Expansion Front Research Articles

Bycatch in local fishery disrupts natural reestablishment of Eurasian otter in western Norway

AbstractLack of proper estimates of nonreported bycatch has made it difficult to evaluate the actual impact of bycatch in many local fisheries. For the Eurasian otter (Lutra lutra), there are no published estimates on the extent to which bycatch is unreported, despite nonreporting being a well‐known issue. Through public outreach efforts in local news and media we collected information from citizen scientists on nonreported otter mortalities of a small‐scale fishery in western Norway along the recent natural expansion front of otters. We compared this to deaths that were properly registered by local authorities. In total, cause of mortality was determined from 218 otters between 2003 and 2018. We found that the chances of a mortality being properly reported varied between type of death. Drowning in fyke and gillnets were the most common cause of mortality, and at least 61 and 69% of these deaths were never reported, respectively. The high level of bycatch occurring in the otters' current region of expansion is likely to have demographic impacts and hinder otter reestablishment in the area. Banning the use of fyke nets and setting a minimum fishing depth for gillnets would enhance otter recovery and simultaneously alleviate bycatch of other nontarget species in the region, while having little economic impact on a fishery which is mostly noncommercial. The case of the otter is a classic example of lack of communication between government agencies which have so far failed to establish platforms where issues like this can be discussed and solved by adequate law implementation.

Dissection of the mutation accumulation process during bacterial range expansions

BackgroundRecent experimental work has shown that the evolutionary dynamics of bacteria expanding across space can differ dramatically from what we expect under well-mixed conditions. During spatial expansion, deleterious mutations can accumulate due to inefficient selection on the expansion front, potentially interfering with and modifying adaptive evolutionary processes.ResultsWe used whole genome sequencing to follow the genomic evolution of 10 mutator Escherichia coli lines during 39 days ( ~ 1650 generations) of a spatial expansion, which allowed us to gain a temporal perspective on the interaction of adaptive and non-adaptive evolutionary processes during range expansions. We used elastic net regression to infer the positive or negative effects of mutations on colony growth. The colony size, measured after three day of growth, decreased at the end of the experiment in all 10 lines, and mutations accumulated at a nearly constant rate over the whole experiment. We find evidence that beneficial mutations accumulate primarily at an early stage of the experiment, leading to a non-linear change of colony size over time. Indeed, the rate of colony size expansion remains almost constant at the beginning of the experiment and then decreases after ~ 12 days of evolution. We also find that beneficial mutations are enriched in genes encoding transport proteins, and genes coding for the membrane structure, whereas deleterious mutations show no enrichment for any biological process.ConclusionsOur experiment shows that beneficial mutations target specific biological functions mostly involved in inter or extra membrane processes, whereas deleterious mutations are randomly distributed over the whole genome. It thus appears that the interaction between genetic drift and the availability or depletion of beneficial mutations determines the change in fitness of bacterial populations during range expansion.

The hidden faces of a biological invasion: parasite dynamics of invaders and natives

One of the primary drivers of emerging infectious diseases (EIDs) is human intervention via host or parasite translocations. A unique opportunity to study host and parasite dispersal during a bio-invasion currently exists in Ireland due to the introduction of the bank vole (Myodes glareolus) in the 1920s. The continuing range expansion of M. glareolus within Ireland presents a natural large-scale perturbation experiment. This study used the Irish M. glareolus model to conduct a spatiotemporal study analysing the parasite dynamics of native and invasive species throughout their range. Myodes glareolus and native Apodemus sylvaticus were trapped in woodlands across Ireland and surveyed for their helminth parasites. Myodes glareolus in Ireland were found to have lower parasite diversity in comparison to records of M. glareolus from across Europe and A. sylvaticus in Ireland. Increased density of M. glareolus resulted in a dilution effect, with significantly lower levels of parasitism overall in native hosts, where M. glareolus has been established longest. However, three helminth parasite species of A. sylvaticus increased in abundance in the presence of M. glareolus. Furthermore, M. glareolus at the expansion front were less parasitised (lower abundance and prevalence of certain parasites and lower parasite diversity) than M. glareolus from the core population. This “enemy release” is believed to be mediating the continued successful spread of the invader across Ireland. Our results identify two important variables, seasonality and the stage of the invasion, which should not be overlooked when investigating or managing the changing distribution of hosts and their parasites. Studies of bio-invasions and parasite transmission have primarily focused on the invasive host species or the native host species in cases where virulent pathogen spillover is observed. Our results demonstrate how the concurrent study of invasive and native hosts, and the careful identification of their parasite communities, allows the dynamic processes influencing the parasite component and intracommunity to be identified.

Dispersal and life-history traits in a spider with rapid range expansion

BackgroundDispersal and reproduction are key life-history traits that jointly determine species’ potential to expand their distribution, for instance in light of ongoing climate change. These life-history traits are known to be under selection by changing local environmental conditions, but they may also evolve by spatial sorting. While local natural selection and spatial sorting are mainly studied in model organisms, we do not know the degree to which these processes are relevant in the wild, despite their importance to a comprehensive understanding of species’ resistance and tolerance to climate change.MethodsThe wasp spider Argiope bruennichi has undergone a natural range expansion - from the Mediterranean to Northern Europe during the recent decades. Using reciprocal common garden experiments in the laboratory, we studied differences in crucial traits between replicated core (Southern France) and edge (Baltic States) populations. We tested theoretical predictions of enhanced dispersal (ballooning behaviour) and reproductive performance (fecundity and winter survival) at the expansion front due to spatial sorting and local environmental conditions.ResultsDispersal rates were not consistently higher at the northern expansion front, but were impacted by the overwintering climatic conditions experienced, such that dispersal was higher when spiderlings had experienced winter conditions as occur in their region. Hatching success and winter survival were lower at the range border. In agreement with theoretical predictions, spiders from the northern leading edge invested more in reproduction for their given body size.ConclusionsWe found no evidence for spatial sorting leading to higher dispersal in northern range edge populations of A. bruennichi. However, reproductive investment and overwintering survival between core and edge populations differed. These life-history traits that directly affect species’ expansion rates seem to have diverged during the recent range expansion of A. bruennichi. We discuss the observed changes with respect to the species’ natural history and the ecological drivers associated with range expansion to northern latitudes.

Vespa velutina: An Alien Driver of Honey Bee Colony Losses

Vespa velutina, or Asian yellow-legged hornet, was accidentally introduced from China to other parts of the world: South Korea in 2003, Europe in 2004, and Japan in 2012. V. velutina represents a serious threat to native pollinators. It is known to be a fierce predator of honey bees, but can also hunt wild bees, native wasps, and other flying insects. When V. velutina colonies are developed, many hornets capture foraging bees which are coming back to their hives, causing an increase in homing failure and paralysis of foraging thus leading to colony collapse. The hornets may enter weak beehives to prey on brood and pillage honey. Unlike Apis cerana, Apis mellifera is unable to cope with the predation pressure of V. velutina. Monitoring the spread of an invasive alien species is crucial to plan appropriate management actions and activities to limit the expansion of the species. In addition, an early detection of V. velutina in areas far away from the expansion front allows a rapid response aimed to remove these isolated populations before the settlement of the species. Where V. velutina is now established, control measures to prevent colony losses must be implemented with an integrated pest management approach.

Evolution of Dispersal Can Rescue Populations from Expansion Load.

Understanding the causes and consequences of range expansions or range shifts has a long history in evolutionary biology. Recent theoretical, experimental, and empirical work has identified two particularly interesting phenomena in the context of species range expansions: (i) gene surfing and the relaxation of natural selection and (ii) spatial sorting. The former can lead to an accumulation of deleterious mutations at range edges, causing an expansion load and slowing down expansion. The latter can create gradients in dispersal-related traits along the expansion axis and cause an acceleration of expansion. We present a theoretical framework that treats spatial sorting and gene surfing as spatial versions of natural selection and genetic drift, respectively. This model allows us to analytically study how gene surfing and spatial sorting interact and derive the probability of fixation of pleiotropic mutations at the expansion front. We use our results to predict the coevolution of mean fitness and dispersal rates, taking into account the effects of random genetic drift, natural selection, and spatial sorting, as well as correlations between fitness- and dispersal-related traits. We identify a "rescue effect" of spatial sorting, where the evolution of higher dispersal rates at the leading edge rescues the population from incurring expansion load.

Cooperation mitigates diversity loss in a spatially expanding microbial population

The evolution and potentially even the survival of a spatially expanding population depends on its genetic diversity, which can decrease rapidly due to a serial founder effect. The strength of the founder effect is predicted to depend strongly on the details of the growth dynamics. Here, we probe this dependence experimentally using a single microbial species, Saccharomyces cerevisiae, expanding in multiple environments that induce varying levels of cooperativity during growth. We observe a drastic reduction in diversity during expansions when yeast grows noncooperatively on simple sugars, but almost no loss of diversity when cooperation is required to digest complex metabolites. These results are consistent with theoretical expectations: When cells grow independently from each other, the expansion proceeds as a pulled wave driven by growth at the low-density tip of the expansion front. Such populations lose diversity rapidly because of the strong genetic drift at the expansion edge. In contrast, diversity loss is substantially reduced in pushed waves that arise due to cooperative growth. In such expansions, the low-density tip of the front grows much more slowly and is often reseeded from the genetically diverse population core. Additionally, in both pulled and pushed expansions, we observe a few instances of abrupt changes in allele fractions due to rare fluctuations of the expansion front and show how to distinguish such rapid genetic drift from selective sweeps.

Introduction history and population genetics of intracontinental scotch broom (Cytisus scoparius) invasion

AbstractAimBiological invasions at the intracontinental scale are poorly studied, and intracontinental invasions often remain cryptic. Here, we investigate the recent range expansion of scotch broom (Cytisus scoparius) into Norway and clarify whether the genetic patterns indicate natural spread or human introduction. Furthermore, we investigate whether plants were moved within the native range and how this influences invasion success. We also infer the level and structuring of genetic diversity within and between the putative native and introduced range.LocationEurope.MethodsWe analysed the chloroplast sequence variation in 267 scotch broom samples from its northern expansion front and from its native range across Europe, including herbarium samples dating back to 1835. For 37 populations, we analysed variation in nuclear single‐nucleotide polymorphic markers to study gene flow and genetic diversity.ResultsWe identified 20 different haplotypes, which lacked spatial and temporal distribution patterns in the recent expansion range in Norway. They also mostly lacked patterns across the native European range of scotch broom. The genetic diversity of nuclear genomic SNP markers across populations in the introduced range was similar to that of populations in the native range, with limited differentiation among populations.Main conclusionsScotch broom is alien to Norway and was introduced by humans on multiple occasions from diverse origins over a long period of time. High propagule pressure has probably maintained the high genetic diversity in the novel range through a combination of genetically diverse source populations and high gene flow among them. Within the native European range, our results suggest the presence of cryptic intraspecific admixture, most likely mediated by humans moving genotypes among the regions occupied by distinct native genotypes. Intracontinental invasions may easily go unnoticed and revealing these invasions and the factors driving them may be of great importance for the management of alien species.

Dynamics of the boundary layer created by the explosion of a dense object in an ambient dilute gas triggered by a high power laser

The dynamics of the boundary layer in between two distinct collisionless plasmas created by the interaction between a dense object modeling a cluster and a short laser pulse in the presence of an ambient gas is studied with two dimensional relativistic particle-in-cell simulations, which are found to be described by three successive processes. In the first phase, a collisionless electrostatic shock wave, launched near the cluster expansion front, reflects the ambient gas ions at a contact surface as a moving wall, which allows a particle acceleration with a narrower energy spread. In the second phase, the contact surface disappears and the compressed surface of the ambient gas ions passes over the shock potential, forming an overlapping region between the cluster expansion front and the compressed surface of the ambient gas. Here, another type of nonlinear wave is found to be evolved, leading to a relaxation of the shock structure, while continuing to reflect the ambient gas ions. The nonlinear wave exhibits a bipolar electric field structure that is sustained for a long timescale coupled with slowly evolving ion dynamics, suggesting that a quasistationary kinetic equilibrium dominated by electron vortices in the phase space is established. In the third phase, a rarefaction wave is triggered and evolves at the compressed surface of ambient gas. This is because some of the ambient gas ions tend to pass over the potential of the bipolar electric field. Simultaneously, a staircase structure, i.e., a kind of internal shock, is formed in the cluster due to the deceleration of cluster ions. Such structure formations and successive dynamics accompanied by the transitions from the shock wave phase through the overlapping phase to the rarefaction wave phase are considered to be a unique nature at the boundary layer created by an explosion of a dense plasma object in an ambient dilute plasma.

The implications of rapid eco‐evolutionary processes for biological control ‐ a review

AbstractNovel environmental conditions experienced by introduced species can drive rapid evolution of diverse traits. In turn, rapid evolution, both adaptive and non‐adaptive, can influence population size, growth rate, and other important ecological characteristics of populations. In addition, spatial evolutionary processes that arise from a combination of assortative mating between highly dispersive individuals at the expanding edge of populations and altered reproductive rates of those individuals can accelerate expansion speed. Growing experimental evidence shows that the effects of rapid evolution on ecological dynamics can be quite large, and thus it can affect establishment, persistence, and the distribution of populations. We review the experimental and theoretical literature on such eco‐evolutionary feedbacks and evaluate the implications of these processes for biological control. Experiments show that evolving populations can establish at higher rates and grow larger than non‐evolving populations. However, non‐adaptive processes, such as genetic drift and inbreeding depression can also lead to reduced fitness and declines in population size. Spatial evolutionary processes can increase spread rates and change the fitness of individuals at the expansion front. These examples demonstrate the power of eco‐evolutionary dynamics and indicate that evolution is likely more important in biocontrol programs than previously realized. We discuss how this knowledge can be used to enhance efficacy of biological control.

Steady‐State Solution of a Solar Wind‐Generated Electron Cloud in a Lunar Crater

AbstractWe formulate an analytic description of a steady‐state electron cloud and affiliated surface charge, formed in the plasma wake generated as the solar wind flows horizontally over a lunar crater. The solution is complementary to the well‐known self‐similar plasma expansion formulation, which breaks down at the plasma wake front and thus fails to capture a substantial region of the crater interior. The present model establishes a theoretical basis for existing simulation results, which suggest that the cavity formed below the wake front is populated mainly by electrons, resulting in a substantial negative surface charge. The electrostatic potential throughout this subwake region is determined by solving Poisson's equation for a Maxwellian electron cloud, bounded above by the self‐similar plasma expansion front and below by the electrostatically charged surface.

Environmental heterogeneity can tip the population genetics of range expansions.

The population genetics of most range expansions is thought to be shaped by the competition between Darwinian selection and random genetic drift at the range margins. Here, we show that the evolutionary dynamics during range expansions is highly sensitive to additional fluctuations induced by environmental heterogeneities. Tracking mutant clones with a tunable fitness effect in bacterial colonies grown on randomly patterned surfaces we found that environmental heterogeneity can dramatically reduce the efficacy of selection. Time-lapse microscopy and computer simulations suggest that this effect arises generically from a local 'pinning' of the expansion front, whereby stretches of the front are slowed down on a length scale that depends on the structure of the environmental heterogeneity. This pinning focuses the range expansion into a small number of 'lucky' individuals with access to expansion paths, altering the neutral evolutionary dynamics and increasing the importance of chance relative to selection.

Ionization and recombination effects on laser ion acceleration in a finite mass plasma expansion

A self-similar multi-fluid model is performed to describe ion acceleration in singly ionized plasma with nonthermal electrons, where ionization and recombination are considered. It is found that ion acceleration in plasma expansion is strongly influenced by the competition of ionization and recombination processes under different nonthermal effects, at electron temperatures, and for various target materials. Two phases of expansion are shown in the profiles. The first one is the strongest collisional dense plasma that is created, spreading smoothly into vacuum near the surface of the target, with low slopes in all ion expansion profiles. The second is the core or the central phase of expansion dominated by recombination processes, with steep slopes up to the expansion front position, where the ion velocities and the electric field amplitude have reached their maximum values. The limit of expansion is determined where the ion density and electric field vanish. The interest of such a study may concern the dynamics of ionization and recombination processes in laser-plasma acceleration where nonthermal, energetic electrons are present.

Long-distance pollen dispersal during recent colonization favors a rapid but partial recovery of genetic diversity in Picea sitchensis.

Tree species in the northern hemisphere are currently subject to rapid anthropogenic climate change and are shifting their ranges in response. This prompts questions about the mechanisms allowing tree populations to respond quickly to selection pressures when establishing into new areas. Focusing on the northern expanding range edge of Picea sitchensis, a widespread conifer of western North America, we ask how genetic structure and diversity develop during colonization, and assess the role of demographic history in shaping the evolutionary trajectory of an establishing population. We combined 500yr of tree-ring and genotyping-by-sequencing data in 639 trees at the expansion front on the Kodiak Archipelago. We show that alleles accumulated rapidly during an increase in recruitment rate in the early 1700s. A shift from foreign to local pollen flow subsequently homogenized genetic structure at the expansion front. Taking advantage of the exceptional longevity of conifers, we highlight the major role of long-distance pollen dispersal in the rapid but incomplete recovery of genetic diversity during the initial stages of colonization. We also warn that slow initial population growth as well as long-lasting dominance of local gene flow by early founders could increase evolutionary load under a rapidly changing climate.

Invasion Dynamics of A Termite, Reticulitermes flavipes, at Different Spatial Scales in France.

Termites are social insects that can also be major pests. A well-known problem species is the subterranean termite, Reticulitermes flavipes. It is invasive in France and is thought to have arrived from Louisiana during the 18th century. While the putative source of French populations has been identified, little is known about how the termite spread following its establishment. Here, we examined expansion patterns at different spatial scales in urban areas to clarify how R. flavipes spread in France. Based on our analyses of phylogeography and population genetics, results suggest a scenario of successive introductions into the Charente-Maritime region, on the Atlantic Coast. Two major expansion fronts formed: one that spread toward the northeast and the other toward the southeast. At the regional scale, different spatial and genetic distribution patterns were observed: there was heterogeneity in Île-de-France and aggregation in Centre-Val de Loire. At the local scale, we found that our three focal urban sites each formed a single large colony that contained several secondary reproductives. Our findings represent a second step in efforts to reconstruct termite’s invasion dynamics. They also highlight the role that may have been played by the French railway network in transporting termites over long distances.

Laser-generated plasmas in length scales relevant for thin film growth and processing: simulation and experiment

In pulsed laser deposition, thin film growth is mediated by a laser-generated plasma, whose properties are critical for controlling the film microstructure. The advent of 2D materials has renewed the interest in how this ablation plasma can be used to manipulate the growth and processing of atomically thin systems. For such purpose, a quantitative understanding of the density, charge state, and kinetic energy of plasma constituents is needed at the location where they contribute to materials processes. Here, we study laser-induced plasmas over expansion distances of several centimeters from the ablation target, which is the relevant length scale for materials growth and modification. The study is enabled by a fast implementation of a laser ablation/plasma expansion model using an adaptive Cartesian mesh solver. Simulation outcomes for KrF excimer laser ablation of Cu are compared with Langmuir probe and optical emission spectroscopy measurements. Simulation predictions for the plasma-shielding threshold, the ionization state of species in the plasma, and the kinetic energy of ions, are in good correspondence with experimental data. For laser fluences of 1–4 J cm−2, the plume is dominated by Cu0, with small concentrations of Cu+ and electrons at the expansion front. Higher laser fluences (e.g. 7 J cm−2) lead to a Cu+ -rich plasma, with a fully ionized leading edge where Cu2+ is the dominant species. In both regimes, simulations indicate the presence of a low-density, high-temperature plasma expansion front with a high degree of ionization that may play a significant role in doping, annealing, and kinetically-driven phase transformations in 2D materials.

Influence of gas adsorption induced non-uniform deformation on the evolution of coal permeability

When adsorbing gas is injected into coal, the gas fills in the fractures quickly and a pressure difference between matrix and fractures is created. Because of this difference, there is a pressure gradient within the matrix. The gradient evolves with time from the initial equilibrium (zero gradient) to the final equilibrium (zero gradient), so does the adsorption–swelling induced matrix deformation. Previous studies have not taken this effect into consideration. In this study, we hypothesize that the pressure gradient affects the expansion of the gas-invaded area/volume with the matrix and the propagation of the expansion front creates a non-uniform deformation within the matrix. Under this hypothesis, a relation between coal permeability and the expansion of the gas-invaded area with the matrix can be established. When the gas-invaded area is localized in the vicinity of the fracture wall, the expansion of the matrix within this area narrows the fracture opening. We define this as local swelling/shrinking. This local swelling/shrinking is controlled primarily by the coal internal structure. When the gas-invaded area is further spread over the matrix, the expansion of the whole matrix may narrow or widen the fracture opening depending on the external boundary conditions. This global swelling/shrinking is controlled primarily by the external boundary conditions. These conceptual understandings are defined through strain rate-based coal permeability models for both the matrix and the fractures. This strain rate based time-dependent permeability model was verified against experimental observations, that couples coal deformation, the gas flow in the matrix system and gas flow in the fracture system.

High genomic diversity and candidate genes under selection associated with range expansion in eastern coyote (Canis latrans) populations.

Range expansion is a widespread biological process, with well‐described theoretical expectations associated with the colonization of novel ranges. However, comparatively few empirical studies address the genomic outcomes accompanying the genome‐wide consequences associated with the range expansion process, particularly in recent or ongoing expansions. Here, we assess two recent and distinct eastward expansion fronts of a highly mobile carnivore, the coyote (Canis latrans), to investigate patterns of genomic diversity and identify variants that may have been under selection during range expansion. Using a restriction‐associated DNA sequencing (RADseq), we genotyped 394 coyotes at 22,935 SNPs and found that overall population structure corresponded to their 19th century historical range and two distinct populations that expanded during the 20th century. Counter to theoretical expectations for populations to bottleneck during range expansions, we observed minimal evidence for decreased genomic diversity across coyotes sampled along either expansion front, which is likely due to hybridization with other Canis species. Furthermore, we identified 12 SNPs, located either within genes or putative regulatory regions, that were consistently associated with range expansion. Of these 12 genes, three (CACNA1C, ALK, and EPHA6) have putative functions related to dispersal, including habituation to novel environments and spatial learning, consistent with the expectations for traits under selection during range expansion. Although coyote colonization of eastern North America is well‐publicized, this study provides novel insights by identifying genes associated with dispersal capabilities in coyotes on the two eastern expansion fronts.

Magnetic Reconnection Detonation in Supernova Remnants

Abstract As a key process that refreshes the interstellar medium (ISM), the dynamics and radiative properties of the supernova remnant (SNR) expansion front not only reflect the physical environment of the old ISM surrounding the supernova, but they also provide information about the refreshed ISM. However the expansion dynamics of SNRs cannot be simply explained by the conventional law of spherical shock wave propagation; on the other hand, the high-energy radiation requires an additional electron acceleration mechanism in the shock front beyond thermal collision. We consider herein the detonation wave description of the SNR expansion, in which magnetic reconnection follows the shock front and transfers the SNR magnetic field energy to both fluid thermal energy and particle kinetic energy. The structure of the magnetic reconnection detonation (MRD) is identified based on the scaling analysis in this paper. By applying the MRD description of the SNR expansion shock to the example of the Crab Nebula, this paper shows that the MRD description can explain both the accelerative expansion of the nebula as well as the origin of the luminous expanding shell.

Den site selection by male brown bears at the population's expansion front.

Brown bears (Ursus arctos) spend about half of the year in winter dens. In order to preserve energy, bears may select denning locations that minimize temperature loss and human disturbance. In expanding animal populations, demographic structure and individual behavior at the expansion front can differ from core areas. We conducted a non-invasive study of male brown bear den sites at the male-biased, low-density western expansion front of the Scandinavian brown bear population, comparing den locations to the available habitat. Compared to the higher-density population core in which intraspecific avoidance may affect den site selection of subordinate bears, we expected resource competition in the periphery to be low, and all bears to be able to select optimal den sites. In addition, bears in the periphery had access to free-ranging domestic sheep during summer. We found that males in the periphery denned on high-elevation slopes, probably providing good drainage, longer periods of consistent, insulating snow cover and fewer melting-freezing events. Forests were the principal denning habitat and no dens were found in alpine areas. The Scandinavian brown bears have a history of intense harvest, including culling at the den. This may have exerted a selection pressure to avoid denning in open alpine habitat which compared to forests provide little cover. The bears denned away from main roads and in steep, rugged terrain, probably limiting human access. The odds for finding a bear den decreased with increasing distance to the population core where females could be found. Previous studies have documented directed movement of male brown bears from the male-biased population periphery toward the core areas during the mating season. In this way, denning males may be trading off between low resource competition and access to sheep in the low-density periphery, and mating opportunities in the higher-density population core.