Reproductive isolation and hybrid pollen disadvantage in Ipomopsis

One cause of reproductive isolation is gamete competition, in which conspecific pollen has an advantage over heterospecific pollen in siring seeds, thereby decreasing the formation of F1 hybrids. Analogous pollen interactions between hybrid pollen and conspecific pollen can contribute to post-zygotic isolation. The herbaceous plants Ipomopsis aggregata and I. tenuituba frequently hybridize in nature. Hand-pollination of I. aggregata with pollen from F1 or F2 hybrids produced as many seeds as hand-pollination with conspecific pollen, suggesting equal pollen viability. However, when mixed pollen loads with 50% conspecific pollen and 50% hybrid pollen were applied to I. aggregata stigmas, fewer than half of the seeds had hybrid sires. Such pollen mixtures are frequently received if plants of the two species and F1 and F2 hybrids are intermixed, suggesting that this advantage of conspecific over hybrid pollen reduces backcrossing and contributes to reproductive isolation.     

Reproductive isolation in Caenorhabditis: terminal phenotypes of hybrid embryos

SUMMARY Several interspecific combinations of the "elegans" group of Caenorhabditis species are cross-fertile. Most F1 hybrids from these crosses arrest during embryogenesis. Developmental defects observed in hybrid embryos include defects in gastrulation initiation, defects in embryonic compaction, and defects in embryonic elongation. These reproductive barriers have arisen multiple times in the evolution of Caenorhabditis.     

Reproductive biology in an invasive plant Solidago canadensis

Solidago canadensis,a perennial Compositae plant originating from North America,was introduced into China as a horticultural plant in 1935.Under natural conditions,S.canadensis allocates large amounts of energy to sexual reproduction and produces many seeds,which reflects an r-strategy with high seed number and small seed size.In addition,naturalized populations have a great capacity to grow clonally with underground stems.S.canadensis has become an invasive weed in eastern China,and has caused serious damages to agricultural production and ecosystems in several provinces in China.In order to understand the reproductive characteristics of S.canadensis and effectively control its spread,we examined soil conditions,seed characteristics,seed germination and the capacity for asexual reproduction in different plant parts.We investigated the population dispersion of S.canadensis in fixed sites for three years,and analyzed the seasonal dynamics of the morphological parameters of the underground parts and the caloric values of different organs of S.canadensis.We also compared differences in the root systems of S.canadensis and composite exotic weeds.The following results were obtained:1)Under natural conditions,the germination season of S.canadensis lasts from March to October,with a peak from April to May.Vegetative growth and asexual reproduction are especially vigorous during summer due to high temperatures and soil drought stress.On the other hand,the rainy season proves suitable for seed germination.Most S.canadensis flower between September and January,and fruit in late October.A mature plant can produce about 20000 seeds.The mean weight of 1000 seeds ranges from 0.045 g to 0.050 g,and the mean seed moisture content ranges from 60% to 80%.The light-winged seeds disperse readily by air,water,vehicles,human activity or through livestock.2)S.canadensis seeds have a wide tolerance for different values of pH,salinity and soil moisture.The mean percent germination of seeds is 30% under suitable conditions.The results of seed germination under various environmental stresses and investigation of soil conditions indicate that well-aerated,slightly acidic soils with low salinity are suitable for the growth of S.canadensis.Additionally,S.canadensis has a high tolerance for contamination by heavy metal elements including Zn,Cu and Pb,but has low accumulation coefficients for these elements.3)S.canadensis reproduces asexually via underground rhizomes and nodes on the stem base to recruit new individuals,and in plants that experience mechanical damage,this reproductive strategy is used to produce clonal shoots.The capacity for asexual reproduction among different plant parts rank as follows:underground parts＞stem-base (20 cm)＞stem-base (30 cm)＞stem-base (45 cm)＞stem.Further,with increasing mechanical damage,the quantity of shoots produced by the plant decreases.4)The morphological parameters of the root system of S.canadensis including length,surface area,volume,and average diameter are greater than for composite exotic weeds.These parameters indicate that S.canadensis has the physiological potential to widely invade China.5)The aboveground growth rate and most of the underground morphological parameters vary remarkably among the seasons,with a peak normally occurring in September.In August,a fraction of the energy in leaves and stems is allocated underground to increase fine root growth and water uptake during hot weather.Additionally,the seasonal dynamics of the underground morphological parameters and the caloric values of different organs of S.canadensis enhance its reproductive ability.Based on the results above,we conclude that S.canadensis has great invasive potential in China.We suggest that urgent measures should be taken to control its further spread,and to minimize its impact on local plant diversity.     

Multiple herbivores and coevolutionary interactions in an Ipomopsis hybrid swarm

Studies focusing on pairwise interactions between plants and herbivores may not give an accurate picture of the overall selective effect of herbivory, given that plants are often eaten by a diverse array of herbivore species. The outcome of such interactions may be further complicated by the effects of plant hybridization. Hybridization can lead to changes in morphological, phenological and chemical traits that could in turn alter plant–herbivore interactions. Here we present results from manipulative field experiments investigating the interactive effects of multiple herbivores and plant hybridization on the reproductive success of Ipomopsis aggregata formosissima X I. tenuituba. Results showed that ungulate herbivores alone had a net positive effect on plant relative fitness, increasing seed production approximately 2-fold. Caterpillars had no effect on plant relative fitness when acting alone, with caterpillar-attacked plants producing the same number of flowers, fruits and seeds as the uneaten controls. Caterpillars, however, significantly reduced flower production of ungulate browsed plants. Flower production in these plants, however, was still significantly greater (approximately 1.7-fold greater) than uneaten controls, likely leading to an increase in reproductive success through the paternal component of fitness given that fruit and seed production was not significantly different from that of herbivore-free controls. Although results suggest that herbivore imposed selection is pairwise, ungulates likely have a large influence on the abundance of, and hence the amount of damage caused by, caterpillar herbivores. Thus, because of the ecological interactions between ungulates and caterpillars, selection on Ipomopsis may be diffuse rather than pairwise, assuming such interactions translate into differential effects on plant fitness as herbivore densities vary. Plant hybridization had no significant effect on patterns of ungulate or caterpillar herbivory; i.e., no significant interactions were detected between herbivory and plant hybridization for any of the fitness traits measured in this study nor did plant hybridization have any significant effect on host preference. These results may be due to patterns of introgression or the lack of species-specific differences between I. aggregate formosissima and I. tenuituba. Plant hybridization per se resulted in lowered reproductive success of white colored morphs due in part to the effects of pollination. Although it appears that there would be strong directional selection favoring darker flower colors due to the lower reproductive success of the white colored morphs in the short run, the natural distribution of hybrids suggest that over the long run selection either tends to average out or there are no fitness differences among morphs in most years due to the additive fitness effects of hawkmoth and hummingbird pollinators.     

Adaptive plasticity and niche expansion in an invasive thistle

Phenotypic differentiation in size and fecundity between native and invasive populations of a species has been suggested as a causal driver of invasion in plants. Local adaptation to novel environmental conditions through a micro‐evolutionary response to natural selection may lead to phenotypic differentiation and fitness advantages in the invaded range. Local adaptation may occur along a stress tolerance trade‐off, favoring individuals that, in benign conditions, shift resource allocation from stress tolerance to increased vigor and fecundity and, therefore, invasiveness. Alternately, the typically disturbed invaded range may select for a plastic, generalist strategy, making phenotypic plasticity the main driver of invasion success. To distinguish between these hypotheses, we performed a field common garden and tested for genetically based phenotypic differentiation, resource allocation shifts in response to water limitation, and local adaptation to the environmental gradient which describes the source locations for native and invasive populations of diffuse knapweed (Centaurea diffusa). Plants were grown in an experimental field in France (naturalized range) under water addition and limitation conditions. After accounting for phenotypic variation arising from environmental differences among collection locations, we found evidence of genetic variation between the invasive and native populations for most morphological and life‐history traits under study. Invasive C. diffusa populations produced larger, later maturing, and therefore potentially fitter individuals than native populations. Evidence for local adaptation along a resource allocation trade‐off for water limitation tolerance is equivocal. However, native populations do show evidence of local adaptation to an environmental gradient, a relationship which is typically not observed in the invaded range. Broader analysis of the climatic niche inhabited by the species in both ranges suggests that the physiological tolerances of C. diffusa may have expanded in the invaded range. This observation could be due to selection for plastic, “general‐purpose” genotypes with broad environmental tolerances.     

Reproductive isolation and the period gene of Drosophila

The identification of genes of large effect on ecologically important traits is an important aim of molecular ecology. The period gene of Drosophila is a candidate for a gene with a large influence on premating isolation between Drosophila species, as it determines species specific aspects of courtship behaviour. Strains of D. melanogaster are available which have been genetically transformed with the period gene of either D. melanogaster or D. simulans. Here we show that D. melanogaster females do not discriminate between two such strains. This suggests that period may only make a small contribution to total premating isolation between these species. We discuss the use of genetically transformed strains in assessing the influence of single genes on complex traits.     

Reproductive isolation among two interfertile Rhododendron species: low frequency of post-F1 hybrid genotypes in alpine hybrid zones

Hybrids between the acid-loving species Rhododendron ferrugineum and the basic soil species Rhododendron hirsutum occur on soils of intermediate pH in the European Alps. Material from two hybrid zones ~500 m apart, and also nearby populations of each parent species, was surveyed for presence/absence of 31 random amplified polymorphic DNA markers that distinguish parents. Based on morphological assessment, the material comprised 51 putative hybrids, 18 putative R. ferrugineum individuals and 26 putative R. hirsutum plants. RAPD data were analysed using a Bayesian approach implemented by the program newhybrids , and also by principal coordinates analysis. The identity of all R. ferrugineum plants examined was confirmed; however, of the putative R. hirsutum individuals examined, two were certainly and 11 possibly hybrid derivatives. Among all hybrid derivatives examined, about half were designated as F₁s or a similar class, otherwise backcrosses to R. hirsutum appeared to be common whereas other hybrid classes were rare and backcrosses to R. ferrugineum possibly absent. Despite this, artificially generated seed of F₂ class and backcrosses in each direction showed greater viability than one parent ( R. hirsutum ). Introgression from R. ferrugineum was also detected in a population that from morphology appeared to contain only R. hirsutum . Hence, the direction of backcrossing might be highly asymmetric within hybrid zones, causing unidirectional gene flow from R. ferrugineum into R. hirsutum . Conversely, the rarity of backcrosses to R. ferrugineum , F₂s and later hybrid generations, which might be due to phenology effects and habitat-mediated selection, could play a part in restricting gene flow towards R. ferrugineum .     

Cultural isolation is greater than genetic isolation across an avian hybrid zone

Elucidating the relationship between genetic and cultural evolution is important in understanding speciation, as learned premating barriers might be involved in maintaining species differences. Here, we test this relationship by examining a widely recognized premating barrier, bird song, in a hybrid zone between black‐throated green (Setophaga virens) and Townsend's warblers (S. townsendi). We use song analysis, genomic techniques and playback experiments to characterize the cultural and genetic backgrounds of individuals in this region, expecting that if song is an important reproductive barrier between these species, there should be a strong relationship between song and genotype. We show that songs in the hybrid zone correspond to the distinctly different songs found in allopatry but that song and genotype are not tightly coupled in sympatry. Allopatric individuals responded only to local songs, indicating that individuals may have learned to respond to songs they commonly hear. We observed discordance between song and genotype clines; a narrower cline suggests that cultural selection on song is stronger than natural selection on genotype. These findings indicate that song is unlikely to play a role in reproductive isolation between these species, and we suggest that spatial variation in song may nonetheless be maintained by frequency‐dependent cultural selection. This decoupling of genes and culture may contribute to hybridization in this region.     

Reproductive isolation in a nascent species pair is associated with aneuploidy in hybrid offspring

Speciation may occur when the genomes of two populations accumulate genetic incompatibilities and/or chromosomal rearrangements that prevent inter-breeding in nature. Chromosome stability is critical for survival and faithful transmission of the genome, and hybridization can compromise this. However, the role of chromosomal stability on hybrid incompatibilities has rarely been tested in recently diverged populations. Here, we test for chromosomal instability in hybrids between nascent species, the ‘dwarf’ and ‘normal’ lake whitefish (Coregonus clupeaformis). We examined chromosomes in pure embryos, and healthy and malformed backcross embryos. While pure individuals displayed chromosome numbers corresponding to the expected diploid number (2n = 80), healthy backcrosses showed evidence of mitotic instability through an increased variance of chromosome numbers within an individual. In malformed backcrosses, extensive aneuploidy corresponding to multiples of the haploid number (1n = 40, 2n = 80, 3n = 120) was found, suggesting meiotic breakdown in their F₁ parent. However, no detectable chromosome rearrangements between parental forms were identified. Genomic instability through aneuploidy thus appears to contribute to reproductive isolation between dwarf and normal lake whitefish, despite their very recent divergence (approx. 15–20 000 generations). Our data suggest that genetic incompatibilities may accumulate early during speciation and limit hybridization between nascent species.     

Reproductive biology in an invasive plant Solidago canadensis

Solidago canadensis, a perennial Compositae plant originating from North America, was introduced into China as a horticultural plant in 1935. Under natural conditions, S. canadensis allocates large amounts of energy to sexual reproduction and produces many seeds, which reflects an r-strategy with high seed number and small seed size. In addition, naturalized populations have a great capacity to grow clonally with underground stems. S. canadensis has become an invasive weed in eastern China, and has caused serious damages to agricultural production and ecosystems in several provinces in China. In order to understand the reproductive characteristics of S. canadensis and effectively control its spread, we examined soil conditions, seed characteristics, seed germination and the capacity for asexual reproduction in different plant parts. We investigated the population dispersion of S. canadensis in fixed sites for three years, and analyzed the seasonal dynamics of the morphological parameters of the underground parts and the caloric values of different organs of S. canadensis. We also compared differences in the root systems of S. canadensis and composite exotic weeds. The following results were obtained:

Multiple mechanisms underlie rapid expansion of an invasive alien plant

? With growing concerns over serious ecological problems, a particular challenge is to reveal the complex mechanisms underlying rapid expansion of invasive species. Ageratina adenophora is of particular interest in addressing this question. ? We used geographic information systems and logistic regression to identify the geographic and environmental factors contributing to the presence of A. adenophora. Join-count spatial statistics with reproduction mode examination were employed to elucidate the spatiotemporal dispersal mechanisms. ? Multiple factors have significantly contributed to the rapid expansion of A. adenophora. Its biological traits, favoring dispersal by water and wind coupled with local spatiotemporally heterogeneous geography and ecology, promote invasion downstream and upstream along river valleys, while other factors associated with human activities facilitate its invasion over high mountains and across river valleys, providing new scope for progressive invasions. We further identified an unusual invasion event of A. adenophora subsequent to a great flood that amplified its dispersal ability from vegetative propagules and seeds. ? These findings suggest that dynamic interactions of multiple factors in heterogeneous ecogeographical environments - a 'combinatorial' invasion mechanism - would generate an unexpected invasion rate of an alien species or a seemingly stochastic invasion event.     

Reproductive success of cultivated Pyrus calleryana (Rosaceae) and establishment ability of invasive, hybrid progeny

?Premise of the study: Pyrus calleryana Decne., an ornamental tree species introduced from China, is a relatively new invasive that has only recently begun to spread across the United States after intraspecific hybridization between cultivars. The function of such hybridization in the evolution of invasiveness is still relatively understudied, especially with respect to the initial establishment and persistence of invasive genotypes. Multiple introductions of genetically divergent populations or cultivars may benefit from new genetic combinations created during hybridization events and/or release from Allee effects in founder populations. ?Methods: We quantified the outcome of intraspecific hybridization between cultivars of P. calleryana in a common garden. Measures of the reproductive success and establishment ability of their early- and advanced-generation hybrid offspring were collected to assess the likelihood of particular cultivar genotypes to establish in invasive populations. These traits also were compared between cultivated and invasive parents to identify any generational differences in invasive potential. ?Key results: Differences were detected in measures of reproductive ability, but no group emerged as consistently more fecund. Advanced-generation hybrids also had significantly less biomass, indicating a reduction in hybrid performance relative to that of the cultivated progeny. ?Conclusion: Ultimately, this study indicates that increased spread of P. calleryana has been initiated by introduction of multiple cultivar types and subsequent widespread planting and is not the result of an inherent fitness advantage of hybrid progeny.     

Reproductive isolation of a new hybrid species, Senecio eboracensis Abbott & Lowe (Asteraceae)

The nature and extent of reproductive isolation was examined between a new self-compatible hybrid species Senecio eboracensis (2n=40) and its parents, self-incompatible S. squalidus (2n=20) and self-compatible S. vulgaris (2n=40). The triploid F(1) of S. eboracensis x S. squalidus exhibited very low seed set (x=0.63%), and F(2) and F(3) progeny were able to recover nominal levels of fertility (x=23.9 and 9.7%), while F(1) and F(2) offspring of S. eboracensis x S. vulgaris showed reduced seed set (x=63.8 and 58.8%). In both cases, evidence from previous work indicates that reduced fertility is associated with meiotic chromosome mispairing, and is a likely consequence of recombining both parental genomes within this new taxon. No hybrid offspring between S. eboracensis and S. squalidus were found in the wild, and only one such hybrid was recorded among 769 progeny produced by S. eboracensis surrounded by S. squalidus on an experimental plot. Natural crossing between S. eboracensis and S. vulgaris was recorded to be very low (between 0 and 1.46%) in the wild, but rose to 18.3% when individuals of S. eboracensis were surrounded by plants of S. vulgaris. It was concluded that strong breeding barriers exist between the new hybrid species and its two parents. Prezygotic isolation between S. eboracensis and S. vulgaris is likely to be largely due to both species reproducing by predominant self-fertilisation. However, differences recorded for germination, seedling survival, time of flowering and characters associated with pollinator attraction, plus significant clumping of juvenile and adult conspecifics in the wild, probably also contribute to reproductive isolation and ecological differentiation.     

Fine-scale genetic structure arises during range expansion of an invasive gecko

Processes of range expansion are increasingly important in light of current concerns about invasive species and range shifts due to climate change. Theoretical studies suggest that genetic structuring may occur during range expansion. Ephemeral genetic structure can have important evolutionary implications, such as propagating genetic changes along the wave front of expansion, yet few studies have shown evidence of such structure. We tested the hypothesis that genetic structure arises during range expansion in Hemidactylus mabouia, a nocturnal African gecko recently introduced to Florida, USA. Twelve highly variable microsatellite loci were used to screen 418 individuals collected from 43 locations from four sampling sites across Florida, representing a gradient from earlier (～1990s) to very recent colonization. We found earlier colonized locations had little detectable genetic structure and higher allelic richness than more recently colonized locations. Genetic structuring was pronounced among locations at spatial scales of tens to hundreds of meters near the leading edge of range expansion. Despite the rapid pace of range expansion in this introduced gecko, dispersal is limited among many suitable habitat patches. Fine-scale genetic structure is likely the result of founder effects during colonization of suitable habitat patches. It may be obscured over time and by scale-dependent modes of dispersal. Further studies are needed to determine if such genetic structure affects adaptation and trait evolution in range expansions and range shifts.     

Modeling future range expansion and management strategies for an invasive squirrel species

Successful management of an invasive species requires in depth knowledge of the invader, the invaded ecosystem, and their interactions. The complexity of the species-system interactions can be reduced and represented in ecological models for better comprehension. In this study, a spatially explicit population model was created using the RAMAS software package to simulate the past and future invasion dynamics of the eastern grey squirrel (Sciurus carolinensis) in the fragmented habitat in case study areas in Ireland. This invasive squirrel species causes economic damage by bark stripping forest crops and is associated with the decline of its native congener (S. vulgaris). Three combinations of demographic and dispersal parameters, which best matched the distribution of the species shortly after introduction, were used to simulate invasion dynamics. Future population expansion was modeled under scenarios of no control and two different management strategies: fatal culls and immunocontraceptive vaccination programmes. In the absence of control, the grey squirrel range is predicted to expand to the south and southwest of Ireland endangering internationally important habitats, vulnerable forest crops, and the native red squirrel. The model revealed that region-wide intensive and coordinated culls would have the greatest impact on grey squirrel populations. Control strategies consisting solely of immunocontraceptive vaccines, often preferred by public interest groups, are predicted to be less effective. Complete eradication of the grey squirrel from Ireland is not economically feasible and strategic evidence-based management is required to limit further range expansion. Ecological models can be used to choose between informed management strategies based on predicted outcomes.     

Reproductive effort in invasive and non-invasive Rubus

We quantified the physiological costs and the total amount of resources allocated to reproduction in two closely related species of Rubus, one of which is invasive. These two species share several morphological and life-history characteristics and grow together in the Pacific Northwestern United States. Reproductive effort was manipulated in canes of both species by removing flower buds. The non-invasive species, R. ursinus, exhibited significantly greater water stress in the reproductive canes, as indicated by lower leaf water potential (O) and reduced stomatal conductance (g_s). This species also showed a reduction in leaf nitrogen concentration ([N]) associated with reproduction. Combined, these factors led to reduced photosynthesis (A) on a diurnal basis, lower water-use efficiency as inferred from '¹³C, and reduced photosynthetic capacity. All of these effects were more pronounced during the fruiting stage than in the flowering stage. The invasive species, R. discolor, showed no changes in water stress, [N], '¹³C, or A associated with reproduction. A model was used to estimate total gross photosynthesis (A_gross) for reproductive and non-reproductive canes of both species over cane lifetime. Reproduction was associated with a greater decline in A_gross for the non-invasive R. ursinus than for the invasive R. discolor. Although R. discolor allocated more resources directly to flowers and fruit than R. ursinus, the invasive species had significantly lower reproductive effort, or total amount of resources diverted from vegetative activity to reproduction, than the non-invasive species. By minimizing the reduction of photosynthesis associated with reproduction, this invasive species may be able to minimize the trade-offs commonly associated with reproduction.     

Rapid adaptive evolution of the diapause program during range expansion of an invasive mosquito

In temperate climates, the recurring seasonal exigencies of winter represent a fundamental physiological challenge for a wide range of organisms. In response, many temperate insects enter diapause, an alternative developmental program, including developmental arrest, that allows organisms to synchronize their life cycle with seasonal environmental variation. Geographic variation in diapause phenology contributing to local climatic adaptation is well documented. However, few studies have examined how the rapid evolution of a suite of traits expressed across the diapause program may contribute to climatic adaptation on a contemporary timescale. Here, we investigate the evolution of the diapause program over the past 35 years by leveraging a “natural experiment” presented by the recent invasion of the Asian tiger mosquito, Aedes albopictus, across the eastern United States. We sampled populations from two distinct climatic regions separated by 6° of latitude (∼700 km). Using common-garden experiments, we identified regional genetic divergence in diapause-associated cold tolerance, diapause duration, and postdiapause starvation tolerance. We also found regional divergence in nondiapause thermal performance. In contrast, we observed minimal regional divergence in nondiapause larval growth traits and at neutral molecular marker loci. Our results demonstrate rapid evolution of the diapause program and imply strong selection caused by differences in winter conditions.