Alkaloid chemotaxonomy of Nordic Papaver sect. Scapiflora (Papaveraceae)

A chemotaxonomic study of the Nordic Papaver species belonging to the Scapiflora group has been carried out by means of gas chromatography (GC), mass spectro-graphy (MS), and thin layer chromatography (TLC). The data obtained were analysed by the aid of hard and fuzzy clustering. Of the alkaloids identified, protopine and allocryptopine were found in all species investigated (i.e. Papaver radicaium - including all subspecies - P. dahlianum, P. lapponicum , and P. laesiadianum ). Protopine and allocryptopine were the only alkaloids found in P. dahlianum, P. lapponicum , and P. laesiadianum . In addition O -methylthalisopavine was found in all investigated subspecies of P. radicatum; reframine in P. radicatum sspp. ovatilobum, gjaerevollii, groevudalense, intermedium, oeksendalense, radicatum, relictum , and subglobosum; reframidine in ssp. hyperboreum; cryptopine in sspp. ovatilobum, gjaerevollii, groevudalense, relictum, subglobosum, macrostigma , and hyperboreum; muramine in sspp. macrostigma and hyperboreum; and amurine in sspp. relictum and subglobosum .
These chemotaxonomic results confirm the systematics of the Nordic Papavers of the sect. Scapiflora introduced by Knaben who based the taxonomy on morphological and cytological characteristics. Moreover, the content of alkaloids in the different isolated Nordic Papaver populations may be of plant geographical importance. There are, for instance, a marked chemical difference between all the South Norwegian subspecies of P. radicatum and the North Norwegian P. radicatum ssp. subglobosum on one hand and the rest of the North Norwegian subspecies on the other.     

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.     

Polyploidy and karyotype variation in Turkish Bellevalia (Hyacinthaceae)

There are 20 Bellevalia species in Turkey, half of which are endemic. Chromosome numbers are known for 15 species. A chromosome survey of 145 Bellevalia individuals showed that the karyotype is remarkably stable. All are based on x  = 4. The majority are diploid with 2 n  = 8, but there is also a polyploid series of 2 n  = 16, 24 and 32. Aneuploidy occurs only at the octoploid level. Eleven individuals had metacentric B chromosomes, one had acrocentric Bs and one had telocentric Bs. Bellevalia pycnantha and B. paradoxa are morphologically similar, with B. pycnantha reduced to a synonom of B. paradoxa .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 87–98.     

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.     

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.     

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.     

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.     

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.     

Morphological and pollen differentiation in Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii

A multivariate analysis to differentiate morphologically the populations of wild potatoes Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii was carried out. An analysis of the morphology and the viability of pollen of these potato populations was also made. The results of the morphometric analysis indicate that both subspecies are phenetically different. The pollen grain shape and size in ssp. cardiophyllum are different in northern and southern populations. Pollen diameter is significantly different between subspecies. Based on these results we propose that these taxa should be considered as two separate species.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 415–426.     

Determination of ploidy and nuclear DNA content in populations of Atriplex halimus (Chenopodiaceae)

Nuclear DNA contents were determined by flow cytometry for 20 populations of the perennial C₄ shrub Atriplex halimus L. (Chenopodiaceae) originating from the Mediterranean basin and Fuerteventura (Canary Islands). Two populations were also analysed for chromosome number: one (from Ibiza, Spain), with a 2C nuclear DNA content of 2.40 pg, was shown to be diploid (2 n  = 2 x  = 18), whilst the other (from Sicily, Italy), with 5.11 pg, was tetraploid (2 n  = 2 x  = 36). With respect to nuclear DNA content, two groups of populations were detected, diploids with 2.40–2.44 pg and tetraploids with 4.77–5.13 pg. The diploid populations were mainly from the western Mediterranean (Spain and France) and Fuerteventura, whereas tetraploids were generally, but not exclusively, from more arid areas in North Africa and the eastern Mediterranean. In general, the diploid and tetraploid populations corresponded to the subspecies halimus and schweinfurthii , respectively. For certain populations having morphologies intermediate between those considered typical of these two subspecies, nuclear DNA contents showed them to be tetraploid. There was significant variation in nuclear DNA content among the tetraploid populations, with greater values in the more easterly populations.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 147 , 441−448.     

Chromosomal variability in the genus Primula (Primulaceae)

Chromosome numbers and measurements were recorded in 47 individuals of ten taxa of Primula , representing eight species and two subspecies from ten populations. The basic numbers of chromosomes were x  = 8, 9 or 11, and they were mostly metacentric, medium-long to medium-small, ranging in length from c . 3.6 µm to 1.6 µm. Diploid chromosome numbers of 2 n  = 2 x  = 24 and 22 were scored for the first time in P. loeseneri Kitag. and P. prenantha Balf. & Sm., respectively. A new ploidy level of 2 n  = 4 x  = 44 was found in P. burmanica Balf. & King. The recorded diploid numbers of the remaining species confirmed earlier reports. Interspecific variability in chromosome numbers was correlated with heterogeneity in their mean length. Comparison of the data with those in the literature revealed that the observed variability of chromosomal characters was compatible with the other taxonomic criteria and supports the current taxonomic delimitation. Chromosomal variation at the diploid level is the predominant feature in Primula evolution, and allopatric speciation has played a major role in its specific diversity. Subgen. Aleuritia could represent the main clade from which the other subgenera have evolved. The origin of the chromosome numbers, geographical distribution and evolution of the species were assessed, and the parallel polyphyletic mode of evolution in the genus was confirmed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 211–219.     

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.     

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.     

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.     

The Deschampsia cespitosa complex (Poaceae: Aveneae) with special reference to Russia

The Deschampsia cespitosa complex in Russia is revised. Related taxa common to Russia and adjacent regions (i.e. northern Europe, Arctic America and Japan) are also considered. Deschampsia cespitosa s.l. is characterized by open to less commonly rather contracted panicles, spikelets (1) 2 (3)-flowered, 4-toothed lemmas with the lateral teeth larger, and awns straight or bent, inserted from the base to the top of the lemmas. Due to extensive overlapping in diagnostic characters, it is not possible to accept the specific status for D. borealis (Traut.) Roshev., D. brevifolia R.Br., D. macrothyrsa Tatew. et Ohwi and D. obensis Roshev; instead, they are treated as subspecies of D. cespitosa (L.) Beauv. Morphological differences are insufficient for a clear differentiation among subspecies, and geographical distributions also need to be considered. A total of 14 subspecies is accepted. Observations on ecology, problems of specific and subspecific delimitation, a distribution map, and considerations of evolution of the complex are provided. One new combination is made, Deschampsia cespitosa ssp. sukatschewii (Popl.) Chiapella & Prob.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 213–228.     

Clarification of the systematic position of Hippophae goniocarpa (Elaeagnaceae)

New information on (1) Hippophae goniocarpa Y.S. Lian et al . ex Swenson et Bartish and (2) H. goniocarpa ssp. litangensis (Lian et X.L. Chen ex Swenson et Bartish) Lian et K. Sun from China is provided and illustrated. Based on morphological characters and distribution, H. litangensis Y.S. Lian et X.L. Chen ex Swenson et Bartish is re-described as a subspecies. H. goniocarpa (including ssp. litangensis ) is a distinct species and should be placed in Section Gyantsensis of Hippophae , although it has been suggested that sspp. litangensis and goniocarpa are the result of hybridization between H. rhamnoides ssp. sinensis and H. neurocarpa ssp. neurocarpa , and H. rhamnoides ssp. yunnanensis and H. neurocarpa ssp. stellatopilosa , respectively. The type subspecies of H. goniocarpa is distributed in Songpan and Hongyuan Counties of Sichuan Province and Qilian County of Qinghai Province, at 3500–3750 m, whereas ssp. litangensis is found only in Jiawa, Litang County, Sichuan Province at c . 3700 m. A key including the four taxa of sect. Gyantsensis is provided.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 425−430.     

Chromosome counts in the genus Festuca section Eskia (Poaceae) in the Iberian Peninsula

Chromosome numbers of taxa of Festuca L. section Eskia Willk. in the Iberian Peninsula are given. The levels of ploidy for five taxa are confirmed. Idiograms and karyotypic formulae of the five taxa are presented for the first time. Two levels of ploidy occur in this section: diploid and tetraploid. One taxon, Festuca elegans ssp. merinoi is tetraploid and two other taxa have diploid and tetraploid populations. The remaining two taxa are solely diploid.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 331–337.