Quantitative Genetic Differentiation Research Articles

Adaptive Trait Variation in Seedlings of Rare Endemic Mexican Spruce Provenances under Nursery Conditions

The distributions of the three Mexican spruces are fragmented, possibly leading to phenological, morphological and genetic differentiation, which is partly caused by local adaptation. In this study, we estimated for the first time the intra- and inter-specific phenotypic variation in 5641 seedlings from provenances of the three Mexican spruces. We examined (i) provenance-related differences in the seedling survival rate, diameter (D), height (H) and seed weight (SW) as quantitative traits, (ii) the association between the survival rate, D, H and SW and climatic and soil variables in the Picea provenances and (iii) (narrow-sense) heritability (within-provenance) based on D and H under the same nursery conditions, assuming that the response can be considered as a proxy for quantitative genetic differentiation between provenances. All Mexican spruce species differed significantly in H, and all eight provenances studied were significantly different in D and H, except for two neighboring provenances of P. mexicana. Very strong, significant correlations (up to R2 = 0.96) were found between H, the survival rate and SW with respect to environmental factors of provenance/seed origin. Additionally, the heritability index explained a high percentage of the provenance-related variance. The use of germplasm for restoration in different sites and with different populations requires collecting seeds from numerous trees from as many provenances as possible, but should be carried out with caution owing to the apparently strong local adaptation in provenances of the Mexican spruces.

Molecular and quantitative genetic variation within and between populations of the declining grassland species Saxifraga granulata.

Formerly common plant species are expected to be particularly susceptible to recent habitat fragmentation. We studied the population genetics of 19 recently fragmented Saxifraga granulata populations (max. distance 61 km) in Luxembourg and neighboring Germany using RAPD markers and a common garden experiment. We assessed (1) the relationships between plant fitness, quantitative genetic variation, molecular genetic variation, and population size; and (2) the relative importance of genetic drift and selection in shaping genetic variation. Molecular genetic diversity was high but did not correlate with population size, habitat conditions, or plant performance. Genetic differentiation was low (F ST=0.079 ± 0.135), and there was no isolation by distance. Longevity, clonality, and the long-lived seed bank of S.granulata may have prevented strong genetic erosion and genetic differentiation among populations. However, genetic distinctness increased with decreasing genetic diversity indicating that random genetic drift occurred in the studied populations. Quantitative and molecular genetic variations were correlated, and their differentiation (Q ST vs. F ST) among S.granulata populations was similar, suggesting that mainly random processes have shaped the quantitative genetic differentiation among populations. However, pairwise quantitative genetic distances increased with geographic and climatic distances, even when adjusted for molecular genetic distances, indicating diversifying selection. Our results indicate that long-lived clonal species may be buffered at least temporarily against the negative effects of fragmentation. The relationship between quantitative genetic and geographic distance may be a more sensitive indicator of selection than Q ST-F ST differences.

Climate-Driven Adaptive Differentiation in Melia azedarach: Evidence from a Common Garden Experiment.

Studies of local adaptation in populations of chinaberry (Melia azedarach L.) are important for clarifying patterns in the population differentiation of this species across its natural range. M. azedarach is an economically important timber species, and its phenotype is highly variable across its range in China. Here, we collected M. azedarach seeds from 31 populations across its range and conducted a common garden experiment. We studied patterns of genetic differentiation among populations using molecular markers (simple sequence repeats) and data on phenotypic variation in six traits collected over five years. Our sampled populations could be subdivided into two groups based on genetic analyses, as well as patterns of isolation by distance and isolation by environment. Significant differentiation in growth traits was observed among provenances and families within provenances. Geographic distance was significantly correlated with the quantitative genetic differentiation (QST) in height (HEIT) and crown breadth. Climate factors were significantly correlated with the QST for each trait. A total of 23 climatic factors were examined. There was a significant effect of temperature on all traits, and minimum relative humidity had a significant effect on the survival rate over four years. By comparing the neutral genetic differentiation (FST) with the QST, the mode of selection acting on survival rate varied, whereas HEIT and the straightness of the main trunk were subject to the same mode of selection. The variation in survival rate was consistent with the variation in genetic differentiation among populations, which was indicative of local adaptation. Overall, our findings provide new insights into the responses of the phenological traits of M. azedarach to changes in the climate conditions of China.

Evolution of plant drought strategies and herbivore tolerance after two decades of climate change.

Ongoing global warming, coupled with increased drought frequencies, together with other biotic drivers may have resulted in complex evolutionary adaptation. The resurrection approach, comparing ancestors raised from stored seeds with their contemporary descendants under common conditions, is a powerful method to test for recent evolution in plant populations. We used 21-26-yr-old seeds of four European plant species - Matthiola tricuspidata, Plantago crassifolia, Clinopodium vulgare and Leontodon hispidus - stored in seed banks together with re-collected seeds from their wild populations. To test for evolutionary changes, we conducted a glasshouse experiment that quantified heritable changes in plant responses to drought and simulated insect herbivory. In three out of the four studied species, we found evidence that descendants had evolved shorter life cycles through faster growth and flowering. Shifts in the osmotic potential and leaf dry matter content indicated that descendants also evolved increased drought tolerance. A comparison of quantitative genetic differentiation (QST ) vs neutral molecular differentiation (FST ) values, using double digest restriction-site associated DNA (ddRAD) genotyping data, suggested that directional selection, and therefore adaptive evolution, was underlying some of the observed phenotypic changes. In summary, our study revealed evolutionary changes in plant populations over the last decades that are consistent with adaptation of drought escape and tolerance as well as herbivory avoidance.

Potential adaptive divergence between subspecies and populations of snapdragon plants inferred from QST -FST comparisons.

Phenotypic divergence among natural populations can be explained by natural selection or by neutral processes such as drift. Many examples in the literature compare putatively neutral (F ST) and quantitative genetic (Q ST) differentiation in multiple populations to assess their evolutionary signature and identify candidate traits involved with local adaptation. Investigating these signatures in closely related or recently diversified species has the potential to shed light on the divergence processes acting at the interspecific level. Here, we conducted this comparison in two subspecies of snapdragon plants (eight populations of Antirrhinum majus pseudomajus and five populations of A. m. striatum) in a common garden experiment. We also tested whether altitude was involved with population phenotypic divergence. Our results identified candidate phenological and morphological traits involved with local adaptation. Most of these traits were identified in one subspecies but not the other. Phenotypic divergence increased with altitude for a few biomass‐related traits, but only in A. m. striatum. These traits therefore potentially reflect A. m. striatum adaptation to altitude. Our findings imply that adaptive processes potentially differ at the scale of A. majus subspecies.

Plasticity to drought and ecotypic differentiation in populations of a crop wild relative

Populations of widely distributed species often exhibit geographic variation in functional traits in response to environmental heterogeneity. Such trait variation may be the result of different adaptive mechanisms, including genetically based differentiation, phenotypic plasticity or a combination of both. Disentangling the genetic and environmental components of trait variation may be particularly interesting in crop wild relatives, since they may provide unique reservoirs of genetic diversity for crop improvement. In this study, we assessed ecotypic differentiation and patterns of plasticity to drought in populations of Lupinus angustifolius, a Mediterranean crop wild relative, from two climatically distinct regions in the Iberian Peninsula. Using an outdoor common garden, we compared phenotypic responses of inbred maternal families to two ecologically meaningful water availability treatments (drought and high-moisture). We measured 18 different functional traits related to growth, morphology, phenology and reproduction. Plants in the drought treatment grew less, had lower leaf chlorophyll content and photochemical efficiency, but also reproduced faster, produced larger seeds and altered leaflet morphology through increased leaflet thickness, higher leaflet dry matter content and lower specific leaf area. We also found significant differences between regions that likely reflect adaptation to climatically distinct environments, with populations from the south showing a faster onset of reproduction, higher leaf thickness and higher seed size, consistent with the drier conditions experienced in southern sites. Plasticity to drought was in most cases in the same direction as quantitative genetic differentiation (i.e. cogradient variation), providing evidence of the adaptive value of the plastic change. Our results show that both genetic differentiation and plasticity can generate adaptive phenotypic variation in L. angustifolius, and help to identify potentially valuable genetic resources to incorporate into breeding programmes.

Adaptive evolution of chestnut forests to the impact of ink disease in Spain

AbstractPhytophthora cinnamomi (Pc) is an extremely destructive soil‐borne pathogen of Asiatic origin responsible for “ink disease” in chestnut. This work assesses the adaptive potential to the impact of Pc of four Spanish populations of Castanea sativa undergoing different selection pressures. To explore the evolvability of C. sativa to Pc in the selected populations, parameters obtained from neutral and functional genetic diversity were compared with estimates of quantitative genetic variability. Nine expressed sequence tags‐simple sequence repeat (EST‐SSR) markers were selected and their transferability and polymorphism in 137 C. sativa individuals were evaluated. To test the potential of EST‐SSR markers for early selection of Pc tolerant plant material, the offspring of selected individuals were challenged with Pc. Expressed sequence tags‐simple sequence repeat markers and seedling life expectancy after Pc inoculation revealed significant different responses of C. sativa populations to Pc. The genetic variability observed within populations showed the potential response capacity of Spanish C. sativa populations to undergo fast adaptive evolution. The heritability value obtained for the “life expectancy” variable (h2 = 0.21 ± 0.11) indicated that selection for resistance to Pc is possible. Genetic patterns reflected two evolutionarily meaningful groupings of populations, corresponding to the different selective pressure of the oomycete between sites. The differentiation coefficient obtained through markers classified as under neutral selection (FST = 0.185) was lower than the quantitative genetic differentiation of “life expectancy” between C. sativa populations (QST = 0.682), providing evidence that selection acted spatially in a heterogeneous manner. A first link has been identified in trees between population structure and adaptive responses to pathogen‐induced selection. The study identified one marker under positive selection that could be used in marker assisted selection to predict resistance to Pc in non‐inoculated C. sativa trees.

Effects of marker type and filtering criteria on Q ST-F ST comparisons.

Comparative studies of quantitative and neutral genetic differentiation (QST-FST tests) provide means to detect adaptive population differentiation. However, QST-FST tests can be overly liberal if the markers used deflate FST below its expectation, or overly conservative if methodological biases lead to inflated FST estimates. We investigated how marker type and filtering criteria for marker selection influence QST-FST comparisons through their effects on FST using simulations and empirical data on over 18 000 in silico genotyped microsatellites and 3.8 million single-locus polymorphism (SNP) loci from four populations of nine-spined sticklebacks (Pungitius pungitius). Empirical and simulated data revealed that FST decreased with increasing marker variability, and was generally higher with SNPs than with microsatellites. The estimated baseline FST levels were also sensitive to filtering criteria for SNPs: both minor alleles and linkage disequilibrium (LD) pruning influenced FST estimation, as did marker ascertainment. However, in the case of stickleback data used here where QST is high, the choice of marker type, their genomic location, ascertainment and filtering made little difference to outcomes of QST-FST tests. Nevertheless, we recommend that QST-FST tests using microsatellites should discard the most variable loci, and those using SNPs should pay attention to marker ascertainment and properly account for LD before filtering SNPs. This may be especially important when level of quantitative trait differentiation is low and levels of neutral differentiation high.

Genetic variation in the constitutive defensive metabolome and its inducibility are geographically structured and largely determined by demographic processes in maritime pine

Abstract Interspecific phenotypic variation in plant secondary metabolites (PSM) is often explained by biotic and abiotic factors. However, patterns of variation within species do not clearly fit the theoretical predictions. Exploring how genetics, environment and demographic processes shape such variation among and within populations is crucial for understanding evolution of PSM, particularly in long‐lived plants such as forest trees. Here, we quantified genetic variation in PSM among and within populations, and explored drivers of local adaptation by studying the role of climate as a source of population differentiation in PSM of maritime pine. Constitutive profile and concentrations of 63 PSM and their herbivory‐associated inducibility were determined in the bark of 130 clonally replicated genotypes with known familial structure from 10 populations covering the distribution range of the species. We compared neutral and quantitative population genetic differentiation of PSM (FST and QST). Also, we accounted for population genetic structure and kinship among individuals when exploring climate–trait relationships. We found large population differentiation and additive genetic variation in constitutive PSM. Many PSM were inducible, although very low genetic variation was observed with respect to their inducibility. QST–FST comparisons suggest that differentiation of most diterpenes, monoterpenes, and phenolics can be explained by neutral demographic processes. Spatially heterogeneous selection across populations leading to local adaptation was only found for total constitutive sesquiterpenes and a few individual PSM. After accounting for population genetic structure, only the constitutive concentration of two sesquiterpenes showing signs of diversifying selection was predicted by climate, with decreasing concentrations along a growth‐prone climatic gradient. Synthesis. Evolutionary patterns of plant secondary metabolites depended on their chemical nature, with neutral differentiation governing most plant secondary metabolites. Evidence of local adaptation was only found for total constitutive sesquiterpenes and a few individual plant secondary metabolites. The low genetic variation in the inducibility of plant secondary metabolites suggests a conserved model of defensive induction in this species. Since population differentiation linked to past demographic history could lead to false positives of adaptive differentiation signals, accounting for the genetic relatedness among populations is required to infer the environmental determinants of intraspecific genetic variation in putatively adaptive traits such as plant defences.

Adaptive diversity and drought tolerance in Castanea sativa assessed through EST-SSR genic markers

Increasing drought conditions in Mediterranean countries are negatively impacting the survival and productivity of Castanea sativa Mill. The study aimed to select EST-SSR markers associated with drought stress developed in Quercus spp. and evaluate their transferability and polymorphism in C. sativa. Eight EST-SSR markers were selected to examine the adaptive potential of four wild populations of C. sativa in relation to drought tolerance. To validate markers, offspring of the study trees were water stressed and their drought tolerance was assessed. EST-SSR markers and leaf wilting of seedlings after drought treatment revealed a north–south gradient of C. sativa populations. The heritability value obtained for the ‘leaf wilting’ trait (h² = 0.26 ± 0.08) indicated that selection for drought tolerance is possible. The differentiation coefficient of markers showing neutral selection (FST = 0.080) was lower than the quantitative genetic differentiation of populations (QST = 0.28), indicating that selection of drought tolerant trees acted spatially in a heterogeneous manner. When assessing the genetic structure of populations, FIR080 was identified as outlier locus under positive selection. When assessing the phenotypic tolerance to drought of offspring, GOT004 and GOT045 were identified as outlier loci under balancing selection and FIR059 was identified as an outlier locus under positive selection. FIR059 showed three private alleles for drought-susceptible individuals and two private alleles for drought-tolerant individuals and could therefore be considered as a candidate marker to predict drought tolerance in unstressed C. sativa trees. Combined use of functional markers and phenotypic traits is a powerful approach to determine genetic variation at the adaptive level in C. sativa. The results illustrate the potential of EST-SSR markers for early selection of drought tolerant plant material.

Mediterranean and Northern Iberian gene pools of wild Castanea sativa Mill. are two differentiated ecotypes originated under natural divergent selection.

Nine wild Iberian provenances of Castanea sativa Mill. grouped in two gene pools, North Iberian Peninsula and Mediterranean, were evaluated for several adaptive traits in two provenance–progeny trials with the aim of evaluating the role of natural selection in shaping adaptive variation and increasing our understanding of the genetic structure of this species, as well as reporting complete information on the genetic variation among and within the studied populations. An annual growth rhythm experiment was evaluated during the first 3 years after establishment for phenology, growth, stem form and survival, and a periodic drought-stress experiment was evaluated for dry weight, growth, survival and other related drought traits in both well-watered and drought-stress treatments. The high genetic variability reported in both trials is largely due to the genetic variation among populations. The significant differences reported between quantitative genetic and neutral marker differentiation indicated the local adaptation of these populations through directional selection, mainly for phenology, growth and biomass allocation. A clinal variation among populations was determined through correlations of phenology with latitude and xerothermic index of the provenances, showing that central and southern Mediterranean populations had earlier phenology than northern populations and that drought played a relevant role in this differentiation. The significant correlation between phenological traits and the ancestry values in the Mediterranean gene pool supported the different pattern of behavior between both gene pools and also indicated the existence of two ecotypes: xeric and mesophytic ecotypes, corresponding to Mediterranean and North Iberian gene pools, respectively. The results obtained in the drought-stress experiment confirmed that, in general terms, xeric populations showed a greater adaptability to drought, with more developed root systems and higher survival than northern populations. Moreover, the genetic variability observed within populations indicated the potential response capacity of Iberian C. sativa populations to undergo fast adaptive evolution.

Geographic clines in wing morphology relate to colonization history in New World but not Old World populations of yellow dung flies.

Geographic clines offer insights about putative targets and agents of natural selection as well as tempo and mode of adaptation. However, demographic processes can lead to clines that are indistinguishable from adaptive divergence. Using the widespread yellow dung fly Scathophaga stercoraria (Diptera: Scathophagidae), we examine quantitative genetic differentiation (QST ) of wing shape across North America, Europe, and Japan, and compare this differentiation with that of ten microsatellites (FST ). Morphometric analyses of 28 populations reared at three temperatures revealed significant thermal plasticity, sexual dimorphism, and geographic differentiation in wing shape. In North America morphological differentiation followed the decline in microsatellite variability along the presumed route of recent colonization from the southeast to the northwest. Across Europe, where S. stercoraria presumably existed for much longer time and where no molecular pattern of isolation by distance was evident, clinal variation was less pronounced despite significant morphological differentiation (QST >FST ). Shape vector comparisons further indicate that thermal plasticity (hot-to-cold) does not mirror patterns of latitudinal divergence (south-to-north), as might have been expected under a scenario with temperature as the major agent of selection. Our findings illustrate the importance of detailed phylogeographic information when interpreting geographic clines of dispersal traits in an adaptive evolutionary framework.

Life history traits, but not body size, vary systematically along latitudinal gradients on three continents in the widespread yellow dung fly

Large‐scale clinal variation in body size and other life‐history traits is common enough to have stimulated the postulation of several eco‐geographical rules. Whereas some clinal patterns are clearly adaptive, the causes of others remain unclear. We present a comprehensive intraspecific population comparison for the cosmopolitan yellow dung fly Scathophaga stercoraria (Diptera: Scathophagidae) to check for consistent world‐wide patterns. Common garden assessment of various life history traits permitted continental comparison of (clinal) quantitative genetic differentiation (Qst) with putatively neutral genetic differentiation (Fst) derived from field‐caught flies. Latitudinal clines in fly development time, growth rate, and overwintering propensity were consistent among North American, European and Japanese populations. Increased winter dormancy incidence and duration at higher latitude, combined with a faster growth rate and shorter development time, suggest that flies are adaptated to season length more than to temperature. The resulting body size clines, in contrast, were not very consistent; importantly, they were not negative, as expected under seasonal constraints, but flat or even positive clines. Quantitative genetic differentiation QST exceeded neutral molecular variation FST for most traits, suggesting that natural selection plays a consistent role in mediating global dung fly life histories. We conclude that faster growth and development in response to shorter growing seasons at higher latitudes may indirectly counteract expected direct effects of temperature on body‐size, potentially resulting in flat or inconsistent body size clines in nature.

Genetic differentiation, local adaptation and phenotypic plasticity in fragmented populations of a rare forest herb.

BackgroundDue to habitat loss and fragmentation, numerous forest species are subject to severe population decline. Investigating variation in genetic diversity, phenotypic plasticity and local adaptation should be a prerequisite for implementing conservation actions. This study aimed to explore these aspects in ten fragmented populations of Physospermum cornubiense in view of translocation measures across its Italian range.MethodsFor each population we collected environmental data on landscape (habitat size, quality and fragmentation) and local conditions (slope, presence of alien species, incidence of the herbivorous insect Metcalfa pruinosa and soil parameters). We measured vegetative and reproductive traits in the field and analysed the genetic population structure using ISSR markers (STRUCTURE and AMOVA). We then estimated the neutral (FST) and quantitative (PST) genetic differentiation of populations.ResultsThe populations exhibited moderate phenotypic variation. Population size (range: 16–655 individuals), number of flowering adults (range: 3–420 individuals) and inflorescence size (range: 5.0–8.4 cm) were positively related to Mg soil content. Populations’ gene diversity was moderate (Nei-H = 0.071–0.1316); STRUCTURE analysis identified five different clusters and three main geographic groups: upper, lower, and Apennine/Western Po plain. Fragmentation did not have an influence on the local adaptation of populations, which for all measured traits showed PST < FST, indicating convergent selection.DiscussionThe variation of phenotypic traits across sites was attributed to plastic response rather than local adaptation. Plant translocation from suitable source populations to endangered ones should particularly take into account provenance according to identified genetic clusters and specific soil factors.

Quantitative genetic differentiation and phenotypic plasticity of European beech in a heterogeneous landscape: Indications for past climate adaptation

Tree growth and species distributions are expected to be altered by climate change. European beech (Fagus sylvatica), one of the major tree species in Central Europe, is considered to be particularly threatened by the expected changes in local water regimes. Basic knowledge on the species⬢ genetic variation, environmental adaptation, and phenotypic plasticity is required to assess its potential for climate change adaptation, but sufficient information is lacking. Here, we describe a seedling common garden study at two field sites incorporating 77 natural populations of European beech from an environmentally heterogeneous mountain region in the center of its distribution. We aimed to identify patterns of genetic variation and phenotypic plasticity in growth and phenology, and to associate these with seed source environments. Population differentiation was greater for phenology than for growth (Qst=0.18⬜0.32 vs. 0.00⬜0.16), but within-population genetic variation was large for all seedling traits. The phenotype-environment associations indicated adaptive divergence in phenology and growth with respect to temperature and water availability, but not to soil characteristics, latitude, longitude, or topography of the seed source locations. Phenotypic plasticity was detected in growth and leaf duration, the magnitude of which differed among populations with different seed source temperatures. We conclude that seedling phenology is key to temperature and drought adaptation in European beech. Changes in local temperature and water regimes might result in local phenological maladaptation of European beech populations, although within-population genetic variation, gene flow, and phenotypic plasticity might mitigate the negative effects of climate change.

Adaptive genetic variability and differentiation of Croatian and Austrian Quercus robur L. populations at a drought prone field trial

Provenance trials, where populations of different geographical origin are tested in a common environment (common garden test), are a tool suited to allow the study of intraspecific adaptive genetic variation. Research of pedunculate oak (Quercus robur L.) adaptive genetic variability through analyses of populations in common garden tests has a long tradition. However, pedunculated oak populations originating south-eastern from the Alps have been scarcely studied in this way. This study addresses the adaptive genetic variability and differentiation of pedunculate oak populations originating from Austria and Croatia in a provenance/progeny field trial. Studied plants were six years old and were growing at the trial for three years. After two years of unusually low precipitations height and survival were analysed. The total mean height of all plants in the trial was 137.8 cm and ranged from 123.0 cm to 151.8 cm. The overall mean survival rate was rather high (0.85). Mean population survival ranged from 0.64 to 0.94. Individual narrow-sense heritabilities (hi2), family mean heritabilities (hf2), the coefficients of additive genetic variation (CVA) and quantitative genetic differentiation coefficients (QST) were calculated. A multivariate regression tree (MRT) analysis was used to determine the pattern of genetic differentiation of the populations. Individual heritabilities for height ranged between 0.00 and 0.39. Family mean heritabilities for height were rather low in most populations as well (<0.5). Family mean heritabilities for survival were higher than for height (ranging between 0.00 and 0.77). Calculated QST coefficients (0.25 for height and 0.14 for survival) indicated between-population genetic differentiation. The populations were separated into two clusters by MRT analysis regarding a climatic variable, namely Hargreaves’ reference evapotranspiration. Populations originating from comparatively more humid habitats were grouped in the first cluster. The first cluster had a lower mean height and survival compared to the second one. The differences between these clusters were highly statistically significant. The observed quantitative genetic differentiation might have been driven by natural selection caused by differences in the relative moisture of the habitats from which the progeny populations originate. The results suggest ecotypic pattern of the quantitative genetic differentiation among studied populations.

Genetic determination of tannins and herbivore resistance in Quercus ilex

Genetic variability of trees influences the chemical composition of tissues. This determines herbivore impact and, consequently, herbivore performance. We evaluated the independent effects of plant genotype and provenance on the tannin content of holm oak (Quercus ilex) and their consequences for herbivory and performance of gypsy moth (Lymantria dispar) larvae. Oak seedlings of 48 open-pollinated families from six populations were grown in a common garden in central Spain. Half the plants were subjected to defoliation by gypsy moth larvae and the other half were destructively sampled for chemical analysis. Tannin content of leaves did not differ significantly among populations but differed significantly among families. Estimates of heritability (h 2) and quantitative genetic differentiation among populations for tannin content (Q ST) were 0.83 and 0.12, respectively. Defoliation was not related to the tannin content of plants nor to spine and trichome densities of leaves, although positive family–mean associations were observed between defoliation and both seed weight and plant height (P < 0.003). Among the oak populations, differential increase in larval weight gain with defoliation was observed. Leaf tannin content in Q. ilex is genetically controlled but does not influence defoliation or predict performance of the larvae. Different efficiencies of food utilisation depending on the oak genotypes indicate that other plant traits are influencing the feeding patterns and fitness of L. dispar and consequent population dynamics.

Capturing neutral and adaptive genetic diversity for conservation in a highly structured tree species.

Preserving intraspecific genetic diversity is essential for long-term forest sustainability in a climate change scenario. Despite that, genetic information is largely neglected in conservation planning, and how conservation units should be defined is still heatedly debated. Here, we use maritime pine (Pinus pinaster Ait.), an outcrossing long-lived tree with a highly fragmented distribution in the Mediterranean biodiversity hotspot, to prove the importance of accounting for genetic variation, of both neutral molecular markers and quantitative traits, to define useful conservation units. Six gene pools associated to distinct evolutionary histories were identified within the species using 12 microsatellites and 266 single nucleotide polymorphisms (SNPs). In addition, height and survival standing variation, their genetic control, and plasticity were assessed in a multisite clonal common garden experiment (16 544 trees). We found high levels of quantitative genetic differentiation within previously defined neutral gene pools. Subsequent cluster analysis and post hoc trait distribution comparisons allowed us to define 10 genetically homogeneous population groups with high evolutionary potential. They constitute the minimum number of units to be represented in a maritime pine dynamic conservation program. Our results uphold that the identification of conservation units below the species level should account for key neutral and adaptive components of genetic diversity, especially in species with strong population structure and complex evolutionary histories. The environmental zonation approach currently used by the pan-European genetic conservation strategy for forest trees would be largely improved by gradually integrating molecular and quantitative trait information, as data become available.

Population size is weakly related to quantitative genetic variation and trait differentiation in a stream fish.

How population size influences quantitative genetic variation and differentiation among natural, fragmented populations remains unresolved. Small, isolated populations might occupy poor quality habitats and lose genetic variation more rapidly due to genetic drift than large populations. Genetic drift might furthermore overcome selection as population size decreases. Collectively, this might result in directional changes in additive genetic variation (VA ) and trait differentiation (QST ) from small to large population size. Alternatively, small populations might exhibit larger variation in VA and QST if habitat fragmentation increases variability in habitat types. We explored these alternatives by investigating VA and QST using nine fragmented populations of brook trout varying 50-fold in census size N (179-8416) and 10-fold in effective number of breeders, Nb (18-135). Across 15 traits, no evidence was found for consistent differences in VA and QST with population size and almost no evidence for increased variability of VA or QST estimates at small population size. This suggests that (i) small populations of some species may retain adaptive potential according to commonly adopted quantitative genetic measures and (ii) populations of varying sizes experience a variety of environmental conditions in nature, however extremely large studies are likely required before any firm conclusions can be made.

Genomic architecture of phenotypic divergence between two hybridizing plant species along an elevational gradient.

Knowledge of the genetic basis of phenotypic divergence between species and how such divergence is caused and maintained is crucial to an understanding of speciation and the generation of biodiversity. The hybrid zone between Senecio aethnensis and S. chrysanthemifolius on Mount Etna, Sicily, provides a well-studied example of species divergence in response to conditions at different elevations, despite hybridization and gene flow. Here, we investigate the genetic architecture of divergence between these two species using a combination of quantitative trait locus (QTL) mapping and genetic differentiation measures based on genetic marker analysis. A QTL architecture characterized by physical QTL clustering, epistatic interactions between QTLs, and pleiotropy was identified, and is consistent with the presence of divergent QTL complexes resistant to gene flow. A role for divergent selection between species was indicated by significant negative associations between levels of interspecific genetic differentiation at mapped marker gene loci and map distance from QTLs and hybrid incompatibility loci. Within-species selection contributing to interspecific differentiation was evidenced by negative associations between interspecific genetic differentiation and genetic diversity within species. These results show that the two Senecio species, while subject to gene flow, maintain divergent genomic regions consistent with local selection within species and selection against hybrids between species which, in turn, contribute to the maintenance of their distinct phenotypic differences.