PATTERNS OF HYBRIDIZATION IN THE PIRIQUETA CAROLINIANA COMPLEX IN CENTRAL FLORIDA: EVIDENCE FOR AN EXPANDING HYBRID ZONE

We have identified a broad zone of hybridization between two morphologically and ecologically distinct herbaceous perennial taxa (morphotypes) within the Piriqueta caroliniana complex, which extends more than 300 km across the central Florida peninsula. Phylogeographic analyses indicate that the caroliniana morphotype has been present in north and central Florida since the early Pleistocene and that the viridis morphotype has immigrated into southern Florida much more recently. We examine the distribution of diagnostic morphological characters and nuclear genetic markers to assess the extent and patterns of introgression in this system. The results are consistent with the hypothesis that this hybrid zone has expanded north in recent history as viridis alleles have introgressed into regions that were previously occupied by populations of caroliniana. Genetic markers diagnostic for caroliniana have consistently high frequencies across the hybrid zone, whereas markers for viridis are extremely variable among populations with frequency reversals in adjacent populations. The latter pattern is probably the result of the combined stochastic effects of dispersal and drift on viridis alleles as they introgressed northward. Additional evidence for the recent expansion of this hybrid zone comes from patterns of variation for morphological and genetic markers. As expected for an expanding hybrid zone, within-population morphological variation was greatest toward the advancing front of introgression and levels of genetic variation for neutral diagnostic markers were greatest in the region of initial contact and lower in areas of recent expansion. The observed patterns of variation suggest that at least some hybrid genotypes have high fitnesses, which has led to the expansion of the hybrid zone via the displacement of parental genotypes in central Florida.     

<Emphasis Type="SmallCaps">P</Emphasis>lastic responses to temporal variation in moisture availability: consequences for water use efficiency and plant performance

The ability to appropriately modify physiological and morphological traits in response to temporal variation should increase fitness. We used recombinant hybrid plants generated by crossing taxa in the Piriqueta caroliniana complex to assess the effects of individual leaf traits and trait plasticities on growth in a temporally variable environment. Recombinant hybrids were used to provide a wide range of trait expression and to allow an assessment of the independent effects of individual traits across a range of genetic backgrounds. Hybrid genotypes were replicated through vegetative propagation and planted in common gardens at Archbold Biological Station in Venus, Florida, where they were monitored for growth, leaf morphological characters, and integrated water use efficiency (WUE) (C isotope ratio; δ¹³C) for two successive seasons. Under wet conditions only leaf area had significant effects on plant growth, but as conditions became drier, growth rates were greatest in plants with narrow leaves and higher trichome densities. Plants with higher WUE exhibited increased growth during the dry season but not during the wet season. WUE during the dry season was increased for plants with smaller, narrower leaves that had higher trichome densities and increased reflectance. Examination of alternative path models revealed that during the dry season leaf traits had significant effects on plant growth only through their direct effects on WUE, as estimated from δ¹³C. Over the entire growing season, plants with a greater ability to produce smaller and narrower leaves with higher trichome densities in response to reduced water availability had the greatest growth rate. These findings suggest that plants making appropriate changes to leaf morphology as conditions became dry had increased WUE, and that the ability to adjust leaf phenotypes in response to environmental variation is a mechanism by which plants increase fitness.     

Expression of anatomical leaf traits in homoploid hybrids between deciduous and evergreen species of Vaccinium

We investigated the anatomical expression of leaf traits in hybrids between evergreen Vaccinium vitis‐idaea and deciduous V. myrtillus. We compared parents from four populations with their respective F1 hybrids and tested whether (i) transgression can be the source of novel anatomical traits in hybrids; (ii) expression of transgressive traits is more probable for traits with similar values in parents and intermediate for more distinct values, as predicted by theory; and (iii) independent origin of hybrids leads to identical trait expression profiles among populations. We found that anatomical leaf traits can be divided into four categories based on their similarity to parents: intermediate, parental‐like, transgressive and non‐significant. Contrary to the common view, parental‐like trait values were equally important in shaping the hybrid profile, as were intermediate traits. Transgression was revealed in 17/144 cases and concerned mainly cell and tissue sizes. As predicted by theory, we observed transgressive segregation more often when there was little phenotypic divergence, but intermediate values when parental traits were differentiated. It is likely that cell and tissue sizes are phylogenetically more conserved due to stabilising selection, whereas traits such as leaf thickness and volume fraction of the intercellular spaces, showing a consistent intermediate pattern across populations, are more susceptible to directional selection. Hybrid populations showed little similarity in expression profile, with only three traits identically expressed across all populations. Thus local adaptation of parental species and specific genetic background may be of importance.     

Pollinator‐mediated selection on floral morphology: evidence for transgressive evolution in a derived hybrid lineage

Hybridization between closely related lineages is a mechanism that might promote substantive changes in phenotypic traits of descendants, resulting in transgressive evolution. Interbreeding between divergent but morphologically similar lineages can produce exceptional phenotypes, but the potential for transgressive variation to facilitate long‐term trait changes in derived hybrid lineages has received little attention. We compare pollinator‐mediated selection on transgressive floral traits in both early‐generation and derived hybrid lineages of the Piriqueta cistoides ssp. caroliniana complex. The bowl‐shaped flowers of morphotypes in this complex have similar gross morphologies and attract a common suite of small insect pollinators. However, they are defined by significant differences in characters that generate pollinator interest and visitation, including floral area and petal separation. In common garden experiments, patterns of pollen deposition in early‐generation recombinant hybrids indicate that Piriqueta's pollinators favour flowers with greater area and reduced petal separation. Changes in floral morphology in derived hybrid lineages are consistent with predictions from selection gradients, but the magnitude of change is limited relative to the range of transgressive variation. These results suggest that hybridization provides variation for evolution of divergent floral traits. However, the potential for extreme transgressive variants to contribute to phenotypic shifts may be limited due to reduced heritability, evolutionary constraints or fitness trade‐offs.     

桉树及其杂交种叶片形态的遗传变异特征

为了解桉树重要性状在杂交种亲本、子代间的遗传变异规律,以尾叶桉(Eucalyptus urophylla)为母本、粗皮桉(E. pellita)和细叶桉(E. tereticornis)为父本、8个全同胞杂交子代为材料,对亲本和子代的叶片形态性状进行了分析。结果表明,亲缘关系近的尾叶桉和粗皮桉的叶片性状大多无显著差异;但他们与亲缘关系远的细叶桉间的差异显著;尾粗桉与父母本间的性状无显著差异或仅与母本差异显著,属偏父本型;尾细桉与双亲间的多数性状都差异显著,且多偏向于母本。在树种内的家系间或个体间,叶片数量性状差异显著,而质量性状仅叶面革质厚度、叶面朝向差异显著;杂交种家系部分性状有超亲现象,且尾细桉家系超亲频率显著高于尾粗桉。叶长、叶长宽比、形状因子、叶色、叶面革质是区别叶片形态特征的决定性状。     

Transgressive character expression in a hybrid sunflower species

Diploid hybrid lineages often are ecologically distinct from their parental species. However, it is unclear whether this niche divergence is typically achieved via hybrid intermediacy, a mixture of parental traits, and/or the evolution of extreme (transgressive) morphological and ecophysiological features. Here we compare an extensively studied hybrid sunflower species, Helianthus anomalus, with its putative parents, H. annuus and H. petiolaris, for 41 morphological and 12 ecophysiological traits. Helianthus anomalus was morphologically intermediate for one trait (2.4%), parental-like for 23 traits (56.1%), and transgressive for 17 traits (41.5%). For ecophysiological traits, H. anomalus was not significantly different from one or both parents for nine traits (75%), and was transgressive for the remaining three (25%). Thus, H. anomalus appears to be a mosaic of parental-like and transgressive phenotypes. Although the fitness effects of the transgressive characters are not yet known, many of these characters are consistent with adaptations reported for other sand dune plants. Genetic studies are currently underway to ascertain whether these extreme characters arose as a direct byproduct of hybridization or whether they evolved via mutational divergence.     

Genetic structure of a hybrid zone between two violets,Viola rossii Hemsl. and V. bissetii Maxim.: dominance of F1 individuals in a narrow contact range

The genetic composition of a hybrid zone can provide insight into the evolution of diversification in plants. We carried out morphological and amplified fragment length polymorphism analyses to investigate the genetic composition of a hybrid zone between two violets, Viola bissetii Hemsl. and Viola rossii Maxim. Our aim was to clarify the formation and maintenance of hybrids between these Viola species. We found that most hybrid individuals (V. bissetii × V. rossii) were of the F₁ generation, with a few of the F₂ generation. We found no backcrosses. The scarcity of post‐F₁ hybrids indicates that a species barrier is established between the parental species. The F₁‐dominated hybrid zone occupied only a narrow, intermediate ecotone between the parental habitats, suggesting that selection by environmental factors against hybrids may help to maintain the current conditions in this hybrid zone.     

Community structure of canopy arthropods associated to Quercus crassifolia×Quercus crassipes complex

Quercus crassifolia and Q. crassipes are dominant species in temperate forests of central Mexico and hybridize between each other when they occur in sympatry. Oak canopies contain a considerable portion of arthropod diversity and the hybrid zones can provide new habitats to epiphyte fauna. We tested if the establishment of hybrids in contact zones with their parental hosts increases the species diversity of canopy arthropods assuming that hybrid trees constitute new genotypes of potential new habitats to small organisms. We examined the effect of hybridization on some community structure parameters (diversity, composition, similarity and density of arthropod fauna) of canopy arthropods compared to their parental species in a hybrid zone located in central Mexico. We employed 17 leaf morphological traits and six diagnostic RAPD primers to identify parental and hybrid plants. The RAPDs marker showed unidirectional introgression towards Q. crassifolia, and were detected hybrid (F₁), backcrosses and introgression individual trees. In total, 30 oak canopies were fogged during rainy and dry season. We recognized 532 taxa of arthropods belonging to 22 orders associated with tree canopies. The taxonomic status of host‐trees may be an important factor in the arthropod community structure and that seasonality (dry and rainy) is not a factor that could modify their organization. Trees of Q. crassipes registered the highest densities of arthropod fauna followed by hybrid hosts (F₁); trees originated by backcrosses towards Q. crassifolia registered a significant less arthropod density than F₁ hybrids; and trees of Q. crassifolia had the lowest density. Hybrid plants and Q. crassipes individuals had higher diversity (H′) of arthropods than Q. crassifolia plants. Hybrid plants had also more rare species in both seasons in comparison with parental species. This study suggests that hybrid oaks act as a center of biodiversity by accumulating arthropods of both parental and different species including a considerable number of rare species.     

Morphological Variation in Wild Marmosets (Callithrix penicillata and C. geoffroyi) and Their Hybrids

Evolutionary theory and observation predict wider phenotypic variation in hybrids than parental species. Emergent phenotypic novelty in hybrids may in turn drive new adaptations or speciation by breaking parental phenotypic constraints. Primate hybridization is often documented through genetic evidence, but knowledge about the primate hybrid phenotype remains limited due to a small number of available studies on hybrid primate morphology. Here, we examine pelage and morphometric variation in two Brazilian marmoset species (Callithrix penicillata and C. geoffroyi) and their hybrids. Hybrids were sampled in an anthropogenic hybrid zone in the municipality of Viçosa, Minas Gerais state, Brazil. We analyzed hybrid facial and body pelage color variation, and compared 13 morphometric measures between hybrids and parental species. Five different hybrid facial morphotypes were observed, varying from intermediate to parental-like. Hybrid facial morphotypes were biased towards C. penicillata, suggesting that the pelage of this species may be dominant to that of C. geoffroyi in this context, and indicating that mate preference, and therefore gene flow/introgression, may be biased towards C. penicillata within the hybrid zone. Hybrid morphometric features were on average intermediate to parental species traits, but transgressive hybrids were also observed, suggesting that morphometric variation for the studied traits is consistent with Rieseberg’s complementary allele model. Finally, we observed a decoupling of facial patterning and size/shape in hybrids, relative to parent phenotypes, suggesting that an important factor driving phenotypic novelty within the Viçosa marmoset hybrid zone might be the loosening of evolutionary constraints on phenotypic trait integration.     

Chromosome study of hybrid and gynogenetic offspring of artificial crosses between members of the catfish families Clariidae and Pangasiidae

Synopsis Artificial hybridization between species of the catfish families Clariidae (Clarias macrocephalus) and Pangasiidae (Pangasius sutchi) resulted in first generation offspring comprising two intermediate morphotypes and one morphotype indistinguishable from its clariid parent. The two intermediate morphotypes apparently correspond to hybrid morphotypes 2 and 4 resulting from earlier hybridization experiments between the same parent species (Tarnchalanukit 1986). We did not obtain morphotypes 1 and 3. Chromosome spreads of the relatively pangasiid-like morphotype 2 reveal a diploid number of 57, presumably comprising one set of Clarias (n=27) and one of Pangasius (n=30) chromosomes. The relatively clariid-like morphotype 4 is a triploid with 84 chromosomes, presumably comprising two sets from Clarias (2n=54) and one from Pangasius (n=30). Finally, the morphotype indistinguishable from Clarias, is a diploid with 54 chromosomes, apparently arising from gynogenesis.     

Geographic occurrence of intermediate type between<Emphasis Type="Italic"> Castanopsis sieboldii</Emphasis> and<Emphasis Type="Italic"> C. cuspidata</Emphasis> (Fagaceae) based on the structure of leaf epidermis

The number of layers of epidermis in the leaves is used as a criterion to distinguish between Castanopsis sieboldii (two layers) and C. cuspidata (one layer). An intermediate type, which has one and two layers within a single leaf, is frequently seen in the field. The origin of the intermediate type has been supposed to be a hybrid between C. sieboldii and C. cuspidata. If the intermediate type is produced by hybridization, we expect that the F₁ seedlings of the intermediate type should occur in the co-occurrence area of those two species. To clarify the geographic occurrence of the intermediate type, we collected nuts of 443 mother trees from throughout the distribution area of the genus Castanopsis in Japan. A total of 7,260 seedlings germinated from these nuts were examined as to their leaf structure. The seedlings of the intermediate type occurred not only in the area where C. sieboldii and C. cuspidata coexisted, but also in the area where only C. sieboldii grows. The leaf structure of intermediate seedlings was independent of the traits of mother trees. These findings suggest that the intermediate seedlings that occurred in the area where only C. sieboldii grows are not hybrid between C. sieboldii and C. cuspidata but are a morphological variation of C. sieboldii. The difference in the structure of leaf epidermis is not always appropriate for the identification of the hybrid.     

Leaf traits in parental and hybrid species of Sorbus (Rosaceae)

? Premise of the study: Knowledge of functional leaf traits can provide important insights into the processes structuring plant communities. In the genus Sorbus, the generation of taxonomic novelty through reticulate evolution that gives rise to new microspecies is believed to be driven primarily by a series of interspecific hybridizations among closely related taxa. We tested hypotheses for dispersion of intermediacy across the leaf traits in Sorbus hybrids and for trait linkages with leaf area and specific leaf area. ? Methods: Here, we measured and compared the whole complex of growth, vascular, and ecophysiological leaf traits among parental (Sorbus aria, Sorbus aucuparia, Sorbus chamaemespilus) and natural hybrid (Sorbus montisalpae, Sorbus zuzanae) species growing under field conditions. A recently developed atomic force microscopy technique, PeakForce quantitative nanomechanical mapping, was used to characterize the topography of cell wall surfaces of tracheary elements and to map the reduced Young's modulus of elasticity. ? Key results: Intermediacy was associated predominantly with leaf growth traits, whereas vascular and ecophysiological traits were mainly parental-like and transgressive phenotypes. Larger-leaf species tended to have lower modulus of elasticity values for midrib tracheary element cell walls. Leaves with a biomass investment related to a higher specific leaf area had a lower density. Leaf area- and length-normalized theoretical hydraulic conductivity was related to leaf thickness. ? Conclusions: For the whole complex of examined leaf traits, hybrid microspecies were mosaics of parental-like, intermediate, and transgressive phenotypes. The high proportion of transgressive character expressions found in Sorbus hybrids implies that generation of extreme traits through transgressive segregation played a key role in the speciation process.     

The origin of ecological divergence in Helianthus paradoxus (Asteraceae): selection on transgressive characters in a novel hybrid habitat

Diploid hybrid speciation in plants is often accompanied by rapid ecological divergence between incipient neospecies and their parental taxa. One plausible means by which novel adaptation in hybrid lineages may arise is transgressive segregation, that is, the generation of extreme phenotypes that exceed those of the parental lines. Early generation (BC2) hybrids between two wild, annual sunflowers, Helianthus annuus and Helianthus petiolaris, were used to study directional selection on transgressive characters associated with the origin of Helianthus paradoxus, a diploid hybrid species adapted to extremely saline marshes. The BC2 plants descended from a single F1 hybrid backcrossed toward H. petiolaris. The strength of selection on candidate adaptive traits in the interspecific BC2 was measured in natural H. paradoxus salt marsh habitat. Positive directional selection was detected for leaf succulence and Ca uptake, two traits that are known to be important in salt stress response in plants. Strong negative directional selection operated on uptake of Na and correlated elements. A significant decrease in trait correlations over time was observed in the BC2 population for Na and Ca content, suggesting an adaptive role for increased Ca uptake coupled with increased net exclusion of Na from leaves. Patterns of directional selection in BC2 hybrids were concordant with character expression in the natural hybrid species, H. paradoxus, transplanted into the wild. Moreover, the necessary variation for generating the H. paradoxus phenotype existed only in the BC2 population, but not in samples of the two parental species, H. annuus and H. petiolaris. These results are consistent with the hypothesis that transgressive segregation of elemental uptake and leaf succulence contributed to the origin of salt adaptation in the diploid hybrid species H. paradoxus.     

Production of interspecific somatic hybrid plants in the genus Medicago through protoplast fusion

Summary Symmetric somatic hybrid plants have been produced by electrofusion of leaf protoplasts of Medicago sativa and callus protoplasts of Medicago coerulea. The selection of hybrid individuals has been performed at the cellular level by recording the positions of single heterocaryons immobilized in a semisolid culture medium. The hybrid nature of the heterokaryons was assessed in fluorescent light on the basis of their color. Hybrid minicalli were picked up manually and grown first on propagating, and then on regenerating, media. Six putative hybrid calli were selected and two of them regenerated several plants. The hybrid nature of the regenerants was confirmed by cytological and isozyme analysis. Among the several morphological traits taken into account for the characterization of somatic hybrid plants, some were intermediate, some lower, and some higher, with respect to the parents. The somatic hybrid plants were fertile and set seed. The production of somatic hybrid plants in the genus Medicago is discussed in relation to the regenerating capability of parental protoplasts.This research was supported by the National Research Council of Italy, Special Project RAISA, Subproject N. 2 paper N. 347     

Quantitative variation within and among populations of Arabis serrata Fr. & Sav. (Brassicaceae)

Phenotypic and genetic variation within and among eight populations of Arabis serrata are documented in this study. This species shows great morphological variation throughout its geographical distribution in Japan. Plants are located in habitats with different types of soils and degree of disturbance. Half-sibs progenies from eight populations were collected and cultivated in a garden experiment. Nine morphological traits representing size and shape of rosette leaves were recorded. Univariate analyses of measured traits showed that phenotypic means differed among populations for all characters. Leaves of plants from disturbed habitats had the longest petioles (lanceolate) and plants from limestone habitats showed the most roundness in leaf shape (ovate). The northernmost populations always revealed the smallest leaves. Multivariate principal component analyses also showed that leaf shape and size varied among populations. The first three principal components explained 98.5% of the variation. Coefficients of variation had a very wide range and differed from one population to another. Some traits (e.g. leaf width/leaf length ratio) were consistently less variable while others (e.g. leaf area and petiole length) were more plastic. All traits had significant genetic variance in all populations. Intra-class correlation coefficients differed for most of the traits and each population presented a different range of values. Most of the leaf traits were intercorrelated in all the populations studied, although some populations were integrated more tightly for some traits. Populations of A. serrata are differentiated in phenotypic means but they display a mosaic of traits with slight morphological differences in each locality (i.e. a quantitative genetic variation). Some traits can be correlated to the habitats that they occupy but for some of them it is difficult to assign an actual adaptive value.     

Transgressive aggression in Sceloporus hybrids confers fitness through advantages in male agonistic encounters

1.  We investigated agonistic behaviour and associated characteristics of Sceloporus woodi (Florida scrub lizard), Sceloporus undulatus (Eastern fence lizard) and their hybrids using staged territorial encounters.
2.  These Sceloporus hybrids exhibit transgressive aggression and transgressive head-girth relative to the parental species and the transgressive aggression was specifically associated with an advantage in agonistic encounters. Our results suggest a hybrid advantage in natural habitats when defending and invading territories against either parental species.
3.  We further analysed general advantages in agonistic encounters across the entire three-group system to elucidate characteristics that may be advantageous under specific circumstances. Individuals with larger body size (SVL) and greater aggression had an overall advantage in agonistic encounters; however, smaller individuals could win when slightly more aggressive and fatter, and less aggressive individuals could win when slightly larger, especially with greater head-girth.
4.  The extreme hybrid phenotypes likely occurred through transgressive segregation, which has been implicated as a process through which homoploid, hybrid speciation can occur. Some form of ecological divergence is necessary, however, to impede parental gene flow. Our data suggest that ecological divergence could manifest in territorial species through transgressive aggression.     

RAPD analysis of genome variability of planted ginseng,Panax ginseng

Genome variability of 23 ginseng plants (Panax ginseng) grown in culture in Primorskii Krai was studied by RAPD method. Eleven arbitrary chosen primers were used to analyze 138 loci of DNA samples, 17 of which appeared to be polymorphic. The OPD-11-1000 fragment was found to be a RAPD marker allowing plants to be differentiated according to their morphotype. Using five primers, it was demonstrated that the genetic polymorphism of the cultivated plants is lower than that in nature (7.6% and 10.6%, respectively). Dendrograms of genetic relatedness are in accord with genetic differences between individuals of plantedP. ginseng belonging to different morphotypes, and demonstrate close relatedness of one of the morphotypes to wild plants. This morphotype could be recommended for reintroduction into natural habitats.     

Transgressive character expression in hybrid zones between the native invasives Tithonia tubaeformis and Tithonia rotundifolia (Asteraceae) in Mexico

Natural hybridization frequently promotes gene introgression among closely related species in sympatric populations, producing complex patterns of morphological variation. Therefore, a detailed understanding of the dynamics of interspecific gene flow and its morphological patterns is of widespread interest. We tested if introgressive hybridization promotes an increase in transgressive characters in comparison with the parental species. A sunflower species complex occurring in Mexico formed by two native invasive species, Tithonia tubaeformis and Tithonia rotundifolia, was analyzed using 46 morphological characters (leaf, flower and fruit) in five hybrid zones (N = 150 individuals) and two pure sites for each parental species (N = 80 individuals). In general, T. tubaeformis differed significantly from T. rotundifolia in all the examined characters, except six foliar and one inflorescence character. Morphological characters support the hypothesis of hybridization in this complex, even though both species remain morphological distinct in mixed stands. Individual hybrids appear to be a mosaic of parent-like (24.8 % of traits), intermediate (26.1 %) and transgressive (37.8 %) phenotypes (the remaining 11.3 % of the traits did not differ significantly from both parental species). Our results suggest that individuals from the same parental species were more similar among themselves than to putative hybrids, indicating occasional hybridization with segregation in hybrid types or backcrossing to parents. Evidence indicates a unidirectional pattern of gene flow toward T. rotundifolia.     

Morphological plasticity and ecophysiological response of ground ivy (Glechoma hederacea,Lamiaceae) in contrasting natural habitats within its native range

Abstract

Some species intrinsically have a high invasiveness capacity, shown by high phenotypic plasticity and rapid growth, enabling a wide distribution across their native habitats and successful invasion in the introduced range. For such species, information from native habitats is critically important. An example is Glechoma hederacea, native to Eurasia but introduced and widespread in the USA. Our main objective was to investigate variation in traits of native G. hederacea populations across contrasting habitats: open, forest edge and understory. Vegetation was sampled and the ecophysiological and morphological traits were measured with accompanying environmental parameters. Results showed that in native habitats environmental conditions cover wide gradients of light and soil moisture. Plants had the highest cover in nutrient-rich, shaded habitats, representing the optimal habitat, indicating shade tolerance of G. hederacea. Plants from forest understory exhibited strong similarities in investigated traits to plants from the forest edge, even though this was a drier, sunnier habitat. Plants from open, sunny habitats experienced stress as indicated by the quantum efficiency of PSII and significantly higher sexual reproduction. Results show that G. hederacea is moderately tolerant simultaneously to shade and drought, a characteristic that has been reported for numerous invasive species, while at the same time it shares some characteristics with weedy plants.