to have a negative impact at the overall community level since NIS abundance was positively correlated with the abundance and richness of other species. This may be due to biogenic structures built by P. paucibranchiata and M. senhousia that enhance the abundances of other macrofauna.     

The persistence of calcareous grassland species in the soil seed bank under developing and established scrub

The relationships between the composition of the soil seed bank, the field layer vegetation, and the scrub canopy were investigated along a 69 m transect, grading from incipient woodland, through scrub, into intensively rabbit-grazed calcareous grassland. The results are used to assess the persistence of species associated with open calcareous grassland in the seed bank under developing scrub. Scrub age, composition and density, changed along the transect from the woodland to open grassland. A total of 35 forb and grass species were found in the field layer. The pattern evident in the scrub layer was also reflected in the herbaceous vegetation. The field layer in the most closed portion of the transect, where the scrub was oldest, was dominated by shade-tolerant species normally associated with woodland habitats. The abundance of these species decreased along the transect as the scrub age declined, and the field layer became increasingly dominated by species typical of open grassland. A total of 47 species germinated from the seed bank. Few species were recorded in the seed bank along the entire length of the transect. Overall, the seed bank was dominated by Hypericum perforatum and Centaurium erythraea, which accounted for 38.2% and 28.6% of emerging seedlings respectively. As with a number of similar studies, the composition of the seed bank had a low correspondence with the composition of the field layer vegetation. The results also emphasise that the composition of the seed bank can be viewed as an ecological palimpsest, with germinable seed of species from each stage of the old-field succession occurring in the soil. The seed bank is an important component in the re-vegetation of an area after disturbance such as scrub removal. This study supports the findings of previous research in showing that relatively few characteristic calcareous grassland species form persistent seed banks. The soil seed bank would therefore appear to be of limited value in the restoration of such grassland following scrub removal.     

Invasion of two exotic terrestrial flatworms to subantarctic Macquarie Island

Invasive species are a serious threat to biodiversity worldwide. The relatively simple ecological systems of the subantarctic have the potential to be significantly damaged by predatory species that invade. Two species of exotic, predatory, terrestrial flatworms were first collected in 1997 from two localities only 2 km apart, in the southeast of subantarctic Macquarie Island. The species were later identified as Kontikia andersoni and Arthurdendyus vegrandis. We report here the results of fieldwork in 2004 that established that both species now occupy about a seventh of the southeast of the island which has a total area of only 170 km² and that there seem to be no barriers to further expansion. The island was first discovered in 1810 and so it is likely the species were introduced by means of human intervention within the last 200 years. We provide evidence to show that both species originated in New Zealand and have probably been on the island for ∼100 years giving an average rate of spread of about 10 m per year. Other species of Arthurdendyus have been introduced from New Zealand to the United Kingdom where they prey on earthworms. The quarantine significance of A. vegrandis for Australia is discussed and recommendations made to reduce the probability of it entering Tasmania where it has the potential to become an agricultural pest.     

盐城滨海滩涂湿地典型植物群落土壤微生物组成与结构特征

土壤微生物是生态系统维持正常结构与功能的重要组成部分,为探究盐城滩涂典型湿地土壤微生物群落结构特征,以江苏盐城滩涂互花米草、藨草、盐地碱蓬、芦苇及淤泥质光滩5种典型群落为对象,采用16S rRNA高通量测序技术分析0—10 cm(表层)、10—30 cm(中层)、30—60 cm(深层)土壤微生物多样性及群落结构。结果表明：(1)几种植物群落间,土壤微生物群落结构差异较大,主要体现在细菌群落结构的差异性,古菌群落结构差异相对较小。光滩与植物群落间,在土壤细菌种类及相对丰度上差异相对较大,互花米草群落与本土植物群落间,在微生物群落的细菌种类组成上存在较大差异;藨草群落土壤表层微生物群落结构与互花米草群落相似,深层与盐地碱蓬、芦苇群落相似。(2)同一群落不同层次土壤微生物群落结构相似,差异小于不同群落间土壤微生物群落的结构差异性;不同群落对应层次间,表深层土壤中五种群落土壤微生物多样性存在显著差异,中层土壤中五种群落微生物多样性差异不显著。总体上,植物群落类型对土壤微生物群落结构的影响大于土壤深度;与本土植物群落相比,互花米草群落土壤主要优势门微生物种类差异较小,但部分优势门微生物相对丰度...     

The influence of dominant shrubs,fire, and time since fire on soil seed banks in mixed chaparral

The composition and density of soil seed banks beneath co-occurring Adenostoma fasciculatum and Ceanothus greggii shrubs from three chaparral stands last burned 9, 35 and 85 years before 1986 were investigated. The overall density of seeds in the soil, as estimated by germinations under greenhouse conditions, increased with time since fire (ca. 8000 to 25000/m²). However, this increase was due entirely to the accumulation of A. fasciculatum seed in the soil (ca. 2000 to 21000/m²). In contrast, the density of C. greggii seed was different in each of the three stands, but was not correlated with time since fire: maximum densities were recorded from the 35 year old stand (ca. 2000/m²).A total of 31 taxa germinated and 17 occurred in sufficient numbers to be analyzed statistically. Germinable seed densities of three herb species were not influenced by soil source (beneath A. fasciculatum or C. greggii), time since fire, or the direct effects of a controlled fire treatment. Germinable seed densities of a further nine species were significantly influenced by the elapsed time since stands last burned. The densities of four decreased and five increased. Four of the species that increased in seed density over the three stands were annuals, suggesting that the chaparral sub-canopy habitat is not as unfavorable for annuals as is often assumed. The fire treatment decreased germinable seed densities of four annual species by 40–70%, but increased the germinable seed densities of the shrubs A. fasciculatum and C. greggii, and the annual Phacelia brachyloba. Our results indicate that seeds of A. fasciculatum will increase in the soil bank for at least 85 years after fire in chaparral where it is dominant. In contrast, seed reserves of C. greggii appear to be influenced primarily by site-specific patterns of seed production and by the intensity of post-dispersal seed predation.     

Successional dynamics of a semiarid grassland: effects of soil texture and disturbance size

We evaluated effects of soil texture and disturbance size on the successional dynamics of a semiarid grassland dominated by the perennial bunchgrass, Bouteloua gracilis (H.B.K.) Lag. ex Griffiths. A spatially-explicit gap dynamics simulation model was used to evaluate recovery patterns. The model simulates establishment, growth, and mortality of individual plants on an array of small plots through time at an annual time step. Each simulated disturbance consisted of a grid of plots of the same soil texture interconnected by processes associated with dispersal of B. gracilis seeds. Soil texture was incorporated into the model as effects on seed germination, seedling establishment, and subsequent growth of B. gracilis. Five soil texture classes and five disturbance sizes were simulated.Soil texture was more important to recovery of B. gracilis than either size of a disturbance or location of plots within a disturbance. Constraints on recruitment of seedlings had a greater effect on recovery than constraints associated with plant growth. Fastest recovery occurred on soils with the largest silt content, the variable that affects seedling establishment. Disturbances with slowest recovery were on soils with low silt contents, and either high or low water-holding capacity, the variable that affects plant growth. Biomass and recovery decreased as disturbance size increased, and as distance from a disturbed plot to the edge of the disturbance increased. In most cases, important interactions between soil texture and disturbance size on recovery were not found.     

Forest edges and fire ants alter the seed shadow of an ant-dispersed plant

Exotic species invade fragmented, edge-rich habitats readily, yet the distinct impacts of habitat edges and invaders on native biota are rarely distinguished. Both appear detrimental to ant-dispersed plants such as bloodroot, Sanguinaria canadensis. Working in northeastern Georgia (USA), an area characterized by a rich ant-dispersed flora, fragmented forests, and invasions by the red imported fire ant, Solenopsis invicta , I monitored the interactions between ants and S. canadensis seeds in uninvaded forest interiors, uninvaded forest edges, invaded forest interiors, and invaded forest edges. I observed 95% of the seed dispersal events that occurred within the 60-min observation intervals. Seed collection rates were similar among all four (habitat × invasion) groups. The presence of invasive ants had a strong effect on seed dispersal distance: S. invicta collected most seeds in invaded sites, but was a poorer disperser than four of five native ant taxa. Habitat type (interior versus edge) had no effect on seed dispersal distance, but it had a strong effect on seed dispersal direction. Dispersal towards the edge was disproportionately rare in uninvaded forest edges, and ants in those habitats moved the average dispersed seed approximately 70 cm away from that edge. Dispersal direction was also skewed away from the edge in uninvaded forest interiors and invaded forest edges, albeit non-significantly. This biased dispersal may help explain the rarity of myrmecochorous plants in younger forests and edges, and their poor ability to disperse between fragments. This is the first demonstration that forest edges and S. invicta invasion influence seed dispersal destination and distance, respectively. These forces act independently.     

Shell strength and primate seed predation of nontoxic species in eastern and southern Africa

The large African primates that eat fruit destroy the seeds of a number of fruiting species. This paper addresses several questions about seed-eating: What is the nature of the dietary niche provided by the nonpoisonous seeds of eastern and southern Africa? How well are these seeds mechanically protected? What other means of reducing seed predation are employed by the plants? and is the niche ecologically stable? Measurements of seed shell strength on 37 species from 17 families reveal a range of values, from <100-kg (numerous species) to over 2000-kg (palm nuts) breaking load. Primates crack open with their teeth seed shells from species exhibiting test strengths less than 600 kg. Variation in shell strength appears to increase dramatically for average species strengths above 100 kg. Plant species are not characterized by specific shell strengths but instead, display envelopes of shell strength overlapping broadly with other species. Taking this into account, adult male baboons (Papio spp.) appear to be dentally capable of preying upon most of the seed species of eastern and southern Africa. The possibility for predation of nonpoisonous seeds exists primarily because the plants periodically produce large crops in synchrony and the hard-shelled seeds are effectively dispersed, sometimes explosively but more often by means of edible fruits. The concomitant primate seed predation is a facultative specialization, of little apparent threat to the community of plants that support it.     

The effect of soil nitrogen on competition between native and exotic perennial grasses from northern coastal California

The invasion of European perennial grasses represents a new threat to the native coastal prairie of northern California. Many coastal prairie sites also experience anthropogenic nitrogen (N) deposition or increased N availability as a result of invasion by N-fixing shrubs. We tested the hypothesis that greater seedling competitive ability and greater responsiveness to high N availability of exotic perennial grasses facilitates their invasion in coastal prairie. We evaluated pairwise competitive responses and effects, and the occurrence of asymmetrical competition, among three common native perennial grasses (Agrostis oregonensis, Festuca rubra, and Nassella pulchra) and three exotic perennial grasses (Holcus lanatus, Phalaris aquatica, and Festuca arundinacea), at two levels of soil N. We also compared the root and shoot biomass and response to fertilization of singly-grown plants, so we could evaluate how performance in competition related to innate plant traits. Competitive effects and responses were negatively correlated and in general varied continuously across native and exotic species. Two exceptions were the exotic species Holcus, which had large effects on neighbors and small responses to them, and competed asymmetrically with all other species in the experiment, and the native grass Nassella, which had strong responses to but little effect on neighbors, and was out-competed by all but one other species in the experiment. High allocation to roots and high early relative growth rate appear to explain Holcus’s competitive dominance, but its shoot biomass when grown alone was not significantly greater than those of the species it out-competed. Competitive dynamics were unaffected by fertilization. Therefore, we conclude that seedling competitive ability alone does not explain the increasing dominance of exotic perennial grasses in California coastal prairie. Furthermore, since native and exotic species responded individualistically, grouping species as ‘natives’ and ‘exotics’ obscured underlying variation within the two categories. Finally, elevated soil N does not appear to influence competition among the native and exotic perennial grasses studied, so reducing soil N pools may not be a critical step for the restoration of California coastal prairie.