Undergoing Range Expansion Research Articles

Boundary Effects Cause False Signals of Range Expansions in Population Genomic Data.

Studying range expansions is central for understanding genetic variation through space and time as well as for identifying refugia and biological invasions. Range expansions are characterized by serial founder events causing clines of decreasing genetic diversity away from the center of origin and asymmetries in the two-dimensional allele frequency spectra. These asymmetries, summarized by the directionality index (ψ), are sensitive to range expansions and persist for longer than clines in genetic diversity. In continuous and finite meta-populations, genetic drift tends to be stronger at the edges of the species distribution in equilibrium populations and populations undergoing range expansions alike. Such boundary effects are expected to affect geographic patterns in genetic diversity and ψ. Here we demonstrate that boundary effects cause high false positive rates in equilibrium meta-populations when testing for range expansions. In the simulations, the absolute value of ψ (|ψ|) in equilibrium data sets was proportional to the fixation index (FST). By fitting signatures of range expansions as a function of ɛ |ψ|/FST and geographic clines in ψ, strong evidence for range expansions could be detected in data from a recent rapid invasion of the cane toad, Rhinella marina, in Australia, but not in 28 previously published empirical data sets from Australian scincid lizards that were significant for the standard range expansion tests. Thus, while clinal variation in ψ is still the most sensitive statistic to range expansions, to detect true signatures of range expansions in natural populations, its magnitude needs to be considered in relation to the overall levels of genetic structuring in the data.

Across two phylogeographic breaks: Quaternary evolutionary history of a mountain aspen (Populus rotundifolia) in the Hengduan Mountains

Biogeographical barriers to gene flow are central to plant phylogeography. In East Asia, plant distribution is greatly influenced by two phylogeographic breaks, the Mekong-Salween Divide and Tanaka-Kaiyong Line, however, few studies have investigated how these barriers affect the genetic diversity of species that are distributed across both. Here we used 14 microsatellite loci and four chloroplast DNA fragments to examine genetic diversity and distribution patterns of 49 populations of Populus rotundifolia, a species that spans both the Mekong-Salween Divide and the Tanaka-Kaiyong Line in southwestern China. Demographic and migration hypotheses were tested using coalescent-based approaches. Limited historical gene flow was observed between the western and eastern groups of P. rotundifolia, but substantial flow occurred across both the Mekong-Salween Divide and Tanaka-Kaiyong Line, manifesting in clear admixture and high genetic diversity in the central group. Wind-borne pollen and seeds may have facilitated the dispersal of P. rotundifolia following prevalent northwest winds in the spring. We also found that the Hengduan Mountains, where multiple genetic barriers were detected, acted on the whole as a barrier between the western and eastern groups of P. rotundifolia. Ecological niche modeling suggested that P. rotundifolia has undergone range expansion since the last glacial maximum, and demographic reconstruction indicated an earlier population expansion around 600 Ka. The phylogeographic pattern of P. rotundifolia reflects the interplay of biological traits, wind patterns, barriers, niche differentiation, and Quaternary climate history. This study emphasizes the need for multiple lines of evidence in understanding the Quaternary evolution of plants in topographically complex areas.

Overwintering conditions affect cold hardiness, survival, and post‐overwintering fitness of the pea leaf weevil

AbstractOverwintering conditions affect the physiological state of ectotherms, and therefore, their cold hardiness and survival. A measure of the lethal and sublethal impacts of overwintering conditions on pest populations is crucial to predict population dynamics and to manage pests the following spring. The impact of winter conditions can be most intense for invasive insects undergoing range expansion. Insect herbivores can display plastic host use behaviours that depend on their body condition following winter. The pea leaf weevil, Sitona lineatus L. (Coleoptera: Curculionidae), is an invasive pest of field peas, Pisum sativum L., and faba bean, Vicia faba L. (Fabaceae). Pea leaf weevil has expanded its range in North America to include the Prairie Provinces of Canada. This study investigated the effects of temperature and microhabitat on overwintering survival and cold hardiness of pea leaf weevil in its expanded range. Further, we investigated the sublethal effect of overwintering temperature and duration on post‐overwintering survival, feeding, and oviposition of pea leaf weevil. We also investigated the role of juvenile hormone in modulating body condition of overwintering weevils. The overwintering survival of pea leaf weevil adults increased with soil temperature and varied with region and microhabitat. More weevils survived winters when positioned near tree shelterbelts compared to open alfalfa fields. The supercooling point of pea leaf weevil varied throughout its expanding range but did not differ for weevils held in the two microhabitats. The average threshold lethal temperature of pea leaf weevil at all three sites was −9.4 °C. Weevils that overwintered for a longer duration and at a higher temperature subsequently fed more on faba bean foliage and laid more eggs compared to those which overwintered for a shorter duration at a lower temperature. Our findings highlight that warm winters would increase overwintering survival and post‐overwintering fitness, facilitating further pea leaf weevil invasion northward in the Prairie Provinces of Canada.

Spatial and risk factor analyses of vector-borne pathogens among shelter dogs in the Eastern United States

BackgroundVector-borne infections pose significant health risks to humans, domestic animals, and wildlife. Domestic dogs (Canis lupus familiaris) in the United States may be infected with and serve as sentinel hosts for several zoonotic vector-borne pathogens. In this study, we analyzed the geographical distribution, risk factors, and co-infections associated with infection with Ehrlichia spp., Anaplasma spp., Borrelia burgdorferi, and Dirofilaria immitis in shelter dogs in the Eastern United States.MethodsFrom 2016 to 2020, blood samples from 3750 shelter dogs from 19 states were examined with IDEXX SNAP® 4Dx® Plus tests to determine the seroprevalence of infection with tick-borne pathogens and infection with D. immitis. We assessed the impact of factors including age, sex, intact status, breed group, and location on infection using logistic regression.ResultsThe overall seroprevalence of D. immitis was 11.2% (n = 419/3750), the seroprevalence of Anaplasma spp. was 2.4% (n = 90/3750), the seroprevalence of Ehrlichia spp. was 8.0% (n = 299/3750), and the seroprevalence of B. burgdorferi was 8.9% (n = 332/3750). Regional variation in seroprevalence was noted: D. immitis (17.4%, n = 355/2036) and Ehrlichia spp. (10.7%, n = 217/2036) were highest in the Southeast while seroprevalence for B. burgdorferi (19.3%, n = 143/740) and Anaplasma spp. (5.7%, n = 42/740) were highest in the Northeast. Overall, 4.8% (n = 179/3750) of dogs had co-infections, the most common of which were D. immitis/Ehrlichia spp. (1.6%, n = 59/3750), B. burgdorferi/Anaplasma spp. (1.5%, n = 55/3750), and B. burgdorferi/Ehrlichia spp. (1.2%, n = 46/3750). Risk factors significantly influenced infection across the evaluated pathogens were location and breed group. All evaluated risk factors were significant for the seroprevalence of D. immitis antigens.ConclusionsOur results demonstrate a regionally variable risk of infection with vector-borne pathogens in shelter dogs throughout the Eastern United States, likely due to varying distributions of vectors. However, as many vectors are undergoing range expansions or other changes in distribution associated with climate and landscape change, continued vector-borne pathogen surveillance is important for maintaining reliable risk assessment.Graphical

Persistent variation in insecticide resistance intensity in container breeding Aedes (Diptera: Culicidae) co-collected in Houston, TX.

As observed in many locations worldwide, resistance to pyrethroids is common in Aedes aegypti (L.) in the southern United States and northern Mexico. Strong resistance in Aedes albopictus (Skuse) is less common and is not as well characterized. These 2 species have been undergoing range expansion and are sympatric in many locations including Houston, Texas. They are often collected from the same locations and lay eggs in the same larval habitats. In this study, we colonized both Ae. aegypti and Ae. albopictus from 4 locations in Houston and characterized insecticide resistance using permethrin as a model pyrethroid. We found differences in resistance intensity between the species at all 4 sites. Within the Ae. aegypti, resistance ratios ranged from 3.5- to 30.0-fold when compared to the ORL1952 laboratory susceptible strain. Expression of several P450s was higher than in the ORL1952 strain, but the pattern was similar between the field strains of Ae. aegypti. Higher resistance ratios did correlate with increasing percentages of the dilocus knockdown resistance (kdr) genotype. In contrast, Ae. albopictus from the 4 locations all had very low resistance ratios (<4-fold) when compared to the same laboratory susceptible strain. Five years later, we performed additional collections and characterization from the most resistant location to assess the temporal persistence of this difference in resistance between the species. The same pattern of high resistance in Ae. aegypti and low resistance in sympatric Ae. albopictus remained 5 yr later and this may have implications for operational efficacy.

Genetic Structure and Colonization of North America by Depressaria depressana (Fabricius 1775) (Lepidoptera: Depressariidae) over 15 Years; Contrasts with Westward Expansion of Depressaria radiella (Goeze, 1783) over 160 Years.

Simple SummaryAs invasive insect species can disrupt agriculture, ecosystem functions, and human health, monitoring the spread of newly invasive insects is a high priority. The purple carrot-seed moth Depressaria depressana, a European species first reported in North America in 2008, specializes on reproductive structures of taxa throughout the family Apiaceae and thus presents a potential threat to native North American umbellifers. We assessed interpopulational genetic diversity of D. depressana across its eastern North American invasion front and compared those values to available data for European populations. We also compared this diversity with genetic diversity estimates for populations of Depressaria radiella (parsnip webworm), a European species introduced to North America more than 160 years ago that is essentially restricted to two apiaceous genera throughout its native and invasive ranges. Documenting the historical spread of D. radiella with museum and literature records, we found D. depressana displays greater genetic diversity than D. radiella and is colonizing North America more rapidly; greater genetic diversity may facilitate faster colonization via establishment on a wider range of hostplants. The contrast in colonization by these two species may inform management practices for invasive insects utilizing a broad versus narrow array of hosts within a single plant family.Depressaria depressana, the purple carrot seed moth, is a Eurasian species first reported in North America in 2008 and currently undergoing range expansion. This invasion follows that of its Eurasion congener Depressaria radiella (parsnip webworm), first documented in North America 160 years ago. Unlike D. depressana, which utilizes hostplants across multiple tribes of Apiaceae, Depressaria radiella is a “superspecialist” effectively restricted in its native and non-indigenous ranges to two closely related apiaceous genera. We investigated the genetic structure of D. depressana populations across latitudinal and longitudinal gradients in the eastern United States by constructing COI haplotype networks and then comparing these with haplotype networks constructed from available COI sequence data from contemporary European D. depressana populations and from European and North American D. radiella populations. Haplotype data revealed higher genetic diversity in D. depressana, indicating high dispersal capacity, multiple introductions, and/or a genetically diverse founding population. Museum and literature records of D. radiella date back to 1862 and indicate that range expansion to the West Coast required more than 50 years. Higher levels of genetic diversity observed in D. depressana compared to its congener may indicate a greater propensity for dispersal, colonization and establishment in its non-indigenous range.

A theoretical framework for multi‐species range expansion in spatially heterogeneous landscapes

Range expansion is the spatial spread of a population into previously unoccupied regions. Understanding range expansion is important for the study and successful management of ecosystems, with applications ranging from controlling bacterial biofilm formation in industrial and medical environments to large scale conservation programmes for species undergoing climate‐change induced habitat disruption. During range expansion, species typically encounter competitors. Moreover, the spatial environment into which expansion takes place is almost always heterogeneous. Nevertheless, the impact of competition and spatial landscape heterogeneities on range expansion remains understudied. In this paper we present a theoretical framework comprising two competing generic species undergoing range expansion and use it to investigate the impact of spatial landscape heterogeneities on range expansion with a particular focus on its effect on competition dynamics. We reveal that the area covered by range expansion is highly variable due to the landscape heterogeneities. Moreover, we report significant variability in competitive outcome (relative abundance of a focal species), but determine that this is induced by low initial population densities and is independent of landscape heterogeneities. We further show that both area covered by range expansion and competitive outcome can be accurately predicted by a Voronoi tessellation with respect to an appropriate metric, which only requires information on the spatial landscape and the response of each species to that landscape. Finally, we reveal that if species interact antagonistically during range expansion, the dominant mode of competition depends on the initial population density; antagonistic actions determine competitive outcome if the initial population density is high, but competition for space is the dominant mode of competition if the initial population density is low. These findings enhance our understanding of how competition for space and antagonistic interactions affect range expansion in spatially heterogeneous environments and provide a predictive tool for future species‐specific approaches.

New Distribution Records of Biting Midges of the Genus Culicoides (Diptera: Ceratopogonidae) Latreille, Culicoides bergi and Culicoides baueri, in Southern Ontario, Canada.

Some species of Culicoides Latreille (Diptera: Ceratopogonidae) can be pests as well as pathogen vectors, but data on their distribution in Ontario, Canada, are sparse. Collecting this baseline data is important given ongoing, accelerated alterations in global climate patterns that may favor the establishment of some species in northern latitudes. Culicoides spp. were surveyed using UV light traps over two seasons in 2017 and 2018 at livestock farms in southern Ontario, Canada. Two Culicoides spp. not previously recorded in Canada were identified, C. bergi and C. baueri, representing new country and provincial records. Unlike some congenerics, these two species are not currently recognized as vectors of pathogens that pose a health risk to humans, livestock or wildlife in North America. However, the possibility that these Culicoides species may have recently expanded their geographic range, potentially in association with climate and/or landscape changes, warrants ongoing attention and research. Furthermore, our results provoke the question of the potential undocumented diversity of Culicoides spp. in Ontario and other parts of Canada, and whether other Culicoides spp. may be undergoing range expansion. The current and future distributions of Culicoides spp., and other potential vectors of human, agricultural, and wildlife health significance, are important to identify for proper disease risk assessment, mitigation, and management.

Mesopredator frugivory has no effect on seed viability and emergence under experimental conditions

AbstractMembers of the orderCarnivoraare a unique and important seed disperser who consume and deposit undamaged seeds while providing regular long‐distance seed dispersal opportunities. Some members ofCarnivora, such as coyotes (Canis latrans), are undergoing range expansions which may help the plant species they consume colonize new locations or replace dispersal services provided by recently extirpated species. In this study, we evaluated aspects of the seed dispersal effectiveness of coyotes and gut passage time to determine the potential dispersal distances for three commonly consumed and commonly occurring plant species (Amelanchier alnifolia, Celtis ehrenbergiana, andJuniperus osteosperma). We also investigated the potential effects of secondary dispersal of seeds away from scats by comparing seedling emergence from whole scats to those where seeds were first removed from scats. We found that seeds generally took between 4 and 24 h to pass through the digestive tract of coyotes, which could result in regular seed dispersal up to 7 km. Gut passage through coyotes had no effect on seed viability or emergence for any of the three plant species, including that gut passage forA. alnifoliaandJ. osteospermadoes not replace cold stratification for breaking physiological dormancy. By simulating secondary dispersal, we found that 22% (±8.2%) moreC. ehrenbergianaseedlings emerged when seeds were removed from scats and those seedlings emerged 7 d earlier (±5 d) compared to seeds that remained in the coyote scat. Coyotes are effective seed dispersers, with the potential for regular long‐distance dispersal services and for providing opportunities for secondary seed dispersal, which could aid in climate migration or serve to replace extirpated dispersal mutualists.

Spatial segregation and cooperation in radially expanding microbial colonies under antibiotic stress

Antibiotic resistance in microbial communities reflects a combination of processes operating at different scales. In this work, we investigate the spatiotemporal dynamics of bacterial colonies comprised of drug-resistant and drug-sensitive cells undergoing range expansion under antibiotic stress. Using the opportunistic pathogen Enterococcus faecalis with plasmid-encoded β-lactamase, we track colony expansion dynamics and visualize spatial patterns in fluorescently labeled populations exposed to antibiotics. We find that the radial expansion rate of mixed communities is approximately constant over a wide range of drug concentrations and initial population compositions. Imaging of the final populations shows that resistance to ampicillin is cooperative, with sensitive cells surviving in the presence of resistant cells at otherwise lethal concentrations. The populations exhibit a diverse range of spatial segregation patterns that depend on drug concentration and initial conditions. Mathematical models indicate that the observed dynamics are consistent with global cooperation, despite the fact that β-lactamase remains cell-associated. Experiments confirm that resistant colonies provide a protective effect to sensitive cells on length scales multiple times the size of a single colony, and populations seeded with (on average) no more than a single resistant cell can produce mixed communities in the presence of the drug. While biophysical models of drug degradation suggest that individual resistant cells offer only short-range protection to neighboring cells, we show that long-range protection may arise from synergistic effects of multiple resistant cells, providing surprisingly large protection zones even at small population fractions.

Long-term stress level in a small mammal species undergoing range expansion

Abstract Rapid climate change is currently altering species distribution ranges. Evaluating the long-term stress level in wild species undergoing range expansion may help better understanding how species cope with the changing environment. Here, we focused on the white-footed mouse (Peromyscus leucopus), a widespread small mammal species in North-America whose distribution range is rapidly shifting northward. We evaluated long-term stress level in several populations of P. leucopus in Quebec (Canada), from the northern edge of the species distribution to more core populations in Southern Quebec. We first tested the hypothesis that populations at the range margin are under higher stress than more established populations in the southern region of our study area. We then compared four measures of long-term stress level to evaluate the congruence between these commonly used methods. We did not detect any significant geographical trend in stress level across our study populations of P. leucopus. Most notably, we found no clear congruence between the four measures of stress level we used, and conclude that these four commonly used methods are not equivalent, thereby not comparable across studies.

The effect of cross-boundary management on the trajectory to commonness in biological invasions

The number of alien species introduced and undergoing range expansion in novel environments is steadily increasing, with important consequences for native ecosystems. The efficacy of management planning and decision making to limit such invasions can be improved by understanding how interventions will impact the population dynamics of recently introduced species. To do so, here we expand on a typological framework that enables the classification of populations over time into 10 categories of commonness, and apply it to a spatially discrete metapopulation with heterogeneous abundance across spatial units (patches). We use this framework to assess the effect of cross-boundary management on the capacity of a metapopulation with different demographic and dispersal characteristics, including time lags in population growth, to become common. We demonstrate this framework by simulating a simple theoretical metapopulation model capable of exploring a range of environments, species characteristics, and management actions. Management can vary in the efficacy of propagule interception between patches, and in the synchronisation of the implementation of these measures across patches (i.e. if management is implemented simultaneously across patches). Simulations show that poor interception efficacy that only modestly reduces the number of propagules entering a given spatial unit cannot be compensated for by strong management synchronisation between spatial units. Management synchronisation will nonetheless result in a reduction in rates of spread once a critical threshold of interception efficacy has been met. Finally, time lags in population growth that may result in delayed spread are an important aspect to be considered in management as they can amplify the efficacy of management. Our results demonstrate how a typological framework of categories of commonness can be used to provide practical insights for the management of biological invasions.

Genetic and phenotypic displacement of an endemic Barbus complex by invasive European barbel Barbus barbus in central Italy

Invasions of alien fishes can result in considerable consequences for native biodiversity, including local extinctions of native species through genetic introgression. In Italy, the alien European barbel Barbus barbus was first detected in 1994. It has since undergone range expansion, raising conservation concerns on their impacts on endemic Barbus species, including Barbus plebejus and Barbus tyberinus. Here, the genetic and phenotypic consequences of B. barbus invasion in the Tyrrhenian and Adriatic basins of central Italy were assessed by comparing ‘invaded’ with ‘uninvaded’ river sections that remain free of B. barbus due to barriers preventing their upstream dispersal. In both basins, uninvaded sites were confirmed as B. barbus free, but the endemic populations had low genetic variability. In the invaded sections, haplotype and nucleotide diversity was relatively high, with introgression skewed towards B. barbus genes, with the barbel populations comprising of only 4% and 23% of pure B. tyberinus and B. plebejus respectively. Relatively high morphological differentiation was apparent between pure B. tyberinus and hybrid forms, whilst differences were less apparent between pure B. plebejus and their hybrid forms. Thus, the endemic Barbus species only persist in areas that remain free of invasive B. barbus, with this only due to river structures that impede their upstream movements. As these structures also limit the effective population size of the endemic species, conservation plans must reconcile B. barbus dispersal prevention with the need to increase the population connectivity of the endemics.

Citizen science reveals the Palaearctic poison hemlock moth Agonopterix alstroemeriana (Clerck) (Lepidoptera: Depressariidae) has established in Australia

ABSTRACT We report the first occurrence in Australia of the poison hemlock moth Agonopterix alstroemeriana (Clerck), a monophage on the environmental weed Conium maculatum L. (Apiaceae). The hostplant, also of European origin, is a familiar toxic weed in southern Australia and the moth may have some potential as a biocontrol agent. It joins a list of other Palaearctic species undergoing range expansion into the southern temperate zone and probably colonised Tasmania via New Zealand where it first established in 1986. The discovery was facilitated by a citizen science application for smart-phones linking images of fauna and flora taken in the field to crowd-sourced identification resources, which can quickly converge to a satisfactory determination. In addition to its potential utility in the control of poison hemlock, the reunion of this host and its defoliating monophage after almost 150 years offers a testbed for theories of insect–hostplant evolution.

Mate-finding dispersal reduces local mate limitation and sex bias in dispersal.

Sex-biased dispersal (SBD) often skews the local sex ratio in a population. This can result in a shortage of mates for individuals of the less-dispersive sex. Such mate limitation can lead to Allee effects in populations that are small or undergoing range expansion, consequently affecting their survival, growth, stability and invasion speed. Theory predicts that mate shortage can lead to either an increase or a decrease in the dispersal of the less-dispersive sex. However, neither of these predictions have been empirically validated. To investigate how SBD-induced mate limitation affects dispersal of the less-dispersive sex, we used Drosophila melanogaster populations with varying dispersal propensities. To rule out any mate-independent density effects, we examined the behavioural plasticity of dispersal in the presence of mates as well as same-sex individuals with differential dispersal capabilities. In the presence of high-dispersive mates, the dispersal of both male and female individuals was significantly increased. However, the magnitude of this increase was much larger in males than in females, indicating that the former shows greater mate-finding dispersal. Moreover, the dispersal of either sex did not change when dispersing alongside high- or low-dispersive individuals of the same sex. This suggested that the observed plasticity in dispersal was indeed due to mate-finding dispersal, and not mate-independent density effects. Strong mate-finding dispersal can diminish the magnitude of sex bias in dispersal. This can modulate the evolutionary processes that shape range expansions and invasions, depending on the population size. In small populations, mate-finding dispersal can ameliorate Allee effects. However, in large populations, it can dilute the effects of spatial sorting.

Eriophyid mite Floracarus perrepae reduces climbing ability of the invasive vine Lygodium microphyllum

Invasive vines often grow rapidly, smothering native vegetation and causing extensive ecosystem damage. Despite these impacts, the growth rates of individual stems of multi-stemmed vines and the effects of biological control agents on these growth rates are poorly understood. Here, we quantified the growth rates of individual rachises of Lygodium microphyllum (Lygodiaceae; Old World climbing fern), a noxious invasive vine undergoing range expansion in Florida. Our objectives were to (1) quantify L. microphyllum rachis growth under varied light conditions and seasons; and (2) determine the effect of damage by the biological control mite Floracarus perrepae (Eriophyidae) to the apical meristem on rachis growth. We experimentally grew plants onto trellises to isolate, track, and measure individual rachises of the plant during 3 seasons (spring, summer, winter). Healthy (undamaged) rachises of L. microphyllum grew fastest in spring (2.22 ± 0.13 cm/day), particularly in shade where rachises grew nearly 1.5 times faster than in full sun. Mite damage reduced rachis growth rates 3.7 fold and greatly increased plant investment in rachises that did not climb. Furthermore, 85% of rachises damaged by mite feeding did not attain normal growth rates following mite damage. Our findings are the first to quantify the rapid climbing ability of individual L. microphyllum rachises, as well as the potential for F. perrepae to substantially curb rachis growth. Importantly, the ability to impede a vine’s climbing may be critical in determining the efficacy of a biological control agent, and future work on vines should focus on developing such agents.

Patterns of growth, brooding and offspring size in the invasive mussel Sinanodonta woodiana (Lea, 1834) (Bivalvia: Unionidae) from an anthropogenic heat island

In freshwater temperate environments, waterbodies warmed by power plants can serve as a stepping stone in the spread of tropical and subtropical species. This offers a unique opportunity to acquire an insight into the life history of organisms undergoing range expansions beyond their native habitat. Here, we studied the patterns of offspring production and growth in the Asian mussel Sinanodonta woodiana inhabiting the thermal plume of the Odra River in Central Europe. Compared to the S. woodiana males, females had more convex shells and had a greater tendency to continue post-maturation growth and thus follow an indeterminate growth pattern. Gravid females were observed all-year round, and individuals with either large or more convex shells brooded more offspring. The proportion of incubating females, brood size and glochidia size were linked to gonadal activity and changed seasonally. Females with a higher amount of nutritive substances in the ovaries produced more and larger offspring. The smallest and the largest offspring occurred in the summer and in the winter, respectively. Sinanodonta woodiana incubated different offspring generations simultaneously, constantly releasing them into the environment. Using a life history evolution perspective, we discuss our results in view of allocation trade-offs between growth and offspring production.

Mixed evidence for shifts to faster carbon capture strategies towards range edges of two coastal invasive plants in eastern Australia

Understanding shifts in traits across the course of an invasion can significantly increase our understanding of mechanisms underpinning range expansion. For example, shifts to traits associated with faster growth may be advantageous in range edge populations of invasive species to decrease generation time and thus promote rapid range expansion. We tested whether populations at the expanding range edges of two coastal plant species invasive in eastern Australia (Gladiolus gueinzii Kunze and Hydrocotyle bonariensis Lam.) possessed different carbon capture strategies compared with range core populations where they were first introduced. Pairwise leaf trait relationships between specific leaf area (SLA), photosynthetic rate (Amass), foliar nitrogen (Nmass) and foliar phosphorus (Pmass) were investigated for range edge and range core populations using standardised major axis (SMA) regression. Across species, SMA regression slopes for range core and range edge populations for all pairwise comparisons did not differ significantly from each other, suggesting that each species has a similar carbon capture strategy across its range. However, at a species level, H. bonariensis displayed significant shifts in trait values along a common regression slope for many pairwise comparisons. Range edge populations were found to have higher values for Nmass, Amass and SLA compared to range core populations and displayed greater nutrient use efficiency, suggesting that range edge populations are positioned further along the leaf economics spectrum towards faster growth strategies. In contrast, for G. gueinzii, leaf traits were positioned along a common regression slope with no difference in the trait values of range core and range edge populations. Our results suggest that there is selection for faster carbon capture strategies at range edges for some introduced species undergoing range expansion and this may be a contributing factor in explaining rapid range advance.

The role of dispersal, selection, and timing of sampling on the false discovery rate of loci under selection during geographic range expansion.

Identifying adaptive loci is important to understand the evolutionary potential of species undergoing range expansion. However, in expanding populations, spatial demographic processes such as allele surfing can create spatial patterns of neutral genetic variation that appear similar to those generated through adaptive processes. As a result, the false discovery rate of adaptive loci may be inflated in landscape genomic analyses. Here, we take a simulation modelling approach to investigate how range expansion affects our ability to correctly distinguish between neutral and adaptive genetic variation, using the mountain pine beetle outbreak system as a motivating example. We simulated the demographic and population genetic dynamics of populations undergoing range expansion using an individual-based genetic model CDMetaPOP. We investigated how the false discovery rate of adaptive loci is affected by (i) dispersal capacity, (ii) timing of sampling, and (iii) the strength of selection on an adaptive reference locus. We found that a combination of weak dispersal, weak selection, and early sampling presents the greatest risk of misidentifying loci under selection. Expanding populations present unique challenges to the reliable identification of adaptive loci. We demonstrate that there is a need for further methodological development to account for directional demographic processes in landscape genomics.

DNA methylation predicts immune gene expression in introduced house sparrows Passer domesticus

Populations undergoing range expansions are often faced with novel selective pressures, and to cope with such challenges, populations must either adapt quickly or exhibit phenotypic plasticity. This latter option allows for rapid phenotypic adjustments and persistence in novel environments, and thus could be advantageous at range‐edges. Our previous research on house sparrows in Kenya – a site of ongoing range expansion – and a growing literature suggests that invasion success is facilitated by epigenetic regulation of gene expression. Previously, we found 1) differences in the expression of a microbial surveillance gene (i.e. Toll‐like receptor 4‐TLR4), and 2) extensive variation in genome‐wide DNA methylation among house sparrows across Kenya. Here, our goal was to investigate whether these two observations are related, specifically whether DNA methylation within a target sequence upstream of the TLR4 transcription start site is associated with variation in TLR4 expression. We found that DNA methylation in the aforementioned region was quite variable among individuals, and variation at one CpG site predicted differences in TLR4 expression. Moreover, we found genetic variation within the same sequence upstream of the TLR4 exon, but this variation did not predict TLR4 expression. To our knowledge, this is the first study to demonstrate an association between DNA methylation and the expression of an ecologically relevant trait in a range‐expanding vertebrate.