Genetic variation and phylogeographic analyses of two species of Carpobrotus and their hybrids in California

Despite the commonality and study of hybridization in plants, there are few studies between invasive and noninvasive species that examine the genetic variability and gene flow of cytoplasmic DNA. We describe the phylogeographical structure of chloroplast DNA (cpDNA) variation within and among several interspecific populations of the putative native, Carpobrotus chilensis and the introduced, Carpobrotus edulis (Aizoaceae). These species co-occur throughout much of coastal California and form several 'geographical hybrid populations'. Two hundred and thirty-seven individuals were analysed for variation in an approximate 7.0 kb region of the chloroplast genome using PCR-RFLP (polymerase chain reaction - restriction fragment length polymorphism) data. Phylogenetic analyses and cpDNA population differentiation were conducted for all morphotypes. Historic geographical dispersion and the coefficient of ancestry of the haplotypes were determined using nested clade analyses. Two haplotypic groupings (I and II) were represented in C. chilensis and C. edulis, respectively. The variation in cpDNA data is in agreement with the previously reported allozyme and morphological data; this supports relatively limited variation and high population differentiation among C. chilensis and hybrids and more wide-ranging variation in C. edulis and C. edulis populations backcrossed with C. chilensis. C. chilensis disproportionately contributes to the creation of hybrids with the direction of gene flow from C. chilensis into C. edulis. The cpDNA data support C. chilensis as the maternal contributor to the hybrid populations.     

Hybridization and introgression in Carpobrotus spp. (Aizoaceae) in California. II. Allozyme evidence

Morphological evidence has indicated that hybridization and introgression are occurring between Carpobrotus edulis L., a nonindigenous, invasive species in California (Bolus), and its putative native congener, C. chilensis. The identification of allozyme markers has enabled us to quantify hybridization and the extent of introgression between these species. Samples from 20 individuals of each of five morphological types (both parent species and three hybrid phenotypes) were collected from 39 populations along the coast of California from the Mexico to Oregon borders. Ten enzyme systems revealed a total of 17 resolvable loci, eight of which were polymorphic for the genus. Five of the polymorphic loci easily differentiate C. edulis and C. chilensis. Allele frequencies among the morphologically defined types are consistent with estimations of allele frequency changes due to first- and second-generation backcrossing. In comparison to long-lived, herbaceous perennials and widespread species, C. edulis and intermediate types have more variation in their populations (P = 41.73, Ap = 2.11, Ho = 0.246, and P = 38.13, Ap = 2.06, Ho = 0.216, respectively) than C. chilensis (P = 11.76, Ap = 2.00, Ho = 0.082). Indirect estimates of gene flow indicate four of the five morphological types are outcrossing. Gene flow between previously allopatric species may have broad implications if it results in an increase in fitness; further experimentation is needed to determine the ultimate ecological consequences of this phenomenon and its possible threat to limited, remaining natural habitat in California.     

Germination and growth responses of hybridizing Carpobrotus species (Aizoaceae) from coastal California to soil salinity

Germination, growth, and physiological responses of hybridizing Carpobrotus from coastal California to soil salinity were studied. Hybrids are presumably the result of hybridization and introgression between the exotic Carpobrotus edulis, a succulent perennial invading coastal habitats, and the native or long-naturalized C. chilensis. Germination responses were investigated at 0, 10, 20, and 50% seawater. Seedling growth and physiology were compared by irrigating seedlings with solutions of the same seawater concentrations and in low and high nutrients. Germination was inhibited in the presence of salt, but recovered after transferring the seeds to fresh water. Seeds exposed to salt had higher final germination rates than control. Growth of Carpobrotus was slightly enhanced by low seawater concentrations but reduced at high salinity at both nutrient regimes. Leaf cell sap osmolarity increased with increasing soil salinity, and taxa did not differ significantly in this physiological adjustment. Leaf carbon isotope ratios (∂¹³C) ranged from −28 to −22‰ and became less negative at higher salinities, indicating an improved water use efficiency in the seedlings at high salt concentrations. In addition, ∂¹³C values were generally less negative at high than at low nutrients. Differences among taxa were generally small. The results show that salinity affects both establishment and growth of hybridizing Carpobrotus. The overall weak species differences in salt tolerance indicate that the exotic C. edulis can occupy the same sites as C. chilensis in terms of salinity. The similarity of hybrids in their response to salinity suggests that they may contribute to the invasion by Carpobrotus.     

Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: I. Genetic diversity and introgression

This study, based on morphological and isozyme analysis, clearly discriminates two invasive Carpobrotus taxa, C. edulis and C. acinaciformis, in the Hyères archipelago off the southeastern coast of France. However, three different allelic combinations demonstrate the presence of intermediate individuals resulting from an introgression of part of the C. edulis genome into that of C. acinaciformis. Both taxa have higher than average genetic (C. edulis: P(0.95)=62.5%, A=2.25+/-0.70, H(o)=0.329+/-0.324; C. acinaciformis: P(0.95)=75%, A=2.38+/-0.42, H(o)=0.645+/-0.109) and clonal diversities (C. edulis: IP=0.37; C. acinaciformis: IP=0.48). Furthermore, C. acinaciformis has an excess of heterozygotes (F=-0.585+/-0.217), probably due to introgression. The relationship between the probability of clonal identity for two individuals and distance indicates that C. acinaciformis relies more on clonal reproduction than on sexual recruitment (seed recruitment/vegetative propagation=u/v=0.027), in contrast to C. edulis, whose probability of clonal identity did not vary with distance. The overwhelming clonal growth and high genetic diversities of C. acinaciformis and the previously recorded invasion potential for C. edulis raises concern for intensified invasion via hybridisation.     

Unexpected morphological and karyological changes in invasive Carpobrotus (Aizoaceae) in Provence (S-E France) compared to native South African species

Hybridization processes can lead to evolutionary changes, particularly in co-introduced congeneric plant species, such as Carpobrotus spp. which are recognized as invasive in Mediterranean climate regions. Morphological and karyological comparisons have therefore been made between native Carpobrotus edulis and C. acinaciformis in South Africa and their invasive counterparts in Provence (C. edulis and C. aff. acinaciformis). Morphological data exhibited the most significant differences in invasive C. aff. acinaciformis that forms a new phenotypic variant. Unexpected chromosomal restructuring has been highlighted for both taxa in Provence, with in particular a clear decrease in asymmetry, an increase in the intraspecific variability, and an interspecific convergence of karyotypes. These changes suggest a drift that has facilitated various crosses, and has been amplified through hybridization/introgression. Furthermore, several morphological and karyological transgressive characters have been found in the two invasive taxa. These results stress the important role and the rapidity of karyological changes in invasive processes.     

靖西长喙木兰，广西木兰科一新种（英文）

描述了产自广西木兰科一新种：靖西长喙木兰(Lirianthe jingxiensis Y. H. Tong&N. H. Xia)。本种形态上与绢毛木兰[L. albosericea(Chun&C. H. Tsoong)N. H. Xia&C. Y. Wu]接近,但区别在于该种植株较矮,幼枝、叶柄和幼叶被黄棕色绢毛,叶柄较宽,叶片较宽,倒卵形或倒卵状椭圆形,先端钝或短渐尖,花被片较大,心皮数目较多,被黄棕色绢毛。     

ROOT PROFILES AND COMPETITION BETWEEN THE INVASIVE,EXOTIC PERENNIAL,CARPOBROTUS EDULIS,AND TWO NATIVE SHRUB SPECIES IN CALIFORNIA COASTAL SCRUB

The mat-forming succulent, Carpobrotus edulis (Aizoaceae), surrounds and grows over many native plant species in California coastal communities. Two shrub species, Haplopappus ericoides and H. venetus var. sedoides, were found to have shallow root systems that occupied the same soil depths as those of C. edulis. In the presence of C. edulis, the normal rooting profiles of the shrubs were displaced downward, although partial overlap with C. edulis remained. Removal of C. edulis from around individuals of both shrub species resulted in higher predawn xylem pressure potentials in shrubs from the removal treatments as compared to controls, suggesting that the surrounding C. edulis was utilizing water that would otherwise have been available to these shrubs. In H. ericoides, predawn xylem pressure potentials of removal treatment shrubs remained higher than those of the controls throughout the remainder of the dry season even though these shrubs showed a marked increase in canopy area after removal of surrounding C. edulis. Removal of C. edulis from around H. venetus also initially led to higher predawn xylem pressure potentials in the removal shrubs as compared to the controls and increased production of new leaves. After this initial period the predawn xylem pressure potentials of the removal shrubs were not significantly different from those of controls, suggesting that the demand for water by the increased leaf areas eventually matched the enhanced water availability resulting from C. edulis removal. Also, the morphology of H. venetus shrubs changed to that more typical of unaffected individuals. Thus, C. edulis significantly affected not only the water relations of these two shrub species but also their shoot sizes and overall morphologies.     

Natural hybridization in the genus Colletia (Rhamnaceae)

Natural hybridization in the genus Colletia (Rhamnaceae). Natural hybridization between two Chilean species of Colletia is assumed on the basis of morphological evidence. The characters of both species, C. hystrix Clos and C. ulicina Gill. & Hook., and those of their putative hybrids are described. The morphological intermediacy of the assumed hybrids is tested through Wells' distance diagram.     

Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: II. Reproductive strategies

This study compares sexually and asexually produced fruit set, seed production, biomass, germination, and seedling size in Carpobrotus acinaciformis and C. edulis following controlled pollination experiments in order to evaluate the potential role of reproductive traits with respect to the invasive potential of these taxa. C. edulis is slightly agamospermic, completely self-fertile, slightly preferentially self-compatible, experiences no inbreeding depression, and has low hybrid vigour. In contrast, C. acinaciformis does not have reliable agamospermy, is only slightly self-fertile and self-compatible, experiences a slight inbreeding depression, and has a strong hybrid vigour. Both taxa have relatively low, although significantly different germination frequencies, and insignificantly different seedling sizes. Owing to the high performance in hybridisation as compared to all other controlled pollinations in C. acinaciformis, as well as a large amount of previously demonstrated introgression, we refer to the population studied on the island of Bagaud (France) as C. affine acinaciformis. We conclude that both C. edulis and C. affine acinaciformis should be considered as harmful invasive plants in the Mediterranean Basin, the former because of the flexibility of its mating system and high seed production, and the latter because of its strong clonality, high hybrid vigour, and potential for continued introgression from C. edulis genes. These differences require different control strategies, while the avoidance of sympatry is a distinct priority.     

Hybridization and asymmetric introgression between Cypripedium tibeticum and C. yunnanense in Shangrila County,Yunnan Province,China

Hybridization and introgression are thought to be important for speciation and adaptation in many plants. However, little is known about the hybridization and introgression among Cypripedium species. To investigate the evidence for hybridization and the pattern of introgression between Cypripedium yunnanense and C. tibeticum in Shangrila County, Yunnan Province, China, morphological characters and amplified fragment length polymorphism (AFLP) data for both the species and their putative hybrids were studied. Hand pollination was also performed to verify the crossability of the putative parents. Principal coordinate analysis based on morphological characters and the AFLP data suggested that the putative hybrids were true hybrids of these two Cypripedium species. Analysis with the NewHybrids software indicated that the putative hybrids were F1 generation individuals and backcrosses to C. yunnanese, but no F2 generation was found. Analysis with the Structure software demonstrated asymmetric introgression from C. tibeticum to C. yunnanense. We conclude that natural hybridization and introgression can occur between these two species and that in situ conservation of the parental species is required before fully assessing the evolutionary potential of hybrids.     

Natural hybridization between Vanilla claviculata (W.Wright) Sw. and V. barbellata Rchb.f. (Orchidaceae): genetic, morphological, and pollination experimental data

Vanilla claviculata (W.Wright) Sw. and K barbellata Rchb.f. grow sympatrically in the southwestern part of herto Rico. At three localities in the range of the two species, individuals with flowers that appeared Merent from previously known species were found. To test whether these were hybrids between V. claviculata and V. barbellata , 11 morphological floral characters were recorded at seven populations and allozyme profiles of the three taxa compared. Principal component analysis on the morphological characters gave three distinct groups with minor overlaps. The first component axis reflects variation in overall size of the flowers, while the second component reflects variation in the size of the distal aperture of the labellum. Allozyme data revealed that the putative hybrids have a significant surplus of heterozygotes ( F _IS = - 0.65) as expected in a F ₁ generation. Both data sets support the hypothesis that the individuals are hybrids, which is further augmented by interspecific pollination experiments between the putative parental species. This is the first case of natural hybridization reported in the genus Vanilla .     

Transport of Radioactivity in Relation to Bracts in the Cotton Plant

[¹⁴C]sucrose was applied to the leaf subtending 14-d-old boll(cotton fruit) with and without bracts, and to bracts alonein var. Suvin (Gossypium barbadense L.). The removal of bractssubstantially enhanced the transport of radioactivity from theleaf to the carpels and seed cotton. When bracts alone weretreated, still larger quantities of radioactivity were incorporatedin the carpels and seed cotton. The bracts appeared to nourishthe developing boll through their own photosynthate and regulatethe transport of assimilate from the leaf. Radioactivity, transport, bracts, boll, carpels, seed cotton     

猕猴桃(Actinidia Lindl.)叶片与果实维生素C含量的相关性研究

以猕猴桃属中华猕猴桃（Actinidia chinensis）32个品种和1个种间杂交后代群体为研究对象,对猕猴桃属植物叶片与果实维生素C含量的相关性进行了研究。结果表明,在中华猕猴桃种内水平上,幼果与成熟果实的果肉维生素C含量间存在极显著的正相关关系;在种间杂交后代群体中成熟叶片和成熟果实的维生素C含量存在极显著正相关关系,为利用早期相关性状开展猕猴桃育种的可行性提供了理论依据。此外,对15个常见中华猕猴桃品种的果实维生素C含量进行了多重比较,为人工杂交时的亲本选择提供了依据。     

Coincidence of small-scale spatial discontinuities in leaf morphology and nuclear microsatellite variation of Quercus petraea and Q. robur in a mixed forest

BACKGROUND AND AIMS: The taxon complex comprising Quercus petraea and Q. robur shows distinct morphologies and ecological preferences, but mostly low differentiation in various types of molecular markers at a broad spatial range. Local, spatially explicit analyses may reveal patterns induced by microevolutionary processes operating mainly over short distances. However, no attempts have been made to date to explore the potential of spatial analyses combining morphological and genetic data of these oaks. METHODS: A mixed oak stand was studied to elucidate the small-scale population genetic structure. All adult individuals were classified and putative hybrids were identified using multivariate discrimination analysis of leaf morphological characters. Likewise, all trees were genotyped with five nuclear microsatellites, and a Bayesian assignment method was applied based on maximum likelihood of multilocus genotypes for taxon and putative hybrid classification. KEY RESULTS: Multivariate analyses of leaf morphological data recognized two groups with few individuals as putative hybrids. These groups were significantly differentiated at the five microsatellites, and genetic taxon assignment coincided well with morphological classification. Furthermore, most putative hybrids were assigned to the taxon found in their spatial neighbourhood. When grouping trees into clusters according to their spatial positions, these clusters were clearly dominated by one taxon. Discontinuities in morphological and genetic distance matrices among these clusters showed high congruence. CONCLUSIONS: The spatial-genetic analyses and the available literature led to the assumption that reproductive barriers, assortative mating, limited seed dispersal and microsite-induced selection in favour of the locally adapted taxon at the juvenile stage may reinforce taxon-specific spatial aggregation that fosters species separation. Thus, the results tend to support the hypothesis that Q. petraea and Q. robur are distinct taxa which share a recent common ancestry. Occasional hybrids are rarely found in adults owing to selection during establishment of juveniles.     

猕猴桃(Actinidia Lindl.)叶片与果实维生素C含量的相关性研究

以猕猴桃属中华猕猴桃(Actinidia chinensis)32个品种和1个种间杂交后代群体为研究对象,对猕猴桃属植物叶片与果实维生素C含量的相关性进行了研究。结果表明,在中华猕猴桃种内水平上,幼果与成熟果实的果肉维生素C含量间存在极显著的正相关关系;在种间杂交后代群体中成熟叶片和成熟果实的维生素C含量存在极显著正相关关系,为利用早期相关性状开展猕猴桃育种的可行性提供了理论依据。此外,对15个常见中华猕猴桃品种的果实维生素C含量进行了多重比较,为人工杂交时的亲本选择提供了依据。     

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.     

凤丹栽培群体的表型变异研究

从我国6个凤丹(Paeonia ostii T.Hong et J.X.Zhang)中心产区,选取15个群体共398个单株,对20个表型性状进行调查和分析。结果显示,枝条数、地径、株高、茎长、冠幅长、冠幅宽、成花枝、芽位数、叶长和叶宽等10个表型性状受植株株龄的影响显著,随着株龄增加,除叶长和叶宽的平均值呈递减趋势外,其他8个性状的平均值都呈现递增趋势。凤丹群体间的表型分化系数为0~50.14%,平均值为27.62%,变化幅度最大的是二年生枝长,最小的为心皮数和二年生枝径。对当年生枝长、芽位高、当年生枝径、顶生小叶长、二年生枝径、顶生小叶宽、复叶数、心皮数、二年生枝长和叶柄长等不受株龄影响的10个表型性状进行主成分分析,结果显示前6个性状是影响表型差异的主要性状。各表型性状间存在一定的相关性,聚类分析结果将受株龄影响的10个性状划分为4组。研究结果表明凤丹表型性状在群体间和群体内的变异非常丰富,可为遗传改良和品种选育储备丰富的植物材料。     

栒子属（Cotoneaster Medikus）3种植物的比较形态与统计分析

用数量统计的方法对栒子属（Cotoneaster Medikus）植物水栒子（C. multiflorus Bge.）、毛叶水栒子（C. submultiflorus Popov.）和黑果栒子（C. melanocarpus Lodd.）的23个形态指标进行了研究。平均值和变异系数表明不同性状在不同种内的变异程度有所不同,黑果栒子的种内变异幅度较小;方差分析结果显示18个性状的种间差异显著,黑果栒子与其它两者的差异更明显;逐步判别分析筛选出了6个种间划分的关键性状（叶下被毛、花梗被毛、叶基部在中脉处的形状、叶顶端在中脉处的形状、上1/4叶缘的弧向、叶柄长/叶长）,并把82份材料明确分为3组;毛叶水栒子的大部分形态性状具有水栒子和黑果栒子的居间性,但其种级地位合理。     

Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

PSEUDOFAGUS IDAHOENSIS,N. GEN. ET SP. (FAGACEAE) FROM THE MIOCENE CLARKIA FLORA OF IDAHO

A new genus and species of Fagaceae are established for compressed leaves and attached fruits which represent one of the dominant taxa in the rich Miocene Clarkia flora of northern Idaho. The leaf and fruit morphology, cuticular and pericarp anatomy, and leaf phytoliths fall within the range of variation of the subfamily Fagoideae. The unique fruit consists of a large, exposed, distinctly keeled, trigonous nut and a diminutive basal cupule. The cupule has an apical frill of free, petiolate, leaflike appendages but lacks other ornamentation, and there is no evidence of a distinct Valvation. As no other genus of the Fagaceae has fruits with this combination of characters, a new genus is established and is referred to the subfamily Fagoideae. The Miocene genus seems most closely related to the extant genus Fagus.