Responses to fertility and disturbance in a low-diversity grassland

We examined variation in species composition in a low-diversity, anthropogenic grassland in response to 11 years of nitrogen (N) manipulation and disturbance. The species-poor grassland (2–3 species/0.5 m²) represents a wide spread vegetation type (>10 million ha in North America) dominated by the introduced perennial grasses Bromus inermis and Agropyron cristatum. Four levels of N and three of soil disturbance were applied in all combinations to plots (5 × 15 m, N = 120) in a completely randomized design each year. Seeds or transplants of 47 species were added to ensure that dispersal was not a barrier to changes in species composition. After 11 years of treatment, all but the most disturbed plots continued to be dominated by B. inermis. The cover of the second-most abundant species, A. cristatum, decreased with disturbance but did not vary significantly with N. Despite the lack of changes in the identity of the dominant species, our environmental manipulations strongly influenced ecosystem characteristics. Added N increased soil available N, and decreased the cover of bare ground and light availability. Soil disturbance decreased aboveground biomass, and increased the cover of bare ground and light availability. Sawdust application, designed to decrease N availability, significantly reduced community biomass, and increased light availability and the cover of bare ground, but did not alter nutrient availability or species composition. The results highlight the difficulty of restoring diversity in species-poor, anthropogenic communities dominated by introduced species, and thus the importance of conserving remnants of diverse natural grasslands.     

Light reduction predicts widespread patterns of dominance between asters and goldenrods

Here we investigate the long-cited pattern that throughout the eastern United States, Solidago species (goldenrods), and in particular S. canadensis displace Aster species and dominate old-field communities. Theory predicts that such a ubiquitous pattern of repeated dominance should be linked to competitive ability for a limiting resource. However, no one has investigated this possibility in old-fields, representing a potentially significant gap in our understanding of a common human-altered environment. We tested the hypothesis that S. canadensis is the superior competitor for light compared to other common co-occurring goldenrod species, and that the goldenrods in general are the superior competitors for light compared to coexisting aster species, which are typically less abundant. We tested this hypothesis by comparing the light attenuation abilities of four goldenrod species, S. canadensis, S. rugosa, S. gigantea, and Euthamia graminifolia, and three aster species, Aster novae-angliae, A. pilosus, and A. prenanthoides. Consistent with our hypothesis, S. canadensis had a greater ability to attenuate light than any of the other goldenrods at higher densities, and the goldenrods overall had a greater ability to attenuate light than the asters. By conducting a census in our study area, we verified that S. canadensis is locally the most abundant goldenrod and that goldenrods are more locally abundant than asters. Furthermore, by conducting a literature survey we found evidence that S. canadensis replaces A. pilosus through time. Thus we found a close correspondence between relative abundance in the field and light attenuation ability in field experiments. These results are consistent with theory predicting that competition for limiting resources, in this case light, explains patterns of dominance and relative abundance in old-field plant communities.     

Recovery of subalpine dwarf shrub heath after neighbour removal and fertilization

We tested if subalpine heath vegetation in northern Italy recovered after experimental perturbation of soil nutrient availability (fertilization) and species composition (removal of co-dominant dwarf shrubs). Species cover was assessed non-destructively before the start of the treatments (1995), at the end of the treatments (1999) and 4 years after the treatments ended (2003). Shrub removal had rather modest effects on heath vegetation, except for mosses which decreased significantly in removal plots. Fertilization decreased the cover of shrubs, mosses, and some graminoids but increased the cover of Festuca rubra. Fertilization converted heath to grassland, but the response of graminoid species was individualistic. Fertilization decreased vascular species richness and evenness, probably through negative effects of shading and litter accumulation on plant growth or recruitment. The vegetation had not recovered completely 4 years after the perturbations had stopped. This suggests that, in contrast to rapid responses to species removal and fertilization, recovery from these perturbations was rather slow, presumably because recovery was affected by long-term biotic interactions and species controls on ecosystem properties.     

Short-term study of effects of fertilisation and cutting treatments on the vegetation dynamics of mountain heathlands in Spain

The influence of management and nutrient availability on the vegetation dynamics of heathlands characterised by Calluna vulgaris and Erica tetralix were studied in three mountain sites in Northern Spain. A total of 90 plots (1 m² each) received different combinations of cutting and twice the estimated background atmospheric deposition of nitrogen (56 kg ha⁻¹ yr⁻¹). One of the two dominant ericaceous species was selectively cut by hand at ground level and their regeneration compared in the presence or absence of the other. The results after 2 years showed significant effects of the fertiliser on the vegetation cover, mainly by favouring perennial herbaceous species. There were less noteworthy effects on the number of flowers and on the annual growth of the ericaceous species. It is concluded that, in the short term, increased nutrients alone, at twice the estimated current atmospheric deposition for the area, will not alter significantly the composition of the mountain heathlands. However, once the stands reach the mature phase, the capacity of the community to regenerate after a severe disturbance diminishes. A drastic impact, such as cutting may not result in re-growth of the same shrub species but in replacement by herbaceous species, which will also benefit from the increased nutrients.     

外来树种日本落叶松和日本花柏对鄂西北森林群落的影响

【目的】研究大面积栽植外来树种日本落叶松和日本花柏对鄂西北森林群落物种多样性的影响,为评价外来树种的入侵风险提供依据。【方法】采用样地调查法对比研究外来树种日本落叶松和日本花柏人工林与当地森林群落特征差异。【结果】日本落叶松人工林样地林下植物种类丰富,样地平均物种数与当地天然林样地物种数相比无显著差异;林下灌木层和草本层多样性指数较高,草本层多样性指数与当地天然林相比无显著差异;林下未发现日本落叶松的更新苗。日本花柏人工林样地平均物种数显著少于当地其他人工林样地;乔木层、灌木层和草本层的4个多样性指数值均小于当地其他人工林样地;在3个样地中出现日本花柏更新苗,更新苗数量达479株·hm~(-2)。【结论】引种栽培日本落叶松不会导致当地森林群落的物种多样性降低,不会引起生物入侵风险;但引种栽培日本花柏会明显降低当地森林群落的物种多样性,对当地森林生态系统的稳定构成一定威胁。     

Nested spatial patterns in seed bank and vegetation of Mediterranean old-fields

A nested sampling design was used to describe the spatial patterns for the species richness and composition in the seed bank and vegetation of three Mediterranean old-fields (1, 7, and 15 yr after the last ploughing). Three scales were examined hierarchically: sampling units within plots of 0.25 m² for the vegetation and of 0.05 m² for the seed bank, 100 m² plots within fields, and fields of 1000 m². In spite of the strong spatial variation among sampling units, species richness and composition of both seed bank and vegetation showed hierarchically structured patterns of heterogeneity, while each old-field was a homogeneous entity. These spatial patterns tended to be partially masked when the data were aggregated at the scale of the plot. Such results stress the use of a nested sampling design for studying variation in species richness and taxonomic composition in both vegetation and seed bank. This design, in combination with CCA, also showed that the vegetation showed a coarser grain than the seed bank, probably in relation to seed clumping.     

外来入侵植物与人工林下层植被结构的关联性

为有效防控外来入侵植物,在广西陆川县人工林区进行植被嵌套样方调查,分析群落物种组成与植被结构和环境因子之间的相关性。Mantel检验表明样方物种组成与样方植被结构的相关性(r=0.208, P=0.002)高于其与环境因子相关性(r=0.084, P=0.051)。MRPP分析表明,有入侵植物阔叶丰花草(Spermacoce alata)、假臭草(Praxelis clematidea)和胜红蓟(Ageratum conyzoides)样方的植被结构与无这3种植物样方的有极显著不同(P 0.01),而有无薇甘菊(Mikainamicrantha)样方的植被结构相似。t检验表明林冠盖度(3.0 m)与这4种植物入侵均无显著相关性(P0.073)。通过增加垂直高度1.5 m以下的植被密度,可以减少阔叶丰花草、假臭草和胜红蓟的入侵,但对抑制薇甘菊入侵无效。因此,植被结构作为过滤器,可能主要作用于植物的种子产生时间、萌发周期和生长习性;以种子方式繁殖的入侵植物能否定植相对更依赖干扰、暴露的裸地;兼具无性和有性繁殖且能攀援和匍匐生长使得薇甘菊能够在不同植被结构的群落中相对随机地定植。     

Similar biotic factors affect early establishment and abundance of an invasive plant species across spatial scales

Research in community invasibiliy has focused on biotic and abiotic factors that influence the establishment of invasive species and whether such factors vary with spatial scale. Here, we investigate the role of both biotic and abiotic factors associated with the initial establishment of Lespedeza cuneata (L. cuneata) and its abundance at three spatial scales: neighborhoods (9-m² plots), communities (50-m² transect) and old fields (5,000–70,000 m²). We asked: (1) Do resource availability and community structure affect the establishment of L. cuneata?, and (2) Are resource availability and community structure associated with patterns of L. cuneata abundance from neighborhood scales to old-field scales? To investigate the first question, we manipulated soil nitrogen (N) availability at three levels in an existing old-field community and tracked emergence and persistence of L. cuneata seedlings, as well as total plant biomass of the community, availability of light, and soil moisture content. To address the second question, we performed surveys in which we estimated L. cuneata foliar cover at community scales (50-m² belt transects) and old-field scales (total area of 28 ha), and assessed the same biotic and abiotic variables as in the field experiment. The experiment revealed that establishment and persistence by L. cuneata seedlings were 15× and 5× lower in N-added plots than in N-reduced plots. Total plant community biomass was 30% greater in N-added plots than in N-reduced plots. Conversely, light and soil moisture were 60 and 20% lower in N-added plots than in N-reduced plots. Surveys of old fields indicated that community biomass was positively associated with L. cuneata cover at old-field scales likely resulting from greater soil N input from nitrogen fixation in fields with greater L. cuneata cover. In sum, these results indicate that biotic factors associated with establishment of a Rank 1 invasive plant species at the community scale are also related to its distribution at the old-field scale, but the direction of such associations changed across scales.     

Vegetation structure and diversity of an ancient temperate deciduous forest in SW Denmark

Bolderslev Skov (113 ha) is the largest contiguous ancient forest remnant in Denmark. The forest has been preserved since 1998 as a strict non-intervention forest reserve. We studied vegetation structure, floristic gradients, and diversity of the forest in 50 plots of 100 m² placed according to a restricted random sampling design. Dominant tree species were Fagus sylvatica, Fraxinus excelsior, Tilia cordata and Quercus robur, which in most parts of the forest form a mixed canopy. Most stands appeared to be of moderate age, 55-80 years old, and large old trees were rare. pH in association with light and thickness of the litter layer were the most important factors in explaining floristic gradients in the forest. Soil moisture (dry to mesic) was not strongly correlated with DCA axes for neither tree nor field layer, but had a significant effect on the distribution of a number of herb species. Forest structure was not important in explaining species distributions. Field layer species richness showed a positive relationship with the pH-gradient. At the scale of 1m² plots we also found a highly significant negative relationship between field layer species richness and the plot-wise Ellenberg indicator value for nitrogen availability. Structure of the tree layer had little effect on field layer species richness. The mixed composition, long continuity, and presence of a high proportion of the regional species pool assigns a high conservation value to Bolderslev Skov and makes it an important site for future studies of the dynamic properties, niche preferences, and inter-specific competition of temperate deciduous forest species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Responses of native and non-native Mojave Desert winter annuals to soil disturbance and water additions

Habitat modification (i.e., disturbance) and resource availability have been identified as possible mechanisms that may influence the invasibility of plant communities. In the Mojave Desert, habitat disturbance has increased dramatically over the last 50 years due to increased human activities. Additionally, water availability is considered to be a main limiting resource for plant production. To elucidate the effects of soil disturbance and water availability on plant invasions, we created experimental patches where we varied the levels of soil disturbance and water availability in a fully crossed factorial experiment at five replicated field sites, and documented responses of native and non-native winter annuals. The treatments did not significantly affect the density (seedlings m⁻²) of the non-native forb, Brassica tournefortii. However, the relationship between silique production and plant height differed among treatments, with greater silique production in disturbed plots. In contrast to Brassica, density of the non-native Schismus spp. increased in soil disturbed and watered plots, and was greatest in disturbed plots during 2009 (the second year of the study). Species composition of the native annual community was not affected by treatments in 2008 but was influenced by treatments in 2009. The native forb Eriophyllum sp. was most dense on water-addition plots, while density of Chaenactis freemontii was highest in disturbed plots. Results illustrate that habitat invasibility in arid systems can be influenced by dynamics in disturbance regimes and water availability, and suggest that invasiveness can differ between non-native annual species and among native annuals in habitats undergoing changing disturbance and precipitation regimes. Understanding the mechanistic relationships between water availability and non-native plant responses will be important for understanding the effects of shifting precipitation and vegetation patterns under predicted climate change in arid ecosystems.     

Assessing the residual effects of Carpobrotus edulis invasion, implications for restoration

We examined whether the residual effects on soil caused by the invasion of Carpobrotus edulis, common iceplant, would inhibit the reestablishment of a native plant species. Carpobrotus edulis interacts both directly by suppressing the growth and establishment of other plants and indirectly by altering soil chemistry. We tested whether the residual effects of C. edulis resulted in lowered germination, survival, growth, and reproduction of Gilia millefoliata, a rare dune annual. We compared G. millefoliata planted in plots previously occupied by C. edulis to G. millefoliata planted in plots that previously had native vegetation. Each plot received three treatments: seed, transplant, and unplanted, and were censused every three weeks until senescence. Carpobrotus edulis had strong negative effects on the germination, survival, growth, and reproduction of G. millefoliata. C. edulis lowers soil pH and increases organic content due to the recalcitrance of tissue to decomposition, which may have evolved as a mechanism to facilitate recolonization and invasion.     

Growth and pigment composition in the red alga <Emphasis Type="Italic">Halymenia floresii</Emphasis> cultured under different light qualities

Halymenia floresii is an edible species consumed in some Asian markets. In the Yucatan peninsula coast of Mexico, H. floresii dominates rocky substrata between 3 and 40 m where it grows up to 50 cm high. After analyzing the seasonal pattern of pigment content on H. floresii, we evaluate if and how the spectral composition of light affects growth and pigment dynamics under laboratory cultivation. Unialgal cultures were exposed to white, blue, red and green light in a 3-week experiment. Green light resulted in the highest algal growth rates. Synthesis of chlorophyll a, α-carotene and lutein, but not of β-carotene, was induced by white or green light. Phycocyanin synthesis was stimulated by blue light and phycoerythrin synthesis by blue or red light. Light quality treatments may be used to manipulate pigment composition in Halymenia floresii cultures.     

Introduced plants on Kilimanjaro: tourism and its impact

Kilimanjaro, a world heritage site and an icon of global change, not only suffers from climatic alterations but also is undergoing a drastic socio-economic upheaval. A strong increase of tourism enhances the risk of introducing alien plant species in particular in the upper zones of Kilimanjaro. One such species is Poa annua L., a cosmopolitan weed of European origin on roadsides and pastures. The aim of this study is to document its distribution, the speed of its propagation and risks for the indigenous vegetation of Kilimanjaro, and to compare the findings with other introduced species on this mountain. Based on a complete survey of the vegetation of Kilimanjaro with about 1,500 vegetation plots, plant communities invaded by Poa annua are determined. As with most of the other neophytes on Kilimanjaro, Poa annua invades only anthropogenic vegetation but not undisturbed natural vegetation. Similar to the situation in middle Europe, this neophyte is on Kilimanjaro a constituent of the vegetation of trampled ground, occurring between about 1,600 and 4,000 m asl along climbing routes or their vicinity. On a newly opened climbing route a rapid invasion (5.6 km in 3 months) was observed, which makes it likely that Poa annua spread on Kilimanjaro during the last 30 years in parallel to the increase of the climbing tourism. Although Poa annua is still in the stage of propagation, an invasion of natural vegetation types seems to be unlikely.     

Impact of ploughing on soil seed bank dynamics in temporary pools

We examined the impact of ploughing on soil seed banks of plant communities living in temporary marshes located in agricultural fields. The quantity, quality and vertical distribution of seeds were quantified under ploughed or unploughed treatment at community level. We also focussed on a typical semi-aquatic species, Damasonium alisma, to investigate the impact of ploughing at population level. We used two complementary techniques to study seed banks: hand sorting and seedling emergence. We found that species richness of seeds, seed abundance and germination ability were strongly affected by ploughing at community level. Concerning D. alisma, most of the seeds (56%) were stored in the two deepest soil layers among the four considered in ploughed pools. Moreover, the germination rate was higher for buried seeds (84%) than for seeds collected at the surface (33.6%). These patterns were almost inverted in unploughed pools. Our results agree with the temporal storage effect generally suggested to describe the seed bank property of plant communities. But in addition, we showed that ploughing induces a spatial storage effect in accumulating species and individuals in the seed banks that favourably influence community dynamics. We conclude that, in contrast to what is usually thought, ploughing disturbance can be of benefit for such ephemeral wetland vegetation.     

Facilitation and competition influence succession in a subtropical old field

The effects of neighboring vegetation and soil fertility on the establishment, survival and growth of tree species were studied in a subtropical old-field area in south Brazil. Seed damage, germination and seedling establishment of four tree species plus growth and survival of two transplanted tree species were monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control). Facilitation was the main process affecting plant performance. The presence of pioneer vegetation significantly improved germination, establishment, growth and survival of most study species. Around 90% of sown seeds were damaged and the removal of pioneer vegetation significantly increased seed damage for all species studied, decreased germination in three out of four species, and decreased establishment in one species. Moreover a significantly higher seedling growth rate of Inga virescens was found after the first year of the experiment in plots where vegetation was present. The presence of vegetation significantly increased seedling survival of I. virescens by protecting seedlings from leaf loss due to winter frosts. Competition was detected by the second year when a higher growth of transplanted seedlings of the species Araucaria angustifolia occurred in plots where vegetation was present and fertilizer were applied. A lower growth rate was detected in plots where vegetation was present but fertilizer was not applied. These results indicate a balance between competition for soil nutrients and protection by neighbor vegetation. Damage of seedlings by leaf cutter ants was an important barrier for plant survival. Damage occurred in 80% of the A. angustifolia seedlings and 58% of these damaged seedlings died. The presence of neighboring vegetation tended to protect seedlings from ant damage. Although competition occurred, facilitation seems to be the main process driving early successional changes in this subtropical old field. This was mainly due to the improvement of local microclimatic conditions and protection against herbivores by neighboring vegetation. Facilitation occurred during establishment and growth phases in a subtropical area that is considered a productive, low stress environment. Our results indicate that facilitation may be more frequent in productive environments than previously thought.     

Creosotebush vegetation after 50 years of lagomorph exclusion

In 1939, an experiment was established on the Jornada Experimental Range to evaluate the effects of shrub removal, rabbit exclusion, furrowing, and seeding in creosotebush [Larrea tridentata (DC.) Cov] vegetation. Sixteen plots (21.3×21.3 m) were laid out in four rows of four plots per row with a buffer zone of 7.6 m between plots and rows. A barbed wire fence excluded cattle and poultry wire fencing excluded lagomorphs. Treatments were factorially applied at two levels. Plant cover in the plots was sampled in 1938 (before treatment), 1947, 1956, 1960, 1967 and 1989 with randomly located, line-intercept transects. Data from all sampling dates were analyzed as a split plot in time and main effects for 1989 tested by analysis of variance for a 2×4 factorial experiment. There were significant (P<0.10) year x treatment interactions. Seeding and furrowing treatments were ineffective but lagomorph exclusion and shrub clearing treatments resulted in significant treatment differences for several species. In 1989, basal area of spike dropseed (Sporobolus contractus A.S. Hitchc.) was 30-fold greater on the lagomorph excluded than on the lagomorph unexcluded treatment. Canopy cover of honey mesquite (Prosopis glandulosa Torr. var. glandulosa), tarbush (Flourensia cernua DC.) and mariola (Parthenium incanum H.B.K.) were affected by lagomorph exclusion. None of the responses were viewed as successional in nature. They principally represented individual species sensitivities to either absence of a primary herbivore or removal of aboveground shrub biomass. Though the physical treatments could be regarded as relatively severe disturbances of the system, the impacts on community vegetation dynamics were relatively insignificant.     

Soil Disturbance as a Grassland Restoration Measure—Effects on Plant Species Composition and Plant Functional Traits

Soil disturbance is recognized as an important driver of biodiversity in dry grasslands, and can therefore be implemented as a restoration measure. However, because community re-assembly following disturbance includes stochastic processes, a focus only on species richness or establishment success of particular species will not inform on how plant communities respond ecologically to disturbance. We therefore evaluated vegetation development following disturbance by quantifying species richness, species composition and functional trait composition. Degraded calcareous sandy grassland was subjected to experimental disturbance treatments (ploughing or rotavation), and the vegetation was surveyed during four subsequent years of succession. Treated plots were compared with control plots representing untreated grassland, as well as nearby plots characterized by plant communities representing the restoration target.Species richness and functional diversity both increased in response to soil disturbance, and rotavation, but not ploughing, had a persistent positive effect on the occurrence of specialist species of calcareous sandy grassland. However, no type of soil disturbance caused the plant species composition to develop towards the target vegetation. The disturbance had an immediate and large impact on the vegetation, but the vegetation developed rapidly back towards the control sites. Plant functional composition analysis indicated that the treatments created habitats different both from control sites and target sites. Community-weighted mean Ellenberg indicator values suggested that the observed plant community response was at least partially due to an increase in nitrogen and water availability following disturbance. This study shows that a mild type of disturbance, such as rotavation, may be most successful in promoting specialist species in calcareous sandy grassland, but that further treatments are needed to reduce nutrient availability. We conclude that a functional trait based analysis provides additional information of the vegetation response and the abiotic conditions created, complementing the information from the species composition.     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Influence of woody invader control methods and seed availability on native and invasive species establishment in a Hawaiian forest

When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai’i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but short-lived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.