New distributional and molecular information call into question the systematic position of the West Asian Viola sintenisii (Violaceae)

The west Asian Viola sintenisii W.Becker, described from temperate woodlands of northern Iran and south-western Turkmenistan on the Caspian coast, was long considered a vicariant geographical subspecies of the mainly south European V. alba Besser. The new findings of V. sintenisii in four geographically separated stations in north-eastern Azerbaijan significantly expands its range of distribution. These occurrences are within the range of V. alba ssp. alba . Here the two taxa frequently co-occur but remain distinct and V. sintenisii is therefore justified at species level. It is likely that the two species are also sympatric in the Hyrcanian region, a range previously attributed uniquely to the latter. Viola sintenisii seems to be fairly common in the eastern parts of the Great Caucasus, but there are reasons to suggest that it has there been misinterpreted as V. suavis . The chromosome number of V. sintenisii , 2 n  = 20, is reported here for the first time.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 91–98.     

Viola hirta (Violaceae) and its relatives in Norway

Viola hirta is widespread in most of Europe but rare in Norway, occurring in four isolated enclaves around the Oslo Fjord. Allozymic and morphological patterns of variation were studied in 320 individuals of Viola hirta (2n = 20) and its relatives V. collina (2n = 20), V. odorata (2n = 20), and V. suavis (2n = 40) from 22 Norwegian populations. Ongoing introgression from both V. collina and V. odorata , as hypothesised by several authors, was not supported by our data. All hybrids proved to be primary ones and, apparently, introgression is rare. However, allozyme markers suggested that V. hirta may have been affected by ancient introgression from V. collina and possibly also from the East European V. ambigua (2n = 40) during its evolutionary history. Geographical differentiation, presumably relatively recent and post-glacial, was evident in V. hirta. Variation occurred also between and within V. odorata populations, most likely a consequence of independent introductions by man. No variation was observed in V. collina or in V suavis. A key to the Norwegian species of subsection Viola and their primary hybrids based on morphological characters is provided in Flora Nordica note No. 26.     

Establishing the phylogenetic origin, history, and age of the narrow endemic Viola guadalupensis (Violaceae)

? Premise of the study: Climate change and shifts in land use are two major threats to biodiversity and are likely to disproportionately impact narrow endemics. Understanding their origins and the extent of their genetic diversity will enable land managers to better conserve these unique, highly localized gene pools. Viola guadalupensis is a narrow endemic of the Guadalupe Mountains (west Texas, USA). Its affinities within Viola section Chamaemelanium have been the subject of some debate. Furthermore, the polyploid and presumably reticulate relationships within this section remain largely unknown. ? Methods: We counted chromosomes for V. guadalupensis. Phylogenies for the chloroplast trnL-F region and the low-copy nuclear gene GPI for 24 Viola taxa were generated and used to produce a polyploid phylogenetic network. Divergence dates were obtained by fossil calibration. ? Key results: Meiotic chromosome counts revealed that V. guadalupensis is tetraploid (n = 12), and the presence of two GPI homoeologs further suggested allotetraploidy. Phylogenetic reconstructions showed that it originated through hybridization between unidentified members of subsection Canadenses (paternal parent) and subsection Nuttallianae (maternal parent). A fossil-calibrated relaxed clock dating analysis of GPI estimated the maximum age of V. guadalupensis to be 8.6 (5.7-11.6) Myr, suggesting the species evolved after the Guadalupe Mountains formed 12-13 Ma. ? Conclusions: Viola guadalupensis originated by intersubsectional hybridization followed by polyploidization. Within section Chamaemelanium, this phenomenon has occurred repeatedly in the last 9 Myr (at least for V. bakeri, V. douglasii, V. glabella, and V. sempervirens). Consequences for the systematics of the section are discussed.     

On the origin of the sweet-smelling Parma violet cultivars (Violaceae): wide intraspecific hybridization, sterility, and sexual reproduction

Parma violets are reputed for their double, fragrant flowers and have been cultivated for centuries in Europe. However, due to a rather atypical morphology their taxonomic affinity has not been clarified. Authors have proposed an origin from three possible species, Viola alba, V. odorata, or V. suavis, or a hybrid origin. Using both ITS sequence variation and allozyme variation in 14 putative loci, we showed that the Parma violet cultivars have their origin within Viola alba and that they are best included in the Mediterranean subsp. dehnhardtii. There is no trace of interspecific hybridization. However, the cultivars appear to have a single origin in a wide hybrid within V. alba, involving parental plants from the eastern and western Mediterranean region; historical literature sources seem to indicate Turkey and Italy, respectively. The Parma violet cultivars possess high levels of allozyme heterozygosity and to some extent also within-individual ITS sequence variation. Losses of heterozygosity and within-individual ITS sequence variation in some of the cultivars indicate subsequent rare events of sexual reproduction, presumably through cleistogamous seed set. We unambiguously identify the closest wild relative of this group of cultivars, allowing growers to develop new selection procedures, and show a peculiar molecular process associated with human selection.     

A new species of Viola (Violaceae) from Turkey

A new species of Viola , V. kizildaghensis M. Dinç & S¸. Ẏlḋṙml̇, from Turkey is described and illustrated. It is found on the rocky slopes of K̇żldǎ, K̇żldǎ National Park, in the vilayet of Isparta, south-west Turkey, at an elevation of 1350–1600 m. It belongs to Viola , subsection Viola , and is similar to the Lebanese endemic V. libanotica Boiss., distinguished by its lanceolate, cuneate based leaves, pinkish-purple flowers with white throats, glabrous lateral petals, and peduncles bearing the mature fruit procumbent and pubescent. It is somewhat similar to another species from south-west Turkey, V. sandrasea Melchior, that is endemic to Sandras Dǎ in the vilayet Mǔla; it is distinguished from this by the characters listed above, its pubescent leaves and peduncles, and its linear-lanceolate to lanceolate stipules which have short, gland-tipped fimbriae. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 483–487.     

Intraspecific variation in Viola suavis in Europe: parallel evolution of white-flowered morphotypes

Background and Aims

Viola species are commonly grown for their ornamental flowers, but their evolutionary history and taxonomy are often complicated and have been poorly explored so far. This is a study of the polymorphic, typically blue-flowered species Viola suavis, concentrating on the white-flowered populations of uncertain taxonomic assignment that occur in Spain and central and south-eastern Europe. The aim was to resolve their origin and taxonomic status and to study the intraspecific structure and (post)glacial history of this species.

Methods

Viola suavis and five close relatives were sampled from multiple locations and subjected to molecular (AFLP, sequencing of nrDNA ITS) and morphometric analyses. Data on ploidy level and pollen fertility were also obtained, to address an assumed hybrid origin of the white-flowered populations.

Key Results

In V. suavis a strong intraspecific genetic split into two groups was observed, indicating that there has been a long-term isolation and survival in distinct glacial refugia. The white-flowered populations could be placed within the variation range of this species, and it is clear that they evolved independently in two distant areas. Their parallel evolution is supported by both morphological and genetic differentiation. The strongly reduced genetic variation and absence of unique AFLP fragments suggest their derived status and origin from the typical, blue-flowered populations.

Conclusions

These results suggest that intraspecific variation in V. suavis has been largely shaped by population isolations during the last glaciation and subsequent recolonizations, although cultivation and vegetative spread by humans have affected the present picture as well.Key words: AFLP, central Europe, flow cytometry, ITS sequences, multivariate morphometrics, parallel evolution, Spain, Violaceae     

Anatomical studies on selected species of Viola (Violaceae)

Cross‐sections of roots, stems, petioles and peduncles were investigated in 12 species of Viola distributed mainly in northern Iran: V. kitaibeliana, V. arvensis, V. occulta, V. tricolor (sect. Melanium), V. somchetica (sect. Plagiostigma), V. spathulata (sect. Spathulidium ined.), V. alba, V. odorata, V. sintenisii (sect. Viola subsect. Viola), V. caspia, V. reichenbachiana and V. rupestris (sect. Viola subsect. Rostratae). General anatomical features of Viola species are discussed. The following characters were found to be taxonomically informative: cross‐section shapes of the aerial stem, petiole and peduncle as well as the number of vascular bundles in the aerial stem. In taxa of sect. Melanium, stem cross‐sections are elliptic with two wings, but they are semi‐circular with two wings in taxa of sect. Viola subsect. Rostratae. Within the latter subsection, the number of vascular bundles in the stem can be used to delimit the species V. caspia, V. reichenbachiana and V. rupestris. Calcium oxalate crystals were observed in all vegetative organs of taxa belonging to sect. Viola. All examined taxa had a secondary structure with the exception of V. somchetica. Our results show that anatomical characters are useful for delimiting species, subsections, and sections within Viola.     

国产堇菜属的分子系统学研究

利用基于ITS序列的分子系统学对国产堇菜属(Viola L.)进行了系统发育重建,选择Hybanthus concolor和鳞隔堇(Scyphellandra pierrei)作为外类群,对国产堇菜属植物54种1亚种共66个序列进行分析。分子系统树中获得强烈支持的中国堇菜属属下组级分类群有Sect.Viola、Sect.Diffusae、Sect.Bilobatae、Sect.Vaginatae。Subgen.Melanium与Sect.Viola和Sect.Trigonocarpae聚在一起,说明它们的亲缘关系较近,但Sect.Trigonocarpae支获得的支持率较低。分子系统树证实了Subgen.Dischidium与Subgen.Chamaemelanium的近缘关系,但不支持根据距的长短将Subgen.Dischidium划分为Sect.Longicalcaratae和Sect.Brevicalcaratae的处理。而Pinnatae类群应该作为Sect.Adnatae下的一个亚组处理。根据分子系统发育和形态学特征对V.lucens、V.magnifica、V.dissecta、V.yunnanensis等的分类地位进行了重新评价。     

A new species of Viola L. (Violaceae) from Sichuan, China

Viola muliensis Y.S. Chen & Q.E. Yang, a new species of Viola L. (Violaceae) from Sichuan, China, is described and illustrated. Viola muliensis is endemic to Muli County, south-western Sichuan. This new species has yellow flowers with very short spurs and belongs to subsection Brevicalcaratae W. Beck., section Dischidium Ging. It is distinguished from other related species by having pedatipartite leaves.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 365–368.     

A new species of Viola L. (Violaceae) from South Anatolia

A new species of Viola L., Viola yildirimlii M. Dinç & Y. Bağcı sp. nov. from South Anatolia is described and illustrated. It is found on the rocky slopes of Aladağ National Park, in the county of Adana, south Turkey, at an elevation of 1800 m. It belongs to Viola , subsect. Viola , and is similar to the Turkish endemics Viola isaurica Contandr. & Quézel and V. kizildaghensis M. Dinç & Ş. Yıldırımlı. Diagnostic morphologic characters for a detailed discrimination from two similar taxa and other Turkish Eflagellatae species are discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 477–482.     

Offspring performance in three cleistogamous Viola species

Seed germination and seedling/juvenile fitness in the cleistogamous perennials Viola hirta, V. mirabilis, and V. riviniana were investigated during three growing seasons, to compare the performance of chasmogamously (CH) and cleistogamously (CL) derived progeny. For V. hirta and V. mirabilis the effects of sibling competition were examined, for V. riviniana the effects of interspecific (grass) competition. Seed abortion and seed weight were also taken into account as fitness measures.In none of the species, seed abortion rate differed between CH and CL capsules. In V. mirabilis and V. riviniana, CL seeds had a lower germination rate than CH seeds. In V. hirta the two seed types did not differ in germinability. Mortality did not differ between the two seedling types in any of the species. In V. hirta and V. riviniana, CL progeny had shorter mean length of largest leaf than CH progeny. In V. mirabilis plant size did not differ between progeny types. Sibling competition had little effect on offspring performance, but grass competition increased mortality and reduced plant size of V. riviniana progeny. The two progeny types did not differ in their response to sibling and grass competition.The differences in performance between progeny types could be attributable to inbreeding depression in the CL phase, but the slightly lower fitness of CL offspring is probably balanced by their lower production costs. It is suggested that a dimorphic reproductive system is maintained in perennial Viola species to maximize total seed output in the face of environmental variation.     

Cleistogamy and phylogenetic position of Viola uliginosa (Violaceae) re‐examined

Viola uliginosa is an endangered species throughout its range in Central Europe. Based on an unusual combination of morphological characters, including a reported lack of cleistogamous (CL) flowers, it was previously placed in the monotypic subsection Repentes, section Viola. Available individual phylogenetic analyses identify V. uliginosa as a lineage deeply embedded in section Viola, but hitherto its exact position was unknown. Here we confirm the occurrence of cleistogamy in V. uliginosa. Unlike other species of section Viola, V. uliginosa produced both chasmogamous and CL flowers at the same time, not at different times of the year (seasonal cleistogamy). The two flower types differed conspicuously in morphology and microstructure, especially in size and shape of the pistil and stamens as confirmed by scanning electron microscopy. Secondly, we resolved the phylogenetic position of V. uliginosa as a lineage deeply embedded in subsection Rostratae by means of a multigene multispecies coalescent analysis using *BEAST. In this analysis, V. mirabilis and V. uliginosa were successive sisters to the rest of subsection Rostratae and, because allopolyploids exist between V. mirabilis and the last one, subsection Rostratae needs to comprise all these taxa to be monophyletic. The evolution of key morphological traits is briefly discussed.     

Infrageneric relationships of Korean Viola based on eight chloroplast markers

Phylogeny and infrageneric relationships were reconstructed for Korean Viola(Violaceae)using eight chloroplast sequences: the atpB-rbcL intergenic spacer,the atpF-H intergenic spacer,the matK gene,the psbA-trnH intergenic spacer,the psbK-I intergenic spacer,the rpl16 intron,the rpoC1 exon 2,and the trnL-trnF regions.The combined analyses of the eight chloroplast regions suggest that sections Dischidium and Chamaemelanium are monophyletic,whereas section Nomimium is paraphyletic; sections Dischidium and Chamaemelanium form a clade with subsections Hypocarpae and Trigonocarpae of section Nomimium,but unresolved within the clade; two subsections(subsects.Bilobatae and Vaginatae)of section Nomimium form a clade and are placed between the stem and stemless species group; and subsection Patellares of section Nomimium is monophyletic,and Viola dissecta is diverged first in the clade with high support(BS = 100,PP = 100).Our present results support the subsectional grouping based on morphological characteristics,although discordance remained at the series level.     

Two new species of Viola (Violaceae) from the Intermountain West,U.S.A.

Viola lithion andV. franksmithii are rock-dwelling endemics that belong to the North American perennial, caulescent, simple-leaved, blue-, purple-, or white-flowered group of species.Viola lithion, from the White Pine and Pilot mountain ranges of eastern Nevada and adjacent Utah, belongs to sectionChamaemelanium subsectionCanadenses. It most closely resembles the reniform-leaved, Olympic MountainV. flettii, together with which it possibly shares an ancestry with the widespread and allopatricV. canadensis. Viola franksmithii, restricted to the Logan Canyon drainage of the Bear River Range in northern Utah, belongs to sectionRostellatae subsectionRosulantes. It comes closest morphologically toV. howellii of the Coast Range, but likely evolved from the widespreadV. adunca or a common ancestor.     

Two new stoloniferous species of Viola (Violaceae) from China

Two new stoloniferous species of Viola (Violaceae) from southern China are described and illustrated. Viola nitida is recognizably different from V. fargesii as the plant is evergreen and glabrous throughout, the leaf blade adaxially nitid, base cuneate or shallowly cordate, and margin serrate. Viola maoershanensis is different from V. diffusa as the leaf blade is serrate, base cordate and not decurrent to petiole, the petiole wingless, the flowers larger, the petals bluish violet or pinkish white, and the lower petal obtuse at the apex. The chromosome numbers of the two new species were counted as 2n = 24 (V. nitida) and 2n = 44 (V. maoershanensis). The taxonomic positions of the two species are discussed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 349–356.     

Competitive responses of the rare Viola elatior and the common Viola mirabilis

Viola mirabilis is abundant in Estonia; Viola elatior is rare. We tested whether these species differ in their competitive responses to a common grass. We used a pot experiment in which individuals of each violet species grew with 0, 2, 4, or 8 individuals of Festuca ovina in natural soil. The response patterns of shoot and root mass and leaf numbers of the two species were similar: the biomass and leaf number of both violet species decreased with increasing grass density. The decrease of root mass of the rare V. elatior was significantly greater, and the decrease of leaf number tended to be greater, than that of the common V.␣mirabilis. We conclude that the stronger competitive response of V. elatior compared to V. mirabilis might be one reason behind its lower regional and local abundance.     

Viola umphangensis (Violaceae), a new species from Thailand

Viola umphangensis S. Nansai, Srisanga & Suwanph., a new species from Thailand, is described and illustrated. Morphological, anatomical and palynological characters of V. umphangensis are compared with a similar species, V. betonicifolia Sm. An amended key to the Viola species in the Flora of Thailand account is provided.     

中国堇菜属美丽堇菜亚属(堇菜科)植物分类学修订

对中国堇菜属美丽堇菜亚属植物进行了系统学修订,确认中国堇菜属美丽堇菜亚属植物共有6种,列出分种检索表,对《中国植物志》51卷中未记载的3种进行了详细的描述,并对塔城堇菜、深紫堇菜及隐距堇菜进行了绘图。     

Viola stoloniflora (Violaceae), a new species from the Ryukyus

Viola stoloniflora Yokota et Higa (Violaceae) is newly described from Okinawa Island, the Ryukyus. This species is characterized by the procumbent stolons, the almost round leaf blades, the densely pilose petioles, the ebarbate white petals with violet veins, and the chromosome number of 2n=22. The present species belongs to Sect.Nomimium, Subsect.Serpentes and is closely related to theV. iwagawai group of Sect.Nomimium, Subsect.Adnatae.     

Inferring species networks from gene trees in high-polyploid North American and Hawaiian violets (Viola, Violaceae)

The phylogenies of allopolyploids take the shape of networks and cannot be adequately represented as bifurcating trees. Especially for high polyploids (i.e., organisms with more than six sets of nuclear chromosomes), the signatures of gene homoeolog loss, deep coalescence, and polyploidy may become confounded, with the result that gene trees may be congruent with more than one species network. Herein, we obtained the most parsimonious species network by objective comparison of competing scenarios involving polyploidization and homoeolog loss in a high-polyploid lineage of violets (Viola, Violaceae) mostly or entirely restricted to North America, Central America, or Hawaii. We amplified homoeologs of the low-copy nuclear gene, glucose-6-phosphate isomerase (GPI), by single-molecule polymerase chain reaction (PCR) and the chloroplast trnL-F region by conventional PCR for 51 species and subspecies. Topological incongruence among GPI homoeolog subclades, owing to deep coalescence and two instances of putative loss (or lack of detection) of homoeologs, were reconciled by applying the maximum tree topology for each subclade. The most parsimonious species network and the fossil-based calibration of the homoeolog tree favored monophyly of the high polyploids, which has resulted from allodecaploidization 9-14 Ma, involving sympatric ancestors from the extant Viola sections Chamaemelanium (diploid), Plagiostigma (paleotetraploid), and Viola (paleotetraploid). Although two of the high-polyploid lineages (Boreali-Americanae, Pedatae) remained decaploid, recurrent polyploidization with tetraploids of section Plagiostigma within the last 5 Ma has resulted in two 14-ploid lineages (Mexicanae, Nosphinium) and one 18-ploid lineage (Langsdorffianae). This implies a more complex phylogenetic and biogeographic origin of the Hawaiian violets (Nosphinium) than that previously inferred from rDNA data and illustrates the necessity of considering polyploidy in phylogenetic and biogeographic reconstruction.