Mitochondrial DNA variation in North American populations of Daphnia obtusa: continentalism or cryptic endemism?

The morphological stasis of many freshwater crustaceans has resulted in the prior delineation of cosmopolitan species and has been explained by their capacity for long-distance dispersal. This study examines the phylogeography of Daphnia obtusa, a cladoceran thought to be widespread in North America. However, sequence variation of the mitochondrial cytochrome c oxidase subunit I gene indicates that this taxon is composed of two morphologically cryptic species, designated D. obtusa NA1 and NA2. NA2 is restricted to the east, whereas NA1 is broadly distributed across the United States, and is subdivided into four phylogroups that show weak genetic differentiation over broad geographical areas, which likely reflects recent long-distance dispersal. The current distributions of the four phylogroups in NA1 can be explained by recent range expansion from different refugia following the last Pleistocene glacial advance. Interestingly, the mitochondrial phylogroups identified in this study do not correspond to lineages detected in a previous allozyme analysis. However, the latter groups are associated with a habitat shift suggesting that natural selection may have played a role in their divergence. The results of this and previous studies illustrate the complicated biogeographical history of freshwater cladocerans.     

Signatures of seaway closures and founder dispersal in the phylogeny of a circumglobally distributed seahorse lineage

Background  

The importance of vicariance events on the establishment of phylogeographic patterns in the marine environment is well documented, and generally accepted as an important cause of cladogenesis. Founder dispersal (i.e. long-distance dispersal followed by founder effect speciation) is also frequently invoked as a cause of genetic divergence among lineages, but its role has long been challenged by vicariance biogeographers. Founder dispersal is likely to be common in species that colonize remote habitats by means of rafting (e.g. seahorses), as long-distance dispersal events are likely to be rare and subsequent additional recruitment from the source habitat is unlikely. In the present study, the relative importance of vicariance and founder dispersal as causes of cladogenesis in a circumglobally distributed seahorse lineage was investigated using molecular dating. A phylogeny was reconstructed using sequence data from mitochondrial and nuclear markers, and the well-documented closure of the Central American seaway was used as a primary calibration point to test whether other bifurcations in the phylogeny could also have been the result of vicariance events. The feasibility of three other vicariance events was explored: a) the closure of the Indonesian Seaway, resulting in sister lineages associated with the Indian Ocean and West Pacific, respectively; b) the closure of the Tethyan Seaway, resulting in sister lineages associated with the Indo-Pacific and Atlantic Ocean, respectively, and c) continental break-up during the Mesozoic followed by spreading of the Atlantic Ocean, resulting in pairs of lineages with amphi-Atlantic distribution patterns.     

The possible significance of long-range dispersal for the biogeography of seaweeds

Indirect evidence of the existence of long-distance dispersal of seaweeds is provided by the fact that mid-oceanic islands of volcanic origin are inhabited by well-developed seaweed floras which could reach these islands only overseas from continental donor areas. For instance, the flora of Tristan da Cunha (S. Atlantic Ocean) was established by long-distance dispersal in less than 1 million years (the approximate age of the island); the seaweed flora of the Faeroes (N. Atlantic Ocean) could be constituted in less than 10,000 years (the end of the Pleistocene ice cover of these islands). There is no evidence for either supporting or discounting the possible role of planktonic stages of seaweeds (spores, propagules, zygotes) in the long-distance dispersal of seaweeds. There is, however, some evidence of long-distance dispersal as floating plants, or as plants attached to floating objects (including floating algae). There are a few examples of “artificial” long-range dispersal by man (possibly on ship hulls, oysters, in ballast water). Long-range dispersal of seaweeds does exist, but it is an exception rather than the rule. If it were the rule, the world’s seaweed floras would show similar latitudinal gradients in species composition in the oceans and on both hemispheres. This is, however, not the case.     

Pushing the Pace of Tree Species Migration

Plants and animals have responded to past climate changes by migrating with habitable environments, sometimes shifting the boundaries of their geographic ranges by tens of kilometers per year or more. Species migrating in response to present climate conditions, however, must contend with landscapes fragmented by anthropogenic disturbance. We consider this problem in the context of wind-dispersed tree species. Mechanisms of long-distance seed dispersal make these species capable of rapid migration rates. Models of species-front migration suggest that even tree species with the capacity for long-distance dispersal will be unable to keep pace with future spatial changes in temperature gradients, exclusive of habitat fragmentation effects. Here we present a numerical model that captures the salient dynamics of migration by long-distance dispersal for a generic tree species. We then use the model to explore the possible effects of assisted colonization within a fragmented landscape under a simulated tree-planting scheme. Our results suggest that an assisted-colonization program could accelerate species-front migration rates enough to match the speed of climate change, but such a program would involve an environmental-sustainability intervention at a massive scale.     

Human-mediated dispersal of seeds over long distances

Human activities have fundamental impacts on the distribution of species through altered land use, but also directly by dispersal of propagules. Rare long-distance dispersal events have a disproportionate importance for the spread of species including invasions. While it is widely accepted that humans may act as vectors of long-distance dispersal, there are few studies that quantify this process. We studied in detail a mechanism of human-mediated dispersal (HMD). For two plant species we measured, over a wide range of distances, how many seeds are carried by humans on shoes. While over half of the seeds fell off within 5m, seeds were regularly still attached to shoes after 5 km. Semi-mechanistic models were fitted, and these suggested that long-distance dispersal on shoes is facilitated by decreasing seed detachment probability with distance. Mechanistic modelling showed that the primary vector, wind, was less important as an agent of long-distance dispersal, dispersing seeds less than 250 m. Full dispersal kernels were derived by combining the models for primary dispersal by wind and secondary dispersal by humans. These suggest that walking humans can disperse seeds to very long distances, up to at least 10 km, and provide some of the first quantified dispersal kernels for HMD.     

Modelling effects of habitat fragmentation on the ability of trees to respond to climatic warming

The ability of trees to migrate in response to climatic warming was simulated under various conditions of habitat availability. The model uses Holocene tree migration rates to approximate maximum migration rates in a forested landscape. Habitat availability and local population size was varied systematically under two dispersal and colonization models. These dispersal models varied in the likelihood of long-distance dispersal events. The first model used a negative exponential function that severely limited the probability of long-distance dispersal. The results of this model indicate that migration rate could decline an order of magnitude where the habitat availability is reduced from 80 to 20% of the matrix. The second model, using an inverse power function, carried a higher probability of long-distance dispersal events. The results from this model predict relatively small declines in migration rates when habitat availability is reduced to 50% of the simulation matrix. Below 50% habitat availability, mean migration rate was similar to the negative exponential model. These results predict a failure of many trees to respond to future climatic change through range expansion.     

Phylogeography of the pine processionary moth Thaumetopoea wilkinsoni in the Near East

Phylogeographic structure of the eastern pine processionary moth Thaumetopoea wilkinsoni was explored in this study by means of nested clade phylogeographic analyses of COI and COII sequences of mitochondrial DNA and Bayesian estimates of divergence times. Intraspecific relationships were inferred and hypotheses tested to understand historical spread patterns and spatial distribution of genetic variation. Analyses revealed that all T. wilkinsoni sequences were structured in three clades, which were associated with two major biogeographic events, the colonization of the island of Cyprus and the separation of southwestern and southeastern Anatolia during the Pleistocene. Genetic variation in populations of T. wilkinsoni was also investigated using amplified fragment length polymorphisms and four microsatellite loci. Contrasting nuclear with mitochondrial data revealed recurrent gene flow between Cyprus and the mainland, related to the long-distance male dispersal. In addition, a reduction in genetic variability was observed at both mitochondrial and nuclear markers at the expanding boundary of the range, consistent with a recent origin of these populations, founded by few individuals expanding from nearby localities. In contrast, several populations fixed for one single mitochondrial haplotype showed no reduction in nuclear variability, a pattern that can be explained by recurrent male gene flow or selective sweeps at the mitochondrial level. The use of both mitochondrial and nuclear markers was essential in understanding the spread patterns and the population genetic structure of T. wilkinsoni, and is recommended to study colonizing species characterized by sex-biased dispersal.     

A discrete space model for continuous space dispersal processes

The simulation of dispersal processes in landscapes over large spatial extents is challenging because of the large difference in geographical scale between overwhelmingly dominant localised dispersal events, and rare long-distance dispersal events which typically drive overall rates of spread. While localised dispersal may point to high resolution individual level models, long-distance dispersal events are likely to involve much coarser grid-based models. In this paper we propose a discrete space (i.e., grid-based) model for dispersal processes in continuous space. We start by illustrating the behaviour of continuous space walks when their movement is discretised to a grid. The importance of short time period cell-to-cell moves which return a walk to its previous grid cell location is identified. A conceptual model which uses a Markov chain buffer phase between cells to replicate the observed behaviour of discretised continuous space walks is proposed. Analysis of the Markov chain shows that it can be parameterised using just two parameters in addition to the dispersal kernel. An algorithm for implementation of the proposed model is presented. Empirical results demonstrate that the proposed mechanism produces good matches to continuous space dispersal processes with both exponential and heavy-tailed dispersal kernels.     

Aerial Dispersal and Multiple-Scale Spread of Epidemic Disease

Disease spread has traditionally been described as a traveling wave of constant velocity. However, aerially dispersed pathogens capable of long-distance dispersal often have dispersal gradients with extended tails that could result in acceleration of the epidemic front. We evaluated empirical data with a simple model of disease spread that incorporates logistic growth in time with an inverse power function for dispersal. The scale invariance of the power law dispersal function implies its applicability at any spatial scale; indeed, the model successfully described epidemics ranging over six orders of magnitude, from experimental field plots to continental-scale epidemics of both plant and animal diseases. The distance traveled by epidemic fronts approximately doubled per unit time, velocity increased linearly with distance (slope ~½), and the exponent of the inverse power law was approximately 2. We found that it also may be possible to scale epidemics to account for initial outbreak focus size and the frequency of susceptible hosts. These relationships improve understanding of the geographic spread of emerging diseases, and facilitate the development of methods for predicting and preventing epidemics of plants, animals, and humans caused by pathogens that are capable of long-distance dispersal.     

Detecting dispersal of Nuphar lutea in river corridors using microsatellite markers

1. River corridors are well-known for their role in plant dispersal. The buoyancy of seeds, the possibility of dispersal by vegetative fragments, and the frequency and efficiency of dispersal among different river catchments determine linear distribution patterns. Little is known about the relative importance of these factors to observed patterns of genetic variation.
2. One hundred and fifty-six Nuphar lutea individuals from forty-four sampling sites in the river catchments of the Cidlina River, the Mrlina River and the Labe River (Czech Republic) were studied using ten microsatellite markers. Interpretation of patterns in genetic variation allowed several conclusions about dispersal mechanisms.
3. Vegetative long-distance dispersal is probably very limited in this species. Only one multilocus genotype was found in more than one sampling site. The distance between the sites was about 75 km.
4. To explain the distribution of Bayesian based clusters of related multilocus genotypes, both along-river and inter-river long-distance dispersals have to be invoked.
5. A marginally significant tendency for higher genetic diversity in the lower part of the river Cidlina was detected. Continuous downstream dispersal of seeds by water currents could be a valid explanation.
6. Significant positive autocorrelation was found among individuals at within-river distances of up to 25 km. Repeated dispersal of seeds over distances in the range of tens of kilometers is common.     

Seasonal and Diurnal Patterns of Spore Release Can Significantly Affect the Proportion of Spores Expected to Undergo Long-Distance Dispersal

Many of the fungal pathogens that threaten agricultural and natural systems undergo wind-assisted dispersal. During turbulent wind conditions, long-distance dispersal can occur, and airborne spores are carried over distances greater than the mean. The occurrence of long-distance dispersal is an important ecological process, as it can drastically increase the extent to which pathogen epidemics spread across a landscape, result in rapid transmission of disease to previously uninfected areas, and influence the spatial structure of pathogen populations in fragmented landscapes. Since the timing of spore release determines the wind conditions that prevail over a dispersal event, this timing is likely to affect the probability of long-distance dispersal occurring. Using a Lagrangian stochastic model, we test the effect of seasonal and diurnal variation in the release of spores on wind-assisted dispersal. Spores released during the hottest part of the day are shown to be more likely to undergo long-distance dispersal than those released at other times. Furthermore, interactions are shown to occur between seasonal and diurnal patterns of release. These results have important consequences for further modelling of wind-assisted dispersal and the use of models to predict the spread of fungal pathogens and resulting population and epidemic dynamics.     

High diversity of oilseed rape pollen clouds over an agro-ecosystem indicates long-distance dispersal

Abstract Estimating the frequency of long-distance pollination is important in cultivated species, particularly to assess the risk of gene transfer following the release of genetically modified crops. For this purpose, we estimated the diversity and origin of fertilizing pollen in a 10 x 10 km French oilseed rape production area. First, the cultivar grown in each field was identified through surveys to farmers and using microsatellite markers. Examination of the seed set in fields indicated high rates of seed contamination (8.7%) and pollination from other sources (5%). Then, male-sterile plants were scattered over the study area and their seed genotyped using the same markers. Most pollination was local: 65% of the seeds had a compatible sire in the closest field, i.e. at 50 or 300 m depending on site, but the nearest compatible field was found more than 1000 m away for 13% of the seeds. To assess the diversity of fertilizing pollen, each seed was assigned to the nearest putative siring cultivar. The observed diversity of pollen was then compared to that predicted by simulations using three empirical dispersal models with increasing proportion of long-distance pollination. The diversity was sensitive to the dispersal kernel used in the simulations, fatter-tailed functions predicting higher diversities. The dispersal kernel that was more consistent with our data predicted more long-distance dispersal than the exponential function.     

Long-distance seed dispersal in plant populations

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.     

Measuring and modelling anthropogenic secondary seed dispersal along roadverges for feral oilseed rape

Seed dispersal of feral crop plants along roadverges is likely to be influenced by numerous anthropogenic vectors in the agroecosystem. Within the context of introducing genetically modified (GM) cultivars, long-distance dispersal of feral seeds associated with the growth of GM feral populations (via a selective advantage due to transgene expression) could make these populations become invasive. Their expansion could then favour the spread of transgenes and modify the composition of roadverge plant communities. Because quantitative data on anthropogenic seed dispersal along roadverges were few, we estimated effective secondary dispersal for oilseed rape, the seeds of which are not adapted to dispersal by wind or biotic agents. A seed deposition experiment showed that secondary dispersal did not systematically occur along roadverges, was correlated with traffic intensity and was local. Low traffic intensity and anthropogenic disturbances (covering of seeds by mown grasses and burial by farming machinery) prevented dispersal on three of the experimental sites. Along a road with higher traffic, secondary dispersal occurred (d_max=21.5 m), probably induced by wind turbulence behind vehicles. The best-fitting dispersal kernel was a mixture of two components: 20% of seeds dispersing over a few metres on average and 80% remaining in the original place. Expansion rates of feral populations of GM herbicide-tolerant oilseed rape were computed using an invasion model and this kernel. They were low (1–4 m yr⁻¹) when only ballistic and/or secondary dispersal were included but higher (4–20 m yr⁻¹) when theoretically rare events of long-distance dispersal by verge mowers were added. This study suggests that secondary seed dispersal is unlikely to have a significant impact on the spread of GM feral oilseed rape populations in highly disturbed and dynamic habitats such as roadverges. Detecting long-distance dispersal events induced by other vectors (e.g. mowers) would require integrative approaches based on genetic and spatial data.     

Modelling the impact of colonisation on genetic diversity and differentiation of forest trees: interaction of life cycle,pollen flow and seed long-distance dispersal

It was shown previously that the long lifespan and juvenile phase of trees strongly attenuate founder effects during colonisation in a diffusive dispersal model. However, this model yielded too slow a colonisation rate in comparison with palynological data for temperate forest trees. Since rare long-distance dispersal events have been shown to increase considerably colonisation rates in population dynamics models, we investigate here the impact of long-distance dispersal on within-population diversity (H(S)) and among-population differentiation (F(ST)) during the colonisation process. We use a stochastic approach and compare several dispersal strategies, ranging from very rare dispersal events of large amplitude to more frequent events of smaller amplitude. Using a simulation approach, which takes into account tree life-history traits, we show that long-distance dispersal events increase colonisation speed, and yield much larger founder effects in comparison with the diffusive model. The two models that include intermediate- and long-distance dispersal events show stronger deviations from experimental F(ST) values during and at the end of the colonisation process than the model with more frequent events of smaller dispersal variance. Furthermore, the introduction of a high level of pollen flow has a much more limited impact on models that include long-distance dispersal than on a diffusive dispersal model. The relatively high H(S) values that were obtained in all models are discussed according to the assumed mutation rate and effective population size. This study is an example of how observed genetic data can provide additional evidence on the best demographic model for a given species or group of species.     

Genetic structure and reproduction dynamics of Salix reinii during primary succession on Mount Fuji,as revealed by nuclear and chloroplast microsatellite analysis

The early stage of volcanic desert succession is underway on the southeastern slope of Mount Fuji. We used markers of nuclear microsatellites (simple sequence repeats; SSR) and chloroplast microsatellites (cpSSR) to investigate the population genetic structure and reproduction dynamics of Salix reinii, one of the dominant pioneer shrubs in this area. The number of S. reinii genets in a patch and the area of the largest genet within the patch increased with patch area, suggesting that both clonal growth and seedling recruitment are involved in the reproduction dynamics of S. reinii. Five polymorphic cpSSR markers were developed for S. reinii by sequencing the noncoding regions between universal sequences in the chloroplast genome. Nineteen different cpSSR haplotypes were identified, indicating that S. reinii pioneer genets were created by the long-distance dispersal of seeds originating from different mother genets around the study site, where all vegetation was destroyed during the last eruption. Furthermore, the clustered distributions of different haplotypes within each patch or plot suggested that newly colonized genets tended to be generated from seeds dispersed near the initially established mother genets. These results revealed that the establishment of the S. reinii population on the southeastern slope of Mount Fuji involved two sequential modes of seed dispersal: long-distance dispersal followed by short-distance dispersal.     

Serological investigation of Theileria sergenti using latex agglutination test in South Korea

Theileria sergenti causes persistent theileriosis in cattle, characterized by fever and chronic anemia. Theileriosis causes losses in feed efficiency and growth retardation through cycling infections in endemic areas. Among several major proteins of T. sergenti merozoites, the surface protein p33 is reported to be the most immunogenic. In this study, we investigated the use of p33 as a diagnostic antigen in a latex agglutination test to monitor antibodies against T. sergenti. When compared with TaqMan polymerase chain reaction, the sensitivity and specificity of the latex agglutination test were 86.5 and 92.5%, respectively. An epidemiological survey using the latex agglutination test was conducted with 1,046 sera collected from 4 slaughterhouses and 2 individual pasture farms throughout South Korea; 27.3% of samples were seropositive, depending on the areas in which the cattle were raised. This study indicated that the latex agglutination test could be used as a convenient tool for epidemiological monitoring of T. sergenti infections in the field.     

Mitochondrial DNA and microsatellite variation in the eider duck (Somateria mollissima) indicate stepwise postglacial colonization of Europe and limited current long-distance dispersal

To unravel the postglacial colonization history and the current intercolony dispersal in the common eider, Somateria mollissima, we analysed genetic variation at a part of the mitochondrial control region and five unlinked autosomal microsatellite loci in 175 eiders from 11 breeding colonies, covering the entire European distribution range of this species. As a result of extreme female philopatry, mitochondrial DNA differentiation is substantial both among local colonies and among distant geographical regions. Our study further corroborates the previous hypothesis of a single Pleistocene refugium for European eiders. A nested clade analysis on mitochondrial haplotypes suggests that (i) the Baltic Sea eider population is genetically closest to a presumably ancestral population and that (ii) the postglacial recolonization progressed in a stepwise fashion via the North Sea region and the Faroe Islands to Iceland. Current long-distance dispersal is limited. Differentiation among colonies is much less pronounced at microsatellite loci. The geographical pattern of this nuclear genetic variation is to a large extent explained by isolation by distance. As female dispersal is very limited, the geographical pattern of nuclear variation is probably explained by male-mediated gene flow among breeding colonies. Our study provides genetic evidence for the assumed prominent postglacial colonization route shaping the present terrestrial fauna of the North Atlantic islands Iceland and the Faroes. It suggests that this colonization had been a stepwise process originating in continental Europe. It is the first molecular study on eider duck populations covering their entire European distribution range.     

Synoptic climatology of the long-distance dispersal of white pine blister rust II. Combination of surface and upper-level conditions

An invasive forest pathogen, Cronartium ribicola, white pine blister rust (WPBR), is believed to have arrived in the Sacramento Mountains of south-central New Mexico about 1970. Epidemiological and genetic evidence supports the hypothesis that introduction was the result of long-distance dispersal (LDD) by atmospheric transport from California. This study applies a method to identify the atmospheric conditions favorable for rust transport and infection. An upper level synoptic classification (ULSC) identifies patterns of upper-level flow favorable for the transport of rust spores from a source to a target. Transport data are coupled with data for surface conditions favorable for infection at a designated target. A resulting calendar lists likelihood classes for establishment by four-times-daily observations during a dispersal season from April through July in the years 1965 to 1974. The single most-favorable period for transport and infection at the New Mexico site was identified as 1-15 June 1969. Five additional sites in the western United States with susceptible white pine populations and known infestation status were then evaluated to verify the model. Only the infested sites exhibit an establishment likelihood of "high" or "very high." This suggests that the methodology correctly identifies locations with elevated establishment likelihood. Finally, likelihoods at nine additional points in the southwestern United States are determined and used to map regional patterns of transport, infection and establishment. The ULSC combined with appropriate surface meteorological data could be used to further investigate transport and infection, identify other areas at risk, assess the potential for gene flow of WPBR and evaluate long-distance dispersal of other pathogens.     

Genetic structure and inferences on potential source areas for Bactrocera dorsalis (Hendel) based on mitochondrial and microsatellite markers

Bactrocera dorsalis (Diptera: Tephritidae) is mainly distributed in tropical and subtropical Asia and in the Pacific region. Despite its economic importance, very few studies have addressed the question of the wide genetic structure and potential source area of this species. This pilot study attempts to infer the native region of this pest and its colonization pathways in Asia. Combining mitochondrial and microsatellite markers, we evaluated the level of genetic diversity, genetic structure, and the gene flow among fly populations collected across Southeast Asia and China. A complex and significant genetic structure corresponding to the geographic pattern was found with both types of molecular markers. However, the genetic structure found was rather weak in both cases, and no pattern of isolation by distance was identified. Multiple long-distance dispersal events and miscellaneous host selection by this species may explain the results. These complex patterns may have been influenced by human-mediated transportation of the pest from one area to another and the complex topography of the study region. For both mitochondrial and microsatellite data, no signs of bottleneck or founder events could be identified. Nonetheless, maximal genetic diversity was observed in Myanmar, Vietnam and Guangdong (China) and asymmetric migration patterns were found. These results provide indirect evidence that the tropical regions of Southeast Asia and southern coast of China may be considered as the native range of the species and the population expansion is northward. Yunnan (China) is a contact zone that has been colonized from different sources. Regions along the southern coast of Vietnam and China probably served to colonize mainly the southern region of China. Southern coastal regions of China may also have colonized central parts of China and of central Yunnan.