Development and localization of microsatellite markers for the sibling species Chironomus riparius and Chironomus piger (Diptera: Chironomidae)

Five variable microsatellite loci are reported for the nonbiting midge species Chironomus riparius and Chironomus piger. All loci show considerable intraspecific variation and species‐specific alleles, which allow to discriminate among the two closely related species and their interspecific hybrids, and to estimate genetic diversity within and between populations. Additionally, the loci were localized on C. riparius polytene chromosomes to verify their single copy status and investigate possible chromosomal linkage. The described markers are used in different studies with regard to population and ecological genetics and evolutionary ecotoxicology of Chironomus.     

Chironomus riparius (Diptera) genome sequencing reveals the impact of minisatellite transposable elements on population divergence

Active transposable elements (TEs) may result in divergent genomic insertion and abundance patterns among conspecific populations. Upon secondary contact, such divergent genetic backgrounds can theoretically give rise to classical Dobzhansky–Muller incompatibilities (DMI), thus contributing to the evolution of endogenous genetic barriers and eventually causing population divergence. We investigated differential TE abundance among conspecific populations of the nonbiting midge Chironomus riparius and evaluated their potential role in causing endogenous genetic incompatibilities between these populations. We focussed on a Chironomus‐specific TE, the minisatellite‐like Cla‐element, whose activity is associated with speciation in the genus. Using a newly generated and annotated draft genome for a genomic study with five natural C. riparius populations, we found highly population‐specific TE insertion patterns with many private insertions. A significant correlation of the pairwise F_ST estimated from genomewide single‐nucleotide polymorphisms (SNPs) and the F_ST estimated from TEs is consistent with drift as the major force driving TE population differentiation. However, the significantly higher Cla‐element F_ST level due to a high proportion of differentially fixed Cla‐element insertions also indicates selection against segregating (i.e. heterozygous) insertions. With reciprocal crossing experiments and fluorescent in situ hybridization of Cla‐elements to polytene chromosomes, we documented phenotypic effects on female fertility and chromosomal mispairings. We propose that the inferred negative selection on heterozygous Cla‐element insertions may cause endogenous genetic barriers and therefore acts as DMI among C. riparius populations. The intrinsic genomic turnover exerted by TEs may thus have a direct impact on population divergence that is operationally different from drift and local adaptation.     

EDENetworks: A user‐friendly software to build and analyse networks in biogeography,ecology and population genetics

The recent application of graph‐based network theory analysis to biogeography, community ecology and population genetics has created a need for user‐friendly software, which would allow a wider accessibility to and adaptation of these methods. EDENetworks aims to fill this void by providing an easy‐to‐use interface for the whole analysis pipeline of ecological and evolutionary networks starting from matrices of species distributions, genotypes, bacterial OTUs or populations characterized genetically. The user can choose between several different ecological distance metrics, such as Bray‐Curtis or Sorensen distance, or population genetic metrics such as F_ST or Goldstein distances, to turn the raw data into a distance/dissimilarity matrix. This matrix is then transformed into a network by manual or automatic thresholding based on percolation theory or by building the minimum spanning tree. The networks can be visualized along with auxiliary data and analysed with various metrics such as degree, clustering coefficient, assortativity and betweenness centrality. The statistical significance of the results can be estimated either by resampling the original biological data or by null models based on permutations of the data.     

The genomic footprint of climate adaptation in Chironomus riparius

The gradual heterogeneity of climatic factors poses varying selection pressures across geographic distances that leave signatures of clinal variation in the genome. Separating signatures of clinal adaptation from signatures of other evolutionary forces, such as demographic processes, genetic drift and adaptation, to nonclinal conditions of the immediate local environment is a major challenge. Here, we examine climate adaptation in five natural populations of the harlequin fly Chironomus riparius sampled along a climatic gradient across Europe. Our study integrates experimental data, individual genome resequencing, Pool‐Seq data and population genetic modelling. Common‐garden experiments revealed significantly different population growth rates at test temperatures corresponding to the population origin along the climate gradient, suggesting thermal adaptation on the phenotypic level. Based on a population genomic analysis, we derived empirical estimates of historical demography and migration. We used an F_ST outlier approach to infer positive selection across the climate gradient, in combination with an environmental association analysis. In total, we identified 162 candidate genes as genomic basis of climate adaptation. Enriched functions among these candidate genes involved the apoptotic process and molecular response to heat, as well as functions identified in studies of climate adaptation in other insects. Our results show that local climate conditions impose strong selection pressures and lead to genomic adaptation despite strong gene flow. Moreover, these results imply that selection to different climatic conditions seems to converge on a functional level, at least between different insect species.     

Polymorphism and Divergence of Multilocus DNA Markers in Sibling Species Chironomus riparius Meigen and Chironomus piger Strenzke (Diptera,Chironomidae)

Intra- and interspecific variation and divergence of multilocus markers for genomic DNA of the sibling species from the thimmi group,Chironomus riparius and C. piger, were studied by PCR with arbitrary primers (RAPD). A high level of RAPD polymorphism was determined in both laboratory and natural populations of these species. The genetic distances were estimated between the C. riparius populations and between the sibling species C. riparius and C. piger. The genetic distance between C. riparius andC. piger was 4 to 5 times higher than that between the C. riparius populations. A comparison of the variation and divergence for the RAPD markers with those for other genomic markers—enzyme-coding genes and chromosomes (gene linkage groups)—showed that different components of the genome differed in their contribution to the genome divergence.     

Genomic pedigree reconstruction identifies predictors of mating and reproductive success in an invasive vertebrate

The persistence of an invasive species is influenced by its reproductive ecology, and a successful control program must operate on this premise. However, the reproductive ecology of invasive species may be enigmatic due to factors that also limit their management, such as cryptic coloration and behavior. We explored the mating and reproductive ecology of the invasive Brown Treesnake (BTS: Boiga irregularis) by reconstructing a multigenerational genomic pedigree based on 654 single nucleotide polymorphisms for a geographically closed population established in 2004 on Guam (N = 426). The pedigree allowed annual estimates of individual mating and reproductive success to be inferred for snakes in the study population over a 14‐year period. We then employed generalized linear mixed models to gauge how well phenotypic and genomic data could predict sex‐specific annual mating and reproductive success. Average snout–vent length (SVL), average body condition index (BCI), and trappability were significantly related to annual mating success for males, with average SVL also related to annual mating success for females. Male and female annual reproductive success was positively affected by SVL, BCI, and trappability. Surprisingly, the degree to which individuals were inbred had no effect on annual mating or reproductive success. When juxtaposed with current control methods, these results indicate that baited traps, a common interdiction tool, may target fecund BTS in some regards but not others. Our study emphasizes the importance of reproductive ecology as a focus for improving BTS control and promotes genomic pedigree reconstruction for such an endeavor in this invasive species and others.     

Known Knowns,Known Unknowns,and Unknown Unknowns: Coverage in MS Experiments

A mathematical model from ecology, namely, the capture–recapture model with a closed population and time‐varying and heterogeneous individual probabilities of capture, is implemented to model the number of protein identifications across the various cycles of a mass spectroscopy experiment. Rcapture, a package available in the R computing environment, can easily provide estimates of the cardinality of the proteome from such experiments. Alternatively, model fitting can be undertaken in other software platforms, such as Matlab, that can accommodate general linear models. It has not escaped our notice that capture–recapture models can be more broadly applied to other settings, so as to estimate the number of missing observations in an experiment.     

Reproductive ecology of Ocotea catharinensis,an endangered tree species

• Ocotea catharinensis (Lauraceae) is an endangered tree species from the Brazilian Atlantic Rainforest. Currently, little is known about the reproductive ecology of this species. Aiming to propose conservation measures, we described aspects related to phenology, floral biology, pollination, seed dispersal and mating system of O. catharinensis.
• We conducted phenological observations in 62 individuals for 2 years. In one reproductive event, we evaluated nectar production, stigmatic receptivity and pollen viability. Floral visitors were observed, identified and classified on a scale of pollination effectiveness. Seed dispersers were observed and identified using camera traps. Finally, the mating system was evaluated through pollen/ovule ratios, experimental pollination treatments and genetic analysis with molecular markers.
• Ocotea catharinensis presented a supra‐annual fruiting pattern with a substantial reduction of reproducing individuals from bud phase to ripe fruit phase. Several mechanisms prompting cross‐fertilisation were identified, such as attractive, herkogamic and protogynic flowers. The main floral visitors and pollinators were from the Diptera order, and all seed dispersers were birds. The species presented a predominantly outcrossed mixed mating system with significant selfing rate (17.3%).
• Although based on restricted evidence, we hypothesised that selfing is an escape mechanism for situations unfavourable to cross‐fertilisation. Specifically, for the studied population selfing is a response to reduced population size, which is caused by the non‐reproduction of all potentially reproductive individuals and by past exploitation events. Therefore, conservation efforts should be able to enhance population sizes, as well as prevent overexploitation.

     

Oviposition habitat selection in response to risk of predation in temporary pools: mode of detection and consistency across experimental venue

Natural selection should favor females that avoid ovipositing where risk of predation is high for their progeny. Despite the large consequences of such oviposition behavior for individual fitness, population dynamics, and community structure, relatively few studies have tested for this behavior. Moreover, these studies have rarely assessed the mode of detection of predators, compared responses in prey species that vary in vulnerability to predators, or tested for the behavior in natural habitats. In an outdoor artificial pool experiment, we tested the oviposition responses of two dipteran species, Culiseta longiareolata (mosquito) and Chironomus riparius (midge), to the hemipteran predator, Notonecta maculata. Both dipteran species have similar life history characteristics, but Culiseta longiareolata larvae are highly vulnerable to predation by Notonecta, while Chironomus riparius larvae are not. As their vulnerabilities would suggest, Culiseta longiareolata, but not Chironomus riparius, strongly avoided ovipositing in pools containing Notonecta. An experiment in natural rock pools assessing oviposition by Culiseta longiareolata in response to Notonecta maculata yielded an oviposition pattern highly consistent with that of the artificial pool experiment. We also demonstrated that the cue for oviposition avoidance by Culiseta longiareolata was a predator-released chemical: Notonecta water (without Notonecta replenishment) repelled oviposition for 8 days. Oviposition avoidance and mode of detection of the predator have important implications for how to assess the true impact of predators and for the use of commercially produced kairomones for mosquito control.     

All in the ears: unlocking the early life history biology and spatial ecology of fishes

Obtaining biological and spatial information of the early life history (ELH) phases of fishes has been problematic, such that larval and juvenile phases are often referred to as the ‘black box’ of fish population biology and ecology. However, a potent source of life‐history data has been mined from the earstones (otoliths) of bony fishes. We systematically reviewed 476 empirical papers published between 2005 and 2012 (inclusive) that used otoliths to examine fish ELH phases, which has been an area of increasing attention over this period. We found that otolith‐based research during this period could be split into two broad themes according to whether studies examined: (i) biological objectives related to intrinsic processes such as larval and juvenile age, growth and mortality, and/or (ii) spatial objectives, such as habitat use, dispersal and migration. Surprisingly, just 24 studies (5%) explored a combined biological–spatial objective by simultaneously exploiting biological and spatial information from otoliths, suggesting much more scope for such integrated research objectives to be answered via the use of multiple otolith‐based techniques in a single study. Mapping otolith analytical techniques across these two approaches revealed that otolith structural analysis was mainly used to investigate biological processes, while otolith chemical analyses were most often applied to spatial questions. Heavy skew in research effort was apparent across biomes, with most (62%) publications specific to marine species, despite comparable levels of species richness and the importance of freshwater taxa (just 15% of papers). Indeed, around 1% (380 species) of a possible 31400+ extant species were examined in our surveyed papers, with a strong emphasis on temperate marine species of commercial value. Potential model species for otolith‐based ELH ecology research are arising, with the eel genus Anguilla (24 studies) and the European anchovy Engraulis encrasicolis (14 studies) attracting more research effort than most other taxa. While there is a preponderance of common techniques (e.g. daily otolith increment counts, increment widths), novel techniques such as transgenerational marking and computed X‐ray tomography, are increasingly being applied in published studies. The application of an integrative approach based on a combination of emerging techniques and traditional methods holds promise for major advances in our understanding of ELH fish ecology and to shine light into the ‘black box’ of fish ecology.     

Leptospira spp., rotavirus,norovirus, and hepatitis E virus surveillance in a wild invasive golden‐headed lion tamarin (Leontopithecus chrysomelas; Kuhl, 1820) population from an urban park in Niterói,Rio de Janeiro,Brazil

The world currently faces severe biodiversity losses caused by anthropogenic activities such as deforestation, pollution, the introduction of exotic species, habitat fragmentation, and climate changes. Disease ecology in altered environments is still poorly understood. The golden‐headed lion tamarin (GHLT, Leontopithecus chrysomelas) is an endangered species that became invasive in an urban park in Niterói, Rio de Janeiro, Brazil. The initially few invasive GHLT individuals became hundreds, adapted to living in proximity to humans and domestic animals. These GHLTs were captured as part of a conservation project; some animals were translocated to Bahia and some were kept in captivity. This study tested 593 GHLT for Leptospira serology; 100 and 95 GHLT for polymerase chain reaction (PCR) toLeptospira and hepatitis E virus genotype 3 (HEV‐3), respectively, and 101 familiar groups for PCR to viruses (rotavirus A, norovirus GI and GII, and HEV‐3). One animal had antibodies for Leptospira serovar Shermani and another for serovar Hebdomadis. One saprophyticLeptospira was found by the 16S PCR and sequencing. Viruses were not detected in samples tested. Findings suggest that the epidemiological importance of such pathogens in this GHLT population is either low or nonexistent. These data are important to understand the local disease ecology, as well as monitoring a translocation project, and to contribute data for species conservation.     

Evolutionary determinants of population differences in population growth rate × habitat temperature interactions in Chironomus riparius

Little is known about intraspecific variation in fitness performance in response to thermal stress among natural populations and how this relates to evolutionary aspects of species ecology. In this study, population growth rate (PGR; a composite fitness measure) varied among five natural Chironomus riparius populations sampled across a climatic gradient when subjected to three temperature treatments reflecting the typical range of summer habitat temperatures (20, 24 and 28 °C). The variation could be explained by a complex model including effects of genetic drift, genetic diversity and adaptation to average temperature during the warmest month, in addition to experimental temperature. All populations suffered a decrease in PGR from 20 to 28 °C and ΔPGR was significantly correlated with the respective average habitat temperature in the warmest month—populations from warmer areas showing lower ΔPGR. This implies that long-term exposure to higher temperatures in the warmest month (the key reproductive period for C. riparius) is likely to be a key selective force influencing fitness at higher temperatures. A comparison of phenotypic divergence and neutral genetic differentiation revealed that one phenotypic trait—the number of fertile egg masses per female—appeared to be under positive selection in some populations. Our findings support a role for response to temperature selection along a climatic gradient and suggest population history is a key determinant of intraspecific fitness variation. We stress the importance of integrating different types of data (climatic, experimental, genetic) in order to understand the effects of global climate change on biodiversity.     

The influence of larval density, food availability and habitat longevity on the life history and population growth rate of the midge Chironomus riparius

Despite long‐standing interest in the forms and mechanisms of density dependence, these are still imperfectly understood. However, in a constant environment an increase in density must reduce per capita resource availability, which in turn leads to reduced survival, fecundity and somatic growth rate. Here we report two population experiments examining the density dependent responses under controlled conditions of an important indicator species, Chironomus riparius. The first experiment was run for 35 weeks and was started at low density with replicate populations being fed three different rations. Increased ration reduced generation time and increased population growth rate (pgr) but had no effect on survival, fecundity and female body weight in the first generation. In the second generation there was a six‐fold increase in generation time, presumably due to the greatly reduced per capita resource availability as the estimated initial densities of the second generation were 300 times greater than the first. Juvenile survival to emergence, fecundity, adult body weight and pgr declined by 90%, 75%, 35% and 99%, respectively. These large between‐generation effects may have obscured the effects of the threefold variation in ration, as only survival to emergence significantly increased with ration in the second generation. These results suggest that some chironomid larvae survive a reduction in resource availability by growing more slowly. In the ephemeral habitats sometimes occupied by C. riparius, the effects of population density may depend crucially on the longevity of the environment. A second experiment was therefore performed to measure pgr from six different starting densities over an eight‐week period. The relationship between pgr and density was concave, viewed from above. At densities above 16 larvae per cm², less than 1% of the population emerged and no offspring were produced. Under the conditions of experiment 2 – an 8‐week habitat lifespan – carrying capacity was estimated as 8 larvae per cm².     

Effects of soil characteristics and human disturbance on tiger beetle oviposition

Abstract 1. Understanding species‐specific habitat requirements is important in insect ecology. Co‐occurrence of closely related species can provide insight into microhabitat segregation. Tiger beetles are well suited for such studies, as they occur in multi‐species assemblages, exhibit species‐specific habitat requirements, and are sensitive to anthropogenic modifications of the environment. 2. This study investigated ovipositional microhabitat choice of two sympatric tiger beetles at Point Reyes National Seashore, California, U.S.A., and evaluates how human activity affects that choice. Cicindela oregona is widespread and common whereas C. hirticollis, an undescribed subspecies, is locally threatened. Laboratory experiments tested oviposition‐site selection of the two species in response to variations in moisture, pH, salinity, and grain size. Field experiments tested larval burrow location in response to experimental shading and experimental human disturbance. 3. Laboratory results indicated that while C. hirticollis females preferred a narrow range of moisture levels and fine sands, C. oregona females had more specific pH and salinity requirements and preferred coarser sands. They also differed in timing of activity. Preferences are probably related to prevention of fungal or bacterial infection and avoidance of desiccation and immersion. 4. In the field, significantly more larval burrows were found under experimental shading for both species. In addition, repeated human compaction of the sand significantly decreased the number of larval burrows. 5. Results suggest that protection of microhabitat, restoration of native plants, and limitation of human traffic are vital in the conservation management of tiger beetles.     

Social structure and nestmate discrimination in two species of Brachyponera ants distributed in Japan

The ponerine ant Brachyponera chinensis was introduced to the USA, where it has become invasive. Although various ecological data have been collected for B. chinensis populations in the USA, most aspects concerning the biology and ecology of native populations in Japan, a presumed origin, remain unknown. Here we investigated the social structure and nestmate discrimination in native populations of B. chinensis and a closely related species, B. nakasujii. Both species showed functional polygyny over seasons. Only in B. nakasujii was there a seasonal change in the numbers of queens and workers per nest. In arena tests, workers of neither species showed aggressive behaviors to conspecific non‐nestmates from the same population, and the mean aggression score did not increase with the distance between nests. However, some differences in non‐aggressive responses were detected between nestmate and non‐nestmate pairs in both species. In an experiment to introduce a single worker into a nest, B. chinensis accepted non‐nestmates with a high probability just like nestmates, whereas in B. nakasujii non‐nestmates were less accepted than nestmates. These findings suggest that native populations of B. chinensis already possess some of the key characteristics shared by many invasive exotic ants in introduced ranges, such as stable polygyny, weak internest aggression and acceptance of non‐nestmates. These tendencies are remarkable in comparison to the closely related B. nakasujii.     

Effect of Ascaridia compar infection on rock partridge population dynamics: empirical and theoretical investigations

Helminth parasites have the potential to significantly affect the dynamics of their hosts. As a consequence, they can dramatically threaten the persistence of endangered species, such as rock partridge Alectoris graeca saxatilis, found in the Province of Trento (northern Italy). The aim of this work was to understand the effect of helminth parasites on rock partridge fitness, and the subsequent potential effects on host population dynamics. In particular, we investigated the hypothesis that infections from Ascaridia compar induce rock partridge population cycles observed in Trentino. In order to support this hypothesis, we compared the predictions obtained from a host–parasite interaction model including variable parasite aggregation with multi‐annual empirical data of A. compar infection in natural host populations. We estimated host demographic parameters using rock partridge census data from Trentino, and the parasitological parameters from a series of experimental infections in a captive rock partridge population. The host–parasite model predicted higher A. compar abundance in rock partridge populations exhibiting cyclic dynamics compared to non‐cyclic ones. In addition, for cyclic host populations, the model predicted an increase in mean parasite burden with the length of cycle period. Model predictions were well‐supported by field data: significant differences in parasite infection between cyclic and non‐cyclic populations and among cyclic populations with different oscillation periods were observed. On the basis of these results, we conclude that helminth parasites can not be ruled out as drivers of rock partridge population dynamics in Trentino and must be considered when planning conservation strategies of this threatened species.     

Estimating population abundance in plant species with dormant life-stages: Fire and the endangered plant Grevillea caleyi R. Br.

Summary Assessment of the conservation significance of a species at a particular site involves estimating the population size. Generally this is based on a single survey. However, where plant species vary greatly in abundance in response to disturbance regimes, there will be uncertainty associated with the use of single estimates of abundance. The interpretation of such estimates is dependent on an understanding of the ecology of the species and the disturbance regimes that impact on it. We examined the usefulness of abundance estimates in the endangered shrub Grevillea caleyi (a fire‐sensitive shrub with a persistent soil seed bank) from south‐eastern Australia, where fire is a major landscape disturbance. Comparisons of estimates of abundance before and after fire showed very large changes in the number of plants of G. caleyi above ground. Changes in abundance of over two orders of magnitude were observed. The longer the site was left unburnt, the greater the magnitude of change in abundance after the next fire. Above‐ground plants may be rare or absent at sites unburnt for over 15–20 years, but were abundant after fire, due to re‐establishment from the soil seed bank. Sites burnt by two fires in quick succession showed declines in population abundance, most likely due to the soil seed bank not being replenished between such short interval fires. Assessments of the conservation significance of remnant sites of G. caleyi and similar species based on a single sample of above‐ground plant abundance at one time are considered inappropriate. The amount of available habitat for G. caleyi, either as area of occupancy or preferably extent of available habitat, was a moderate predictor of the likely magnitude of abundance in the species after fire. However, the usefulness of these measures for species whose biology is comparable to Grevillea caleyi, will be limited due to factors relating to the degree of species‐specific habitat requirements, local site fire history and the impact of any one fire on resultant post‐fire germination levels. Any assessment of conservation significance will require the interpretation of available information in relation to the ecology of a species.     

What do we know about armadillos? An analysis of four centuries of knowledge about a group of South American mammals,with emphasis on their conservation

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Ecologically relevant stress resistance: from microarrays and quantitative trait loci to candidate genes — A research plan and preliminary results using<Emphasis Type="Italic">Drosophila</Emphasis> as a model organism and climatic and genetic stress as model stresses

We aim at studying adaptation to genetic and environmental stress and its evolutionary implications at different levels of biological organization. Stress influences cellular processes, individual physiology, genetic variation at the population level, and the process of natural selection. To investigate these highly connected levels of stress effects, it is advisable - if not critical - to integrate approaches from ecology, evolution, physiology, molecular biology and genetics. To investigate the mechanisms of stress resistance, how resistance evolves, and what factors contribute to and constrain its evolution, we use the well-defined model systems ofDrosophila species, representing both cosmopolitan species such asD. melanogaster with a known genome map, and more specialized and ecologically well described species such as the cactophilicD. buzzatii. Various climate-related stresses are used as model stresses including desiccation, starvation, cold and heat. Genetic stress or genetic load is modelled by studying the consequences of inbreeding, the accumulation of (slightly) deleterious mutations, hybridization or the loss of genetic variability. We present here a research plan and preliminary results combining various approaches: molecular techniques such as microarrays, quantitative trait loci (QTL) analyses, quantitative PCR, ELISA or Western blotting are combined with population studies of resistance to climatic and genetic stress in natural populations collected across climatic gradients as well as in selection lines maintained in the laboratory.     

Contrasting diversity and demographic signals in sympatric narrow‐range endemic shrubs of the south‐west Western Australian semi‐arid zone

Comparative studies of sympatric species that integrate both phylogeographical and population genetic approaches provide insight into how demographic events and life history traits shape adaptive potential and drive species persistence. Such studies are rare for species‐rich and strongly structured environments, especially those of the southern hemisphere. For two sympatric, perennial shrubs of the south‐west Western Australian semi‐arid zone, Grevillea globosa and Mirbelia sp. Bursarioides, we assessed historical and contemporary genetic diversity and structure, demographic processes and ratios of pollen to seed dispersal. Phylogeographical structure was not detected and haplotype networks were star‐like. Number of haplotypes, nucleotide diversity, haplotype diversity, and allelic diversity were statistically significantly lower for G. globosa than for M. sp. Bursarioides. Levels of haplotype divergence and more contemporary genetic divergence and expected heterozygosity were lower for G. globosa than for M. sp. Bursarioides, but differences were not statistically significant. Both species exhibited signals of isolation by distance and low pollen to seed dispersal ratios (5.26:1 and 6.88:1). Grevillea globosa displayed signals of historical and contemporary demographic expansion. Results imply an important role for aspects of seed ecology that impact population demography, as well as direct dispersal and a significant contribution of seed dispersal to genetic connectivity in a semi‐arid landscape.