Running off the road: roadside non-native plants invading mountain vegetation

Prevention is regarded as a cost-effective management action to avoid unwanted impacts of non-native species. However, targeted prevention can be difficult if little is known about the traits of successfully invading non-native species or habitat characteristics that make native vegetation more resistant to invasion. Here, we surveyed mountain roads in seven regions worldwide, to investigate whether different species traits are beneficial during primary invasion (i.e. spread of non-native species along roadside dispersal corridors) and secondary invasion (i.e. percolation from roadsides into natural adjacent vegetation), and to determine if particular habitat characteristics increase biotic resistance to invasion. We found primary invasion up mountain roads tends to be by longer lived, non-ruderal species without seed dispersal traits. For secondary invasion, we demonstrate that both traits of the non-native species and attributes of the receiving natural vegetation contribute to the extent of invasion. Non-native species that invade natural adjacent vegetation tend to be shade and moisture tolerant. Furthermore, non-native species invasion was greater when the receiving vegetation was similarly rich in native species. Our results show how mountain roads define which non-native species are successful; first by favouring certain traits in mountain roadsides (the key dispersal pathway to the top), and secondly by requiring a different set of traits when species invade the natural adjacent vegetation. While patterns in species traits were observed at a global level, regional abiotic and biotic variables largely generated region-specific levels of response, suggesting that management should be regionally driven.     

云南南部和中部地区公路旁紫茎泽兰土壤种子库分布格局

公路边缘生境促进外来植物的入侵,土壤种子库作为植被天然更新的物质基础对未来植被的构成至关重要。研究路旁生境中紫茎泽兰土壤种子库的分布格局,对揭示路旁紫茎泽兰种群的补充和更新、探讨公路与入侵植物之间的关系以及科学制定紫茎泽兰防除对策具有重要意义。本文在紫茎泽兰入侵的云南南部和中部地区选择三种不同级别的9条公路的23个取样地点,沿垂直于公路方向设置55条样线,采得374个10 cm?10 cm?10 cm土样。采用温室萌发法,研究了紫茎泽兰土壤种子库从公路沿线到邻近景观的储量、影响因子以及分布格局。结果表明：9条公路旁的土壤中贮藏着丰富的紫茎泽兰种子,其种子库密度变动于3 152～25 225 粒?m-2,占所有有效种子密度的平均比例为48.7%。公路级别、路旁景观类型和海拔对路旁紫茎泽兰种子库密度有显著影响,密度随公路级别的提高而增加;不同路旁景观类型中的种子库密度排序为：稀树林>稀树灌木林>撂荒地和荒坡>森林;海拔1 700～1 900 m范围内的种子库密度最大。不同样线的种子库密度值随垂直于公路的距离变化格局略有不同,平均密度值和平均种子数量占样线总数的比例值都呈单峰变化,在距离公路最近端已经有很高密度,大多数样线的高峰值出现在9 m以内。公路旁已经分布有一个紫茎泽兰种子带,因而提高路旁本地植物的盖度及郁闭度有利于控制紫茎泽兰通过种子更新进一步扩散。     

A case study of human exacerbation of the invasive species problem: transport and establishment of polygyne fire ants in Tallahassee, Florida, USA

Understanding how exotic invasive species are spread is fundamental for ecology and conservation biology. Human transport has become one of the primary modes of dispersal for exotic species. We examined how the multiple queen, or polygyne, social form of the fire ant Solenopsis invicta is spread along roadsides in Tallahassee, Florida, USA. We then determined the likely source of this expanding population, which was a central soil depot. A survey of road maintenance practices in counties of several southeastern states and Texas revealed that the use of a central soil depot is a common practice. Road maintenance therefore may be the primary source for the establishment of new polygyne fire ant populations in this region and elsewhere. Control efforts focused on the soil depots will help to limit further spread of polygyne fire ants and perhaps other invasive organisms, particularly invasive weeds.     

A keystone ant species promotes seed dispersal in a “diffuse” mutualism

In order to understand the dynamics of co-evolution it is important to consider spatial variation in interaction dynamics. We examined the relative importance of ant activity, diversity and species identity in an ant seed dispersal mutualism at local, regional and continental scales. We also studied the determinants of seed dispersal rates and dispersal distances at eight sites in the Eneabba sandplain (29.63 S, 115.22 E), western Australia to understand local variation in seed dispersal rate and distance. To test the generality of the conclusions derived from the eight local sites, we established 16 sites along a 1650-km transect in western Australia, covering 11° of latitude and a six-fold increase in rainfall, at which we sampled the ant assemblage, estimated ant species richness and ant activity and observed the removal rate of myrmecochorous seeds. We also assessed the importance of ant species identity at a continental scale via a review of studies carried out throughout Australia which examined ant seed dispersal. Among the eight sandplain shrubland sites, ant species identity, in particular the presence of one genus, Rhytidoponera, was associated with the most dispersal and above average dispersal distances. At the landscape scale, Rhytidoponera presence was the most important determinant of seed removal rate, while seed removal rate was negatively correlated with ant species richness and latitude. Most ant seed removal studies carried out throughout Australia reinforce our observations that Rhytidoponera species were particularly important seed dispersers. It is suggested that superficially diffuse mutualisms may depend greatly on the identity of particular partners. Even at large biogeographic scales, temporal and spatial variation in what are considered to be diffuse mutualisms may often be linked to variation in the abundance of particular partners, and be only weakly – or negatively – associated with the diversity of partners.     

Sampling Bias in Roadsides: The Case of Galling Aphids on Pistacia Trees

Sampling along roadsides is convenient and is widely practiced in insect population researches. Ecological conditions in road verges are very different than those prevailing in natural habitats and they affect the annual growth of plants in semi-arid and arid regions. This in turn may improve development, survival and abundance of insects feeding on plants growing in roadsides. These trends may bias the results of sampling. To verify this assertion, we quantified the effects of growing in roadside on annual growth of Pistacia atlantica trees and Pistacia palaestina shrubs and compare two demographic indexes of nine gall-inducing aphid species on trees growing along roads with trees in the open landscape, in Israel. The annual growth of the two host plants was significantly more vigorous in roadsides than away from roads. Tests of Combined Probabilities showed that the likelihood of P. atlantica and P. palaestina to be parasitized by more galls of Fordini species is higher in roadsides than away from roads. Moreover, in the semi-dry regions of Israel, three aphid species on P. atlantica and five species on P. palaestina induced more galls in plants growing along roads than away from roads, while in the rainy Northern region, the difference was not significant between the two habitats. These results indicate a biased evaluation of population size in roadside habitat, which has to be accounted in insect–plant relation researches.     

Rapid continental spread of a salt-tolerant plant along the European road network

The spread of alien species with the expansion of road networks and increasing traffic is a well-known phenomenon globally. Besides their corridor effects, road maintenance practices, such as the use of de-icing salts during winter facilitate the spread of halophyte (salt tolerant) species along roads. A good example is Plantago coronopus, a mainly coastal halophyte which has started spreading inland from the Atlantic and Mediterranean coastal habitats, recently reaching even Central European countries (e.g. Hungary). Here we studied the spread of this halophyte and tried to identify factors explaining its successful dispersion along roads, while also comparing native and non-native roadside occurrences with regard to altitude of the localities, size of roadside populations and frequency of roadside occurrences. We completed a comprehensive literature review and collected more than 200 reports of occurrence from roadsides spanning a total of 38 years. During systematic sampling the frequency of the species along roads was significantly higher in the Mediterranean (native area), than along Hungarian (non-native area) roads, however the average number of individuals at the sampling localities were very similar, and no significant difference could be detected. Using a germination experiment, we demonstrate that although the species is able to germinate even at high salt concentrations, salt is not required for germination. Indeed salt significantly decreases germination probability of the seeds. The successful spread of the species could most likely be explained by its remarkably high seed production, or some special characteristics (e.g. seed dimorphism) and its ability to adapt to a wide range of environmental conditions. Considering the recent and rapid eastward spread of P. coronopus, occurrences in other countries where it has not been reported yet can be predicted in coming years.

     

Soil disturbance effects on the composition of seed-dispersing ants in roadside environments

Myrmecochory (the dispersal of seeds by ants) is a significant ecological process in sclerophyll woodlands, but habitat disturbance is known to alter the extent and success of this mutualism. We investigated the influence of soil disturbance on the composition of the seed-dispersing ant community. Surveys were conducted in roadside verges where soils are regularly disturbed by road maintenance activities. Using a ‘cafeteria’ bait station approach, we selected 24 roads of different widths to investigate ant composition and abundance in relation to soil disturbance. We found ant species richness was greater in non-disturbed than disturbed zones, where road verge width significantly influenced results. The composition and abundance of individual seed-dispersing ant species varied between disturbed and non-disturbed zones. Rhytidoponera metallica were more abundant in non-disturbed sites, whereas Melophorus bruneus and Monomorium rothseini were more frequently recorded in disturbed areas. Commonly found Iridomyrmex purpureus was significantly more abundant in disturbed zones in narrow roadsides and vice versa in wide roadsides, and strongly influenced total community composition. Variation in the abundance of commonly recorded Iridomyrmex and Monomorium genera were related more to site conditions (roadside width and habitat) than soil disturbance. The rich composition of seed dispersing ants in roadside environments, and the effects of soil disturbances on these ant communities that we describe, provide a key insight to important seed dispersal vectors occurring in fragmented rural landscapes.     

How did common ragweed (Ambrosia artemisiifolia L.) spread in Québec? A historical analysis using herbarium records

Aim To reconstruct the spread of common ragweed (Ambrosia artemisiifolia L.; Asteraceae) using herbarium specimens to document whether the habitat preferences of this plant have shifted through time. Location Southern Québec. Methods Herbarium specimens stored in the main herbaria of southern Québec were used to reconstruct the spread of common ragweed. All data (sampling location, year of sampling, habitat characteristics, etc.) were incorporated into a geographical information system. Maps indicating the spatial distribution of common ragweed were produced for four time periods. The cumulative number of locations was plotted against time to construct invasion curves. The sequence of habitats where herbarium specimens were collected was also reconstructed. Results A data base incorporating 707 common ragweed herbarium specimens was constructed for this study. The spread of common ragweed in most regions of southern Québec was initiated at the beginning of the 20th century. Herbarium specimens suggest that common ragweed first spread along river corridors. Specimens of common ragweed were not collected in agricultural fields before the mid‐1920s, nor along roads and railways before the mid‐1930s. The colonization of a large number of agricultural fields by common ragweed probably began with seed‐contaminated crops, but was certainly accelerated by the dispersal of seeds from populations growing along nearby roads. Main conclusions Herbarium specimens suggest that common ragweed has been present in southern Québec for at least 200 years, but the species was probably restricted to the Montréal area during the 19th century. It is likely that the development of the road network in Québec since the mid‐1930s significantly contributed to the spread of common ragweed. Controlling common ragweed solely in agricultural fields would not prevent the re‐infestation of crops, because roadsides would act as refuges for the weed.     

Green Frogs Show Reduced Foraging Success in Habitats Invaded by Japanese knotweed

Habitat loss is causing amphibian population declines worldwide, so there is increased attention to forces that degrade remaining habitats. Terrestrial habitats surrounding wetlands are critical foraging areas for temperate anurans. We investigated plant community changes in two old fields invaded by Japanese knotweed (Fallopia japonica) and the foraging success of Green frogs (Rana clamitans) in invaded and non-invaded portions of those fields. Within each field, vegetation data were recorded in quadrats located along two transects bisecting the invasion fronts. We placed frogs in ‘foraging buckets’ along transects and measured their change in mass over a 38 h period. There were significant changes in vegetation structure and composition associated with Japanese knotweed invasion. Diverse assemblages of native plants that covered non-invaded plots were absent from areas invaded by Japanese knotweed. There was also a significant change in vegetation architecture between invaded and non-invaded habitats. Change in frog mass declined significantly along transects, with most frogs in non-invaded plots gaining mass and no frogs in invaded plots gaining mass. Most frogs from non-invaded plots but only two from invaded plots defecated shortly after removal from foraging buckets (verification of recent feeding). We hypothesize that Japanese knotweed invasions degrade terrestrial habitat quality for frogs by indirectly reducing arthropod abundance. Nonnative plant invasions may be another factor contributing to amphibian population declines.     

Acorn dispersal estimated by radio-tracking

Bird-dispersed seeds are difficult to track, especially in the case of long-distance dispersal events. To estimate the oak dispersal distance and the seed shadow generated by the European jay (Garrulus glandarius), we inserted radio-transmitters in 239 acorns, placed them in bird-feeders and then located them by radio-tracking. Using this methodology we located the exact caching site of 94 Quercus ilex and 54 Q. suber acorns and determined the caching habitat characteristics (vegetation type, distance, spatial distribution). The results show that: (1) there is no differences in the dispersal distance distribution between the different acorn species or sizes, (2) dispersal distances range from approximately 3 m up to approximately 550 m (mean = 68.6 m; median = 49.2 m), (3) recently abandoned fields and forest tracks were the sites preferred by jays to cache acorns, whereas fields and shrublands were avoided and (4) seed shadows showed acorn aggregation zones (i.e. clusters of caches) close to the feeder as well as isolated caches at longer distances. The results also suggest that radio-transmitters are a cheap and reliable way to determine seed shadows and quantify both seed dispersal and post-dispersal seed predation for medium to large seeds.     

Habitat modification inhibits conspecific seedling recruitment in populations of an invasive ecosystem engineer

Invasive species that strongly modify their physical habitat are a particular management concern. Theoretical models predict that habitat modification could speed spread rates or allow invasion of sites that would otherwise resist invasion. There are few empirical tests of this hypothesis, however. We tested whether habitat modification by invading Spartina alterniflora populations facilitates conspecific seedling recruitment and spatial spread in Willapa Bay, WA, USA. Established S. alterniflora individuals strongly modified their local physical environment. Hydrologic flow, porewater salinity, and light availability were decreased while sediment NH₄ ⁺ increased with increasing S. alterniflora stem density. The S. alterniflora seed bank was greater and spring seedlings were denser within meadows of S. alterniflora than on unvegetated tideflats. However, almost all seedling recruitment after 1 year occurred on tideflats or on meadow edge plots where the above ground S. alterniflora biomass had been removed. Instead of facilitating invasive spread, ecosystem engineering in this system appears to create conditions that inhibit local seedling recruitment. These results suggest that the influence of ecosystem engineering on invasive spread is highly contingent on the relative spatial scales of habitat modification, environmental heterogeneity, and propagule availability. Control activities could change these spatial relationships, however, inadvertently promoting invasive recruitment.     

Human-mediated dispersal via rural road maintenance can move invasive propagules

Although preventing introduction is the best way to avoid invasive species problems, the dynamics of spread into new areas are not well understood. Invasive plant presence has frequently been associated with roads, but quantification of the mechanisms behind this phenomenon is lacking. Although some invasive plants may experience more rapid natural dispersal in a roadside habitat, scaling up this increase often cannot account for regional spread. Typically human activities drive long-distance dispersal, which is critical for predicting large-scale spread rates. We followed the movement of seeds by routine rural road maintenance as a potential factor explaining rapid regional spread of invasives. We found that 23.5% of seeds were not moved by road grading, 33.1% moved short distances (between 0 and 10 m), 41.8% moved intermediate distances (10–50 m), and very few moved long distances (more than 50 m, 1.6%). The furthest movement observed was 273 m. Nearly 1/3 of seeds were moved moderate distances; this intermediate distance dispersal is likely a key driver of the presence of invasives along the entirety of many roadsides and poses a large threat to the forest interior. Propagule spread via grading is a fairly stochastic process; nonetheless, a small but potentially important portion of propagules are moved long distances, accelerating spread rates. We discuss these results from the perspective of the rapid invasion of Microstegium vimineum in central Pennsylvania, USA. To slow the spread of seeds via road maintenance, managers should consider shortening grading passes, inspecting vehicles for invasive propagules, and otherwise ensuring that seeds are not transported long distances. Greater attention should be paid to human aided dispersal of invasive plant propagules particularly in light of an increasing forest roads network associated with logging and natural gas development in some parts of the United States.     

Edge Effects on Species Composition and Exotic Species Abundance in the North Carolina Piedmont

Edges between forest and non-forest habitats often have significant effects on forest microclimate and resource availability, with corresponding effects on species composition and abundance. Exotic species are often increased in abundance near forest edges. This increase in abundance could be either because of the increase in resource availability near edges, or because of increased dispersal into forest edges. We measured species composition and a set of geospatial variables on transects at 66 edges in the North Carolina Piedmont in an attempt to distinguish between these two factors. Mantel tests show that species composition is significantly different in forest edges than in the forest interior, but that this effect only penetrates about 5 m into the forest. Indicator species analysis finds several species that are indicative of edge communities, including trumpet vine (Campsis radicans), two drought-tolerant oak species (Quercus stellata and Q. falcata), a serviceberry (Amelanchier arboreum), and a common exotic species, tree-of-heaven (Ailanthus altissima). Poisson regression techniques showed that in both the seedling and tree strata of the forest, exotic species increased in abundance on flat sites with a high potential seed source. Mapping predicted exotic species abundance onto the landscape. We find that large-scale variation in exotic species abundance is due mostly to variation in potential seed sources, while small-scale variation relates more to edaphic factors. Our results stress that both dispersal and environmental filters are important for determining exotic species abundance, but potentially the filters operate at different spatial scales.     

Processes at multiple scales affect richness and similarity of non‐native plant species in mountains around the world

Aim To investigate how species richness and similarity of non‐native plants varies along gradients of elevation and human disturbance. Location Eight mountain regions on four continents and two oceanic islands. Methods We compared the distribution of non‐native plant species along roads in eight mountainous regions. Within each region, abundance of plant species was recorded at 41–84 sites along elevational gradients using 100‐m² plots located 0, 25 and 75 m from roadsides. We used mixed‐effects models to examine how local variation in species richness and similarity were affected by processes at three scales: among regions (global), along elevational gradients (regional) and with distance from the road (local). We used model selection and information criteria to choose best‐fit models of species richness along elevational gradients. We performed a hierarchical clustering of similarity to investigate human‐related factors and environmental filtering as potential drivers at the global scale. Results Species richness and similarity of non‐native plant species along elevational gradients were strongly influenced by factors operating at scales ranging from 100 m to 1000s of km. Non‐native species richness was highest in the New World regions, reflecting the effects of colonization from Europe. Similarity among regions was low and due mainly to certain Eurasian species, mostly native to temperate Europe, occurring in all New World regions. Elevation and distance from the road explained little of the variation in similarity. The elevational distribution of non‐native species richness varied, but was always greatest in the lower third of the range. In all regions, non‐native species richness declined away from roadsides. In three regions, this decline was steeper at higher elevations, and there was an interaction between distance and elevation. Main conclusions Because non‐native plant species are affected by processes operating at global, regional and local scales, a multi‐scale perspective is needed to understand their patterns of distribution. The processes involved include global dispersal, filtering along elevational gradients and differential establishment with distance from roadsides.     

New mutualism for old: indirect disruption and direct facilitation of seed dispersal following Argentine ant invasion

The indirect effects of biological invasions on native communities are poorly understood. Disruption of native ant communities following invasion by the Argentine ant (Linepithema humile) is widely reported to lead indirectly to the near complete collapse of seed dispersal services. In coastal scrub in southeastern Australia, we examined seed dispersal and handling of two native and two invasive alien plant species at Argentine ant-invaded or -uninvaded sites. The Argentine ant virtually eliminates the native keystone disperser Rhytidoponera victoriae, but seed dispersal did not collapse following invasion. Indeed, Argentine ants directly accounted for 92% of all ant-seed interactions and sustained overall seed dispersal rates. Nevertheless, dispersal quantity and quality among seed species differed between Argentine ant-invaded and -uninvaded sites. Argentine ants removed significantly fewer native Acacia retinodes seeds, but significantly more small seeds of invasive Polygala myrtifolia than did native ants at uninvaded sites. They also handled significantly more large seeds of A. sophorae, but rarely moved them >5 cm, instead recruiting en masse, consuming elaiosomes piecemeal and burying seeds in situ. In contrast, Argentine ants transported and interred P. myrtifolia seeds in their shallow nests. Experiments with artificial diaspores that varied in diaspore and elaiosome masses, but kept seed morphology and elaiosome quality constant, showed that removal by L. humile depended on the interaction of seed size and percentage elaiosome reward. Small diaspores were frequently taken, independent of high or low elaiosome reward, but large artificial diaspores with high reward instead elicited mass recruitment by Argentine ants and were rarely moved. Thus, Argentine ants appear to favour some diaspore types and reject others based largely on diaspore size and percentage reward. Such variability in response indirectly reduces native seed dispersal and can directly facilitate the spread of an invasive alien shrub.     

Geographical ecology and variation of plant-seed disperser interactions: southern Spanish junipers and frugivorous thrushes

Spatial and temporal predictability in the mutual selective pressures of plants and frugivorous birds is a prerequisite for coevolution to occur. I examine the interaction patterns of strongly frugivorous thrushes (Turdus spp.) and their major winter food plants (Juniperus spp., Cupressaceae) and how they vary in space and time. Spatial congruency, rarely considered in seed dispersal studies, is studied at three spatial scales: 1) the total species range; 2) regional distribution; and 3) local abundance and its variation between seasons. Southern Spanish frugivorous thrushes and junipers show very low congruence in distribution patterns at each of these scales. Most juniper species show geographic distributions that are nested within the geographic ranges of thrush species. Bird species showed greater habitat breadth values than plants and were found in a greater percentage of localities. The local bird abundance was strongly correlated across years and sites with the local availability of juniper cones. Cone production varied markedly between years, but the rankings for different species in different years were statistically concordant at mid-elevation and lowland sites. Both bird abundance and cone production showed greater temporal than spatial variability. Variation of cone productions at both temporal and spatial scales was greater than variability in bird abundance. Species with strong interactions of mutual dependence showed very low values of biogeographic congruence, caused by differences in geographic range and habitat specificity. This obviously limits the possibilities for pairwise, specific coevolution to occur. However, mutual effects of species groups are possible to the extent that the component species are ecologically interchangeable in their selective effects and other constraints on coevolution are not operating. The approach used here to examine the patterns of species interactions at different biogeographic scales might prove useful in comparative studies of plant-animal interactions.     

Long‐term persistence and spatial assortment of nonnative plant species in second‐growth forests

Long‐term conservation of forested areas requires an estimate of nonnative species’ impact on a scale of centuries. However, long‐term changes in forest structure have not been included in previous succession studies, which have typically covered only 20–50 yr. To estimate persistence of nonnative plant species on a longer time scale, and to examine the development of spatial structure in their distributions, we selected thirty‐seven second‐growth forest sites in southeastern Ohio, USA, to form a chronosequence spanning 160 yr. At each site nonnative species were surveyed in 100 contiguous plots forming a 50‐m belt transect. Nineteen nonnative species were encountered, largely consisting of shade‐intolerant herbs typical of abandoned agricultural land. Nonnative species richness, abundance, and frequency declined through the chronosequence with most species dropping out after the first twenty years, although three persisted 120–140 yr. Most species appeared to be residual populations from the open‐habitat stage. Presence of Rosa multiflora and Polygonum hydropiper in stands older than their date of arrival in the region indicate their capacity to colonize long‐established forest. Distributions of four species showed dependence on micro‐environmental gradients. All four were positively autocorrelated at scales up to 2–8 m, but pattern did not vary among age classes. We conclude that pattern in nonnative species’ distributions arises very early, possibly in the open‐habitat stage. Most species encountered in these sites are not likely to become long‐term components of the forest community and do not require active management. Small‐gap disturbances have not allowed colonization of long‐established forest stands, nor released pre‐existing populations from competitive suppression. Management should focus on the small number of shade‐tolerant species able to persist in long‐established forest.     

Spatial structure and genetic diversity of three tropical tree species with different habitat preferences within a natural forest

Analyses of the spatial distribution pattern, spatial genetic structure and genetic diversity were carried out using a 33-ha plot in a hill dipterocarp forest for three dipterocarps with different habitat preferences, i.e. Shorea curtisii on the ridges, Shorea leprosula in the valleys and Shorea macroptera both on the ridges and in the valleys. The significant spatial aggregation in small-diameter trees of all the three species was explained by limited seed dispersal. At the large-diameter trees, only S. macroptera showed random distribution and this might further prove that S. macroptera is habitat generalist, whilst S. curtisii and S. leprosula are habitat specific. The levels of genetic diversity estimated based on five microsatellite loci were high and comparable in all the three studied species. As the three studied species reproduced mainly through outcrossing, the observed high levels of genetic diversity might support the fact that the plant mating system can be used as guideline to infer the levels of genetic diversity, regardless of whether the species is habitat specific or habitat generalist. The lack of spatial genetic structure but significant aggregation in the small-diameter trees of all the three species might indicate limited seed dispersal but extensive pollen flow. Hence, if seed dispersal is restricted but pollen flow is extensive, significant spatial aggregation but no spatial genetic structure will be observed at the small-diameter trees, regardless of whether the species is habitat specific or habitat generalist. The inferred extensive pollen flow might indicate that energetic pollinators are involved in the pollination of Shorea species in the hill dipterocarp forests.     

Grassland connectivity by motor vehicles and grazing livestock

In addition to habitat loss and fragmentation, agricultural change has led to a change in seed dispersal processes in the rural landscape through a loss of structural and functional connectivity. Here, human‐mediated dispersal vectors are prevalent, and we explored whether the loss of connectivity via free‐ranging livestock could be mitigated by the increase in roads and motor vehicles. We found that structurally, 39% of all valuable semi‐natural grassland habitats in southern Sweden are adjacent to public road verges, which in the rural landscape are often considered to be suitable habitat for grassland species. Additionally, by collecting mud attached to cars and farming machinery and manure from livestock (cattle, horse, sheep) grazing semi‐natural grassland pasture, we found that motor vehicles are also capable seed dispersers. A similar number of species were dispersed by both vectors, although the composition of samples was quite different. Motor vehicles dispersed more grassland specialists than invasive species, although in much lower abundances than did grazing livestock. Despite these differences, motor vehicles were found to be able to disperse species with the same kinds of dispersal traits as livestock. A high number of seeds, species and specialists in manure samples means that greater movement of livestock is desirable to increase functional grassland connectivity. However, effective management could improve the suitability of roadsides as grassland corridors and increase the availability of seeds for long‐distance human‐mediated dispersal via cars and tractors. Our results suggest that in many rural landscapes, connectivity by road networks could help mediate habitat loss and fragmentation of grasslands. However, such effects can be context dependent, and the connectivity provided by roads could have serious negative consequences in other regions.