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In 83 species of the families Monotropaceae, Apocynaceae, Oleaceae, Scrophulariaceae, Lentibulariaceae, Bignoniaceae, Martyniaceae, Myoporaceae, Verbenaceae, Lamiaceae, Campanulaceae, and Liliaceae, protein bodies in the cell nuclei have been found, in 68 of these species for the first time. On the basis of their structure in accordance with morphological characters the genera Burgsdorfia, Hesiodia, Olisia, and Phlomoides of the Lamiaceae are accepted; Lamium is divided into Lamium, Lamiastrum and Orvala; two new combinations are established: Kickxia campyloceras ( Rech. fil. & Esfandiari) Speta and Etornotus papilionaceus ( Burm. in L.) Speta. Deviating shape or lack of protein bodies corroborate former taxonomic decisions, e.g. the transfer of Monotropa to Monotropaceae or the separation of Galeopsis and Ladanum; Veronica schmidtiana should not be included in Pseudolysimachion. Systematic affinities are discussed primarily within Scrophulariaceae because nuclear protein bodies have been studied already in many species of this family. For Campanula patula two 2 x populations are reported. [ABSTRACT FROM AUTHOR]

This study presents an analysis of plastomes to assess relationships among the seven genera of tribe Myoporeae (Scrophulariaceae). Sampling included individuals from type species for all seven genera included in Myoporeae (Bontia, Calamphoreus, Diocirea, Eremophila, Glycocystis, Myoporum, Pentacoelium), one additional species of the large genus Eremophila, one representative of the sister tribe Leucophylleae (Leucophyllum), and the previously published plastome of Scrophularia takesimensis as an outgroup. Phylogenetic analyses were performed using maximum parsimony and Bayesian inference analyses based on > 7000 variable nucleotide positions. Maximum parsimony analysis produced a single tree with the same topology as the Bayesian consensus tree, and all nodes were fully resolved with 100% bootstrap support and 1.0 Bayesian posterior probability. Tribe Myoporeae was supported as monophyletic and sister to Leucophylleae. Two lineages are identified within Myoporeae; one containing the Australian endemic genera Calamphoreus, Diocirea, Glycocystis and the type species of Eremophila (E. oppositifolia), the second containing species with distributions outside of Australia (Bontia, Myoporum, Pentacoelium) and a species of Eremophila (E. gibbifolia). Eremophila is thus supported as polyphyletic. The application of genome skimming and assembly of plastomes has resolved generic relationships in tribe Myoporeae and revealed variable regions of the genome that will be useful in further genetic study of the group. [ABSTRACT FROM AUTHOR]

Cynoglossinae with ca. 200 spp. is one of the taxonomically most challenging subtribes of tribe Cynoglosseae with regard to generic delimitations. Here, we present a phylogenetic analysis based on nuclear ITS and cpDNA trnL-F and rps16 sequences of 270 accessions including 102 newly sequenced ones, representing all currently recognized genera of Cynoglossinae to clarify relationships among Cynoglossum and allied genera. We conducted maximum likelihood, maximum parsimony, and Bayesian inference analyses on separate and combined datasets. Cynoglossinae is recovered as a monophylum of closely related genera. Subtribe Cynoglossinae falls into three major groups: (1) Clade I including Microparacaryum spp., (2) Clade II comprising two species of Lindelofia, several Cynoglossum and Paracynoglossum spp., and (3) a larger clade (Clades III + IV) consisting of Solenanthus, Rindera, Trachelanthus, Paracaryum, Pardoglossum, Mattiastrum, and the remainder of Cynoglossum. None of the genera were monophyletic in the combined nuclear-plastid and nuclear datasets, except for Microparacaryum. A close relationship between Solenanthus apenninus and Cynoglossum officinale is shown, which suggests transferring S. apenninus to Cynoglossum. Moreover, Pardoglossum atlanticum, the type of the genus, is placed within Mediterranean Cynoglossum corroborating its synonymy under Cynoglossum. Lindelofia olgae is distantly related to the type of the genus (L. longiflora), but more closely connected with Solenanthus core-group. The majority of species currently recognized as members of Rindera do not form a clade with the type of the genus (R. tetraspis). The genus Paracaryum as currently circumscribed is paraphyletic and includes some species of Mattiastrum and Rindera. [ABSTRACT FROM AUTHOR]

Locally adapted ecotypes can constitute an important part of the biodiversity, especially in young floras with few endemic species. However, the origins, distinctness and conservation value of many ecotypes remain uncertain because genetic data are lacking or no common-garden study has been carried out. In the present study, we evaluated the distinctness and genetic structure of a phenotypically deviating morph of Rhinanthus angustifolius, growing in calcareous spring fens on the Baltic island of Gotland. Using data from a common-garden experiment and analyses of nuclear microsatellite variation, we compared fen populations on Gotland with conspecific populations from habitats more typical of the study species. We also included the fen specialist R. osiliensis from the Baltic island of Saaremaa in the molecular analyses to make further inferences about the origin of the Gotlandic fen morph. Our data indicate that the Gotlandic fen populations constitute a phenotypically and genetically distinct ecotype that most likely has evolved at least two times on Gotland after the last ice age. In congruence with previous studies, we also infer that fen ecotypes have evolved independently on Gotland and Saaremaa. We propose a varietal status for the Gotlandic fen ecotype and give recommendations for the conservation of this taxon. [ABSTRACT FROM AUTHOR]

The size and shape of the pollen grains are studied in a hybridogeneous complex of the genus Onosma. This plant system encompasses the hybrid taxa O. helvetica and O. pseudoarenaria with bimodal karyotypes that comprise large and small chromosomes and their ancestral lineages, paternal O. fastigiata and Asterotricha taxa belonging to a maternal lineage. The pollen grains are investigated under light microscopy, and the size and shape parameters are retrieved based on image analysis of the pollen grain outlines. The shape of the pollen grain is analyzed by applying elliptic Fourier transform coupled with principal component analysis. Extensive size and shape variations of the pollen grain within cytotypes as well as differences between cytotypes are recorded. The cytotypes are mostly differentiated due to the size parameters of the pollen grain that are correlated with ploidy level and the proportion of large and small chromosomes. Despite significant differences being observed between cytotypes, the precise identification of the ploidy level of an individual based on the pollen size and shape analysis is not possible in most of the cases, mostly due to extensive intracytotype variation. Allometric analyses exhibit a different relation between the pollen shape and size in case of cytotypes. The ratio of polar to equatorial axes length is, generally, a good approximation of the pollen shape. However, the study clearly exhibits that the relation between pollen shape and the ratio of polar to equatorial axes length is different for cytotypes, which justifies the application of outline analysis in palynological studies. [ABSTRACT FROM AUTHOR]

Onosma atila- ocakii sp. nova [Boraginaceae, Sect. Asterotricha (Boiss.) Gürke] is described and illustrated as a new species. It grows in a steppe on magnesite rocks in Eskişehir province (Central Anatolia, Turkey). Its description, photographs, and an identification key including related species ( O. roussaei DC. and O. aucheriana DC.) are given. Diagnostic morphological and palynological characters of closely related species are compared and discussed. The characteristic features of indumentum and pollen structure are studied using a light microscope. The new species distinctly differs from related species because of its setose and adpressed retrorse–pilose indumentum, 2–3 rows of setules on foliar tubercles, 10–14 mm white petals and 2.1–2.6 × 1.7–2 mm dark brown nutlets. The International Union for Conservation of Nature threat category and observations on the ecology of the populations are noted. The distribution map of this new species and closely related species in Turkey is presented. [ABSTRACT FROM AUTHOR]

Lappula Moench includes ca. 70 species and exhibits a wide range of nutlet variation. Currently, the evolutionary relationships among species of Lappula have not been examined; therefore, to elucidate phylogenetic relationships and morphological evolution within Lappula and related genera, we conducted phylogenetic analyses with matrices that include 48 species as well as four DNA regions (ITS, trnL- trnF, rpS16 and psbA- trnH) and 18 morphological characters. These matrices were analyzed using maximum parsimony, maximum likelihood, and Bayesian inference methods. Analyses of the combined molecular and morphological data result in a phylogeny that is better resolved than that based solely on molecular sequence data. Phylogenetic results suggest that the current infrageneric classification of Lappula, at least at the subsectional and series level, is artificial. The evolutionary patterns of 18 morphological characters are investigated in a phylogenetic context. In Lappula, nutlet homomorphism and small corollas are resolved as ancestral, while nutlet heteromorphism and larger corollas are derived. [ABSTRACT FROM AUTHOR]

Onosma is a species-rich genus with complicated patterns of morphological and karyological variation, and controversial taxonomic treatments. In the present study we focused on the Asterotricha group, one of three major infrageneric groups, in the area of central and southern Europe. Ribosomal internal transcribed spacer (ITS) sequences and amplified fragment length polymorphism (AFLP) markers were used to assess species differentiation and relationships, and to gain insight into the evolutionary history of the group. ITS data supported the monophyly of the asterotrichous taxa. Six genetically differentiated entities were resolved by AFLP markers, corresponding to O. echioides, O. heterophylla s.l., O. thracica including O. rigida, O. malkarmayorum, O. taurica, and O. stellulata. Onosma stellulata is identified as the most differentiated lineage within the Asterotricha group, which is consistent with its unique chromosome number. Within O. heterophylla s.l., recognition of O. viridis as a species distinct from O. heterophylla s.str. is supported. On the other hand, O. tornensis appears to be conspecific with O. viridis. Inferring from the patterns of genetic diversity and divergence, the evolutionary history of O. heterophylla s.l. is discussed. Genetic variation within the polymorphic Apennine-Dalmatian O. echioides is almost continuous and shaped by isolation by distance, thus none of the previous intraspecific taxonomic concepts are valid. The assumed hybrid origin of O. malkarmayorum is supported, and its affinity to potential ancestors, O. heterophylla s.l. and O. thracica incl. O. rigida, is indicated. Future studies of asterotrichous Onosma species should focus on diversity centers in the eastern Mediterranean and Asia Minor, to build up a comprehensive taxonomic concept of the group and to clarify its evolution. [ABSTRACT FROM AUTHOR]

Introns and spacers are a rich and well-appreciated information source for evolutionary studies in plants. Compared to coding sequences, the mutational dynamics of introns and spacers is very different, involving frequent microstructural changes in addition to substitutions of individual nucleotides. An understanding of the biology of sequence change is required for correct application of molecular characters in phylogenetic analyses, including homology assessment, alignment coding, and tree inference. The widely used term “indel” is very general, and different kinds of microstructural mutations, such as simple sequence repeats, short tandem repeats, homonucleotide repeats, inversions, inverted repeats, and deletions, need to be distinguished. Noncoding DNA has been indispensable for analyses at the species level because coding sequences usually do not offer sufficient variability. A variety of introns and spacers has been successfully applied for phylogeny inference at deeper levels (major lineages of angiosperms and land plants) in past years, and phylogenetic structure R in intron and spacer data sets usually outperforms that of coding-sequence data sets. In order to fully utilize their potential, the molecular evolution and applicability of the most important noncoding markers (the trnT– trnF region comprising two spacers and a group I intron; the trnS–G region comprising one spacer and a group II intron in trnG; the group II introns in petD, rpl16, rps16, and trnK; and the atpB– rbcL and psbA– trnG spacers) are reviewed. The study argues for the use of noncoding DNA in a spectrum of applications from deep-level phylogenetics to speciation studies and barcoding, and aims at outlining molecular evolutionary principles needed for effective analysis. [ABSTRACT FROM AUTHOR]

Awareness of the complex structure and evolutionary dynamics of noncoding DNA has improved both noncoding sequence alignment and the use of microstructural changes as characters in phylogenetic analysis. The next step is to consider improvements in the use and selection of phylogenetic models for noncoding sequence data. Models of character evolution are central to phylogeny estimation, but the use of an inadequate model can mislead topology selection and branch length estimations. This is particularly likely when sequence divergence is either limited (nearly invariable, as in population-level or species-level studies) or extreme (nearly saturated, as in deep-level studies that focus on conserved secondary structures). Noncoding data sets are often at these extremes, and they can be particularly awkward for model definition and model selection. This paper introduces the goals of model use in phylogenetics and identifies ten issues that arise from the application of models to noncoding sequence data. It is concluded that most of these issues derive from small data set sizes, very low or very high sequence variability, limitations of current phylogenetic models, and possibly character definition and nonindependence. Recommendations are made that should help to improve alignment, character quality, model selection, and phylogeny estimation based on noncoding sequence data. [ABSTRACT FROM AUTHOR]

Our study (survey, atlas of 136 microphotographs and 67 drawings) points out the actual chromosome numbers of 82 taxa of the genus Pinguicula L. They were gathered from literature and critically examined. In addition, numerous counts are published for the first time. They represent about 80% of all the taxa known. The basic chromosome numbers are x = 6, 8, 9, 11, and 14; the ploidy levels are 2 n (diploid), 4 n (tetraploid), 8 n (octoploid) and 16 n (hexadecaploid). The basic number x = 6 is a one-off, x = 8 and 11 are the most frequent in the genus; x = 14 indicates a hybridogenous differentiation process in the past. The caryological differentiation—chromosome numbers and ploidy level—is discussed with regard to distribution pattern, growth type, and infrageneric classification (at the level of sections). [ABSTRACT FROM AUTHOR]

This study investigates the genetic differentiation within the Central European Armeria maritima (Mill.) Willd. complex with special reference to the metallicolous populations using AFLP markers. Our sampling comprised all metallicolous (ssp. halleri, hornburgensis, bottendorfensis, eifeliaca, calaminaria), and non-metallicolous taxa (ssp. maritima, elongata, alpina). Geographical and genetic distances between populations were moderately positively correlated. Genetic variability of metallicolous and non-metallicolous populations was not significantly different. Lowland populations were clearly differentiated from the alpine populations. Within the lowland group metallicolous and non-metallicolous populations were not genetically differentiated. All lowland populations show a regional differentiation and close relationships to ssp. elongata. Thus, the metallicolous taxa should not be maintained as subspecies. Likewise, their treatment as varieties of a ssp. halleri s.l. is critical because this taxon cannot be consistently characterized throughout its geographical range and may be an artefact itself. If a taxonomical recognition should be considered necessary it is advisable to treat the microendemics as varieties of ssp. elongata. [ABSTRACT FROM AUTHOR]

We assessed nectar and honey potential of Phacelia tanacetifolia, an American plant, under Mediterranean conditions. Its flowering occurred during the major flowering season unless intensely irrigated, whereas duration and flower life span were shorter compared to continental conditions. Mediterranean climate limitations to nectar secretion were assessed on plants grown under natural conditions (xeric) vs. regular irrigation. May flowering xeric plants produced less nectar per flower than the irrigated ones, but had the same nectar potential per surface area. On the contrary, at the cost of intense irrigation, July flowering plants produced much higher nectar per flower and per surface area compared to xeric ones. In all flowering periods and sections honeybees were the most constant and numerous visitors, visiting the flowers mainly for nectar, whereas solitary bees were scarce. Based on our results, we suggest that although the plant may be a good nectar source for honeybees in some cases, we have serious reservations for a generalised use in the Mediterranean. [ABSTRACT FROM AUTHOR]

There is much taxonomic confusion in the genus Gentianella section Gentianella, five species of which are found in Britain. Gentianella anglica is a British endemic restricted in its distribution to the chalk downland of southern England. It is considered to be threatened because of habitat loss, and due to hybridisation with the closely related, widely distributed G. amarella. The Red Data Book species, G. uliginosa, shares morphological similarities with the other two species but has a very restricted distribution. Individual plants from populations across the range of these three species were analysed using AFLPs and universal cpSSRs to determine the degree of genetic variability within and between them, and to look for evidence of hybridisation. Samples of the two other Gentianella section Gentianella species found in Britain, G. campestris and G. germanica, were also included in the study. The analysis of chloroplast SSRs was not informative for G. anglica, G. amarella and G. uliginosa, while G. campestris and G. germanica differed from each other and the other three species at a single locus. Principal co-ordinate analysis of the AFLP data revealed only three distinct groups: one group contained G. campestris samples, another contained all samples of G. germanica, and a third contained all individuals from the other three species. In mixed populations of G. anglica and G. amarella, individuals of the two species were genetically more similar to each other than they were to individuals of the same species from other populations. It is proposed that G. anglica may be an early flowering form of G. amarella that has been created and maintained as a consequence of former grassland management practices. [ABSTRACT FROM AUTHOR]

A data matrix of 143 morphological and chemical characters for 142 genera of euasterids according to the APG system was compiled and complemented with rbcL and ndhF sequences for most of the genera. The data were subjected to parsimony analysis and support was assessed by bootstrapping. Strict consensus trees from analyses of morphology alone and morphology + rbcL + ndhF are presented. The morphological data recover several groups supported by molecular data but at the level of orders and above relationships are only superficially in agreement with molecular studies. The analyses provide support for monophyly of Gentianales, Aquifoliales, Apiales, Asterales, and Dipsacales. All data indicate that Adoxaceae are closely related to Dipsacales and hence they should be included in that order. The trees were used to assess some possible morphological synapomorphies for euasterids I and II and for the orders of the APG system. Euasterids I are generally characterised by opposite leaves, entire leaf margins, hypogynous flowers, “early sympetaly” with a ring-shaped corolla primordium, fusion of stamen filaments with the corolla tube, and capsular fruits. Euasterids II often have alternate leaves, serrate-dentate leaf margins, epigynous flowers, “late sympetaly” with distinct petal primordia, free stamen filaments, and indehiscent fruits. It is unclear which of these characters represent synapomorphies and symplesiomorphies for the two groups, respectively, and there are numerous expections to be interpreted as reversals and parallelisms. [ABSTRACT FROM AUTHOR]

Based on results from a cladistic analysis of molecular characters using parsimony and Bayesian inference, the tribe Selagineae (Scrophulariaceae, sometimes Selaginaceae or Globulariaceae), is found to have arisen within Manuleeae (Scrophulariaceae). The inclusion of Selagineae into Manuleeae is therefore proposed. The result is corroborated by morphological characters, and indicates that the typical Selagineae condition with one ovule per ovary locule has arisen several times within the Manuleeae. Two former Manuleeae taxa are excluded from Manuleeae; Antherothamnus and Camptoloma. The former shows affinity with Scrophularia and Verbascum, and the latter groups with Buddleja and related genera. The controversy between parsimony and likelihood based methods are briefly discussed as is the possibility of historical bias in prior studies. [ABSTRACT FROM AUTHOR]

Leaf samples of mostly herbarium specimens (237 species of 172 genera) were cleared. Internal secretory structures of large size or unusual shape were detected and observed. Selected samples were processed into resin and sectioned for light microscopy or prepared for scanning electron microscopy. Adding results from two earlier publications, our survey includes 365 species from 174 genera. Five types of internal secretory structure, mostly unknown previously in the traditional Scrophulariaceae, occur in only nine genera: 1) single-celled subepidermal idioblasts, empty at maturity, in Scrophularia and Verbascum (Lersten and Curtis 1997), 2) 2–16 (or more)-celled nodules, with fibrous contents, in mid-mesophyll strata of Graderia and Radamaea, 3) epithelium-lined oil cavities in Leucophyllum (Lersten and Beaman 1998) and Capraria, 4) Kranz-type enlarged bundle sheath in Anticharis, and 5) paraveinal mesophyll (PVM) in Picria and Bonnaya. [ABSTRACT FROM AUTHOR]

We studied the interrelation between nectary structure (13 parameters), nectar characteristics (yield, chemical composition), and flower size of 11 Labiatae species in a Mediterranean shrub community near Athens, Greece. We also explored whether the above attributes are affected by the Mediterranean summer drought constraints. Our findings show that among all nectary parameters studied, nectary size and stomatal opening are the most important in (positively) shaping nectar secretion, nectary size being the most meaningful. Nectary structure is correlated to quantity of the nectar secreted, not its quality. Wide flowers bear wide nectaries with large stomatal openings, whereas deep flowers are not related to any nectary size. Corolla size (both length and width) and nectary stomatal opening decrease with flowering time. This applies also to nectary size, nectar volume and sugar content of the perennials (9 species). All above cases of time dependence show that there is a constraint effect of Mediterranean climate on floral and nectary structure, reflected also as a decrease in nectar secretion. Nectary structure in Labiatae is largely shaped by both phylogenetic and climate constraints. On the other hand, although nectar is largely influenced by nectary structure, it is to a large extent ecologically biased, implying that, in addition to phylogeny, there are many other ecological parameters interfering in its secretion such as time within the season, life history, and light requirements. [ABSTRACT FROM AUTHOR]

Bees require pollen for their reproduction and pollen comprises the basic larval food for bees. Most bees acquire pollen passively during flower visitation, but female bees may also collect pollen actively with the aid of various structural and behavioral adaptations. Most bees have evolved adaptations to concentrate pollen into discrete loads and transport pollen back to their nests. The various structural and behavioral adaptations of female bees for acquiring and transporting pollen are the basis of this review. [ABSTRACT FROM AUTHOR]

The ever-larger data matrices resulting from continuing improvements in DNA sequencing techniques require faster and more efficient methods of phylogenetic analysis. Here we explore a promising new method, parsimony jackknifing, by analyzing a matrix comprising 2538 sequences of the chloroplast gene rbcL. The sequences included cover a broad taxonomic range, from cyanobacteria to flowering plants. Several parsimony jackknife analyses were performed, both with and without branch-swapping and multiple random addition sequences: 1) including all positions; 2) including only first and second codon positions; 3) including only third positions; and 4) using only transversions. The best resolution was obtained using all positions. Removal of third positions or transitions led to massive loss of resolution, although using only transversions somewhat improved basal resolution. While branch-swapping improved both resolution and the support found for several groups, most of the groups could be recovered by faster simple analyses. Designed to eliminate groups poorly supported by the data, parsimony jackknifing recognizes 1400 groups on the basis of all rbcL positions. These include major taxa such as green plants, land plants, flowering plants, monocots and eudicots. We include appendices of supported angiosperm families, as well as larger groups. [ABSTRACT FROM AUTHOR]

Sequences of rbcL and ndhF were analysed independently and in combination to resolve phylogenetic relationships in Labiatae s. l. Monophyly of Labiatae s. l was supported by all three analyses. Congea tomentosa ( Symphoremataceae) is nested within Labiatae s. l. in the rbcL analysis, but emerges as the sister group of Labiatae s. l. in the ndhF and combined analyses. Four noteworthy clades of Labiate s. l. also are supported by all analyses corresponding to subfamilies Nepetoideae, Lamioideae, Pogostemonoideae and Scutellarioideae. Monophyly of subfamilies Chloanthoideae and Viticoideae is not supported. A clade comprising Teucrioideae plus Ajuga is supported by ndhF and the combined analysis. [ABSTRACT FROM AUTHOR]

The familiar limits of Verbenaceae are controversial, ranging from a sensu lato arrangement of seven subfamilies until a sharply defined sensu stricto group consisting of only some genera from subfam. Verbenoideae. Iridoid contents of genera belonging to Verbenaceae s. l. were examined through a cladistic analysis, using outgroup analysis to evaluate character polarity within the ingroup. The cladogram obtained shows the distribution of the species according to their iridoid content. [ABSTRACT FROM AUTHOR]

Morphological and cytological investigations were carried out in Stachys beckeana Dörfler & Hayek and S. recta L. subsp. sarajevensis sensu Hayek. These two taxa have the same morphological characteristics (shape and indumentum of leaves; morphology of sepals, petals and fruits) and both have 2n = 2x = 34 chromosomes. They cannot be considered as different taxa; S. beckeana Dörfler & Hayek is the valid name (synonyms: S. recta L. subsp. sarajevensis sensu Hayek, S. hayekii Malý, S. recta L. subsp. hayekii Malý). [ABSTRACT FROM AUTHOR]

At least 560 species of flowering plants distributed in 64 genera and 16 families native to Southern Western Australia are pollinated by birds. This represents 15% of the total flora, and indicates adaptation to bird pollination is a major evolutionary force in this region. [ABSTRACT FROM AUTHOR]