Occurrence of tetraploid and octoploid cytotypes in Senecio jacobaea ssp. jacobaea (Asteraceae) in Pannonia and the Carpathians

Four different cytotypes have been reported for Senecio jacobaea L. ssp. jacobaea throughout Europe, with the most common occurrence of tetraploids (2 n  = 40). Here we present a survey of previously published chromosome number data on this subspecies and its geographical distribution, and focus on populations from Pannonia and the Carpathians. Two ploidy levels have been determined in the study area, using chromosome counting and flow cytometry: tetraploid (2 n  = 40) and octoploid (2 n  = 80). Fifty-one populations originating from Slovakia, Austria, the Czech Republic, Hungary, Ukraine and Romania have been analysed. Multivariate morphometric analyses have been performed on 39 populations to study morphological differentiation between these two cytotypes. Despite slight morphological tendencies expressed on the level of populations, tetraploid and octoploids cannot be reliably distinguished morphologically and they are not taxonomically classified formally here.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 231–242.     

Iris stenophylla Hausskn. & Siehe ex Baker from Turkey and its cytology

A morphological study, both in the field and from herbarium material, showed that there are no convincing characteristics separating the Turkish endemics Iris stenophylla ssp. stenophylla and ssp. allisonii . A cytological survey revealed chromosome numbers of 2 n  = 24, 26 and 26 + 1B. No variation in either chromosome number or karyotype was found in ssp. allisonii , which always had 2 n  = 26, although the chromosome number and karyotypes of ssp. stenophylla were variable. Since neither morphological nor cytological differences were sufficient to separate the two subspecies, I. stenophylla ssp. allisonii is therefore reduced to a synonym of I. stenophylla .  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 115−127.     

Intraspecific taxonomy and ecology characterize morphological divergence among cutthroat trout (Oncorhynchus clarkii ssp. Richardson) populations

We compared the proportion of morphological variation accounted for by subspecies categories with the proportion encompassed by ecologically based categories in cutthroat trout ( Oncorhynchus clarkii ssp.), as a means of assessing the relative importance of each approach in identifying intraspecific diversity. We used linear and geometric morphometrics to compare measures of body shape, fin length, and head features between and within subspecies of cutthroat trout. Both categories accounted for a significant proportion of the variation between and within the subspecies; however, the larger proportion was explained by subspecific differences, with the greatest morphological divergence between coastal cutthroat trout ( Oncorhynchus clarkii clarkii ) and interior subspecies. Ecotypic categories within each subspecies also explained significant morphological differences: stream populations had longer fins and deeper, more robust bodies than lake populations. The largest ecotypic differences occurred between stream and lake populations of Yellowstone cutthroat trout ( Oncorhynchus clarkii bouvieri ). Given that many cutthroat trout subspecies are of conservation concern, our study offers a better understanding of intraspecific variation existing within the species, providing precautionary evidence of incipient speciation, and a framework of describing phenotypic diversity that is correlated with ecological conditions.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 266–281.     

Morphological differentiation among closely related species with disjunct distributions: a case study of Mediterranean Cyclamen L. subgen. Psilanthum Schwarz (Primulaceae)

In Cyclamen subgenus Psilanthum , previous authors have recognized three species ( C. repandum, C. balearicum and C. creticum ), with C. repandum subdivided into three subspecies (ssp. repandum , ssp. peloponnesiacum and ssp. rhodense ), and two varieties (var. vividum and var. peloponnesiacum ) within C. repandum ssp. peloponnesiacum . To examine the validity of this classification and the degree of morphological differentiation among closely related endemic taxa with disjunct distributions, we undertook analyses of three quantitative and eight qualitative morphological traits of flowers and leaves, most of which have previously been used without statistical comparison in the classification of the different taxa. Cyclamen balearicum showed high levels of morphological differentiation from all other taxa in subgenus Psilanthum consistent with its specific status. In contrast, morphological differentiation between C. creticum and subspecies of C. repandum was similar to that among the three subspecies of C. repandum . When compared with a recent molecular phylogeny of this subgenus, our results suggest that C. creticum may best be described as a geographical subspecies of C. repandum . The two varieties of C. repandum ssp. peloponnesiacum showed few significant differences for individual traits and their overall morphology is very similar to each other. The morphology of plants at three sites on Corsica strongly suggest, consistent with molecular data, the occurrence of hybrid populations between C. balearicum (outside of its previously recognized distribution) and local C. repandum .  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 133–144.     

Patterns of genetic diversity in related taxa of Antirrhinum L. assessed using allozymes

Allozyme diversity was studied within and among populations of five related taxa of Antirrhinum L. endemic to the Iberian Peninsula ( A. graniticum Rothm. ssp. graniticum , ssp. brachycalyx Sutton and ssp. ambiguum (Lange) Mateu & Segarra, A. boissieri Rothm. and A. onubensis (Fdez. Casas) Fdez. Casas). All of the studied taxa are obligate outcrossing endemic perennial herbs which form isolated populations. However, the taxa vary in range and population sizes, and are found on different soil types. The level and distribution of allozyme diversity differed widely between taxa: A. graniticum ssp. brachycalyx had the lowest level of allozyme diversity (H_T = 0.09), whilst the highest level was detected in A. boissieri (H_T = 0.25). Total variation was partitioned into within- and among-population variation. The proportion attributable to variation within populations varied from about 67% up to 84.3% and 89.5% in A. graniticum ssp. brachycalyx and A. graniticum ssp. ambiguum , respectively. Both these subspecies also showed little population divergence (G_ST = 0.10 and 0.09, respectively) and had high levels of estimated gene flow (Nm = 2.18 and 2.62, respectively). These results are discussed in relation to geographical proximity of populations and habitat continuity. Isolation by distance was not detected in any of the studied taxa. This result suggests that divergence among populations is due to random genetic drift.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 79 , 299–307.     

Paeonia daurica Andrews or P. mascula ssp. triternata (Pall. ex DC.) Stearn & P. H. Davis (Paeoniaceae)?

The peony in the Crimea of Ukraine and its allied populations have been variously taxonomically treated, as Paeonia daurica Andrews or P. mascula ssp. triternata (Pall. ex DC.) Stearn & P. H. Davis. Supported by the National Geographical Society, we have conducted extensive field observations and population sampling of this group in Turkey. In addition, relevant herbarium specimens from the herbaria B, BEO, BM, BUCA, E, G, GZU, K, P, SA, SOM, UPA, and WU were critically examined. Principal coordinate analysis was performed using MVSP-Version 3.13b analysis software. As a result, P. daurica was shown to be clearly differentiated from P. mascula in the number of leaflets/segments of the lower leaves and the shape of the terminal leaflets. P. daurica is diploid, except for three local tetraploids in the Caucasus, whereas P. mascula is consistently tetraploid. The two units were not found growing together, even in southern Turkey, where they are sympatric. P. daurica is considered to be a good species, which ranges from Croatia to Iran through Turkey and the Caucasus, and comprises six subspecies.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 1–11.     

Taxonomic status and relationships of bracken ferns (Pteridium: Dennstaedtiaceae) of Laurasian affinity in Central and North America

Detailed quantitative comparisons confirm and extend the discrimination of four major morphotypes amongst brackens of Laurasian affinity in Central and North America. These are recognized here at subspecies level as: Pteridium aquilinum sspp. feei , pubescens , latiusculum , and pseudocaudatum . Measurements of spore size indicate that sporophytes of P. aquilinum ssp. feei are diploid (2 n  = 104), as are sspp. pubescens , latiusculum , and pseudocaudatum . Phenetic cluster analysis based on DNA fingerprinting by arbitrarily primed polymerase chain reaction groups these four subspecies as genomically more similar to each other than to any taxa from other geographical regions. The chloroplast haplotype of ssp. feei is the same as that of sspp. latiusculum , pseudocaudatum , and pubescens with respect to the absence of both of the short direct repeats in the rps 4– trn S region (haplotype A), whereas the European ssp. aquilinum (haplotype B) has one of these repeats, and the Southern Hemisphere brackens P. arachnoideum and P. esculentum (haplotype C) have the other.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 1–17.     

Cyanogenesis variation in the Turnera sidoides L. polyploid complex (Turneraceae)

This study constitutes the first investigation of cyanogenesis variation within the Turnera sidoides autopolyploid complex. The qualitative and quantitative analysis carried out in 38 populations showed that the five subspecies of the complex are cyanogenic, although the levels of cyanide released varied both between and within subspecies. Between the subspecies, carnea , holosericea and integrifolia showed higher values of HCN released than subspecies pinnatifida and sidoides . Within subspecies, there is a tendency toward an increment of mean cyanide values with ploidy levels and different hypotheses are considered in order to explain this variation. The study of cyanogenesis in populations from different geographical areas cultivated under uniform greenhouse conditions showed that, in the widespread subspecies pinnatifida , HCN variation may also reflect the action of different selection pressures, since populations having dissimilar cyanide levels grow in different phytogeographical provinces with different climatic regimes. On the other hand, the fact that the variation of HCN released was greater between populations from different geographical areas than that observed between diploid and tetraploid populations of the same area provides additional evidence for the multiple origins of polyploids of the T. sidoides complex.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 85–94.     

The effects of loss of sex in clonal populations of an endangered perennial Coptis teeta (Ranunculaceae)

The reproductive biology of two subspecies of Coptis teeta was studied. The species is an endangered, herbaceous perennial, endemic to the Mishmi Hills in Eastern Himalaya . We investigated reproductive strategies of the subspecies, their breeding systems and reproductive output. They had contrasting reproductive strategies; ssp. teeta reproduces sexually and produces fewer ramets, while ssp. lohitensis propagates mostly vegetatively through a higher number of ramets . There is a resource adjustment between vegetative and sexual functions with a greater allocation to the former, particularly in ssp. lohitensis. Manual crosses showed that the species is self- and cross-compatible, but is essentially an autogamous selfer. Seed set in the species is generally low, with higher output in ssp. teeta and lower in ssp. lohitensis. The low fecundity is ascribed to a number of pre- and postfertilization events, including unsuitability of floral features for attracting pollinators, high pollen sterility and high rates of fruit and seed abortion. Sexual reproduction does not contribute significantly towards population increase in this species. This could be an important factor contributing to its rarity.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 47−54.     

Genetic diversity and differentiation of allopolyploid Dactylorhiza (Orchidaceae) with particular focus on the Dactylorhiza majalis ssp. traunsteineri/lapponica complex

Genetic differentiation of Dactylorhiza majalis ssp. traunsteineri from the Alps, Scandinavia, and Britain was studied and compared with other allotetraploid members of the systematically challenging genus Dactylorhiza . One-hundred and eleven populations from altogether 18 taxa were analysed for eight polymorphic plastid markers and two size-variable fragments from the nuclear internal transcribed spacer (ITS) region. In total, 60 plastid haplotypes and six ITS alleles were found among the 737 individuals analysed. No clear differentiation between populations of ssp. traunsteineri from the three regions was revealed. However, ssp. traunsteineri was genetically differentiated from Dactylorhiza baumanniana , Dactylorhiza elata , and D. majalis ssp. sphagnicola , although the majority of allotetraploid taxa remained inseparable. Judging from the degree of concerted evolution in ITS, D. majalis ssp. alpestris may be regarded as a relatively old allotetraploid, whereas ssp. baltica and ssp. purpurella may be considerably younger. Based on plastid data, the Alp region had the highest genetic diversity followed by Scandinavia and Britain. The geographic distribution of haplotypes provided support for possible refugial areas around the Alps and for several independent immigration routes into Scandinavia after the last ice age.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 52–67.     

The taxonomic status and relationships of bracken ferns (Pteridium: Dennstaedtiaceae) from sub-Saharan Africa

Two morphotypes of bracken fern in the genus Pteridium Gled. ex Scop. occur in sub-Saharan Africa. The more common and widespread of these is assigned in contemporary systematic treatments to the European subspecies, P. aquilinum (L.) Kuhn ssp. aquilinum . We show that this African form is separable morphometrically and genomically from the European subspecies, and reinstate its earlier name P. aquilinum ssp. capense (Thunb.) C.Chr. The second African bracken, with a more localized tropical distribution mainly in the drainage basins of the Congo and Zambezi River systems, is confirmed as P. aquilinum ssp. centrali-africanum Hieron. ex R.E. Fr. We reject suggestions that this taxon be treated as a full species. Phenetic cluster analysis based on use of Arbitrarily Primed Polymerase Chain Reaction (A-P PCR) and Inter Simple Sequence Repeat (ISSR) fingerprinting of the nuclear genome provides evidence that the two African brackens are more closely related to each other than to other taxa, and sister to a grouping of the European sspp. aquilinum and pinetorum . The two African subspecies share solely with ssp. aquilinum a distinctive chloroplast haplotype carrying a 5-base direct repeat in the rps 4 –trn S region, confirming the close phyletic relationship between sspp. aquilinum , capense and centrali-africanum .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 311–321.     

Morphometric study and taxonomy of Genista pulchella Vis. s.l. (Fabaceae), a south European species

A morphometric study of the known populations of Genista pulchella has been undertaken, based on herbarium specimens and field research. This has made it possible to provide a new taxonomic outline as the disjunct populations of G. pulchella (western part of the Balkan peninsula, central Italy and southern France) seem to be distinct. A new taxon is described: G. pulchella ssp. aquilana ssp. nov. (central Italy), a new combination G. pulchella ssp. villarsiana comb. nov. is proposed for the population from southern France, while G. pulchella ssp. pulchella can be considered endemic to the Balkan Peninsula. Genista pulchella Vis., G. villarsii Clementi and G. villarsiana Jord. are lectotypified.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 245–254.     

Classification of Primula sect. Auricula (Primulaceae) based on two molecular data sets (ITS, AFLPs), morphology and geographical distribution

On the basis of the study of c . 1100 herbarium specimens, field observations and molecular (ITS, AFLPs) evidence, Primula sect . Auricula is classified into two subsections, 25 species and six subspecies. Primula auricula L. ssp. widmerae (Pax) L. B. Zhang stat. nov. , P. auricula Linn. ssp. tatriaca L. B. Zhang ssp. nov. , and P. latifolia Lapeyr. ssp. cynoglossifolia (Widmer) L. B. Zhang stat. nov. are newly described or combined; P. auricula s.l. is divided into two species: P. auricula Linn. and P. balbisii Lehm., representing the northern and the southern populations of P. auricula s.l. resolved in the molecular study, respectively; P. cottia Widmer and P. balbisii Lehm. are recovered from synonymy; P. grignensis D. M. Moser and P. pedemontana E. Thomas ex. Gaudin ssp. iberica Losa et P. Monts. are synonymized with P. hirsuta All. and P. pedemontana , respectively; P. clusiana is postulated to be an allopolyploid species. Phylogenetic relationships in the section are discussed based on morphological and molecular variation and geographical distribution.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 1–26.     

Karyotype differentiation between Koelreuteria bipinnata and K. elegans ssp. formosana (Sapindaceae) based on chromosome banding patterns

Koelreuteria Laxm., a genus in the tribe Koelreuterieae (Sapindaceae), includes three tree species that are widely recognized as having horticultural merit. The two more closely related species, K. bipinnata Franch. and K. elegans (Seem.) A. C. Sm., are easily distinguished from K. paniculata Laxm. by their bipinnate leaves. In this study, both species were investigated cytogenetically and their karyotypes and heterochromatic patterns were compared. Koelreuteria bipinnata and K. elegans ssp. formosana (Hayata) F. G. Mey. both have 2 n  = 32 but their karyotypes present slight morphological differences when observed using conventional staining. Chromosome banding patterns are reported for the first time for this genus. Both species exhibit terminal heterochromatic blocks, as revealed by C-Giemsa and C-CMA₃, but the band size varies between the species. Koelreuteria bipinnata has larger heterochromatic blocks and more GC-rich segments, while in K. elegans ssp. formosana these bands are smaller. The relationship between the karyotype features in these closely related species is discussed.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 451–455.     

Synopsis of Turkish Acantholimon Boiss. (Plumbaginaceae)

A synopsis of Acantholimon in Turkey is given on the basis of a revisional study conducted in Turkey by the authors. An account of 52 species, ten subspecies, and 17 varieties is given, including keys to the species, subspecies, and varieties, and a general indication of their distribution, as well as their conservation status in Turkey. Seven new taxa, namely A .  ekimii sp. nov. , A. capitatum ssp. sivasicum ssp. nov. , A. calvertii var. glabrum var. nov. , A. huetii var. breviscapum var. nov. , A. acerosum ssp. longibracteolorum ssp. nov. , A. lycaonicum ssp. cappadocicum ssp. nov. , and A. halophilum var. coloratum var. nov. , are described, together with a few new combinations. A. wiedemannii , A. iconicum , and A. lycaonicum , described from Turkey in the past, but placed under other species as synonyms in the Flora of Turkey , are recognized once again as good species. A. multiflorum and A. parviflorum are given a species status. In Turkey, the presence of A. latifolium , A. araxanum , A. lepturoides , A. hohenackerii , A. senganense , A. fominii , A. petraeum , and A. tragacanthinum is also confirmed.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 397–419.     

Numerical taxonomy study of Salvia sect. Salvia (Labiatae)

Multivariate analysis was carried out in order to elucidate the taxonomic relationships between Salvia officinalis L., the type species of the genus, and S. fruticosa Mill., both taxa included in section Salvia (Labiatae). Seventy-five different herbarium specimens from all over the Mediterranean Region, belonging to these two taxa, were analysed. Twenty-four specimens belonging to S. lavandulifolia Vahl. s.l. , were used as outgroups. Twenty-six morphological characters were measured and a data matrix was constructed prior to multivariate analysis by means of R software. Multiple correspondence analysis was used to obtain a single dendrogram, applying Ward's minimum variance algorithm. This tree was used as a basis to propose a key for the determination of the species and subspecies studied. S. lavandulifolia is separated in two groups, one belonging to S. officinalis and another one representing what was called S. blancoana . Salvia officinalis L. ssp. gallica (W. Lippert) Reales, D. Rivera & Obón and S. officinalis L. ssp. oxyodon (Webb & Heldr.) Reales, D. Rivera & Obón are discussed as new combinations. In addition, a hybrid between S. officinalis and S. fruticosa is identified and its importance in gardening and cultivation is discussed. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 353–371.