Seed dispersal of fleshy-fruited invasive plants by birds: contributing factors and management options

The ecology of seed dispersal by vertebrates has been investigated extensively over recent decades, yet only limited research has been conducted on how suites of invasive plants and frugivorous birds interact. In this review, we examine how plant fruit traits (morphology, colour and display, nutritional quality, accessibility and phenology), avian traits (fruit handling techniques, gut passage time and effect, bird movements and social behaviour and dietary composition) and landscape structure (fruit neighbourhood, habitat loss and fragmentation and perch tree effects) affect frugivory and seed dispersal in invasive plants. This functional approach could be used to develop generic models of seed dispersal distributions for suites of invasive plant species and improve management efficiencies. Four broad research approaches are described that could direct management of bird‐dispersed invasive plants at the landscape scale, by manipulating dispersal. First, research is needed to quantify the effect of biological control agents on dispersal, particularly how changes in fruit production and/or quality affect fruit choice by frugivores, dispersal distributions of seed and post‐dispersal processes. Second, we explore how seed dispersal could be directed, such as by manipulating perch structures and/or vegetation density to attract frugivorous birds after they have been foraging on invasive plant fruits. Third, the major sources of seed spread could be identified and removed (i.e. targeting core or satellite infestations, particular habitats and creating barrier zones). Fourth, alternative food resources could be provided for frugivores, to replace fruits of invasive plants, and their use quantified.     

Propagule dispersal and the scales of marine community process

Benthic marine organisms are characterized by a bipartite life history in which populations of sedentary adults are connected by oceanic transport of planktonic propagules. In contrast with the terrestrial case, where ‘long distance dispersal’ (LDD) has traditionally been viewed as a process involving rare events, this creates the possibility for large numbers of offspring to travel far relative to the spatial scale of adult populations. As a result, the concept of LDD must be examined carefully when applied in a marine context. Any measure of LDD requires reference to an explicit ‘local’ scale, often defined in terms of adult population demography, habitat patchiness, or the average dispersal distance. Terms such as ‘open’ and ‘closed’ are relative, and should be used with caution, especially when compared across different taxa and systems. We use recently synthesized data on marine propagule dispersal potential and the spread of marine invasive species to draw inferences about average and maximum effective dispersal distances for marine taxa. Foremost, our results indicate that dispersal occurs at a wide range of scales in marine communities. The nonrandom distribution of these scales among community members has implications for marine community dynamics, and for the implementation of marine conservation efforts. Second, in agreement with theoretical results, our data illustrate that average and extreme dispersal scales do not necessarily covary. This further confounds simple classifications of ‘short’ and ‘long’ dispersers, because different ecological processes (e.g. range expansion vs. population replenishment) depend on different aspects of the dispersal pattern (e.g. extremes vs. average). Our findings argue for a more rigorous quantitative view of scale in the study of marine dispersal processes, where relative terms such as ‘short’ and ‘long’, ‘open’ and ‘closed’, ‘retained’ and ‘exported’ are defined only in conjunction with explicit definitions of the scale and process of interest. This shift in perspective represents an important step towards unifying theoretical and empirical studies of dispersal processes in marine and terrestrial systems.     

Comparing the rate of invasion by Heracleum mantegazzianum at continental, regional, and local scales

This paper compares the rate of invasion of Heracleum mantegazzianum (Apiaceae), a Caucasian species invading Europe, at three spatial scales (continental, regional, and local). The rate of invasion was evaluated using inclusion curves, by plotting the cumulative number of invaded countries against time on the continental scale of Europe, number of occupied grid cells at the regional scale of the Czech Republic, and invaded area inferred from a series of aerial photographs taken at the local scale over a period of 49 years in the Slavkovký les region, Czech Republic. Time of 50% inclusion (with 95% confidence intervals, CI) of invaded countries, occupied grid cells, and invaded area was assessed. The invasion was slowest at the continental scale (62 years, CI = 53–70) and did not differ significantly between regional (16 years, CI = 10–20) and local (22 years, CI = 19–24) scales. Our results indicate that there are two different mechanisms of spread acting together in this system, namely human influences and natural spread, and the relative influence of these mechanisms appears to change in an inverse proportion from the largest to the smallest scale. At the local scale, under suitable habitat conditions, the process is driven by biological traits of the species related to dispersal. At the continental and regional scales, humans played a crucial role in the invasion of H. mantegazzianum by planting it as a garden ornamental. At these scales, human-mediated dispersal seems to have been the major driver of spread, responsible for creating dispersal foci in the initial phases of invasion. Species traits played an important role in local spread, resulting in the colonization of new sites.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

Combined modelling of distribution and niche in invasion biology: a case study of two invasive Tetramorium ant species

Spatial modelling of species distributions has become an important tool in the study of biological invasions. Here, we examine the utility of combining distribution and ecological niche modelling for retrieving information on invasion processes, based on species occurrence data from native and introduced ranges. Specifically, we discuss questions, concerning (1) the global potential to spread to other ranges, (2) the potential to spread within established invasions, (3) the detectability of niche differences across ranges, and (4) the ability to infer invasion history through data from the introduced range. We apply this approach to two congeneric pavement ants, Tetramorium sp.E (formerly T. caespitum (Linnaeus 1758)) and T. tsushimae Emery 1925, both introduced to North America. We identify (1) the potential of both species to inhabit ranges worldwide, and (2) the potential of T. sp.E and T. tsushimae, to spread to 23 additional US states and to five provinces of Canada, and to 24 additional US states and to one province of Canada, respectively. We confirm that (3) niche modelling can be an effective tool to detect niche shifts, identifying an increased width of T. sp.E and a decreased width of T. tsushimae following introduction, with potential changes in niche position for both species. We make feasible that (4) combined modelling could become an auxiliary tool to reconstruct invasion history, hypothesizing admixture following multiple introductions in North America for T. sp.E, and a single introduction to North America from central Japan, for T. tsushimae. Combined modelling represents a rapid means to formulate testable explanatory hypotheses on invasion patterns and helps approach a standard in predictive invasion research.     

Extreme long-distance dispersal of the lowland tropical rainforest tree Ceiba pentandra L. (Malvaceae) in Africa and the Neotropics

Many tropical tree species occupy continental expanses of rainforest and flank dispersal barriers such as oceans and mountains. The role of long-distance dispersal in establishing the range of such species is poorly understood. In this study, we test vicariance hypotheses for range disjunctions in the rainforest tree Ceiba pentandra, which is naturally widespread across equatorial Africa and the Neotropics. Approximate molecular clocks were applied to nuclear ribosomal [ITS (internal transcribed spacer)] and chloroplast (psbB-psbF) spacer DNA sampled from 12 Neotropical and five West African populations. The ITS (N=5) and psbB-psbF (N=2) haplotypes exhibited few nucleotide differences, and ITS and psbB-psbF haplotypes were shared by populations on both continents. The low levels of nucleotide divergence falsify vicariance explanations for transatlantic and cross-Andean range disjunctions. The study shows how extreme long-distance dispersal, via wind or marine currents, creates taxonomic similarities in the plant communities of Africa and the Neotropics.     

Towards a mechanistic understanding of dispersal evolution in plants: conservation implications

Aim A species’ dispersal characteristics will play a key role in determining its likely fate during a period of environmental change. However, these characteristics are not constant within a species – instead, there is often both considerable interpopulation and interindividual variability. Also changes in selection pressures can result in the evolution of dispersal characteristics, with knock‐on consequences for a species’ population dynamics. Our aim here is to make our theoretical understanding of dispersal evolution more conservation‐relevant by moving beyond the rather abstract, phenomenological models that have dominated the literature towards a more mechanism‐based approach. Methods We introduce a continuous‐space, individual‐based model for wind‐dispersed plants where release height is determined by an individual’s ‘genotype’. A mechanistic wind dispersal model is used to simulate seed dispersal. Selection acts on variation in release height that is generated through mutation. Results We confirm that, when habitat is fragmented, both evolutionary rescue and evolutionary suicide remain possible outcomes when a mechanistic dispersal model is used. We also demonstrate the potential for what we term evolutionary entrapment. A population that under some conditions can evolve to be sufficiently dispersive that it expands rapidly across a fragmented landscape can, under different conditions, become trapped by a combination of limited dispersal and a large gap between patches. Conclusions While developing evolutionary models to be used as conservation tools is undoubtedly a challenge, we believe that, with a concerted collaborative effort linking the knowledge and methods of ecologists, evolutionary biologists and geneticists, it is an achievable aim.     

Personality-dependent dispersal in the invasive mosquitofish: group composition matters

Understanding/predicting ecological invasions is an important challenge in modern ecology because of their immense economical and ecological costs. Recent studies have revealed that within-species variation in behaviour (i.e. animal personality) can shed light on the invasion process. The general hypothesis is that individuals' personality type may affect their colonization success, suggesting that some individuals might be better invaders than others. We have recently shown that, in the invasive mosquitofish (Gambusia affinis), social personality trait was an important indicator of dispersal distance, with more asocial individuals dispersing further. Here, we tested how mean personality within a population, in addition to individual personality type, affect dispersal and settlement decisions in the mosquitofish. We found that individual dispersal tendencies were influenced by the population's mean boldness and sociability score. For example, individuals from populations with more asocial individuals or with more bold individuals are more likely to disperse regardless of their own personality type. We suggest that identifying behavioural traits facilitating invasions, even at the group level, can thus have direct applications in pest management.     

Modelling population redistribution in a leaf beetle: an evaluation of alternative dispersal functions

1. Dispersal is a fundamental ecological process, so spatial models require realistic dispersal kernels. We compare five different forms for the dispersal kernel of the tansy beetle Chrysolina graminis moving between patches of its host-plant (tansy Tanacetum vulgare) in a riparian landscape. 2. Multi-patch mark-recapture data were collected every 2 weeks over 2 years within a large network of patches and from 2226 beetles. Dispersal was common (28.4% of 880 recaptures after a fortnight) and was more likely over longer intervals, out of small patches, for females and during flooding. Interpatch movement rates did not differ between years and exhibited no density dependence. Dispersal distances were similar for males and females, in both years and over all intervals, with a median dispersal distance of just 9.8 m, although a maximum of 856 m was recorded. 3. A model of dispersal, where patches competed for dispersers based on their size and distance from the beetle's source patch (scaled by the dispersal kernel) was fitted to the field data with a maximum likelihood procedure and each of five alternative kernels. The best fitting had relatively extended tails of long-distance dispersal, while Gaussian and negative exponential kernels performed worst. 4. The model suggests that females disperse more commonly than males and that both are strongly attracted to large patches but do not differ between years, which are consistent with the empirical results. Model-predicted emigration and immigration rates and dispersal phenologies match those observed, suggesting that the model captured the major drivers of tansy beetle dispersal. 5. Although negative exponential and Gaussian kernels are widely used for their simplicity, we suggest that these should not be the models of automatic choice, and that fat-tailed kernels with relatively higher proportions of long-distance dispersal may be more realistic.     

Impact of avian frugivores on dispersal and recruitment of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in an agricultural landscape

Although seed dispersal is considered to be a key process determining the spatial structure and spread of non-native plant populations, few studies have explicitly addressed the link between dispersal vector behaviour, seed distribution and seedling recruitment to gain insight into the process of exotic species invasion within a fragmented landscape context. The present study analyses the relationship between avian frugivory and spatial patterns of seed deposition and seedling recruitment for an expanding population of the invasive Prunus serotina in a hedgerow network landscape in Flanders, Belgium. We quantified fruit production, observed frugivores, and determined the spatial distribution of bird droppings and P. serotina seedlings. A relatively diverse assemblage of frugivores visited P. serotina seed trees, with Columba palumbus and Turdus merula being by far the most important dispersers. Landscape structure strongly affected dispersal vector behaviour and the spatial distribution of perching birds, droppings and seedlings. Frugivorous birds non-randomly dispersed seeds to perching sites and an association between perching birds, seed deposition and seedling recruitment was found. Results indicate that landscape structure contributes to non-random seed deposition of P. serotina by common local frugivores. Cutting the larger seed trees is proposed as the most feasible measure to slow down the invasion rate.     

Range expansion by invasive marine algae: rates and patterns of spread at a regional scale

Aim  Introduced macroalgae are widespread in the world's oceans and, despite increasing awareness and attempts to limit the phenomenon, the number of species introductions in coastal waters has increased exponentially over time. Little is known about the rates and mechanisms of spread, even among species that have received the most attention. We compare patterns of range expansion for nine species of invasive algae across eight geographic regions.
Location  World-wide.
Methods  We compiled records of introduced algae from the scientific literature, herbaria, and by contacting experts to reconstruct chronologies for 22 algal invasions. These were used to map patterns of spread at a regional scale (thousands of km).
Results  Range size tended to increase linearly with time, often after an initial lag. Range expansion occurred at rates of tens to hundreds of kilometres per year, often with large infrequent increases. Rates of range expansion differed significantly between species within the same region, and between regions for the same species.
Main conclusions  Our results suggest that anthropogenic vectors likely play a key role in the spread of introduced macroalgae at a regional scale, although natural long-distance dispersal also may be important for some species. The lack of consistency in the rates within individual species and regions suggests that multiple interacting factors (e.g. algal traits, characteristics of invaded communities, environmental conditions and anthropogenic activities) determine where propagules of introduced algae are delivered and whether they become established.     

Seed production,dispersal and germination in Cryptostegia grandiflora and Ziziphus mauritiana,two invasive shrubs in tropical woodlands of northern Australia

Abstract Cryptostegia grandiflora and Ziziphus mauritiana are exotic shrubs that are invading tropical woodlands of northern Australia. Although they are widespread, they currently occupy only small proportions of their respective potential ranges. Large C. grandiflora can produce more than 8000 wind-dispersed seeds in a single reproductive episode and can set seed at least twice per year. More than 90% of seeds will germinate within 10 days of moisture becoming available. Few, if any, seeds survive for more than 12 months in the soil. Large Z. mauritiana can produce more than 5000 fruits per year. The fruit is a drupe that contains a single seed (sometimes two) enclosed in a woody endocarp. Less than 10% of fresh seeds will germinate without removal of the endocarp. Removal of the endocarp increased germination to 56%. Less than 10% of seeds of Z. mauritiana remain viable after burial in the soil for 12 months. Seeds of Z. mauritiana are dispersed by several mammalian vectors, including wallabies and feral pigs, but domestic cattle are probably the major means by which large numbers of seeds reach new sites. Propagules pass intact through the digestive tract of cattle, feral pigs and wallabies and contain viable seeds that germinate more readily than seed in fresh intact fruits. Controlling movement of cattle that have had recent access to mature fruit of Z. mauritiana could significantly reduce the likelihood of new infestations. For both species, management that reduces seed production will be important for containing spread.     

Evolution of fruit characters and dispersal modes in the tropical family Rubiaceae

We investigated the evolution of fruit characters, animals versus abiotic dispersal modes, life forms and geographical distribution, in the large, mostly tropical, family Rubiaceae. As a basis for our analysis we used a phylogenetic tree derived from chloroplast DNA variation. Fleshy fruits have evolved independently at least 12 times in the family. Most of these originations appear to have occurred during Eocene to Oligocene, i.e. the radiation period for some animal taxa (bird families, mammal orders) comprising most extant dispersers of Rubiaceae fruits. Changes of dispersal modes may be of more recent origin in a few cases, e.g. evolution of drupes in some lineages, and shifts from drupes to nuts. The distribution of fruit characters suggested that in several lineages animal-dispersed fruits, such as berries and drupes have remained largely unaltered since the time of origination. This is in contrast to the occurrence of winged seeds in capsules, and pterophylls, i.e. enlarged calyx lobes promoting wind dispersal of fruits, which apparently have shifted more frequently during evolution, indicating a difference in 'phylogenetic plasticity' between modes of animal and wind dispersal.
Animal dispersal was over-represented among genera dominated by shrubs, whereas abiotic dispersal was most prevalent among herbaceous genera. Drupes were over-represented in groups with transoceanic distributions, and on islands, indicating dispersal over long distances, probably by birds. In contrast, no evidence was found to support the view that animal dispersal in general enhances long distance dispersal. We also analysed geographical patterns on the tribal level but these were too complex to yield any resolved area cladograms due to the occurrence of many widespread taxa and area redundancy.     

Surveillance protocols for management of invasive plants: modelling Chilean needle grass (Nassella neesiana) in Australia

Aim To develop a surveillance support model that enables prediction of areas susceptible to invasion, comparative analysis of surveillance methods and intensity and assessment of eradication feasibility. To apply the model to identify surveillance protocols for generalized invasion scenarios and for evaluating surveillance and control for a context‐specific plant invasion. Location Australia. Methods We integrate a spatially explicit simulation model, including plant demography and dispersal vectors, within a Geographical Information System. We use the model to identify effective surveillance protocols using simulations of generalized plant life‐forms spreading via different dispersal mechanisms in real landscapes. We then parameterize the surveillance support model for Chilean needle grass [CNG; Nassella neesiana (Trin. & Rupr.) Barkworth], a highly invasive tussock grass, which is an eradication target in south‐eastern Queensland, Australia. Results General surveillance protocols that can guide rapid response surveillance were identified; suitable habitat that is susceptible to invasion through particular dispersal syndromes should be targeted for surveillance using an adaptive seek‐and‐destroy method. The search radius of the adaptive method should be based on maximum expected dispersal distances. Protocols were used to define a surveillance strategy for CNG, but simulations indicated that despite effective and targeted surveillance, eradication is implausible at current intensities. Main conclusions Several important surveillance protocols emerged and simulations indicated that effectiveness can be increased if they are followed in rapid response surveillance. If sufficient data are available, the surveillance support model should be parameterized to target areas susceptible to invasion and determine whether surveillance is effective and eradication is feasible. We discovered that for CNG, regardless of a carefully designed surveillance strategy, eradication is implausible at current intensities of surveillance and control and these efforts should be doubled if they are to be successful. This is crucial information in the face of environmentally and economically damaging invasive species and large, expensive and potentially ineffective control programmes.     

Going against the flow: a case for upstream dispersal and detection of uncommon dispersal events

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