Historical and ecological divergence among populations of Monttea chilensis (Plantaginaceae), an endemic endangered shrub bordering the Atacama Desert, Chile

Species delimitations by morphological and by genetic markers are not always congruent. Magnolia kobus consists of two morphologically different varieties, kobus and borealis. The latter variety is characterized by larger leaves than the former. For the conservation of M. kobus genetic resources in natural forests, the relationships between morphological and genetic variation should be clarified. We investigated variations in nuclear microsatellites, chloroplast DNA (cpDNA) sequences and leaf morphological traits in 23 populations of M. kobus over the range of species. Two genetically divergent lineages, northern and southern were detected and their geographical boundary was estimated to be at 39°N. The northern lineage consisted of two genetic clusters and a single cpDNA haplotype, while the southern one had multiple genetic clusters and cpDNA haplotypes. The northern lineage showed significantly lower genetic diversity than the southern. Approximate Bayesian computation indicated that the northern and southern lineages had experienced, respectively, population expansion and long-term stable population size. The divergence time between the two lineages was estimated to be 565,000 years ago and no signature of migration between the two lineages after divergence was detected. Ecological niche modeling showed that the potential distribution area in northern Japan at the last glacial maximum was very small. It is thus considered that the two lineages have experienced different population histories over several glacial-inter-glacial cycles. Individuals of populations in the central to northern part of Honshu on the Sea of Japan side and in Hokkaido had large leaf width and area. These leaf characteristics corresponded with those of variety borealis. However, the delimitation of the northern and southern lineages detected by genetic markers (39°N) was not congruent with that detected by leaf morphologies (36°N). It is therefore suggested that variety borealis is not supported genetically and the northern and southern lineages should be considered separately when identifying conservation units based not on morphology but on genetic markers.

Floral traits are commonly thought to be more canalized than vegetative ones. In addition, floral and vegetative traits are hypothesized to be genetically decoupled, enabling vegetative structures to respond plastically to environmental heterogeneity, and to evolve in response to selection without disrupting the reproductive function of flowers. To test these hypotheses, we evaluate the genetic architecture of floral and vegetative traits in natural populations of Arabidopsis thaliana raised under variable light-quality environments. Plants were grown either under high or low ratios of red to far-red (R:FR) light, an aspect of light quality that varies with neighbor proximity and regulates competitive shade-avoidance responses. Across environments, we detected significant genetic variation for the average expression of all measured floral traits (petal length and width, stamen length, pistil length, stigma-anther separation, and exsertion of both the stamen and pistil beyond the corolla). Light quality significantly influenced the absolute size of several floral traits as well as the allometry (i.e., relative scaling) of all floral traits, and genotypes differed in the plasticity of floral traits to the light treatments. Exposure to low relative to high R:FR resulted in significantly greater elongation in the vegetative trait, petiole length, and genotypes again differed in the plasticity of this trait to R:FR. Consistent with prior studies, most floral traits were less plastic than the vegetative trait; herkogamy (i.e., stigma-anther separation) was the exception and expressed more variable trait values across environments than petiole length, apparently as a consequence of the independent responses of stamens and pistils. Flowers also showed strong phenotypic integration; genotypic correlations were significantly positive among floral traits within each light treatment. Although floral-vegetative correlations were not significant in the high R:FR light treatment, significant correlations were detected between petal traits, pistil length, and petiole length under low R:FR, in contrast to the widely held hypothesis that floral and vegetative traits are genetically independent. Finally, we detected selection for reduced herkogamy in the low R:FR light treatment. The observed correlation between functional trait groups suggest that vegetative plasticity may affect the expression of floral traits in some environments, and that environment-specific constraints may exist on the evolution of floral and vegetative traits.

Nearly forty years ago R. L. Berg proposed that plants with specialized pollination ecology evolve genetic and developmental systems that decouple floral morphology from phenotypic variation in vegetative traits. These species evolve separate floral and vegetative trait clusters, or as she termed them, "correlation pleiades." The predictions of this hypothesis have been generally supported, but only a small sample of temperate-zone herb and grass species has been tested. To further evaluate this hypothesis, especially its applicability to plants of other growth forms, we examined the patterns of phenotypic variation and covariation of floral and vegetative traits in nine species of Neotropical plants. We recognized seven specific predictions of Berg's hypothesis. Our results supported some predictions but not others. Species with specialized pollination systems usually had floral traits decoupled (weak correlation; Canna and Eichornia) or buffered (relationship with shallow proportional slope; Calathea and Canna) from variation in vegetative traits. However, the same trend was also observed in three species with unspecialized pollination systems (Echinodorus, Muntingia, and Wedelia). One species with unspecialized pollination (Croton) and one wind-pollinated species (Cyperus) showed no decoupling or buffering, as predicted. While species with specialized pollination usually showed lower coefficients of variation for floral traits than vegetative traits (as predicted), the same was also true of species with unspecialized or wind pollination (unlike our prediction). Species with specialized pollination showed less variation in floral traits than did species with unspecialized or wind pollination, as predicted. However, the same was true of the corresponding vegetative traits, which was unexpected. Also in contrast to our prediction, plants with specialized pollination systems did not exhibit tighter phenotypic integration of floral characters than did species with generalized pollination systems. We conclude that the patterns of morphological integration among floral traits and between floral and vegetative traits tend to be species specific, not easily predicted from pollination ecology, and generally more complicated than R. L. Berg envisaged.

The modern coast of China is characterized by complicated hydrology, which might be a potential factor shaping genetic structure of marine organisms. Tridentiger barbatus is a widespread benthic goby in coastal China whose range expansion might depend on larval dispersal in its spawning season. We explored the genetic structure of T. barbatus and its interrelationships with coastal currents, planktonic larval duration, and Pleistocene climate fluctuations. Genealogies of Cytb, Rh, and RAG1 reveal two lineages in T. barbatus, namely the southern and northern lineages. The southern lineage is distributed through the South China Sea to the South Yellow Sea but is absent from the North Yellow Sea and Bohai Sea. The HYCOM average flow field indicates an eddy between the North and South Yellow Seas, which might cause larval retention and limit dispersal of the southern lineage into the North Yellow Sea and the Bohai Sea. The northern lineage is distributed through the South Yellow Sea to the Bohai Sea but is absent from the South China Sea and East China Sea. The distribution of the northern lineage fits the eastward deflection of the Yellow Sea Coastal Current at the boundary of the East China Sea and South Yellow Sea, which might limit southward larval dispersal of the northern lineage into the East China Sea. We also uncovered two Cytb haplotype groups within the southern lineage, namely groups A and B, whose distributions might be shaped by the northward Taiwan Strait Current. The coincidence between the genetic structure of T. barbatus and the coastal currents and eddies strongly implies significant roles of larval dispersal and current transportation in shaping the genetic structure of T. barbatus. In addition, significant isolation by distance is found in the southern lineage, which coincides with its short planktonic larval duration (20–30 days). The inference on divergence time suggests that the observed lineages and haplotype groups in T. barbatus originated in the Pleistocene glaciations. Therefore, the coastal currents, planktonic larval duration, and historical glacial isolation are proposed as important factors shaping the genetic structure of T. barbatus.

BackgroundUncovering how populations of a species differ genetically and ecologically is important for understanding evolutionary processes. Here we combine population genetic methods (microsatellites) with phylogenetic information (mtDNA) to define genetic population clusters of the wide-spread Neotropical túngara frog (Physalaemus pustulosus). We measure gene flow and migration within and between population clusters and compare genetic diversity between population clusters. By applying ecological niche modeling we determine whether the two most divergent genetic groups of the túngara frog (1) inhabit different habitats, and (2) are separated geographically by unsuitable habitat across a gap in the distribution.ResultsMost population structure is captured by dividing all sample localities into two allopatric genetic lineages. The Northern genetic lineage (NW Costa Rica) is genetically homogenous while the Southern lineage (SW Costa Rica and Panama) is sub-divided into three population clusters by both microsatellite and mtDNA analyses. Gene flow is higher within the Northern lineage than within the Southern lineage, perhaps due to increased landscape heterogeneity in the South. Niche modeling reveals differences in suitable habitat between the Northern and Southern lineages: the Northern lineage inhabits dry/pine-oak forests, while the Southern lineage is confined to tropical moist forests. Both lineages seem to have had little movement across the distribution gap, which persisted during the last glacial maximum. The lack of movement was more pronounced for the Southern lineage than for the Northern lineage.ConclusionsThis study confirms the finding of previous studies that túngara frogs diverged into two allopatric genetic lineages north and south of the gap in the distribution in central Costa Rica several million years ago. The allopatric distribution is attributed to unsuitable habitat and probably other unknown ecological factors present across the distribution gap. Niche conservatism possibly contributes to preventing movements across the gap and gene flow between both groups. Genetic and ecological data indicate that there is the potential for ecological divergence in allopatry between lineages. In this context we discuss whether the Northern and Southern lineages should be recognized as separate species, and we conclude that further studies of pre- and post-zygotic isolation are needed for a final assessment. Identified population clusters should motivate future behavioral and ecological research regarding within-species biodiversity and speciation mechanisms.

Theory predicts that tighter correlation between floral traits and weaker relationship between floral and vegetative traits more likely occur in specialized flowers than generalized flowers, favoring by precise fit with pollinators. However, traits and trait correlations frequently vary under different environments. Through detecting spatiotemporal variation in phenotypic traits (floral organ size and vegetative size) and trait correlations in four Ranunculaceae species, we examined four predictions. Overall, our results supported these predictions to a certain degree. The mean coefficient of variation (CV) of floral traits in two specialized species (Delphinium kamaonense and Aconitum gymnandrum) was marginally significantly lower than that of another two generalized species (Trollius ranunculoides and Anemone obtusiloba). The two specialized species also showed marginally significantly smaller CV in floral traits than vegetative size across the two species. The absolute mean correlation between floral and vegetative traits, or that between floral traits in species with specialized flowers was not significantly lower, or higher than that in generalized plants, weakly supporting the predictions. Furthermore, we documented a large variation in trait correlations of four species among different seasons and populations. Study of covariance of floral and vegetative traits will benefit from the contrast of results obtained from generalized and specialized pollination systems.

Damage and recovery from drift of synthetic-auxin herbicide dicamba depends on concentration and varies among floral, vegetative, and lifetime traits in rapid cycling Brassica rapa

Ecological niche evolution can promote or hinder the differentiation of taxa and determine their distribution. Niche-mediated evolution may differ among climatic regimes, and thus, species that occur across a wide latitudinal range offer a chance to test these heterogeneous evolutionary processes. In this study, we examine (a) how many lineages have evolved across the continent-wide range of the Eurasian nuthatch (Sitta europaea), (b) whether the lineages' niches are significantly divergent or conserved and (c) how their niche evolution explains their geographic distribution. Phylogenetic reconstruction and ecological niche models (ENMs) showed that the Eurasian nuthatch contained six parapatric lineages that diverged within 2Myr and did not share identical climatic niches. However, the niche discrepancy between these distinct lineages was relatively conserved compared with the environmental differences between their ranges and thus was unlikely to drive lineage divergence. The ENMs of southern lineages tended to cross-predict with their neighbouring lineages whereas those of northern lineages generally matched with their abutting ranges. The coalescence-based analyses revealed more stable populations for the southern lineages than the northern ones during the last glaciation cycle. In contrast to the overlapping ENMs, the smaller parapatric distribution suggests that the southern lineages might have experienced competitive exclusion to prevent them from becoming sympatric. On the other hand, the northern lineages have expanded their ranges and their current abutting distribution might have resulted from lineages adapting to different climatic conditions in allopatry. This study suggests that niche evolution may affect lineage distribution in different ways across latitude.

Phylogenetic relationships inferred from sequences of the mitochondrial 12S rRNA, tRNA val , and 16S rRNA genes and nuclear POMC and RAG-1 genes revealed that fanged frogs from Thailand usually associated with Limnonectes kuhlii are monophyletic and are collectively sister to the clade containing three Chinese and Japanese species. Within the Thai clade, the northern lineage, the southern lineage, and a population originally assigned to L. megastomias show unresolved relationships with each other, but are separated by genetic distances that correspond to values found among species of the Chinese-Japanese clade. Hybridization and past gene introgression are not detected among these three lineages of fanged frogs from Thailand. Adult specimens of the northern and southern lineages are phenotypically similar to each other, but can be separated by the combination of several morphometric characters. From the genetic and morphological evidence, they are considered to represent taxonomically different species. We therefore describe the northern lineage as L. taylori sp. nov. and the southern lineage as L. jarujini sp. nov. Taxonomic identity of the Loei population of L. megastomias requires future morphological investigation. The distribution pattern of fanged frogs within Thailand is discussed and the significance of the Three Pagodas Fault Zone is noted.

Clines in traits compared over two decades in a plant hybrid zone.

To investigate genetic diversity and phylogeography of the Asian leopard cat (Felis bengalensis), mitochondrial DNA (mtDNA) sequences were determined for 39 individuals from various areas. Sequences combining the complete cytochrome b gene (1,140 bp) with the partial control region (646-810 bp) were classified into 24 haplotypes: 21 types from 21 animals, one from eight animals from Tsushima Islands, one from eight animals from Iriomote Island, and one from two animals from Southeast Asia. Phylogenetic trees of the 24 haplotypes clearly showed three clades: a Northern Lineage and Southern Lineages 1 and 2. The Northern Lineage consisted of animals from Tsushima Islands, the Korean Peninsula, the continental Far East, Taiwan, and Iriomote Island. Within the Northern Lineage, genetic contacts could have occurred between geographically neighboring populations before isolation by straits. Southern Lineage 1, comprising Southeast Asian animals, showed higher genetic diversity. Southern Lineage 2 had large genetic distances from other lineages. Within the control region, the Asian leopard cats shared two to four repetitive motifs, and the number of motifs and their constitution were highly variable among individuals. The motifs were polymorphic even within individuals and could be classified into 31 types. Finally, males of mtDNA Southern Lineage 1 had either of two types of the Y-chromosomal gene ZFY, whereas all males of Northern Lineage shared only one type. Our results indicate that the diversity of southern populations is higher and that genetic differentiation among northern local populations reflects past geographical isolation.

BackgroundFor the genus Thylamys, the rivers have been reported as barriers to dispersal, limiting current and historical distribution of its lineages. We hypothesized that the Maipo river has affected the genetic structure of northern and southern lineages of Thylamys elegans, recovering a phylogenetic relationships with reciprocally monophyletic sister groups on opposite river banks. We evaluated the role of other rivers in the Mediterranean zone of Chile as historical and recent modulators of the biogeographic processes of this species.MethodsWe applied a phylogeographic approach, using the cytochrome-b mitochondrial gene for 93 individuals of T. elegans, from 37 localities in a latitudinal gradient between 21°25’ and 35˚56’S, encompassing a geographic area between the Atacama Desert and most of the Mediterranean Chilean zone.ResultsThe phylogenetics results recovered six lineages within T. elegans: Thylamys elegans elegans, Thylamys elegans coquimbensis, the Loa lineage and three other lineages not described previously (Aconcagua, South 1 and South 2). We suggest that following rivers play a role like primary barrier: the Maipo river in the genetic differentiation of northern and southern ancestral lineages, and the Mataquito river and its tributary Teno river for the South 1 and South 2 lineages. On the other hand, the Quilimarí river preserve the genetic divergence in T. e. coquimbensis and Aconcagua lineage and the Aconcagua river in Aconcagua lineage and T. e. elegans acting like secondary barriers.ConclusionsWe concluded that the genetic diversity and biogeographic history of T. elegans was shaped by mountain glaciers, changes in river water levels during the Pleistocene glaciations and hyperaridity, promoting the differentiation and persistance of the T. elegans lineages.

Major dimensions of plant ecological strategies have been widely studied bringing forward the concept of 'economic spectra' of plants. Sexual reproductive traits, 'floral traits', have been largely neglected in this context, despite their strong link to fitness. Here, we aimed at integrating floral traits into the dimensionality of plant form and function so far dominated by vegetative traits. We used principal component analyses and constructed trait networks to assess the correlation structure of leaf, belowground, plant size-related, and floral traits. We studied forbs within two independent datasets; one compiled from central European trait databases and one sampled in the Austrian Alps. Floral traits defined the second dimension of trait variability within both datasets, while plant size determined the first dimension. Floral traits were largely independent from the leaf economic spectrum. Flower size, however, positively scaled with plant size and leaf size. Mating system was the most well-connected trait across modules of plant tissue/organ types. The independence of floral traits was consistent also after accounting for phylogenetic relationships between species. Floral traits explained a unique part of the variation in plant form and function and thus, likely play a distinctive ecological role within the whole plant economic spectrum.

Accurate species delimitation is the key to precise estimation of species diversity and is fundamental to most branches of biology. Unclear species boundaries within species complexes could lead to the underestimation of species diversity. However, species delimitation of species complexes remains challenging due to the continuum of phenotypic variations. To robustly examine species boundaries within a species complex, integrative approaches in phylogeny, ecology, and morphology were applied to theStewartia sinensiscomplex (Theaceae) endemic to China. Multispecies coalescent‐based species delimitation using 572 nuclear ortholog sequences (anchored enrichment) supported reciprocal phylogenetic monophyly of the northern lineage (NL) and southern lineage (SL), which were not sister clades. Niche equivalency and similarity tests demonstrated significant climatic niche differentiation between NL and SL with observed Warren et al.'sI = 0.0073 and Schoener'sD = 0.0021. Species distribution modeling also separated their potential distribution. Morphometric analyses suggested significant interlineage differentiation of multiple traits including the ratio of length and width, leaf width, and pedicel length, although overall similarity did not differ. Based on the integrative species concept, two distinct species were proposed with legitimate names ofStewartia gemmatafor SL andS. sinensisfor NL. Our empirical study of theS. sinensiscomplex highlights the importance of applying multiple species criteria, in particular the underappreciated niche differentiation, to species delimitation in species complexes pervasive in plants.

AimTo investigate the intra‐specific genetic divergence and phenotypic diversification in Oophaga granulifera, analysing phylogeographical patterns, population structure, coloration and traits potentially involved in reproductive isolation.LocationCosta Rica.MethodsOur phylogeographical analysis used the mitochondrial genes cytochrome b and 16S. We sequenced the two markers in 28 individuals belonging to 10 populations and 16 individuals from nine populations, respectively. The population structure was analysed based on seven polymorphic microsatellite loci in 7–15 individuals from 11 populations using maximum likelihood and Bayesian analyses. The colour phenotypes of 189 frogs and the reflectance spectra of 15 frogs from five populations were measured. In nine populations, we recorded the advertisement calls of 59 males and measured 10 morphological variables from 147 frogs.ResultsOur phylogeographical reconstruction supported the divergence into southern and northern lineages in O. granulifera. In a colour transition area between the main red and green morphs, we describe novel gradual intermediate phenotypes occurring in polymorphic populations. The colour distribution was asymmetrical compared to the genetic variation: the southern lineage included only monomorphic red populations while the northern lineage contained red, green and polymorphic populations. The two lineages differed significantly in advertisement calls and morphology. Microsatellites confirmed the clear inter‐lineage divergence. In the northern lineage, the red morph differed from the intermediate and green morphs in the pulse rate of the advertisement call and in body size. In the colour transition area, there was no simple genetic or colour cline.Main conclusionsOophaga granulifera has evolved into two genetic lineages. The reduced gene flow and diversification in advertisement calls and morphology point to reproductive isolation between lineages. Within the northern lineage, the presence of gradual intermediate morphs and the population structure support the presence of gene flow across the colour transition area while the red versus intermediate–green divergence in both calls and morphology reveals the potential for assortative mating. Our data all congruently show that O. granulifera presents deep intra‐specific divergence. The lack of genetic diversification among colour morphs and the genetic connectivity among the populations defining the colour transition area suggest that the colour diversification in the species is a recent phenomenon.