On the relative contributions of wind vs. animals to seed dispersal of four Sierra Nevada pines

Selective pressures that influence the form of seed dispersal syndromes are poorly understood. Morphology of plant propagules is often used to infer the means of dispersal, but morphology can be misleading. Several species of pines, for example, have winged seeds adapted for wind dispersal but owe much of their establishment to scatter-hoarding animals. Here the relative importance of wind vs. animal dispersal is assessed for four species of pines of the eastern Sierra Nevada that have winged seeds but differed in seed size: lodgepole pine (Pinus contorta murrayana, 8 mg); ponderosa pine (Pinus ponderosa ponderosa, 56 mg); Jeffrey pine (Pinus jeffreyi, 160 mg); and sugar pine (Pinus lambertiana, 231 mg). Pre-dispersal seed mortality eliminated much of the ponderosa pine seed crop (66%), but had much less effect on Jeffrey pine (32% of seeds destroyed), lodgepole pine (29%), and sugar pine (7%). When cones opened most filled seeds were dispersed by wind. Animals removed > 99% of wind-dispersed Jeffrey and sugar pine seeds from the ground within 60 days, but animals gathered only 93% of lodgepole pine seeds and 38% of ponderosa pine seeds during the same period. Animals gathered and scatter hoarded radioactively labeled ponderosa, Jeffrey, and sugar pine seeds, making a total of 2103 caches over three years of study. Only three lodgepole pine caches were found. Caches typically contained 1-4 seeds buried 5-20 mm deep, depths suitable for seedling emergence. Although Jeffrey and sugar pine seeds are initially wind dispersed, nearly all seedlings arise from animal caches. Lodgepole pine is almost exclusively wind dispersed, with animals acting as seed predators. Animals treated ponderosa pine in an intermediate fashion. Two-phased dispersal of large, winged pine seeds appears adaptive; initial wind dispersal helps to minimize pre-dispersal seed mortality whereas scatter hoarding by animals places seeds in sites with a higher probability of seedling establishment.     

Release thresholds strongly determine the range of seed dispersal by wind

The effect of the seed abscission process on the dispersal distance of seeds has never been studied explicitly and is often ignored in studies that aim to estimate the seed shadows of species. To examine the importance of the abscission process for the seed shadow we used a seed trajectory model that keeps track of the release threshold dynamics of the individual seeds on mother plant. We defined the release threshold as the critical wind speed that induces a mechanical force that is just large enough to release a seed from its mother plant. The model used real wind speed sequences and seed appearance over time on the mother plant.Several calculations were performed to investigate the effect of release thresholds dynamics on seed shadow of two herbaceous species with contrasting terminal velocity values (V_t): Centaurea jacea (V_t = 4.1 m s⁻¹) and Hypochaeris radicata (V_t = 0.49 m s⁻¹).Release thresholds were responsible for a two-fold increase of median dispersal distances in both species. Tails of the seed shadows, the fraction of seeds that travel furthest, were even more sensitive and increased with a factor 4.5 for Centaurea and 7.0 for Hypochaeris. Our work indicates that the abscission process appears to be very important and suggests that dispersal distance of plants is currently severely underestimated, which, in turn, has major consequences for our current understanding of the distribution, metapopulation dynamics and survival of plant species.     

Effects of matrix characteristics and interpatch distance on functional connectivity in fragmented temperate rainforests

The connectivity of remnant patches of habitat may affect the persistence of species in fragmented landscapes. We evaluated the effects of the structural connectivity of forest patches (i.e., distance between patches) and matrix class (land-cover type) on the functional connectivity of 3 bird species (the White-crested Elaenia [Elaenia albiceps], the Green-backed Firecrown Hummingbird [Sephanoides sephaniodes], and the Austral Thrush [Turdus falklandii]). We measured functional connectivity as the rate at which each species crossed from one patch to another. We also evaluated whether greater functional connectivity translated into greater ecological connectivity (dispersal of fruit and pollen) by comparing among forest patches fruit set of a plant pollinated by hummingbirds and abundance of seedlings and adults of 2 plants with bird- and wind-dispersed seeds. Interpatch distance was strongly associated with functional connectivity, but its effect was not independent of matrix class. For one of the bird-dispersed plants, greater functional connectivity for White-crested Elaenias and Austral Thrushes (both frugivores) was associated with higher densities of this plant. The lack of a similar association for the wind-dispersed species suggests this effect is linked to the dispersal vector. The abundance of the hummingbird-pollinated species was not related to the presence of hummingbirds. Interpatch distance and matrix class affect animal movement in fragmented landscapes and may have a cascading effect on the distribution of some animal-dispersed species. On the basis of our results, we believe effort should be invested in optimizing patch configuration and modifying the matrix so as to mitigate the effects of patch isolation in fragmented landscapes.     

Effect of Edge Structure on the Flux of Species into Forest Interiors

Abstract: A key goal of conservation biology is to prevent the spread of exotic species. Previous work on exotic invasion has two limitations: (1) the lack of a spatially explicit approach and (2) a primary focus on the net effect of invasion by examining invasive species already present in the community. We address these limitations by focusing on the arrival of a potential invader into a community and use a spatially explicit approach to quantify the flow of seeds from the surrounding landscape into the interior of a forest. We hypothesize that the structure of forest-edge vegetation influences how the edge mediates seed flux. To test our hypothesis, we experimentally altered vegetation structure within 20 m of the edge to create two edge treatments: thinned and intact. We quantified the flux of seeds moving into the forest interior across the two treatments. We used seed traps randomly arrayed on transects from 5 to 50 m into the forest. More seeds crossed the thinned treatment than crossed the intact treatment to reach the forest interior. In addition, seeds that crossed the thinned treatment dispersed farther into the forest than those that crossed the intact treatment. These results were consistent throughout the period of maximum autumn dispersal, including periods before and after leaf drop. Our results show that the structure of vegetation on the edge interacts with the flux of wind-dispersed seed across the edge. We demonstrated that an edge with intact vegetation can function as a physical barrier to seed dispersal. Therefore, the structure of vegetation on edges can influence the function of edges as barriers to seed flux into the forest interior.     

Recovery Patterns of Understory Herbs and Their Use as Indicators of Deciduous Forest Regeneration

Abstract: Habitat fragmentation has reduced the richness of native species of forests in northeastern North America. Despite recent large-scale increases in forest cover, studies indicate that understory herbaceous plant communities may take many decades to recover. We studied recovery patterns of vegetation following up to 35 years of forest regeneration in restored former cottage and road sites at Point Pelée National Park, Ontario, Canada, to assess the vulnerability of the understory herbaceous species. Overall, there were no significant differences in the diversity of native species between restored and relatively undisturbed reference sites. There was, however, significant among-site variation in the composition of the native species component of these plant communities. When only restored sites were examined, variation in native species composition was associated with time since site restoration, soil moisture, canopy cover, and distance to continuous forest. Native species were assigned vulnerability rankings according to their relative occurrence in reference and restored sites. Spring-flowering herbs, with ant- or gravity-dispersed seeds, were absent from restored sites and were defined as highly vulnerable. In contrast, summer- and fall-flowering herbs, with vertebrate- and wind-dispersed seeds, dominated restored sites and were less vulnerable. Species of low and intermediate vulnerability had colonized restored sites successfully, and the latter should function as indicators of recovery. In contrast, species with high vulnerability rankings had not recovered at all and, because of their limited dispersal ranges, may recolonize restored sites only if they are actively reintroduced.     

Meta‐Analysis of the Effects of Forest Fragmentation on Interspecific Interactions

Forest fragmentation dramatically alters species persistence and distribution and affects many ecological interactions among species. Recent studies suggest that mutualisms, such as pollination and seed dispersal, are more sensitive to the negative effects of forest fragmentation than antagonisms, such as predation or herbivory. We applied meta‐analytical techniques to evaluate this hypothesis and quantified the relative contributions of different components of the fragmentation process (decreases in fragment size, edge effects, increased isolation, and habitat degradation) to the overall effect. The effects of fragmentation on mutualisms were primarily driven by habitat degradation, edge effects, and fragment isolation, and, as predicted, they were consistently more negative on mutualisms than on antagonisms. For the most studied interaction type, seed dispersal, only certain components of fragmentation had significant (edge effects) or marginally significant (fragment size) effects. Seed size modulated the effect of fragmentation: species with large seeds showed stronger negative impacts of fragmentation via reduced dispersal rates. Our results reveal that different components of the habitat fragmentation process have varying impacts on key mutualisms. We also conclude that antagonistic interactions have been understudied in fragmented landscapes, most of the research has concentrated on particular types of mutualistic interactions such as seed dispersal, and that available studies of interspecific interactions have a strong geographical bias (arising mostly from studies carried out in Brazil, Chile, and the United States). Meta‐Análisis de los Efectos de la Fragmentación del Bosque sobre las Interacciones Interespecíficas     

Soil vs. canopy seed storage and plant species coexistence in species-rich Australian shrublands

The fire-prone shrublands of southwestern Australia are renowned for their high plant species diversity and prominence of canopy seed storage (serotiny). We compared species richness, abundance, and life history attributes for soil and canopy seed banks in relation to extant vegetation among four sites with different substrate conditions and high species turnover (50-80%) to identify whether this unusual community-level organization of seed storage might contribute to maintenance of high species richness. Soil seed bank (SSB) densities were low to moderate (233-1435 seeds/m2) compared with densities for other Mediterranean-type vegetation and were lowest for sites with highest canopy seed bank (CSB) species richness and lowest nutrient availability, but not richness or abundance of resprouters. Annuals were infrequent in the lowest nutrient sites, but there was no evidence that small SSB size was due to low seed inputs or a trade-off between seed production/storage and seed size in response to low nutrient availability. Sorensen's similarity between SSB and extant vegetation was 26-43% but increased to 54-57% when the CSB was included, representing levels higher than reported for most other ecosystems. Resprouting species were well represented in both the SSB and CSB, and there was no evidence for lower seed production in resprouters than in non-sprouters overall. The SSB and CSB held no species in common and were characterized by markedly different seed dispersal attributes, with winged or small seeds in the CSB and seeds dispersed by ants, birds, and wind (though none with wings) in the SSB. There was no evidence of spatial differentiation in the distribution of seeds of SSB species between vegetated and open microsites that might facilitate species coexistence, but most woody non-sprouters showed aggregation at scales of 1-2 m, implying limited seed dispersal. High similarity between overall seed bank (SSB + CSB) and extant species composition, high number of resprouting species, and seed dispersal processes before (SSB) and after fire (CSB) leading to differential spatial aggregation of post-fire recruits from the two seed bank types may buffer species composition against rapid change and provide a mechanism for maintaining species coexistence at the local scale.     

天津臭氧浓度与气象因素的相关性及其预测方法

气象因素在影响夏季臭氧浓度水平和变化特征方面扮演着重要作用.通过对2008年夏季天津地面臭氧体积浓度和气象因素的相关分析,揭示高浓度臭氧发生时的典型气象特征,并初步建立了预测地面臭氧浓度的气象学方法.结果表明:影响臭氧浓度的主要气象因素是气温、相对湿度和风速、风向,当14时气温大于30℃,相对湿度低于60%,风向为偏西或偏南时,高浓度臭氧的发生概率较高.采用14时气温、相对湿度和风速等气象参数拟合臭氧体积浓度,效果良好.     

Periodically driven circulation near the shore of a lake

Solutions are found for a linear model of the circulation near the shore of a lake that is subject to two diurnal forcing mechanisms. The first is the day/night heating/cooling induced horizontal pressure gradient. The second is an unsteady surface stress modelling a sea breeze/gully wind pattern. The two forcing mechanisms can oppose or reinforce each other depending on their relative phase. The interplay of different dynamic balances at different times and locations in the domain lead to complex circulation patterns especially during the period of flow reversal.     

Effects of site-selection bias on estimates of biodiversity change

Estimates of biodiversity change are essential for the management and conservation of ecosystems. Accurate estimates rely on selecting representative sites, but monitoring often focuses on sites of special interest. How such site-selection biases influence estimates of biodiversity change is largely unknown. Site-selection bias potentially occurs across four major sources of biodiversity data, decreasing in likelihood from citizen science, museums, national park monitoring, and academic research. We defined site-selection bias as a preference for sites that are either densely populated (i.e., abundance bias) or species rich (i.e., richness bias). We simulated biodiversity change in a virtual landscape and tracked the observed biodiversity at a sampled site. The site was selected either randomly or with a site-selection bias. We used a simple spatially resolved, individual-based model to predict the movement or dispersal of individuals in and out of the chosen sampling site. Site-selection bias exaggerated estimates of biodiversity loss in sites selected with a bias by on average 300–400% compared with randomly selected sites. Based on our simulations, site-selection bias resulted in positive trends being estimated as negative trends: richness increase was estimated as 0.1 in randomly selected sites, whereas sites selected with a bias showed a richness change of −0.1 to −0.2 on average. Thus, site-selection bias may falsely indicate decreases in biodiversity. We varied sampling design and characteristics of the species and found that site-selection biases were strongest in short time series, for small grains, organisms with low dispersal ability, large regional species pools, and strong spatial aggregation. Based on these findings, to minimize site-selection bias, we recommend use of systematic site-selection schemes; maximizing sampling area; calculating biodiversity measures cumulatively across plots; and use of biodiversity measures that are less sensitive to rare species, such as the effective number of species. Awareness of the potential impact of site-selection bias is needed for biodiversity monitoring, the design of new studies on biodiversity change, and the interpretation of existing data.     

Responses of the whipscorpion, Mastigoproctus liochirus (Arachnida, Uropygi) to environmental humidity

Experiments were conducted to determine the responses of third-instar nymphs of the whipscorpion Mastigoproctus liochirus to various moisture (relative humidity) levels under constant temperature conditions, using a linear humidity gradient apparatus. No previous data exist on the water relations of this uropygid. Under saturated conditions (100% RH) animals showed no preference for any section of the chamber. When a humidity gradient was established (20 to 100% RH), whipscorpions exhibited a marked preference for an area of the chamber characterized by 70 to 80% RH.     

Laboratory investigation of <Emphasis Type="Italic">Fallopia</Emphasis> × <Emphasis Type="Italic">bohemica</Emphasis> fruits dispersal by watercourses

Seed and fruit dispersal along watercourses favours the long-distance migration of invasive species, not only for aquatic or wetland species, but also for terrestrial wind-dispersed plants, like the Japanese knotweed. The present paper aims at investigating the role of watercourses in the dispersal of the knotweed due to its frequent occurrence on riverbanks and production of fertile achenes (type of fruit of the Japanese knotweed). This dispersal occurs along two steps after the fruits deposit on the water surface: floatation first and then sinking towards the bottom of the watercourse. Regarding the first step, the effects of agitation of the water, temperature, surface tension and luminosity on the achenes floatability are experimentally studied. While no influence of luminosity is observed, an increase of temperature greatly decreases the floating time. Floating time also decreases as the contact between water and the fruit is enhanced (through submersion of achenes, agitation of the water or lower surface tension). Regarding the second step, the fall velocity of the fruits in water at rest is measured and appears to be independent of the seed history (floating time). 3D helical motions are systematically observed with constant tangential velocity with respect to the falling velocity. The trajectory of the fruits in a shear flow is then measured and the evolution of their velocity components along the sinking process is discussed. Finally, the contribution of both steps to the long-distance migration of the seeds is estimated.     

Experimental manipulation of seed shadows of an Afrotropical tree determines drivers of recruitment

The loss of animals in tropical forests may alter seed dispersal patterns and reduce seedling recruitment of tree species, but direct experimental evidence is scarce. We manipulated dispersal patterns of Manilkara mabokeensis, a monkey-dispersed tree, to assess the extent to which spatial distributions of seeds drive seedling recruitment. Based on the natural seed shadow, we created seed distributions with seeds deposited under the canopy ("no dispersal"), with declining density from the tree ("natural dispersal"), and at uniform densities ("good dispersal"). These distributions mimicked dispersal patterns that could occur with the extirpation of monkeys, low levels of hunting, and high rates of seed dispersal. We monitored seedling emergence and survival for 18 months and recorded the number of leaves and damage to leaves. "Good dispersal" increased seedling survival by 26%, and "no dispersal" decreased survival by 78%, relative to "natural dispersal." Using a mixed-effects survival model, we decoupled the distance and density components of the seed shadow: seedling survival depended on the seed density, but not on the distance from the tree. Although community seedling diversity tended to decrease with longer dispersal distances, we found no conclusive evidence that patterns of seed dispersal influence the diversity of the seedling community. Local seed dispersal does affect seedling recruitment and survival, with better dispersal resulting in higher seedling recruitment; hence the loss of dispersal services that comes with the reduction or extirpation of seed dispersers will decrease regeneration of some tree species.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

A method for building spatial model of annual weed seed dispersal from experimental data and its application to simulating Bromus sterilis population dispersal

Spatial model of annual weed seed dispersal, in this article, was theoretically derived. According to the requirements of building the spatial model, we designed and done an indoor experiment of weed seed dispersal by wind. In the experiment, the seeds of Bromus sterilis were released at 100 cm height under different wind velocity conditions. Based on the experimental data, the spatial models of seed dispersal of the weed species were built, which were divided into three types according to the coefficient β < 0, β = 0, β > 0. The results showed that dispersal of annual weed seed in any direction obeyed an approximate Gaussian distribution; under the experimental conditions, spatial distribution type of weed seed dispersal changed with variation of wind velocity. Well-known Howard et al.'s model (Howard et al., 1991) of Bromus sterilis seed dispersal is an especial example of the model built in this article. The result of model analysis indicated that the distribution type described by Howard's model was similar to that of seed dispersal of the weed species at the height of 100 cm under the condition of lower wind velocity (about 2.18 m/s). Using CA simulation analysis we found that mean control agent applying to a cell with weed should have a decrease with an increase of wind velocity to prevent weed with the initial configuration from spreading, which implicated less herbicide needs spraying in every cell with weed on average when wind velocity increases.     

Seed dispersal of desert annuals

We quantified seed dispersal in a guild of Sonoran Desert winter desert annuals at a protected natural field site in Tucson, Arizona, USA. Seed production was suppressed under shrub canopies, in the open areas between shrubs, or both by applying an herbicide prior to seed set in large, randomly assigned removal plots (10-30 m diameter). Seedlings were censused along transects crossing the reproductive suppression borders shortly after germination. Dispersal kernels were estimated for Pectocarya recurvata and Schismus barbatus from the change in seedling densities with distance from these borders via inverse modeling. Estimated dispersal distances were short, with most seeds traveling less than a meter. The adhesive seeds of P. recurvata went farther than the small S. barbatus seeds, which have no obvious dispersal adaptation. Seeds dispersed farther downslope than upslope and farther when dispersing into open areas than when dispersing into shrubs. Dispersal distances were short relative to the pattern of spatial heterogeneity created by the shrub and open space mosaic. This suggests that dispersal could contribute to local population buildup, possibly facilitating species coexistence. Overall, these results support the hypothesis that escape in time via delayed germination is likely to be more important for desert annuals than escape in space.     

Long-distance GM pollen movement of creeping bentgrass using modeled wind trajectory analysis.

The importance of understanding the role of atmospheric conditions in pollen dispersal has grown in recent years with increased field-testing of genetically modified (GM) crop plants. An atmospheric model was used to characterize wind trajectories at 10 m and 100 m above GM pollen source fields located within a 4452-ha "control" area north of Madras, Oregon, USA, designated by the Oregon Department of Agriculture (ODA). The area was used in 2003 for the growth of GM creeping bentgrass (Agrostis stolonifera) engineered to be resistant to glyphosate herbicide. The presence of the GM gene (CP4 EPSPS) provided a distinct selectable marker for pollen-mediated gene flow to sentinel and resident Agrostis spp. plants. Linkage of GM gene presence with wind flow characteristics over the "control" area became essential to understand the timing and processes leading to long-distance transport of this pollen. Wind trajectories showed a general pattern of northwest to southeast air movement. Trajectory travel distances calculated hourly from 06:00 hours to 15:00 hours during the 2003 pollination period (15 June-15 July) showed movement up to 15 km from the "control" area's center by the first hour. Maximum travel distances increased to 40 and 55 km after two and three hours from release, respectively. Calculated wind trajectory positions corresponded with observed long-distance pollen-mediated gene flow in the seedlings of sentinel and resident plants. The highest correlations were found during the late morning hours. Back-calculated wind trajectories from sentinel and resident locations with GM-gene-positive progeny suggested that most successful fertilizations occurred in the direction of prevailing winds during late June 2003. The occurrence of positive progeny from sentinel plants, upwind of the "control" area during this period, indicated the additional influence of local topography on pollen dispersal.     

Slow recovery of tropical old‐field rainforest regrowth and the value and limitations of active restoration

There is current debate about the potential for secondary regrowth to rescue tropical forests from an otherwise inevitable cascade of biodiversity loss due to land clearing and scant evidence to test how well active restoration may accelerate recovery. We used site chronosequences to compare developmental trajectories of vegetation between self‐organized (i.e., spontaneous) forest regrowth and biodiversity plantings (established for ecological restoration, with many locally native tree species at high density) in the Australian wet tropics uplands. Across 28 regrowth sites aged 1–59 years, some structural attributes reached reference rainforest levels within 40 years, whereas wood volume and most tested components of native plant species richness (classified by species’ origins, family, and ecological functions) reached less than 50% of reference rainforest values. Development of native tree and shrub richness was particularly slow among species that were wind dispersed or animal dispersed with large (>10 mm) seeds. Many species with animal‐dispersed seeds were from near‐basal evolutionary lineages that contribute to recognized World Heritage values of the study region. Faster recovery was recorded in 25 biodiversity plantings of 1–25 years in which wood volume developed more rapidly; native woody plant species richness reached values similar to reference rainforest and was better represented across all dispersal modes; and species from near‐basal plant families were better (although incompletely) represented. Plantings and regrowth showed slow recovery in species richness of vines and epiphytes and in overall resemblance to forest in species composition. Our results can inform decision making about when and where to invest in active restoration and provide strong evidence that protecting old‐growth forest is crucially important for sustaining tropical biodiversity.     

Seed terminal velocity, wind turbulence, and demography drive the spread of an invasive tree in an analytical model

Little is known about the relative importance of mechanistic drivers of plant spread, particularly when long-distance dispersal (LDD) events occur. Most methods to date approach LDD phenomenologically, and all mechanistic models, with one exception, have been implemented through simulation. Furthermore, the few recent mechanistically derived spread models have examined the relative role of different dispersal parameters using simulations, and a formal analytical approach has not yet been implemented. Here we incorporate an analytical mechanistic wind dispersal model (WALD) into a demographic matrix model within an analytical integrodifference equation spread model. We carry out analytical perturbation analysis on the combined model to determine the relative effects of dispersal and demographic traits and wind statistics on the spread of an invasive tree. Models are parameterized using data collected in situ and tested using independent data on historical spread. Predicted spread rates and direction match well the two historical phases of observed spread. Seed terminal velocity has the greatest potential influence on spread rate, and three wind properties (turbulence coefficient, mean horizontal wind speed, and standard deviation of vertical wind speed) are also important. Fecundity has marginal importance for spread rate, but juvenile survival and establishment are consistently important. This coupled empirical/theoretical framework enables prediction of plant spread rate and direction using fundamental dispersal and demographic parameters and identifies the traits and environmental conditions that facilitate spread. The development of an analytical perturbation analysis for a mechanistic spread model will enable multispecies comparative studies to be easily implemented in the future.