Seed germination and seedling allogamy in Rosmarinus officinalis: the costs of inbreeding

• Self‐pollination by geitonogamy is likely in self‐compatible plants that simultaneously expose a large number of flowers to pollinators. However, progeny of these plants is often highly allogamous. Although mechanisms to increase cross‐pollination have been identified and studied, their relative importance has rarely been addressed simultaneously in plant populations.
• We used Rosmarinus officinalis to explore factors that influence the probability of self‐fertilisation due to geitonogamy or that purge its consequences, focusing on their effects on seed germination and allogamy rate. We experimentally tested the effect of geitonogamy on the proportion of filled seeds and how it influences germination rate. During two field seasons, we studied how life history and flowering traits of individuals influence seed germination and allogamy rates of their progeny in wild populations at the extremes of the altitudinal range. The traits considered were plant size, population density, duration of the flowering season, number of open flowers, flowering synchrony among individuals within populations and proportion of male‐sterile flowers.
• We found that most seeds obtained experimentally from self‐pollination were apparently healthy but empty, and that the proportion of filled seeds drove the differences in germination rate between self‐ and cross‐pollination experiments. Plants from wild populations consistently had low germination rate and high rate of allogamy, as determined with microsatellites. Germination rate related positively to the length of the flowering season, flowering synchrony and the ratio of male‐sterile flowers, whereas the rate of allogamous seedlings was positively related only to the ratio of male‐sterile flowers.
• Rosemary plants purge most of the inbreeding caused by its pollination system by aborting the seeds. This study showed that the rates of seed germination and allogamy of the seedlings depend on a complex combination of factors that vary in space and time. Male sterility of flowers, length of the flowering season and flowering synchrony of individuals within populations all favour high rates of cross‐pollination, therefore increasing germination and allogamy rates. Flowering traits appear to be highly plastic and respond to local and seasonal conditions.

     

Ecotypes of an ecologically dominant prairie grass (Andropogon gerardii) exhibit genetic divergence across the U.S. Midwest grasslands' environmental gradient

Big bluestem (Andropogon gerardii) is an ecologically dominant grass with wide distribution across the environmental gradient of U.S. Midwest grasslands. This system offers an ideal natural laboratory to study population divergence and adaptation in spatially varying climates. Objectives were to: (i) characterize neutral genetic diversity and structure within and among three regional ecotypes derived from 11 prairies across the U.S. Midwest environmental gradient, (ii) distinguish between the relative roles of isolation by distance (IBD) vs. isolation by environment (IBE) on ecotype divergence, (iii) identify outlier loci under selection and (iv) assess the association between outlier loci and climate. Using two primer sets, we genotyped 378 plants at 384 polymorphic AFLP loci across regional ecotypes from central and eastern Kansas and Illinois. Neighbour‐joining tree and PCoA revealed strong genetic differentiation between Kansas and Illinois ecotypes, which was better explained by IBE than IBD. We found high genetic variability within prairies (80%) and even fragmented Illinois prairies, surprisingly, contained high within‐prairie genetic diversity (92%). Using Bayenv 2, 14 top‐ranked outlier loci among ecotypes were associated with temperature and precipitation variables. Six of seven BayeScan F_ST outliers were in common with Bayenv 2 outliers. High genetic diversity may enable big bluestem populations to better withstand changing climates; however, population divergence supports the use of local ecotypes in grassland restoration. Knowledge of genetic variation in this ecological dominant and other grassland species will be critical to understanding grassland response and restoration challenges in the face of a changing climate.     

Standing variation boosted by multiple sources of introduction contributes to the success of the introduced species, Lotus corniculatus

Although ecological differences between native and introduced ranges have been considered to drive rapid expansion of invasive species, recent studies suggest that rapid evolutionary responses of invasive species to local environments may also be common. Such expansion across heterogeneous environments by adaptation to local habitats requires genetic variation. In this study, we investigated the source and role of standing variation in successful invasion of heterogeneous abiotic environments in a self-incompatible species, Lotus corniculatus. We compared phenotypic and genetic variation among cultivars, natives, and introduced genotypes, and found substantial genetic variation within both native and introduced populations. Introduced populations possessed genotypes derived from both cultivars and native populations, and had lower population differentiation, indicating multiple sources of introduction and population admixture among the sources in the introduced range. Both cultivars and introduced populations had similarly outperforming phenotypes on average, with increased biomass and earlier flowering compared with native populations, but those phenotypes were within the range of the variation in phenotypes of the native populations. In addition, clinal variation within introduced populations was detected along a climatic gradient. Multiple introductions from different sources, including cultivars, may have contributed to pre-adaptive standing variation in the current introduced populations. We conclude that both introduction of cultivar genotypes and natural selection in local environments contributed to current patterns of genetic and phenotypic variation observed in the introduced populations.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

The influence of population structure on gene expression and flowering time variation in the ubiquitous weed Capsella bursa‐pastoris (Brassicaceae)

Population structure is a potential problem when testing for adaptive phenotypic differences among populations. The observed phenotypic differences among populations can simply be due to genetic drift, and if the genetic distance between them is not considered, the differentiation may be falsely interpreted as adaptive. Conversely, adaptive and demographic processes might have been tightly associated and correcting for the population structure may lead to false negatives. Here, we evaluated this problem in the cosmopolitan weed Capsella bursa‐pastoris. We used RNA‐Seq to analyse gene expression differences among 24 accessions, which belonged to a much larger group that had been previously characterized for flowering time and circadian rhythm and were genotyped using genotyping‐by‐sequencing (GBS) technique. We found that clustering of accessions for gene expression retrieved the same three clusters that were obtained with GBS data previously, namely Europe, the Middle East and Asia. Moreover, the three groups were also differentiated for both flowering time and circadian rhythm variation. Correction for population genetic structure when analysing differential gene expression analysis removed all differences among the three groups. This may suggest that most differences are neutral and simply reflect population history. However, geographical variation in flowering time and circadian rhythm indicated that the distribution of adaptive traits might be confounded by population structure. To bypass this confounding effect, we compared gene expression differentiation between flowering ecotypes within the genetic groups. Among the differentially expressed genes, FLOWERING LOCUS C was the strongest candidate for local adaptation in regulation of flowering time.     

Can plants evolve tolerance mechanisms to heterospecific pollen effects? An experimental test of the adaptive potential in Clarkia species

Flowering plants do not occur alone and often grow in mixed‐species communities where pollinator sharing is high and interactions via pollinators can occur at pre‐ and post‐pollination stages. While the causes and consequences of pre‐pollination interactions have been well studied little is known about post‐pollination interactions via heterospecific pollen (HP) receipt, and even less about the evolutionary implications of these interactions. In particular, the degree to which plants can evolve tolerance mechanisms to the negative effects of HP receipt has received little attention. Here, we aim to fill this gap in our understanding of post‐pollination interactions by experimentally testing whether two co‐flowering Clarkia species can evolve HP tolerance, and whether tolerance to specific HP ‘genotypes’ (fine‐scale local adaptation to HP) occurs. We find that Clarkia species vary in their tolerance to HP effects. Furthermore, conspecific pollen performance and the magnitude of HP effects were related to the recipient's history of exposure to HP in C. xantiana but not in C. speciosa. Specifically, better conspecific pollen performance and smaller HP effects were observed in populations of C. xantiana plants with previous exposure to HP compared to populations without such exposure. These results suggest that plants may have the potential to evolve tolerance mechanisms to HP effects but that these may occur not from the female (stigma, style) but from the male (pollen) perspective, a possibility that is often overlooked. We find no evidence for fine‐scale local adaptation to HP receipt. Studies that evaluate the adaptive potential of plants to the negative effects of HP receipt are an important first step in understanding the evolutionary consequences of plant–plant post‐pollination interactions. Such knowledge is in turn crucial for deciphering the role of plant–pollinator interactions in driving floral evolution and the composition of co‐flowering communities.     

Climate envelope modelling reveals intraspecific relationships among flowering phenology, niche breadth and potential range size in Arabidopsis thaliana

Species often harbour large amounts of phenotypic variation in ecologically important traits, and some of this variation is genetically based. Understanding how this genetic variation is spatially structured can help to understand species' ecological tolerances and range limits. We modelled the climate envelopes of Arabidopsis thaliana genotypes, ranging from early- to late-flowering, as a function of several climatic variables. We found that genotypes with contrasting alleles at individual flowering time loci differed significantly in potential range size and niche breadth. We also found that later flowering genotypes had more restricted range potentials and narrower niche breadths than earlier flowering genotypes, indicating that local selection on flowering can constrain or enhance the ability of populations to colonise other areas. Our study demonstrates how climate envelope models that incorporate ecologically important genetic variation can provide insights into the macroecology of a species, which is important to understand its responses to changing environments.     

Geographic variation of reproductive traits and competition for pollinators in a bird‐pollinated plant

Geographic variation in the reproductive traits of animal‐pollinated plants can be shaped by spatially variable selection imposed by differences in the local pollination environment. We investigated this process in Babiana ringens (Iridaceae), an enigmatic species from the Western Cape region of South Africa. B. ringens has evolved a specialized perch facilitating cross‐pollination by sunbirds and displays striking geographic variation in perch size and floral traits. Here, we investigate whether this variation can be explained by geographic differences in the pollinator communities. We measured floral and inflorescence traits, and abiotic variables (N, P, C, and rainfall) and made observations of sunbirds in populations spanning the range of B. ringens. In each population, we recorded sunbird species identity and measured visitation rates, interfloral pollen transfer, and whether the seed set of flowers was pollen limited. To evaluate whether competition from co‐occurring sunbird‐pollinated species might reduce visitation, we quantified nectar rewards in B. ringens and of other co‐flowering bird‐pollinated species in local communities in which populations occurred. Variation in abiotic variables was not associated with geographical variation of traits in B. ringens. Malachite sunbirds were the dominant visitor (97% of visits) and populations with larger‐sized traits exhibited higher visitation rates, more between‐flower pollen transfer and set more seed. No sunbirds were observed in four populations, all with smaller‐sized traits. Sunbird visitation to B. ringens was not associated with local sunbird activity in communities, but sunbird visitation was negatively associated with the amount of B. ringens sugar relative to the availability of alternative nectar sources. Our study provides evidence that B. ringens populations with larger floral traits are visited more frequently by sunbirds, and we propose that visitation rates to B. ringens may be influenced, in part, by competition with other sunbird‐pollinated species.     

On the potential strength and consequences for nonrandom gene flow caused by local adaptation in flowering time

Gene flow is generally considered a random process, that is the loci under consideration have no effect on dispersal success. Edelaar and Bolnick (Trends Ecol Evol, 27, 2012 659) recently argued that nonrandom gene flow could exert a significant evolutionary force. It can, for instance, ameliorate the maladaptive effects of immigration into locally adapted populations. I examined the potential strength for nonrandom gene flow for flowering time genes, a trait frequently found to be locally adapted. The idea is that plants that successfully export pollen into a locally adapted resident population will be a genetically biased subset of their natal population – they will have resident‐like flowering times. Reciprocally, recipients will be more migrant‐like than the resident population average. I quantified the potential for biased pollen exchange among three populations along a flowering time cline in Brassica rapa from southern California. A two‐generation line cross experiment demonstrated genetic variance in flowering time, both within and among populations. Calculations based on the variation in individual flowering schedules showed that resident plants with the most migrant‐like flowering times could expect to have up to 10 times more of the their flowers pollinated by immigrant pollen than the least migrant‐like. Further, the mean flowering time of the pollen exporters that have access to resident mates differs by up to 4 weeks from the mean in the exporters’ natal population. The data from these three populations suggest that the bias in gene flow for flowering time cuts the impact on the resident population by as much as half. This implies that when selection is divergent between populations, migrants with the highest mating success tend to be resident‐like in their flowering times, and so, fewer maladaptive alleles will be introduced into the locally adapting gene pool.     

Positive effects of flower abundance and synchronous flowering on pollination success,and pollinia dispersal in rewardless Changnienia amoena (Orchidaceae)

Pollination success and pollen dispersal in natural populations depend on the spatial‐temporal variation of flower abundance. For plants that lack rewards for pollinators, pollination success is predicted to be negatively related to flower density and flowering synchrony. We investigated the relationships between pollination success and flower abundance and flowering synchrony, and estimated pollinia dispersal distance in a rewardless species, Changnienia amoena (Orchidaceae). The results obtained in the present study revealed that male pollination success was negatively influenced by population size but was positively affected by population density, whereas female pollination success was independent of both population size and density. Phenotypic analysis suggested that highly synchronous flowering was advantageous through total pollination success, which is in contrast to previous studies. These results indicate that pollination facilitation rather than competition for pollinator visits occurs in this rewardless plant. The median distance of pollinia dispersal was 11.5 m (mean distance = 17.5 m), which is comparable to that of other rewardless plants but longer than for rewarding plants. However, pollen transfer occured mainly within populations; pollen import was a rare event. Restricted gene flow by pollinia and seeds probably explains the previous population genetic reporting a high degree of genetic differentiation between populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 477–488.     

Breeding system and population genetics of the vulnerable plant Dillwynia tenuifolia (Fabaceae)

Dillwynia tenuifolia Sieber ex DC. is a vulnerable species endemic to the Sydney region in Australia. This investigation focused on the breeding system and genetic structuring of the species in locations encompassing its entire geographical range. A field pollination experiment showed that D. tenuifolia produced fruit from both self‐ and cross‐pollination events, and thus has a facultative breeding system. Fruiting success from self‐ pollination was approximately half that observed from cross‐pollination. The genetic structure of D. tenuifolia populations was investigated by using allozyme electrophoresis. A survey of 16 loci revealed a high level of genetic variation with approximately two‐thirds of the loci being polymorphic and a mean of 1.84 alleles per loci. The genotype frequencies deviated from the Hardy–Weinberg equilibrium, with the observed level of heterozygosity significantly less than expected. The inbreeding coefficient was 0.31. There was very little divergence between populations (F_ST = 0.04) but the fixation indices were high within populations. Spatial autocorrelation, using Moran’s I, showed that neighbouring plants were closely related (I > 0.4). The genetic neighbourhood and effective population size in the species were estimated at one location as 1.1 ha and 10⁴ plants, respectively.     

Reproductive ecology of three endangered African violet (Saintpaulia H. Wendl.) species in the East Usambara Mountains, Tanzania

Knowledge of the reproductive biology of endangered plants is essential for their effective conservation. It also provides important information for understanding the evolutionary processes that affect speciation, thus helping the definition of proper units for conservation in endangered plants with problematic taxonomy. We studied the reproductive potential and possibility for hybridization in the endangered genus Saintpaulia (Gesneriaceae) by examining flowering phenology, flower and seed production and pollination of three sympatric cross‐compatible Saintpaulia species in the East Usambara Mts., Tanzania. The synchrony observed in flowering in S. confusa and S. difficilis may enable hybridization between these two species, whereas partial phenological separation may contribute to the integrity of S. grotei. Although the level of flower abortion is high in S. confusa, each pollinated flower yields about 1000 seeds. Saintpaulia confusa produces fruits following both self‐ and cross‐pollination but spontaneous self‐pollination seems not to occur. Thus, seed production depends on sufficient pollinator service. Floral heteromorphy (i.e. enantiostyly) and bee pollination are likely to further enhance cross‐pollination, suggesting that the genus predominantly outcrosses. Thus, Saintpaulia populations are likely to suffer from negative effects of inbreeding if they become small and isolated.     

Landscape genetic patterns of the rainbow darter Etheostoma caeruleum: a catchment analysis of mitochondrial DNA sequences and nuclear microsatellites

Catchment population structure and divergence patterns of the rainbow darter Etheostoma caeruleum (Percidae: Teleostei), an eastern North American benthic fish, are tested using a landscape genetics approach. Allelic variation at eight nuclear DNA microsatellite loci and two mitochondrial DNA regions [cytochrome (cyt) b gene and control region; 2056 aligned base pairs (bp)] is analysed from 89 individuals and six sites in the Lake Erie catchment (Blanchard, Chagrin, Cuyahoga and Grand Rivers) v. the Ohio River catchment (Big Darby Creek and Little Miami River). Genetic and geographic patterning is assessed using phylogenetic trees, pair‐wise F_ST analogues, AMOVA partitioning, Mantel regression, Bayesian assignment, 3D factorial correspondence and barrier analyses. Results identify 34 cyt b haplotypes, 22 control region haplotypes and 137 microsatellite alleles whose distributions demonstrate marked genetic divergence between populations from the Lake Erie and Ohio River catchments. Etheostoma caeruleum populations in the Lake Erie and Ohio River catchments diverged c. 1·6 mya during the Pleistocene glaciations. Greater genetic separations characterize the Ohio River populations, reflecting their older habitat age and less recent connectivity. Divergence levels within the Lake Erie catchment denote more recent post‐glacial origins. Notably, the western Lake Erie Blanchard River population markedly differs from the three central basin tributary samples, which are each genetically distinguishable using microsatellites. Overall relationships among the Lake Erie sites refute a genetic isolation by geographic distance hypothesis. Etheostoma caeruleum populations thus exchange few genes and have low migration among tributaries and catchments.     

Habitat restoration and native grass conservation: a case study of switchgrass (Panicum virgatum)

Switchgrass (Panicum virgatum) has been planted extensively for habitat restoration across the United States, such as with the Conservation Reserve Program (CRP). However, genetic profiles of these populations have never been studied nor compared with those of remnant prairies or cultivars. In this study, we sampled 16 CRP and 17 prairie populations across eastern Kansas. We assessed ploidy levels of all populations and compared genetic diversity and structure of 10 prairies, 10 CRP areas, and 5 standard cultivars, using nine simple sequence repeat (SSR) DNA markers. All CRP and prairie populations were octaploid (8x), except two prairies with both hexaploid (6x) and octaploid (8x) individuals. Based on the results of SSR analyses, there were no significant differences between CRP and prairie populations in genetic diversity, and 94% of total variation was partitioned within populations. Similarities among prairie and CRP populations were also observed in Bayesian clustering algorithms and principal coordinate analysis, suggesting that they had similar genetic compositions. In addition, positive spatial autocorrelations were detected up to 42 and 46 km among prairie and among CRP populations, respectively. To conclude, the CRP and prairie populations shared similar genetic profiles. However, remnant prairies still harbored unique genotypes and a high level of genetic diversity, highlighting the importance of seed sources for restoration efforts, that is using local wild seeds or cultivars from the same geographical region. A popular tetraploid (4x) cultivar known as “Kanlow” was genetically distinct from the prairie populations and therefore is not recommended for habitat restoration in this region.     

Phenotypic selection on flowering phenology and pollination efficiency traits between Primula populations with different pollinator assemblages

Floral traits have largely been attributed to phenotypic selection in plant–pollinator interactions. However, the strength of this link has rarely been ascertained with real pollinators. We conducted pollinator observations and estimated selection through female fitness on flowering phenology and floral traits between two Primula secundiflora populations. We quantified pollinator‐mediated selection by subtracting estimates of selection gradients of plants receiving supplemental hand pollination from those of plants receiving open pollination. There was net directional selection for an earlier flowering start date at populations where the dominant pollinators were syrphid flies, and flowering phenology was also subjected to stabilized quadratic selection. However, a later flowering start date was significantly selected at populations where the dominant pollinators were legitimate (normal pollination through the corolla tube entrance) and illegitimate bumblebees (abnormal pollination through nectar robbing hole which located at the corolla tube), and flowering phenology was subjected to disruptive quadratic selection. Wider corolla tube entrance diameter was selected at both populations. Furthermore, the strength of net directional selection on flowering start date and corolla tube entrance diameter was stronger at the population where the dominant pollinators were syrphid flies. Pollinator‐mediated selection explained most of the between‐population variations in the net directional selection on flowering phenology and corolla tube entrance diameter. Our results suggested the important influence of pollinator‐mediated selection on floral evolution. Variations in pollinator assemblages not only resulted in variation in the direction of selection but also the strength of selection on floral traits.     

Trait variation in response to varying winter temperatures,diversity patterns and signatures of selection along the latitudinal distribution of the widespread grassland plant Arrhenatherum elatius

Across Europe, genetic diversity can be expected to decline toward the North because of stochastic and selective effects which may imply diminished phenotypic variation and less potential for future genetic adaptations to environmental change. Understanding such latitudinal patterns can aid provenance selection for breeding or assisted migration approaches. In an experiment simulating different winter temperatures, we assessed quantitative trait variation, genetic diversity, and differentiation for natural populations of the grass Arrhenatherum elatius originating from a large latitudinal gradient. In general, populations from the North grew smaller and had a lower flowering probability. Toward the North, the absolute plastic response to the different winter conditions as well as heritability for biomass production significantly declined. Genetic differentiation in plant height and probability of flowering were very strong and significantly higher than under neutral expectations derived from SNP data, suggesting adaptive differentiation. Differentiation in biomass production did not exceed but mirrored patterns for neutral genetic differentiation, suggesting that migration‐related processes caused the observed clinal trait variation. Our results demonstrate that genetic diversity and trait differentiation patterns for A. elatius along a latitudinal gradient are likely shaped by both local selection and genetic drift.     

Two reproductive traits show contrasting genetic architectures in Plantago lanceolata

In many species, temperature‐sensitive phenotypic plasticity (i.e., an individual's phenotypic response to temperature) displays a positive correlation with latitude, a pattern presumed to reflect local adaptation. This geographical pattern raises two general questions: (a) Do a few large‐effect genes contribute to latitudinal variation in a trait? (b) Is the thermal plasticity of different traits regulated pleiotropically? To address the questions, we crossed individuals of Plantago lanceolata derived from northern and southern European populations. Individuals naturally exhibited high and low thermal plasticity in floral reflectance and flowering time. We grew parents and offspring in controlled cool‐ and warm‐temperature environments, mimicking what plants would encounter in nature. We obtained genetic markers via genotype‐by‐sequencing, produced the first recombination map for this ecologically important nonmodel species, and performed quantitative trait locus (QTL) mapping of thermal plasticity and single‐environment values for both traits. We identified a large‐effect QTL that largely explained the reflectance plasticity differences between northern and southern populations. We identified multiple smaller‐effect QTLs affecting aspects of flowering time, one of which affected flowering time plasticity. The results indicate that the genetic architecture of thermal plasticity in flowering is more complex than for reflectance. One flowering time QTL showed strong cytonuclear interactions under cool temperatures. Reflectance and flowering plasticity QTLs did not colocalize, suggesting little pleiotropic genetic control and freedom for independent trait evolution. Such genetic information about the architecture of plasticity is environmentally important because it informs us about the potential for plasticity to offset negative effects of climate change.     

Patterns of introduction and adaptation during the invasion of Aegilops triuncialis (Poaceae) into Californian serpentine soils

Multiple introductions can play a prominent role in explaining the success of biological invasions. One often cited mechanism is that multiple introductions of invasive species prevent genetic bottlenecks by parallel introductions of several distinct genotypes that, in turn, provide heritable variation necessary for local adaptation. Here, we show that the invasion of Aegilops triuncialis into California, USA, involved multiple introductions that may have facilitated invasion into serpentine habitats. Using microsatellite markers, we compared the polymorphism and genetic structure of populations of Ae. triuncialis invading serpentine soils in California to that of accessions from its native range. In a glasshouse study, we also compared phenotypic variation in phenological and fitness traits between invasive and native populations grown on loam soil and under serpentine edaphic conditions. Molecular analysis of invasive populations revealed that Californian populations cluster into three independent introductions (i.e. invasive lineages). Our glasshouse common garden experiment found that all Californian populations exhibited higher fitness under serpentine conditions. However, the three invasive lineages appear to represent independent pathways of adaptation to serpentine soil. Our results suggest that the rapid invasion of serpentine habitats in California may have been facilitated by the existence of colonizing Eurasian genotypes pre‐adapted to serpentine soils.     

The evolution of novel host use is unlikely to be constrained by trade‐offs or a lack of genetic variation

The genetic and ecological factors that shape the evolution of animal diets remain poorly understood. For herbivorous insects, the expectation has been that trade‐offs exist, such that adaptation to one host plant reduces performance on other potential hosts. We investigated the genetic architecture of alternative host use by rearing individual Lycaeides melissa butterflies from two wild populations in a crossed design on two hosts (one native and one introduced) and analysing the genetic basis of differences in performance using genomic approaches. Survival during the experiment was highest when butterfly larvae were reared on their natal host plant, consistent with local adaptation. However, cross‐host correlations in performance among families (within populations) were not different from zero. We found that L. melissa populations possess genetic variation for larval performance and variation in performance had a polygenic basis. We documented very few genetic variants with trade‐offs that would inherently constrain diet breadth by preventing the optimization of performance across hosts. Instead, most genetic variants that affected performance on one host had little to no effect on the other host. In total, these results suggest that genetic trade‐offs are not the primary cause of dietary specialization in L. melissa butterflies.     

Annual variation in floral phenology and pollen production in a 25‐year‐old plantation of Tectona grandis

Teak is a timber tree that is widely distributed in the tropics. Several studies on pollination and reproductive biology have been conducted, but generally information on flowering phenology and annual variation in total pollen production per tree is lacking. The reproductive phenology as well as flower‐ and pollen grain production of individuals in a population is important to theoreticians, field biologists and plant breeders, as they determine the distribution of genotypes within populations and influences the degree of differentiation among populations. This study reports flowering phenology and variation in total flower, fruit‐ and pollen production per tree in teak in a 25‐year‐old plantation across three consecutive years (2006–2008). The results show that the date of onset and end of flowering was highly variable across years. The longest flowering period of 93 days was observed in 2006. There was an asynchrony in the number of open flowers due to differences in time of anthesis among individuals (± 2 days) and inflorescences within individuals (± 6 h). The production of pollen grains per tree in 2007 was 33%, i.e. 16% more compared to 2006 and 2008. The fruit production per tree was 42% and 27% higher in 2007 compared to 2006 and 2008. Concentration of pollen grains (both on jelly‐coated microscopic slides and stigmas) were highest between noon and 2 pm. At this time, the stigmatic pollen load ranged between 4–8 pollen grains per stigma, which is sufficient for fruit development. The study concludes that the asynchronization of the flower opening might give rise to a high amount of self‐pollination in the stand, ultimately leading to poor fruit setting. Also, the large production of flowers and pollen per tree induced geitonogamy and decreased female fitness, as T. grandis is preferentially an out‐crossing species.