Grazing and community structure as determinants of invasion success by Scotch broom in a New Zealand montane shrubland

Abstract. Scotch broom ( Cytisus scoparius (L.) Link; Fabaceae) is a problematic invasive plant in many countries, and while attention has been paid to traits that make it a successful invader, there has been less focus on the properties of ecosystems that it invades. We conducted an experiment in a New Zealand montane shrubland with tussock grasses that has been invaded by Scotch broom to determine features that rendered it susceptible to invasion. We planted broom seedlings into the shrubland (control) and into three treatments: (1) resident shrubs removed, (2) tussocks removed and (3) shrubs and tussocks removed. We measured broom seedling mortality and growth over two growing seasons. The site was grazed by sheep in the first season, and scarcely grazed in the second, wetter season. Survivorship across all treatments after 19 months was 42%, and was lowest where shrubs were retained but tussocks removed. Broom seedlings grew taller and had greater leaf areas in treatments that retained shrubs. Neighbouring (within 49 cm) shrubs had no effects on survivorship or growth of broom seedlings. Neighbouring tussocks increased survivorship of broom seedlings but depressed their growth. Grazing by sheep was the most important determinant of survivorship and growth of broom seedlings, and effects were uniform regardless of experimental treatments. Initial high mortality of seedlings (48% in the first 3 months) was due to grazing, and height growth was often negative during periods of grazing. In the second growing season when the site was less grazed and there was greater rainfall, there was a rapid increase in height across all treatments. Continued grazing of the site by sheep is likely to be the chief means of retarding the invasion.     

Feeding range of Sitona regensteinensis Hbst. (Coleoptera: Curculionidae), a potential agent for biological control of Cytisus scoparius (L.) link (Broom) in New Zealand

The weevil Sitona regensteinensis is being considered for biological control of Cytisus scoparius in New Zealand. Adult feeding tests demonstrated that while foliage of C scoparius was preferred, weevils fed also on Chamaecytisus palmensis and Lupinus arboreus. Tests with the root‐feeding larval stage have yet to be completed.     

辽东山区次生林木本植物空间分布

森林木本植物的空间格局有助于揭示群落结构的形成机制与潜在的生态学过程,且对林分经营具有一定指导意义。在0—50 m尺度范围内综合分析了辽东山区4 hm2温带次生林样地多度10的树种空间格局。研究发现:(1)在完全随机零模型下,大部分树种呈现聚集格局,聚集格局树种的比例随尺度增加而降低;在32 m的较大尺度下,随尺度增加,随机和规则格局成为树种分布的主要形式;(2)在异质性泊松过程零模型下,55.9%的树种呈现随机格局,其余大部分树种在10 m的尺度下呈现聚集格局,且随尺度增加,规则格局成为主要形式;(3)在完全随机零模型下,树种属性(林层、径级和多度)显著地影响种群聚集度,而在异质性泊松过程零模型下,树种属性对种群聚集度不存在显著影响。综上,生境异质性、扩散限制和树种属性部分解释了辽东山区次生林木本植物空间分布格局,相对而言,生境异质性的效应更为突出。研究结果有助于揭示次生林群落生物多样性的维持机制。     

碱化草地群落恢复演替空间格局动态分析

对松嫩平原碱化草地1989年开始围栏封育的1hm^2样地9a间的空间格局进行了分析,结果表明：（1）不同物种斑块空间格局动态不同,在恢复演替各个阶段中的地位与作用也不同。（2）结论基本上支持了斑块化－稳定性假说。     

Plant composition associated with environmental gradients in tropical montane forests (Cueva de Los Guacharos National Park,Huila, Colombia)

Niche differentiation among tropical forest plants can generate species turnover along gradients of soil, topography, climate, and land use history. In this study we explore the relative importance of these variables as drivers of floristic composition in Cueva de Los Guacharos National Park. We established twenty 0.1‐ha plots, within which trees, lianas, and shrubs (diameter ≥ 2.5 cm) were censused. We selected plot locations in primary and disturbed forests, and we measured topography and soil variables. Despite their structural similarity, primary and disturbed forests differed floristically, and also differed in environmental variables measured. A NMDS ordination showed that variation in the floristic composition across plots is highly correlated to the exchangeable acidity, elevation, temperature, and magnesium availability. Variance partitioning analysis shows that together spatial and environmental variables explain 24.2 percent of the variation in species composition. ‘Pure environmental’ variables were more important in explaining compositional variability than ‘pure spatial’ processes (9.8% and 1.4%, respectively). Residual variance may be attributed to stochastic process or non‐measured biotic effects.     

A modelling approach to estimate the effect of exotic pollinators on exotic weed population dynamics: bumblebees and broom in Australia

The role of mutualisms in contributing to species invasions is rarely considered, inhibiting effective risk analysis and management options. Potential ecological consequences of invasion of non‐native pollinators include increased pollination and seed set of invasive plants, with subsequent impacts on population growth rates and rates of spread. We outline a quantitative approach for evaluating the impact of a proposed introduction of an invasive pollinator on existing weed population dynamics and demonstrate the use of this approach on a relatively data‐rich case study: the impacts on Cytisus scoparius (Scotch broom) from proposed introduction of Bombus terrestris. Three models have been used to assess population growth (matrix model), spread speed (integrodifference equation), and equilibrium occupancy (lattice model) for C. scoparius. We use available demographic data for an Australian population to parameterize two of these models. Increased seed set due to more efficient pollination resulted in a higher population growth rate in the density‐independent matrix model, whereas simulations of enhanced pollination scenarios had a negligible effect on equilibrium weed occupancy in the lattice model. This is attributed to strong microsite limitation of recruitment in invasive C. scoparius populations observed in Australia and incorporated in the lattice model. A lack of information regarding secondary ant dispersal of C. scoparius prevents us from parameterizing the integrodifference equation model for Australia, but studies of invasive populations in California suggest that spread speed will also increase with higher seed set. For microsite‐limited C. scoparius populations, increased seed set has minimal effects on equilibrium site occupancy. However, for density‐independent rapidly invading populations, increased seed set is likely to lead to higher growth rates and spread speeds. The impacts of introduced pollinators on native flora and fauna and the potential for promoting range expansion in pollinator‐limited ‘sleeper weeds’ also remain substantial risks.     

Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.     

Components of spatial patterning in a serpentine grassland

We analyzed the components of spatial patterning in species abundance in a diverse grassland dominated by early season, annual forbs. Species abundance, soil depth and gopher disturbances were measured by means of a nested spatial design on two 8m×20m plots that differed in the amount of larger scale variability in soil depth. Species distributions at the scale of the 10cm×10cm quadrat were highly clumped, with a decline in clumping with mean abundance. Even with the effect of abundance on clumping removed, species showed differences in the degree of clumping and these differences were consistent between the two plots. The abundance of the species and the frequency of disturbance correlated weakly to moderately with soil depth. Semivariance analyses indicated that the most common species all showed complex spatial patterning at a range of scales. Some of this variation corresponded to patterns of soil depth variation and patterns of gopher disturbance; however, a large amount of the spatial patterning in species abundance remains unexplained.     

科尔沁沙地草地植被恢复演替进程中群落优势种群空间分布格局研究

通过对处于流动沙地、半固定沙地和固定沙地等不同恢复演替阶段的群落的取样调查 ,研究了科尔沁沙地草地植被恢复进程中群落优势种群的空间分布格局及其动态变化。运用扩散系数、聚集指数、平均拥挤度、聚块性指数、Green指数、聚集强度、Poisson分布和负二项分布的 χ2 拟合检验等 7项指标 ,对群落优势种群的分布格局的研究结果基本一致 ,大多数物种的空间分布呈显著的聚集分布。在演替进程中种群格局强度的变化和群落总体聚集程度的变化与种群动态和群落结构的空间异质性相对应 ,较高的聚集程度是种群分布斑块化和群落结构的空间异质性的反映 ,随着演替进展 ,优势种的种群数量扩展 ,但种群分布的聚集强度下降 ,群落结构的空间异质性降低。在 7种测定分布格局的指标中 ,聚块性指数、Green指数和聚集强度是无量纲的指标 ,其大小不受种群大小的影响 ,其变化可以较好地指示格局强度本身的变化 ,并与种群动态和群落结构变化相联系     

BIODIVERSITY RESEARCH: Experimental introduction of the alien plant Hieracium lepidulum reveals no significant impact on montane plant communities in New Zealand

Aim There is debate over whether alien plants necessarily alter the communities they invade or can coexist with native species without discernable impacts. We followed the fate of montane plant communities in response to the experimental sowing of the alien weed Hieracium lepidulum, looking for changes in plant community composition and structure over 6 years. Location Craigieburn Range, New Zealand. Methods We used a replicated randomised block design, with 30 × 30 cm plots (n = 756) subdivided into 5 × 5 cm cells to examine and compare the effects of H. lepidulum at 0.09 m² (plot) and 0.0025 m² (cell) scales. Plots were sown with between 0 and 15,625 H. lepidulum seeds in 2003, forming gradients of invader density and cover. Measurements comprised community richness, evenness and diversity along with H. lepidulum density and cover at both scales. The relationships between the invader and local community attributes were modelled using hierarchical mixed‐effect models. Results Plant communities differed in the extent to which they became invaded, with H. lepidulum cover in the plots ranging from 0% to 52%, with a mean of only 1.89%. Plot species richness increased from 2003 to 2009, with a component of this increase (+0.002 species per year) associated with increasing H. lepidulum density. Other relationships between the plant community and H. lepidulum were generally non‐significant. Main conclusions In these montane plant communities, it appears H. lepidulum coexists with the native community with no measurable negative effects after 6 years on species richness, evenness or diversity, even where density and cover of the invader are highest. We suggest H. lepidulum has persisted preferentially at those sites with abiotic conditions sufficient to support a species‐rich assemblage.     

Coexistence and invasibility in a two-species competition model with habitat-preference

The outcome of competition among species is influenced by the spatial distribution of species and effects such as demographic stochasticity, immigration fluxes, and the existence of preferred habitats. We introduce an individual-based model describing the competition of two species and incorporating all the above ingredients. We find that the presence of habitat preference—generating spatial niches—strongly stabilizes the coexistence of the two species. Eliminating habitat preference—neutral dynamics—the model generates patterns, such as distribution of population sizes, practically identical to those obtained in the presence of habitat preference, provided an higher immigration rate is considered. Notwithstanding the similarity in the population distribution, we show that invasibility properties depend on habitat preference in a non-trivial way. In particular, the neutral model results more invasible or less invasible depending on whether the comparison is made at equal immigration rate or at equal distribution of population size, respectively. We discuss the relevance of these results for the interpretation of invasibility experiments and the species occupancy of preferred habitats.     

Ecological succession in new and old swiddens of montane Papua New Guinea

Fieldwork conducted in new and old swiddens in the Maring territory of the Bismarck Mountains of Papua New Guinea indicated a successional sequence of crop composition, diversity, and biomass regulated by human and ecological processes. With increasing swidden age, the number of cultivated species, individuals, varieties, and net productivity decreased. In contrast to new swiddens, old swiddens were characterized by a simplified crop composition and a distinct vertical stratification. While a 3-month-old garden contained 10 cultivar species, 43.5 plantings of taro, a cultivar biomass content of 2.54 kg/25 m², and a net productivity of 1.13 gm/m²/day, a 2-year-old garden contained six cultivar species, no taro plantings, a cultivar biomass content of 13.80 kg/25 m², and a net productivity of 0.77 gm/m²/day. Differences in the life histories of cultivars help determine cultivar succession. This study quantitatively defines these changing characteristics of old and new swiddens. It concludes that the Kauwatyi system of subsistence, which emphasizes diversity and conservation practices, such as the planting and protection of certain forest regrowth species, ensures the regeneration of forest.     

Early herbaceous succession along a topographical gradient on forest clear-felling sites in mountainous terrain,central Japan

Early successional patterns of herbaceous communities in forest clear-felling sites were investigated along a topographical gradient, which included ridge, slope and valley habitat types, in warm-temperate evergreen forest regions of central Japan for 5 years. Three dominant species with wind-dispersed seeds played a major role in the succession: an annual,Crassocephalum crepidioides, a biennial,Erigeron canadensis, and a perennial,Miscanthus sinensis. Pioneer herbs that have a seed-bank strategy, which are common in old field succession, were not found in the mountainous sites. The persistence of the annual or biennial dominants during the very early stages of secondary succession was different in the three topographical habitat types. In the ridge habitat,M. sinensis dominated from the first year state, andC. crepidioides andE. canadensis were less prominent.Crassocephalum crepidioides andE. canadensis became dominant as one moved down the slope. In the valley habitat,C. crepidioides dominated in the first year stage, was succeeded byE. canadensis in the second year, and thenM. sinensis gradually replaced it in later years. As all three wind-dispersed dominants simultaneously invaded in all the habitat types after clear-felling, the different successional patterns along the topographical gradient might have resulted from differences in the establishment ability and the growth rate of the three dominants depending on the three habitat types.     

辽东山区次生林乔木幼苗分布格局与种间空间关联性

在辽东山区次生林建立4 hm^2样地(200 m×200 m),研究0~50 m尺度范围内乔木幼苗分布格局及种间空间关联性.结果表明:在完全随机零模型下,0~20 m尺度上,95%的树种呈现聚集分布格局;0~16 m尺度上,19个树种呈现聚集分布;随着尺度的增加,聚集分布树种的比例逐渐减少,50 m尺度上,随机分布成为树种分布的主要形式;在异质性泊松过程零模型下,0~24 m尺度上,5%的树种呈现聚集分布,26~50 m尺度上,42%和58%的树种呈现随机和均匀分布.在完全随机零模型下,正相关树种对比例最高,且在50 m尺度下呈现正相关、负相关、无相关3种相关性的树种对比例相同;在异质性泊松过程零模型下,树种对主要呈现负相关,且随尺度增大,负相关的树种对比例逐渐升高.种子扩散限制和生境异质性在某种程度上解释了乔木幼苗的聚集分布格局,乔木幼苗强烈的聚集分布又促使种间空间关联性密切,更新群落稳定性较差.     

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case–control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.     

Experimental slab salvage and reinstatement after pipeline construction in a threatened grassland community

The protection and management of grasslands is a priority to conserve biodiversity, particularly in fragmented landscapes subject to human‐induced disturbance. We examined the success of salvage and reinstatement of multiple 90 m² slabs of modified temperate grassland in a pipeline construction corridor in south‐eastern Australia and compared it with a reinstated topsoil treatment (i.e. standard pipeline reinstatement practice) and an undisturbed control. Thirty‐six plots (each 9 × 10 m) representing the three treatments were monitored across six locations, pre‐ and postconstruction every 3 months for 2 years. We sampled 18 vegetation/habitat variables to examine the effect of treatments over time. Introduced grass cover was consistently higher than native grasses for both treatments, and the cover of introduced grasses increased over time in both treatments. Conversely, control plots had considerably higher native grass cover than the slab and reinstated topsoil treatments. Native species richness was not negatively affected by the slab reinstatement process. Our results suggest that in this instance, slab replacement was an ineffective way to reinstate native grasslands, as it appeared to confer minimal benefit when compared to the standard practice of reinstating topsoil. This was most likely due to the high degree of disturbance to slabs during the reinstatement process.