Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass

Although numerous studies have identified mechanisms that either resist or facilitate biological invasions, few studies have explicitly tested how resisting and facilitating mechanisms interact to drive invasion success. In California, USA, undisturbed Mediterranean-type shrublands have resisted invasion by the perennial tussock grass Cortaderia jubata. In some cases, however, this resistance has been spectacularly breached even in the absence of large-scale disturbance. I tested the hypothesis that these invasions are facilitated by local reductions in the strength of biotic resistance. I evaluated invasive success using C. jubata seed and seedling additions at different microhabitats: the edge of a chaparral stand, under shrub canopy at different distances from the stand edge, and in canopy gaps within the stand. When left exposed to mammalian herbivores, seedling survivorship decreased sharply from nearly 40% on the stand edge to zero just 10 m into the stand. When transplants were protected from herbivory, however, distance from the edge had no significant influence on transplant survivorship. Seedling emergence was also greater on the edge and in canopy gaps than under the canopy, but these differences were not caused by differences in herbivory. The flux of invasive propagules reaching the soil surface was immense and greater along the edge and within gaps than under the stand canopy. Mirroring these patterns, naturally occurring seedling abundance declined dramatically with distance from the stand edge, and seedlings were far more common within stand gaps than would be expected given gap frequency within the stand. Despite strong biotic resistance to invasion within the stand, the cover of C. jubata has increased 20% over the last 9 years. These results suggest that the relative amount of susceptible edge habitat and the supply of invasive propagules can facilitate invasion even in the face of strong local biotic resistance.     

Snowpack,fire, and forest disturbance: interactions affect montane invasions by non‐native shrubs

Montane regions worldwide have experienced relatively low plant invasion rates, a trend attributed to increased climatic severity, low rates of disturbance, and reduced propagule pressure relative to lowlands. Manipulative experiments at elevations above the invasive range of non‐native species can clarify the relative contributions of these mechanisms to montane invasion resistance, yet such experiments are rare. Furthermore, global climate change and land use changes are expected to cause decreases in snowpack and increases in disturbance by fire and forest thinning in montane forests. We examined the importance of these factors in limiting montane invasions using a field transplant experiment above the invasive range of two non‐native lowland shrubs, Scotch broom (Cytisus scoparius) and Spanish broom (Spartium junceum), in the rain–snow transition zone of the Sierra Nevada of California. We tested the effects of canopy closure, prescribed fire, and winter snow depth on demographic transitions of each species. Establishment of both species was most likely at intermediate levels of canopy disturbance, but at this intermediate canopy level, snow depth had negative effects on winter survival of seedlings. We used matrix population models to show that an 86% reduction in winter snowfall would cause a 2.8‐fold increase in population growth rates in Scotch broom and a 3.5‐fold increase in Spanish broom. Fall prescribed fire increased germination rates, but decreased overall population growth rates by reducing plant survival. However, at longer fire return intervals, population recovery between fires is likely to keep growth rates high, especially under low snowpack conditions. Many treatment combinations had positive growth rates despite being above the current invasive range, indicating that propagule pressure, disturbance, and climate can all strongly affect plant invasions in montane regions. We conclude that projected reductions in winter snowpack and increases in forest disturbance are likely to increase the risk of invasion from lower elevations.     

<Emphasis Type="Italic">PRUNUS</Emphasis>: a spatially explicit demographic model to study plant invasions in stochastic,heterogeneous environments

To model the invasion of Prunus serotina invasion within a real forest landscape we built a spatially explicit, non-linear Markov chain which incorporated a stage-structured population matrix and dispersal functions. Sensitivity analyses were subsequently conducted to identify key processes controlling the spatial spread of the invader, testing the hypothesis that the landscape invasion patterns are driven in the most part by disturbance patterns, local demographical processes controlling propagule pressure, habitat suitability, and long-distance dispersal. When offspring emigration was considered as a density-dependent phenomenon, local demographic factors generated invasion patterns at larger spatial scales through three factors: adult longevity; adult fecundity; and the intensity of self-thinning during stand development. Three other factors acted at the landscape scale: habitat quality, which determined the proportion of the landscape mosaic which was potentially invasible; disturbances, which determined when suitable habitats became temporarily invasible; and the existence of long distance dispersal events, which determined how far from the existing source populations new founder populations could be created. As a flexible “all-in-one” model, PRUNUS offers perspectives for generalization to other plant invasions, and the study of interactions between key processes at multiple spatial scales.     

Traits linked with species invasiveness and community invasibility vary with time,stage and indicator of invasion in a long‐term grassland experiment

Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.     

Dynamics of temperate rainforest from fine resolution pollen analysis,Upper Ringarooma River,northeastern Tasmania

Abstract The relationship between sclerophyll forest and temperate rainforest in Tasmania is believed to be driven by climate, soils and fire regime; however, this has been tested infrequently on time frames relevant to the longevity of the major forest trees involved. This study uses sediments in four small hollows from within a rainforest stand in the Upper Ringarooma valley in northeastern Tasmania, Australia, to analyse vegetation dynamics and disturbance over the last 1000 years. Pollen-vegetation relationships were investigated by comparing pollen rain and current vegetation, and the chronology was established by radiocarbon and lead-210 dates. The lead-210 data indicate that the two Sphagnum humus profiles were not always closed systems for inorganic inwash; however, the pollen and charcoal records are generally consistent indicating that a mixed Eucalyptus-Nothofagus forest shifted to a Kothofagus dominated rainforest as fire became a less prominent component of the environment. About 400 years ago Leptospermum lanigerum invaded the community as canopy gaps developed, possibly as the older Nothofagus Cunninghamii trees died. There is no strong evidence of invasion by Atherosperma moschatum as suggested by some models of forest dynamics in these forests in Tasmania. In this stand of forest, fire and erosion following flooding have been the major causes of disturbance.     

The importance of disturbance by fire and other abiotic and biotic factors in driving cheatgrass invasion varies based on invasion stage

Disturbances create fluctuations in resource availability that alter abiotic and biotic constraints. Exotic invader response may be due to multiple factors related to disturbance regimes and complex interactions between other small- and large-scale abiotic and biotic processes that may vary across invasion stages. We explore how cheatgrass responds to both frequency and season of prescribed burning for a 10-year period in ponderosa pine forested stands. To understand interactions of fire disturbance, other abiotic factors, biotic resistance, and propagule pressure, we use long-term data from different spatial scales representing different invasion stages (local establishment or spread and broader scale extent/impact) to model cheatgrass dynamics. We found that after 10 years, cheatgrass cover increased with fall burning regardless of burn frequency (1 burn vs. 3 burns). There was no evidence that cheatgrass invasion is decreasing through time even in areas burned only once. Factors important for explaining local fine-scale cheatgrass establishment and spread, and broader scale extent/impact varied. The spatial extent of the first burns facilitated fine-scale cheatgrass establishment while bare soil cover constrained establishment. Biotic resistance, in the form of native annual forb cover, constrained fine-scale cheatgrass spread. Initial cheatgrass abundance in 2002, a factor related to propagule pressure, was key for explaining the broader scale extent/impact of cheatgrass by 2012. Biotic resistance, in the form of native perennial grass cover, constrained extent/impact but only when initial cheatgrass abundance was low. Our findings regarding factors affecting invasion dynamics may be useful to consider for future restoration and conservation efforts in burned ponderosa pine forests.     

管涔山青扦(Picea wilsoni)天然林年龄结构及其动态的研究

地种群年龄的研究表明,虽经人为频繁干扰,管涔肝扦天然林仍表现出异龄林结构特征,立木年龄范围超过一个级期,根据年龄结构特征值可分为相对龄林、相对异龄林和异龄林３种类型林下新一代种群的数量和结构受林冠郁闭度和结构的影响。具垂直郁闭型林冠的异龄林,林下更新数量充足,幼苗幼树年龄结构合理;而服闭型林冠,不利于一代种群的发生和发展,青扦种群年龄结构受种群发生和自疏两个过程的控制,林下种群的发生以小规模林冠空     

Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Disturbance, invasion and re-invasion: managing the weed-shaped hole in disturbed ecosystems

We aim to develop a simple model to explore how disturbance and propagule pressure determine conditions for successful invasion in systems where recruitment occurs only in disturbed sites. Disturbance is often thought to favour invaders as it allows recruitment; however, the effects of disturbance are more complicated when it results in mortality of the invader. When disturbance rates in both invader occupied and unoccupied sites are the same, recruitment and mortality effects are exactly balanced, and successful invasion is independent of the disturbance regime. Differences in the disturbance rates between invader occupied and unoccupied sites can occur through invader modification or management of disturbance. Under these conditions, we found a novel mechanism for the generation of an Allee effect, which occurs when the invader promotes disturbance in sites it already occupies. When Allee effects occur one-off, large-scale disturbances can result in permanent, dramatic shifts in invader abundance; and conversely, reducing the population below a critical threshold can cause extinction.     

Propagule pressure and disturbance interact to overcome biotic resistance of marine invertebrate communities

Propagule pressure is fundamental to invasion success, yet our understanding of its role in the marine domain is limited. Few studies have manipulated or controlled for propagule supply in the field, and consequently there is little empirical data to test for non-linearities or interactions with other processes. Supply of non-indigenous propagules is most likely to be elevated in urban estuaries, where vessels congregate and bring exotic species on fouled hulls and in ballast water. These same environments are also typically subject to elevated levels of disturbance from human activities, creating the potential for propagule pressure and disturbance to interact. By applying a controlled dose of free-swimming larvae to replicate assemblages, we were able to quantify a dose–response relationship at much finer spatial and temporal scales than previously achieved in the marine environment. We experimentally crossed controlled levels of propagule pressure and disturbance in the field, and found that both were required for invasion to occur. Only recruits that had settled onto bare space survived beyond three months, precluding invader persistence in undisturbed communities. In disturbed communities initial survival on bare space appeared stochastic, such that a critical density was required before the probability of at least one colony surviving reached a sufficient level. Those that persisted showed 75% survival over the following three months, signifying a threshold past which invaders were resilient to chance mortality. Urban estuaries subject to anthropogenic disturbance are common throughout the world, and similar interactions may be integral to invasion dynamics in these ecosystems.     

管涔山青扦(Picea wilsoni)天然林年龄结构及其动态的研究

对种群年龄结构的研究表明,虽经人为频繁干扰,管涔山青扦天然林仍表现出异龄林结构特征,立木年龄范围超过一个龄级期,根据年龄结构特征值可分为相对同龄林、相对异龄林和异龄林３种类型。林下新一代种群的数量和结构受林冠郁闭度和结构的影响。具垂直郁闭型林冠的异龄林,林下更新数量充足,幼苗幼树年龄结构合理;而水平郁闭型林冠,不利于新一代种群的发生和发展。青扦种群年龄结构受种群发生和自疏两个过程的控制,林下种群的发生以小规模林冠空隙干扰下的连续更新为主。青扦华北落叶松混交林,在其共同适生范围内是某种干扰格局控制下的稳定群落     

Treefall gaps required for establishment,but not survival,of invasive Rubus phoenicolasius in deciduous forest,Maryland, USA

Although plant invasions are often associated with disturbance, localized disturbances can promote invasion either by: (i) creating sites where individuals establish; or (ii) enabling an invader to colonize the entire stand. The former is expected when both establishment and survival to reproductive age require disturbed conditions, whereas the latter should occur in systems when either establishment or survival are limited to disturbed sites. We investigated the role of localized disturbance, specifically treefalls, in the invasion of the Asian Rubus phoenicolasius in a deciduous forest in Maryland, USA. We investigated the density and demography of R. phoenicolasius in treefall gaps of various sizes, but identical age to non‐gap areas, using Classification and Regression Tree (CART) analyses to identify the most important predictors. To explore how the demography of established individuals responds to disturbed versus undisturbed conditions, we carried out a garden experiment with three different levels of shade (5, 12 and 22% full sun). We found vegetative and sexual reproduction, and seedling establishment, to be limited to large gaps in an old stand, but not in a stand in an earlier age of succession. However, in the garden experiment, established plants were able to survive and grow under all shade treatments. These findings indicate that R. phoenicolasius requires disturbances such as treefalls to establish in forests, but established plants will survive canopy closure, leading to stand‐wide invasion. Managers should be able to prevent invasion, however, by inspecting large gaps for new recruits every 3 years.     

Inferring process from pattern in plant invasions: a semimechanistic model incorporating propagule pressure and environmental factors

Propagule pressure is intuitively a key factor in biological invasions: increased availability of propagules increases the chances of establishment, persistence, naturalization, and invasion. The role of propagule pressure relative to disturbance and various environmental factors is, however, difficult to quantify. We explored the relative importance of factors driving invasions using detailed data on the distribution and percentage cover of alien tree species on South Africa's Agulhas Plain (2,160 km2). Classification trees based on geology, climate, land use, and topography adequately explained distribution but not abundance (canopy cover) of three widespread invasive species (Acacia cyclops, Acacia saligna, and Pinus pinaster). A semimechanistic model was then developed to quantify the roles of propagule pressure and environmental heterogeneity in structuring invasion patterns. The intensity of propagule pressure (approximated by the distance from putative invasion foci) was a much better predictor of canopy cover than any environmental factor that was considered. The influence of environmental factors was then assessed on the residuals of the first model to determine how propagule pressure interacts with environmental factors. The mediating effect of environmental factors was species specific. Models combining propagule pressure and environmental factors successfully predicted more than 70% of the variation in canopy cover for each species.     

Disturbance history of an old-growth subalpine larch forest in the Qinling Mountains,north-central China

Knowledge on how historical disturbances shaped the long-term development of forests is essential for understanding the present forest structure and for predicting the future forest ecosystem dynamics. In this study, dendroecological methods were used to reconstruct the disturbance history of an old-growth subalpine larch (Larix chinensis) forest in the Qinling Mountains of north-central China. Growth patterns of 690 and 582 increment cores extracted respectively from two climatically and topographically different larch stands in the northern and southern slope of the Qinling Mountains were examined for abrupt increases in radial growth indicating formation of past canopy gaps and for rapid early growth rates indicating recruitment in former canopy gaps. The findings demonstrated that there were no large-scale, stand-replacing disturbances during the past more than two centuries. Low- and medium-severity disturbance events predominated, which were probably caused by windthrows due to strong winds. The stand was unevenly aged, and the recruitment pulses associated with disturbance peaks could be distinguished. There were considerable spatio-temporal differences in disturbance dynamics of the subalpine larch stand between the topographically and climatically different sites, manifesting that the larch stand in the northern slope experienced frequent moderate but rare major disturbance events, contrasting with frequent major and moderate disturbance events in the southern slope. This study provided strong evidences that there were substantial variations in the intensity and frequency of disturbance dynamics, leading to considerable differences in the size and age structures of the subalpine larch forest.     

Temporal change in the diversity-invasibility relationship in the presence of a disturbance regime

Disturbance can affect both the diversity and invasibility of communities. Many field studies have found correlations between diversity and susceptibility to invasion, but if both factors independently respond to disturbance then spurious non-causal relationships may be observed. Here, we show that disturbance can cause a temporal shift in the diversity-invasibility relationship. In a field experiment using sessile marine communities, disturbance strongly affected both diversity and invasion such that they were highly correlated. Disturbance facilitated initial invasion, creating a negative diversity-invasibility relationship when the invader first arrived. Over time, disturbance hindered the persistence of invaders, creating a positive diversity-invasibility relationship. We suggest that temporal changes in the diversity-invasibility relationship may have contributed to the 'invasion paradox', a term for the contrasting patterns of experimental and observational studies of the diversity-invasibility relationship.     

Biological control as an invasion process: disturbance and propagule pressure affect the invasion success of Lythrum salicaria biological control agents

Understanding the mechanisms behind the successful colonization and establishment of introduced species is important for both preventing the invasion of unwanted species and improving release programs for biological control agents. However, it is often not possible to determine important introduction details, such as date, number of organisms, and introduction location when examining factors affecting invasion success. Here we use biological control introduction data to assess the role of propagule pressure, disturbance, and residence time on invasion success of four herbivorous insect species introduced for the control of the invasive wetland plant, Lythrum salicaria, in the Columbia River Estuary. Two sets of field surveys determined persistence at prior release sites, colonization of new sites, and abundance within colonized sites. We quantified propagule pressure in four ways to examine the effect of different measurements. These included three measurements of introduction size (proximity to introduction site, introduction size at a local scale, and introduction size at a regional scale) and one measure of introduction number (number of introduction events in a region). Disturbance was examined along a tidal inundation gradient (distance from river mouth) and as habitat (island or mainland). Statistical models and model averaging were used to determine which factors were driving invasion success. In this study we found: (1) sparse evidence for the positive influence of propagule pressure on invasion success; (2) disturbance can negatively affect the invasion success of herbivorous insects; (3) the effects of disturbance and propagule pressure are species specific and vary among invasion stages, and (4) not all measures of propagule pressure show the same results, therefore single measures and proxies should be used cautiously.     

Does Prunus serotina act as an aggressive invader in areas with a low propagule pressure?

Since most studies on Prunus serotina in Western Europe focused on heavily invaded areas, we wondered whether P. serotina also acts as an aggressive invader in areas with a low propagule pressure. Based on long-term data for the Liedekerke forest reserve, we found that connectivity to seed sources and light availability were the major drivers of P. serotina presence: long-distance dispersal events and ‘windows of opportunity’ seem to direct P. serotina colonization. In the studied forest, P. serotina could not be considered an aggressive invader since its spread slowed down rather quickly and did not hamper the establishment of native tree species. Furthermore, understory P. serotina showed low growth and seed production, while the high Rubus cover hampered germination and establishment. Nonetheless, calamities opening up the canopy layer in the few areas with high P. serotina sapling density might alter the course of the invasion process.     

Relationship of Ditylenchus dipsaci and Harvest Practices to the Persistence of Alfalfa

Persistence of dormant Ranger and nondormant Moapa alfalfas, both susceptible to Ditylenchus dipsaci, varied with stand age and cutting frequency. Stand reduction increased with cutting frequency. In D. dipsaci-infested soil, stand reductions in Ranger 1, 4, and 5 years old exceeded reductions in stands 2 and 3 years old; persistence was greatest in 2-year-old stands. In Moapa alfalfa, D. dipsaci reduced stands the most in years 2 and 3; whereas persistence was greatest in 1-year-old stands. Harvesting Ranger alfalfa one, two, three, and four times during the growing season reduced 2-year-old stands by 10, 14, 19, and 29% in D. dipsaci-infested soil and by 2, 4, 4, and 7% in uninfested soil, respectively. Comparable reductions in Moapa alfalfa were 13, 16, 18, and 38% in infested soil and 0, 2, 4, and 6% in uninfested soil. Cutting frequency had less effect on persistence of resistant semidormant Lahontan grown in D. dipsaci-infested soil relative to susceptible cultivars. Increasing the number of cuttings per year decreased storage of total nonstructural carbohydrate and adversely affected persistence of alfalfa stands and yields; the greatest negative effects occurred on both resistant and susceptible alfalfa in D. dipsaci-infested soil.     

Invaders' control on post-disturbance succession in coastal mangroves

Aims In recent years, coastal mangroves have been frequently affected by large disturbances (cyclones, hurricanes, flooding and tsunamis) and post-disturbance vegetation is often dominated by small stature mangrove, mangrove-associate and non-mangrove species potentially affecting ecosystem functioning. Knowledge on the processes of mangrove vegetation development and recovery (succession) following normal and large disturbances will benefit practitioners in designing robust ecosystem management/restoration plans. Here we propose a conceptual model of disturbance-mediated succession in mangroves.Methods Based on field observations and species' life history traits, we develop conceptual models of mangrove succession under normal disturbance regime and recently experienced increased frequency of large disturbances. We evaluate our conceptual models by conducting a scenario testing experiment.Important findings We suggest two predominant processes affecting mangrove succession after disturbance: propagule limitation due to damage of seed producing mature trees and dispersal barrier resulting from biological invasion associated with large disturbance. We argue that large disturbances affect mature trees more than the small-stature non-tree (shrubs, herbs and climbers) species creating a larger propagule shortage for mangrove tree species than non-tree species. Secondly, large disturbances facilitate invasion of free-floating aquatics, which may interfere with the flow-facilitated propagule dispersal and seedling establishment of mangrove species. In a scenario testing experiment, we have shown that similar levels of disturbance impact vegetation development and recovery differently depending on the presence or absence of invasive species. We conclude that since biological invasion is one of the major drivers of post-disturbance mangrove succession, the dimension of biological invasion should be included in prediction, management and restoration of mangrove forests.