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Patterns and relationships between and within the sections Alchemilla and Ultravulgares of the genus Alchemilla (Rosaceae) in Estonia
Nordic Journal of Botany
Abstract
Nine Estonian Alchemilla species belonging to the sections Ultravulgares and Alchemilla, or considered being close to them, were analysed. When the analysed specimens were divided into sections, the latter were statistically distinct and formed separable groups in the character space. When the specimens were grouped on a species level, A. cymatophylla, A. subcrenala and A. heptagona were insignificantly distinct, but, in the character space, specimens of A. heptagona were visually well distinguished from the other two. Specimens of A. acutiloba, A. micans, and A. xanthochlora formed confidently distinct (p < 0.01) species-clusters, but at the same time they formed a joint cloud in the character space, indicating the compactness of section Alchemilla. Specimens of A. semilunaris were close to A. lindbergiana, and not, as has previously been supposed, to the section Ultravulgares. A. semilunaris should possibly be kept in a separate section Decumbentes. Specimens of A. lindbergiana were on one hand close to A. semilunaris. on the other hand close to the species of section Alchemilla; including it in the latter section is still doubtful. A. subglobosa was indistinct from A. subcrenata but could generally be separated from sections Alchemilla and Ultmvulgares. Of the 41 tested characters, 35 were useful for species discrimination; counts, nominal and ratio characters were better than metric ones. According to the cluster analysis, specimens of A. acutiloba and A. micans formed one big cluster, all other specimens belonged to another.
... (Juzepczuk, 1941 (Czapik, 1996;Bradshaw, 1962;Izmailow, 1981 (Asker & Jerling, 1992;Sepp & Paal, 2001;Horandi, 2004; . (Juzepczuk, 1941;Davis, 1972;Fröhner, 1969Fröhner, , 1990Fröhner, , 1995Khatamsaz, 1992 ...
Floral morphology of 24 species of genus Alchemilla in Iran, were studied using light and digital microscopies. Flower samples were collected from different species and their morphological traits including hypanthium, calyx and epicalyx, pedicels, ovaries and stamens measured. Among them, calyx shape and length, trichomes position on calyx and pedicle were the most useful features for separating species. Principal component analysis (PCA) and cluster analysis (CA) were used to determine the potential contribution of flower morphological characters in expressing the species relationships. A total of 17 qualitative and quantitative characters were included in both analyses. The results of UPGMA and PCA methods were consistent with each other and current classifications....
... We used the leaves of diploid P. lactucella as a reference standard. Relative fluorescence intensity of DAPI stained nuclei (1500–5000 particles) was measured by Partec Cyflow instrument equipped with a HBO lamp.Sneath & Sokal, 1973) has been successfully used in several taxonomically difficult genera with widespread apomictic reproduction (e.g., Crataegus : Smith & Phipps, 1988; Alchemilla : Sepp & Paal, 2000; Ranunculus : Hörandl & al., 2009; Sorbus : Lepší & al., 2009 ), including the closely related genus Hieracium (Chrtek & al., 2007; Tyler, 2006). Surprisingly, as far as we know this approach has never been applied in the genus Pilosella, except still unpublished work of Urfus (Urfus, 2006). ...
Members of the Pilosella alpicola species group (Asteraceae) are distributed throughout the alpine region of the European mountains (Alps, Carpathians, Balkan mountains). Like other Pilosella species groups (Hieracium subg. Pilosella ), the taxonomy and species' relationships are poorly understood mostly due to widespread facultative apomixis, frequent hybridization and polyploidization—the most important phenomena substantially involved in the evolutionary history of the genus. We assessed morphology, ploidy level, variation in breeding system and molecular variation within the P. alpicola group to provide a new taxonomic concept and to clarify evolutionary relationships among species and origin of polyploids. Multivariate morphometric analyses (UPGMA, CDA, PCA) applied on 324 plants originated from 21 populations revealed existence of four well-separated clusters corresponding to four allopatric taxa: P. alpicola s. str. (Alps), P. rhodopea (Balkan Peninsula, Southern Carpathians), P. serbica (Serbia and Montenegro) and P. ullepitschii (Carpathians). In total, four ploidy levels (2 x, 3 x, 4 x, 5 x ) were detected among 557 plants analysed from 19 populations by classical chromosome counting and flow cytometric analysis. The tetra- and pentaploids of P. alpicola s. str. have an allopatric distribution (Wallis Alps vs. Dolomites, respectively). Four ploidy levels with complex cytogeographic pattern and high frequency of mixed ploidy populations (75%) indicating a primary contact zone were recorded inP. rhodopea. Pilosella ullepitschii andP. serbica are exclusively diploid and are both sexually reproducing. In spite of a clear morphological separation, a molecular analysis (ITS and cpDNA sequences) suggests close relationships and rather recent origin of all studied taxa, except P. alpicola s. str. The latter taxon is an agamospermic allopolyploid that likely originated polytopi cally from a hybridization between P. rhodopea from the Balkans and P. glacialis from the Alps. In contrast to P. alpicola s. str., our data strongly support an autopolyploid origin of P. rhodopea polyploids which reproduced strictly sexually. Pilosella petraea , sometimes treated as a member of the P. alpicola group, differs from the remaining taxa by its conspicuous morphology, ecology and ITS polymorphism and should be removed from the group. Range shifts and extinctions were likely involved in shaping the evolutionary and modern distributional pattern of the group. Our combined methodological approach enabled us to propose a new taxonomic circumscription for the P. alpicola group and revealed auto- and allopolyploidization events.
The distinctions between Alchemilla ser. Calycinae and A. ser. Elatae are emended, treating the latter group as relict, connected to the Tertiary broadleaved forests of the eastern Mediterranean Basin. Alchemilla ser. Calycinae (in renewed circumscription) is subdivided into seven provisional aggregates according to the specially developed coordinate system (with two axes — a ratio of central zone width to leaf length vs. number of leaf teeth in total). The original key included all the species of the series occurring in the Caucasus and Asia Minor was compiled.
Gametophytic apomixis (asexual seed formation without syngamy of female and male gametes) is a highly interesting mechanism for researchers in plant biotechnology, genetics, evolutionary biology, and taxonomy. Apomixis evolved repeatedly and independently in the evolution of multiple genera. It is an effective reproduction barrier and, consequently, conserved apomictic genotypes may become overrepresented in nature. Apomictic plants may easily colonize free niches with only one or a few individuals and outcompete outcrossing plants. In spite of the indisputable pros of asexual reproduction, apomixis also has several cons. One of the most commonly mentioned is the accumulation of deleterious mutations in asexual lineages and decreased genetic variation. However, apomicts in general can be genetically highly diverse. The most common sources of this variation are the accumulation of mutations, hybridization with sexual plants, and facultative apomixis. Facultative apomicts are highly variable in their level of residual sexuality, which increases their genotypic and phenotypic variation. Even in the case of obligate apomicts, gene flow is possible due to functional male meiosis and the production of viable pollen grains by apomicts. Apomixis occurs in plant genera in which hybridization together with polyploidization play an important role in diversification and causes severe problems in taxonomy. How to accommodate apomictic taxa in taxonomic treatments, and understanding what should and what should not be referred to as a species are intriguing questions. This review aims to provide an overview of the main characteristics of “apomictic genera” and the approaches used to treat apomictic taxa within these genera. To achieve this aim, the review was divided into several parts. Firstly, the distinctive features of apomictic reproduction and apomictic taxa are described together with issues related to the taxonomic evaluation of apomictic taxa. The second part discusses approaches in the known apomictic genera, and the final part presents the authors’ view on important points, which need to be taken into account in the classification of apomictic taxa.
Apomictic microspecies, for example those of the genus Alchemilla, are often difficult to distinguish. Still, their differences are thought to be persistent due to apomixis. Apomicts are argued
to have general-purpose genotypes. The present study aims to assess the variation of morphological characters of Alchemilla depending on environmental conditions both in nature and experimentally; to evaluate the efficacy of characters for identifying
the different microspecies; to assess the similarity of different microspecies; and to determine if Alchemilla microspecies have general-purpose genotypes that are not dependent on environmental conditions. The variability of selected
characters in seven microspecies of Alchemilla was studied in nature and five microspecies were grown in a common garden experiment. The growing conditions of the latter
were subjected to various manipulations (fertilization, shading and irrigation). Typical natural habitats of Alchemilla exhibited only small differences. Environmental conditions therefore had little effect on morphological characters. Neither
cultivation in the common garden nor manipulation of conditions therein had a significant impact on the discrimination of
microspecies. However, the metric characters were larger in the garden relative to those observed in nature, particularly
under fertilization. Fertilization affected most characters, whereas shading and irrigation did not. The most effective characters
for discriminating between microspecies were the ratios of the metric characters. Two species pairs: A. vulgaris and A. micans, as well as A. glaucescens and A. hirsuticaulis were morphologically close; however, the species within these pairs could still be distinguished. The morphological characters
of Alchemilla microspecies were only slightly dependent on the environmental conditions.
Väitekirja elektrooniline versioon ei sisalda publikatsioone. Käesolevas töös analüüsiti võrdlevalt hariliku koldrohu (Anthyllis vulneraria) ja hariliku kortslehe (Alchemilla vulgaris) liigikomplekside morfoloogilist ja geneetilist varieeruvust. Hariliku koldrohu puhul mõõdeti kokku 26 tunnust looduslikest populatsioonidest kogutud isendeil ja lisaks herbaarmaterjalil Eesti erinevatest herbaariumidest. Saadud mõõtmistulemusi analüüsiti statistiliste meetodite abil. Ka hariliku kortslehe morfomeetrilisteks analüüsideks kasutati looduslikest populatsioonidest kogutud isendeid, kuid analüüsiti geograafilisi ja ökoloogilisi mõjusid morfoloogilistele tunnustele. Lisaks hinnati hariliku koldrohu geneetilist mitmekesisust molekulaarsete tunnuste alusel kasutades valikulist restriktsiooni fragmentide amplifikatsiooni polümeraasi ahelreaktsioonil (AFLP). Töö tulemusel võib öelda, et nii A. vulneraria s. l. kui A. vulgaris (coll.) on liigikompleksid, mida iseloomustavad keerukas paljunemise süsteem, üleminevate seisunditega tunnused, kompleksi kuuluvate taksonite raskestimääratavus ning koldrohtude puhul ka arvatavate hübriidide esinemine. Kortslehtede puhul on tegemist püsivate tunnustega erinevates keskkonnatingimustes ning mikroliikide eristamine seega antud tulemuste põhjal igati õigustatud. Koldrohtude liigikompleksis aga on liikide eristumine vähem tõestatud – morfomeetrilisel analüüsil saadud tulemused toetavad viie mitte kaheksa liigisisese taksoni eristamist. Samade taksonite molekulaarne analüüs aga ei erista neist ühtegi. Seega on ilmselt vajalikud edasised uuringud morfoloogiliste tunnuste stabiilsuse hindamiseks sarnastel tingimustel või erinevate manipulatsioonide korral. The present study was performed to investigate the morphological variation of A. vulneraria s. l. and A. vulgaris (coll.). As visually subtaxa of these two are hardly separated, a morphometrical study is presented in order to address questions about the variability of taxa. The efficacy of different morphological characters to identify the taxa is estimated. To assess the plasticity of morphological characters under different environmental and experimental conditions Alchemilla microspecies were grown under manipulated conditions in a common garden experiment. As molecular data are often recommended for distinguishing hardly recognizable taxa within species complexes, a molecular study of Anthyllis vulneraria s. l. was carried out. The level of AFLP variation between and within seven intraspecific taxa was investigated and the genetic differences of the taxa were quantified between contrasting edaphic and climatic conditions. The results of morphometrical analysis of this work support the fact that both Anthyllis vulneraria s. l. and Alchemilla vulgaris (coll.) can be considered species complexes, while intraspecific taxa of these two are morphometrically complex and continuous. It can be said that the current work supports the existing system for Alchemilla microspecies and these taxa are morphologically rather distinct. In contrast, obtained results support the deduction of taxa distinguished within A. vulneraria s. l. It was found reasonable to distinguish at the subspecies or even variety level five morphological taxa. Still, similar common garden experiment and manipulations as used on A. vulgaris species complex would make a better basis for final conclusions.
The multivariate structure of the Scutellinla umbrorum complex (feuzales, Ascomycetes), based on the morphometrical parameters of 81 specimens from five ecotopes in the Netherlands, was analysed. According to conventional expert estimation, five putative taxa resp. species were established: S. patagontca (Rehm) Gamundi, S. aff. subhirtella Svrček, S. umbrorum (Fr.) Lambotte, S. parvispora J. Moravec and S. subhirtella s. Kullman. These taxa form a taxonomic continuum hardly separable by traditional taxonomy. Five clusters obtained by UPGMA (with the generalized J-distance for mixed data as a measure of resemblance) are more distinct and in good accordance with ecological factors; some of them, however, are statistically not well separated. The revision of the clusters' structure by k-means approach yields highly discontinuous clusters. The morphometric characters of specimens differ when going from open habitats to the forest. Differences are also revealed in phenology: the growing season starts in the forest later than in open habitats. The data are divided into two subsets according to spore ornamentation and spore width, the withingroup variation of either subset is caused mainly by the length of marginal hairs. On the basis of several statistical methods a supposition was introduced that the S. umbrorum complex probably consists of two polymorphic species, S. umbrorum (Fr.) Lambotte and S. subhirtella Svrček s.l., with the mean value of marginal hairs longer than 450 μm and shorter than 450 μm, respectively. The UPGMA clusters can be interpreted as ecodemes of respective species.
On the Karula Upland, South Estonia, the forest moss layer was analysed using a transect of 726 contiguous 0.2×0.2 m plots.
The sample plots were classified according to a multistage clustering procedure based on the sequential use of algorithms
with different criteria. Several obtained clusters are rather similar in species composition, but abundance relationships
among dominant species are distinctly different. For detailed analysis of mutual relations among societies a formal definition
of adjacency is proposed, and two aspects of the cluster continuum — transitionality and distinctness — are estimated. It
appears that almost all resulting societies are very distinct (P<0.05), but at the same time can be continual in the sense of transitionality. Spatial changes in vegetation along transects
are also discontinuous. The null hypothesis assuming the independency of neighbouring sample plots type was refuted withP=0.01. The spatial extent of different synusiae is typically several times larger than it should be if there were no structure
in moss vegetation.
This new release of the package contains 27 FORTRAN and 4 BASIC programs written for IBM-PC and compatible machines and Apple Macintosh II, Plus and SE personal computers. Widely used standard multivariate methods and specific data analytical techniques, some of them suggested by the author, are represented in the package. The procedures programmed include hierarchical and non-hierarchical clustering, fuzzy classifications, block clustering, ordination, character ranking, comparison of classifications and ordinations, consensus methods, Monte Carlo simulation of distributions of partition agreement, simulated sampling based on digitized point patterns, and information theory functions for evaluating species assemblages. This paper gives general information on the programs; technical details are presented in the user’s manual.
Alchemilla L. (Rosaceae) contains numerous agamospermous microspecies, which are often treated as species. However, many of them are not clearly morphologically distinct, and their genetic variability is practically not investigated. In the present study, we used RAPD analysis to assess the genetic relatedness between Alchemilla microspecies. In all, 51 plants from 12 Alchemilla microspecies were analysed, and 116 characters were considered (68 RAPD bands over three primers and 48 morphological characters). Phylogenetic trees were constructed by the unweighted pair-group method, neighbour-joining and maximum parsimony methods. The genetic data supported most Frohner's system of sections. Despite the use of a limited set of data in the investigation and weak support values, some tentative conclusions could be based on congruence of the RAPD analysis and morphological data. Alchemilla acutiloba Opiz and A. micans Buser should be united as a single microspecies, A. micans; section Plicatae should be divided into two series Pubescentes and Barbulatae; and A. heptagona Juz. may be separated in Exuentes series of Ultravulgares.
23 widespread apomictic Alchemilla microspecies occurring in Estonia are analyzed to investigate whether the species and higher rank taxa are distinct, how variable these taxa are and which characters distinguish them better. Cluster analysis, discriminant analysis, analysis of variance, principal component analysis and continuum analysis are used for data processing. The characters form four correlative groups, describing (i) vegetative and (ii) generative parts of the plant body, (iii) hairiness characters and, (iv) leaf teeth measurements. The best characters according to analysis of variance for disünguishing species are hairiness characters, but often they distinguish only few species very clearly and cannot be used for the remaining ones. Hence the other characters cannot be excluded. From the studied species only A. plicata, A. semilunaris and A. lindbergiana are significantly distinct from all others. The remaining ones form a complicated network of mutually indistinct pairs. Higher rank taxa — sections and series according to Rothmaler, Fröhner and Yuzepchuk are better separated, containing very few mutually indistinct pairs. Results from species centroids' clustering are most congruent with Fröhner's system, but still some changes seem to be necessary and a corrected system is proposed here. Section Plicatae is split into two series: Pubescentes and Barbulatae, sections Alchemilla, Ultravulgares and Decumbentes are joined as three series of section Hirsutae, and A. filicaulis is moved from section Plicatae to section Coriaceae. Coriaceae should also be split into three series: Exuentes (A. filicaulis), Glabricaules (A. glabricaulis) and Coriaceae (other species).
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