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As part of an ongoing project for the investigation of the Algerian-Tunisian flora, six names of species of Caprifoliaceae subfam. Dipsacaceae, endemic to central and western North Africa, described in the 19th and 20th Centuries under the genus Scabiosa, are here lectotypified. They are: Scabiosa camelorum Cosson & Durieu, Scabiosa cartenniana Pons & Quézel, Scabiosa daucoides Desf., Scabiosa farinosa Cosson, Scabiosa parielii Maire, and Scabiosa robertii Barratte. Taxonomic notes, distributisson and new unpublished data about plant phenology of these taxa are given.

In this article, four species of the genus Hemaris (Lepidoptera, Sphingidae) are recognized from the Russian Far East: Hemaris ottonis (Rothschild and Jordan, 1903), Hemaris affinis (Bremer, 1861), Hemaris fuciformis (Linnaeus, 1758), and Hemaris radians (Walker, 1856). An identification key, redescriptions, distributions, and overviews of the ecology of Hemaris species in the Russian Far East are provided. The larval host plant of H. ottonis is recognized for the first time as Weigela praecox (Caprifoliaceae). The morphology and biology of preimaginal stages of H. ottonis have been studied for the first time. Larvae of this species are very different from other Hemaris species. The life cycle of H. radians is given for the first time, with its larval host plant in the southern part of Primorsky Krai determined to be Scabiosa lachnophylla (Dipsacaceae). Keywords: Ecology, Hemaris, Preimaginal stages, Russian Far East, Sphingidae

Wetlands are among the most fragile habitats on Earth and have often undergone major environmental changes. As a study case in this context, the present work aims at increasing the floristic knowledge of a reclaimed land now turned into an agricultural lowland with scarce patches of natural habitats. The study area is named Piana di Rosia, and it is located in southern Tuscany (Italy). The compiled checklist consists of 451 specific and subspecific taxa of vascular plants. The life-form spectrum shows a predominance of hemicryptophytes, followed by therophytes. The chorological spectrum highlights a co-dominance of Euri-Mediterranean and Eurasian species along with many widely distributed species. The checklist includes seven species of conservation concern, three Italian endemics (Crocus etruscus Parl., Polygala vulgaris L. subsp. valdarnensis (Fiori) Arrigoni, and Scabiosa uniseta Savi), 41 alien species, 21 segetal species, and 11 aquatic macrophytes of which five helophytes and six hydrophytes. This study suggests that irreversible land-use changes in wetlands can lead towards a simplification of the flora. However, despite the deep transformations that the former wetland has undergone, the presence of some aquatic and protected taxa is interesting. From a conservation point of view, the natural value of this agricultural area could be enhanced and its current management partly reconsidered, thus preserving the remnants of naturalness present.

In this study, the chemotaxonomic status and fatty acid compositions of eleven species of Scabiosa L., naturally growing in Iran, were determined using gas chromatography. The main compounds were found to be palmitic acid (16:0; 4.63–23%), behenic acid (22:0; 2.40–35.52%), lignoceric acid (24:0; 1.91–34.02%) and linoleic acid (18:3n6; 0.73–13.95%). Principal Component and Principal Coordinate analyses revealed the segregation of the studied Scabiosa species in two groups. This was consistent with the traditional classification presented by Rechinger. However, there is a lot of debate about the taxonomy of the Scabiosa species. Based on pitted epicalyx, Greuter and Raus have removed members of the Scabiosa sect. Asterocephalus to the genus Lomelosia. Our results showed that the fatty acid profiles of the species belonging to the sect. Scabiosa were distinct and confirmed the opinion supported by Greuter and Raus, that these species should be moved to the Lomelosia genus.

Pollen morphology of nine species representing four genera: Cephalaria Schrad, Dipsacus L., Pterocephalus Vaill. and Scabiosa L. of the family Dipsacaceae in Iran has been investigated by means of scanning electron microscopy (SEM). The results showed that pollen grains were triporate and tricolpate. The pollen type of Scabiosa rotata Bieb. (tri- and tetraporate) is the first report in the world. The sizes of pollen grains fall into the classification group magna (pollen grain diameter 50–100 μm). Pollen shapes vary from preoblate to prolate and their polar views were triangulate and lobate. The exine ornamentation varies from gemmate in S. rotata to spinulate in the rest studied species. Species of Scabiosa have been dispersed in UPGMA tree that this confirmed the previous studies about taxonomic problems and species complexity in this genus. These results show the transfer of the some Scabisoa species to Lomelosia Raf. based on palynological characters. Pollen morphology of the family is helpful at the generic and specific level. Bangladesh J. Plant Taxon. 24 (2): 129–136, 2017 (December)

The pollen morphology of six species of Scabiosa L. (Caprifoliaceae) from Iran has been examined by scanning electron microscopy (SEM). Pollen grains were tricolpate in S. columbaria and triporate in the rest studied species. Two types of exine ornamentation were revealed: spinulate and gemmate. The pollen shape in polar view varied from triangular to circular among investigated taxa. Statistical analysis showed that some quantitative morphological features such as polar axis (P), equatorial axis (E) and aperture diameter were main characters in identification of the taxa studied. These taxa were separated from each other using cluster analysis and placed within two clusters. Our result based on UPGMA analysis is in agreement with morphological classification and recent findings on taxonomic position of the Scabiosa.Bangladesh J. Plant Taxon. 23(2): 215-222, 2016 (December)

Non-rewarding orchids rely on various ruses to attract their pollinators. One of the most common is for them to resemble flowers sought by insects as food sources. This can range from generalized food deception to the mimicry of specific sympatric food plants. We investigated the basis of pollinator deception in the European food-deceptive orchid Traunsteinera globosa, which has unusually compact flowerheads resembling those of sympatric rewarding species of Knautia and Scabiosa (Dipsacaceae), and Valeriana (Caprifoliaceae). Visual signals of T. globosa are similar in both fly and bee vision models to those of the sympatric food plants used in the choice experiments, but scent signals are divergent. Field experiments conducted in Austria and the Czech Republic showed that both naive and experienced (with respect to visitation of T. globosa) insect species approached the orchids at the same rate as food plants, but direct contact with orchid flowers was taxon specific. Flies were most easily duped into probing the orchid, and, in doing so, frequently received and deposited pollinaria, whereas most bees and butterflies avoided landing on orchid flowers. We conclude that T. globosa is a mimic of a guild of fly-pollinated plants, but the ecological dependence of the orchid on its models remains to be fully tested. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180, 269–294.

We developed ten polymorphic microsatellite markers for the herbaceous species Scabiosa canescens Waldst. & Kit. The species is currently not assessed under the IUCN Red List but has been declining historically due to intensification in agriculture and is recently declining due to habitat loss caused by agricultural abandonment of steppe-like dry grasslands. The microsatellite loci were tested for polymorphism on DNA templates extracted from 26 specimens collected across the species’ range in Germany. Across the ten loci, two to eight alleles were detected with an observed and expected heterozygosity ranging from 0.192 to 0.769 and 0.265 to 0.768, respectively. These newly developed markers provide a valuable tool for population genetic investigations required to design effective conservation measures.

We carried out a quantitative assessment of the consumption of herbaceous plants by Opatrum sabulosum (Linnaeus, 1761) – a highly significant agricultural pest species. We researched the feeding preferences of this pest species with respect to 33 uncultivated and 22 cultivated plant species. This species of darkling beetle feeds on many uncultivated plant species, including those with hairy leaves and bitter milky sap, such as Scabiosa ucrainca (5.21 mg/specimen/24 hours), Euphorbia virgata (3.45), Solanum nigrum (3.32), Centauria scabiosa (2.47), Lamium album (2.41), Aristolochia clematitis (1.76), Chenopodium album (1.73), Arctium lappa (1.51), Asperula odorata (1.20). A high rate of leaf consumption is also characteristic for cultivated species, for example, Perilla nankinensis (5.05 mg/specimen/24 hours), Lycopersicon esculentum (3.75), Tropaeolum majus (3.29), Nicotiana tabacum (2.66), Rumex acetosa (1.96), Beta vulgaris (1.27). Opatrum sabulosum is capable of feeding on plants which are poisonous to cattle. This species of darkling beetle consumes 95.5% of the cultivated and 48.5% of the uncultivated herbaceous plants researched.

Tragopogon comprises approximately 150 described species distributed throughout Eurasia from Ireland and the UK to India and China with a few species in North Africa. Most of the species diversity is found in Eastern Europe to Western Asia. Previous phylogenetic analyses identified several major clades, generally corresponding to recognized taxonomic sections, although relationships both among these clades and among species within clades remain largely unresolved. These patterns are consistent with rapid diversification following the origin of Tragopogon, and this study addresses the timing and rate of diversification in Tragopogon. Using BEAST to simultaneously estimate a phylogeny and divergence times, we estimate the age of a major split and subsequent rapid divergence within Tragopogon to be ~2.6 Ma (and 1.7-5.4 Ma using various clock estimates). Based on the age estimates obtained with BEAST (HPD 1.7-5.4 Ma) for the origin of crown group Tragopogon and 200 estimated species (to accommodate a large number of cryptic species), the diversification rate of Tragopogon is approximately 0.84-2.71 species/Myr for the crown group, assuming low levels of extinction. This estimate is comparable in rate to a rapid Eurasian radiation in Dianthus (0.66-3.89 species/Myr), which occurs in the same or similar habitats. Using available data, we show that subclades of various plant taxa that occur in the same semi-arid habitats of Eurasia also represent rapid radiations occurring during roughly the same window of time (1.7-5.4 Ma), suggesting similar causal events. However, not all species-rich plant genera from the same habitats diverged at the same time, or at the same tempo. Radiations of several other clades in this same habitat (e.g. Campanula, Knautia, Scabiosa) occurred at earlier dates (45-4.28 Ma). Existing phylogenetic data and diversification estimates therefore indicate that, although some elements of these semi-arid communities radiated during the Plio-Pleistocene period, other clades sharing the same habitat appear to have diversified earlier.

Aim To reconstruct the temporal and biogeographical history of Old World disjunctions in Scabiosa (Dipsacaceae) and the timing of diversification in the Mediterranean Basin, in order to evaluate the importance of biogeographical and climatological history (particularly the onset of a mediterranean climate) in shaping Scabiosa distributions. Location Europe and the Mediterranean Basin, southern Africa and eastern Asia. Methods This study uses maximum-likelihood and Bayesian phylogenetic analyses of chloroplast DNA (atpB–rbcL, trnL–trnF, trnS–trnG, psbA–trnH) and nuclear ribosomal DNA [internal transcribed spacer (ITS) and external transcribed spacer (ETS)] from 24 out of c. 37 ingroup taxa, beast molecular dating, and the dispersal–extinction–cladogenesis method (Lagrange) to reconstruct ancestral geographical ranges and the timing of diversification of the major clades of Scabiosa. Results Biogeographical and divergence time reconstructions showed that Scabiosa originated during the Miocene and diversified in Europe, followed by independent movements into Asia and Africa. Several of the major clades were inferred to have radiated sometime between the late Miocene and early Pleistocene, a timeframe that encompasses the onset of the mediterranean climate in Europe. More recent middle–late Pleistocene radiations in the Mediterranean Basin and southern Africa have played a large role in Scabiosa diversification. Main conclusions Members of Scabiosa appear to have capitalized on adaptations to montane and/or dry conditions in order to colonize similar habitats in different biogeographical regions. The formation of the East African Rift mountains is potentially of great importance in explaining the southward migration of Scabiosa. The initial diversification of Scabiosa in Europe during the Miocene is not consistent with the initiation of the mediterranean climate, but may instead be associated with increased aridity and the retreat of subtropical lineages during this time. However, the radiation of some of the major subclades within Scabiosa may have been associated with an emerging mediterranean climate. More recent and rapid radiations in both the Mediterranean Basin and southern Africa highlight the probable importance of Pleistocene climate fluctuations in Scabiosa diversification.

Lectotypes are designated for names published by Josef Freyn from the Iberian Peninsula in Bulletin de l’Herbier Boissier in 1893: Genista anglica var. pilosa, Leontodon reverchonii, Linaria supina var. glaberima, Scabiosa tomentosa var. cinerea, Trifolium hervieri, Valerianella willkommii and Vicia lusitanica. Relectotypification is done for Thymus portae. Names Arabis reverchonii Freyn and Astragalus aragonensis Freyn from the same publication are mentioned as well.Se designan lectótipos para los nombres publicados por Josef Freyn para la Península Ibérica en la revista Bulletin de l’Herbier Boissier en 1893: Genista anglica var. pilosa, Leontodon reverchonii, Linaria supina var. glaberima, Scabiosa tomentosa var. cinerea, Trifolium hervieri, Valerianella willkommii y Vicia lusitanica. Se vuelve a lectotipificar Thymus portae. Se comentan además los nombres Arabis reverchonii Freyn y Astragalus aragonensis Freyn, publicados en la misma revista.

Pollen morphology of nine species belonging to four genera: Cephalaria, Pterocephalus, Scabiosa, and Lomelosia of Dipsacaceae in Egypt has been investigated by light microscope (LM) and scanning electron microscopy (SEM). Pollen shape, size, exine ornamentation, number and characters of apertures present powerful characters for distinguishing between species. The pollen grains were tricolpate and triporate. Their shapes vary from spheroidal to oblate-spheroidal. Regarding the position of apertures, three types can be recognized viz., circumaperturate, angulaperturate, and planaperturate. Concerning sculpturing of the exine in proximal face, two different pollen types can be distinguished viz., spinulate and gemmate types. The exine ornamentation was found useful to distinguish between closely related genera such as Scabiosa and Lomelosia. A diagnostic key is given for all studied taxa based on palynomorphological characters (© 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Lomelosia caucasica, a common and widespread species native in the Caucasus area, was first collected on the SW margin of its distribution area and illustrated as early as 1702. Subsequently, Tournefort published a polynomial for it that was neither taken up by Linnaeus nor by later taxonomists. Even before a Linnaean name was published by Marschall von Bieberstein, this montane to subalpine plant was cultivated in the Loddiges nursery in Hackney in 1808 and in a few other botanical gardens. The details of the first findings of L. caucasica in the wild and its rapid spread in cultivation are elucidated and the pertinent herbarium specimens examined. In addition, the reasons for having proposed a conserved type are presented. A complete list of synonyms with references to protologues are given as well as notes on the distribution and ecology of L. caucasica. Citation: Lack H. W. 2018: The discovery and naming of Lomelosia caucasica (Dipsacaceae) with notes on its nomenclature and its early cultiv...

In the general characteristics of the family Valerianaceae the occurrence of “stellate hairs” described by the authors of Valeriana tomentosa Kunth and V. malvaceae Graebn. still remains unconsidered. Beyond their diagnostic value, however, it is interesting, that at least the specific ontogeny and anatomical structure of the “stellate hairs” of V. tomentosa corresponds very well with similar trichomes known for several other taxa of Dipsacales, namely species of Viburnum (Viburnaceae) and some Dipsacaceae (Cephalaria, Scabiosa). (© 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Mimicry, as an adaptive explanation for the resemblance between organisms, is not always readily distinguishable from, inter alia, coincidence, shared ancestry, or convergent evolution. We tested the hypothesis that two rare South African orchid taxa Brownleea galpinii ssp. major (nectar-producing) and Disa cephalotes ssp. cephalotes (non-rewarding) are mimics of the nectar-producing flowers of a relatively common species, Scabiosa columbaria (Dipsacaceae), with which they always occur sympatrically. Flowers of the orchids were apparently unscented and had similar dimensions and almost identical spectral reflectance to the flowers of Scabiosa. The orchids were pollinated exclusively by long-proboscid flies (Tabanidae and Nemestrinidae) that feed mainly on nectar in Scabiosa flowers. Choice experiments showed that these flies did not discriminate between the orchids and Scabiosa when alighting on their flat-topped inflorescences. However, flies were not attracted to related orchids dissimilar to Scabiosa, or to inflorescences of B. galpinii that had been artificially reconstructed in the shape of a spike, rather than a flat-topped capitulum. A phylogenetic analysis showed traits that give the orchids a resemblance to Scabiosa, such as a flat-topped inflorescence and cream floral colouration with dark spots and short spurs, to be mostly apomorphic features, and therefore likely to be relatively recent adaptations for mimicry. We caution that the term mimic should not be applied to species whose resemblance to another species is due entirely to plesiomorphic traits that, in all likelihood, evolved prior to the ecological association. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 289–304.

Floral nectary characteristics of five species of Centaurea and Inula (Inula ensifolia, I. salicina, I. spiraeifolia, Centaurea micranthos, C. scabiosa) were examined for using light microscopy and scanning electron microscopy, and the sugar contents were studied in relation to nectary morphology and plant size. The differences in the nectaries, plant size and nectar sugar content of the studied species indicate that the larger plants may reflect a greater allocation to sugar content.

Naive male and female silver Y moths, Autographa gamma (Lepidoptera: Noctuidae), were attracted in a flight tunnel assay to potted creeping thistle, Cirsium arvense (Asteraceae), butterfly-orchid, Platanthera bifolia (Orchidaceae), soapwort, Saponaria officinalis (Caryophyllaceae), greater knapweed, Centaurea scabiosa (Asteraceae), red clover, Trifolium pratense (Fabaceae), and catnip, Nepeta faasseni (Labiatae), plants with flowers. The most attractive plants were C. arvense, P. bifolia and S. officinalis that elicited 87, 78 and 65% source contacts, respectively. C. scabiosa was less attractive eliciting 43% response. T. pratense and N. faasseni showed the least attraction eliciting 28 and 26% source contacts, respectively. A cotton plant used as control, was not attractive. Floral volatiles from the investigated plant species were collected using headspace sampling technique. Samples were analysed using gas chromatography coupled with electroantennographic detection, and electrophysiologically active compounds were identified by coupled gas chromatography/mass spectrometry. Consistent electrophysiological responses were elicited by twelve compounds from headspace of C. arvense, thirteen compounds from P. bifolia, eleven compounds from S. officinalis, nine from C. scabiosa, ten from T. pratense and two from N. faasseni. Most of the active compounds were specific for one or two species, while benzyl benzoate was present in four and benzaldehyde and benzyl alcohol in three species. Floral scents of C. arvense, P. bifolia and S. officinalis, the most attractive flowers, were dominated by aromatic compounds that were not abundant in the scent of other flowers. To conclude, the results demonstrate the absence of a common denominator of odours present in flowers of different plants visited by A. gamma.

Developmental instability and fluctuating asymmetry (FA) have become important topics in evolutionary biology. For example, it has been suggested that FA could be a useful tool for identification of genetic and environmental stress factors. This study used plants from each of six populations of Scabiosa canescens and Scabiosa columbaria grown under greenhouse conditions. I tested whether there was a relationship between petal FA and allozyme heterozygosity, the heritabilities of eight traits, and population size. Flowers displayed no directional asymmetry or antisymmetry. The rare species S. canescens had significantly higher FA values than S. columbaria, but only the latter demonstrated interpopulation differentiation for the expression of FA levels. There was no evidence for an association between population-level FA and genetic variation when compared with the allozyme heterozygosity or with the heritabilities of the quantitative traits. A tendency for a negative association between FA and population size was found for both species, but it was not significant when adjusted for multiple comparison. Hence, flower FA should not be considered a reliable indicator of the amount of genetic variation in populations of S. canescens and S. columbaria.

Four species of Scabiosa L. are transferred to Lomelosia Raf. (Dipsacaceae) to complete the relocation of all scabious taxa with a pitted epicalyx into Lomelosia. The new combinations are: L. deserticola (Rech. f.) P. Caputo & Del Guacchio, L. poecilocarpa (Rech. f.) P. Caputo & Del Guacchio, L. schimperiana (Boiss. & Buhse) P. Caputo & Del Guacchio, and L. transcaspica (Rech. f.) P. Caputo & Del Guacchio.

In the current study, we used bootstrap analyses and the common principal component (CPC) method of Flury (1988) to estimate and compare the G-matrix of Scabiosa columbaria and S. canescens populations. We found three major patterns in the G-matrices: (i) the magnitude of the (co)variances was more variable among characters than among populations, (ii) different populations showed high (co)variance for different characters, and (iii) there was a tendency for S. canescens to have higher genetic (co)variances than S. columbaria. The hypothesis of equal G-matrices was rejected in all comparisons and there was no evidence that the matrices differed by a proportional constant in any of the analyses. The two 'species matrices' were found to be unrelated, both for raw data and data standardized over populations, and there was significant between-population variation in the G-matrix in both species. Populations of S. canescens showed conservation of structure (principal components) in their G-matrices, contrasting with the lack of common structure among the S. columbaria matrices. Given these observations and the results from previous studies, we propose that selection may be responsible for some of the variation between the G-matrices, at least in S. columbaria and at the between-species level. (Less)

A comparative chemometric study of three taxa of the genus Scabiosa L. s. lato (Dipsacaceae) from Greece, treated so far as separate genera by some authors and as separate sections of a single genus by others, was carried out. The HPLC chemical fingerprint, based on several phenolic compounds (flavonoids and caffeic acid), was investigated in the three a forementioned taxa. The results support the contention that there is a close relationship between S. atropurpurea subsp. maritima and S. tenuis, and that S. argentea seems to be well separated from the other two taxa.

Fruits ofDipsacaceae are single-seeded, have bristle-shaped calyx segments and are tightly enclosed by four fused bracts forming an epicalyx. Comparative morphological and anatomical studies reveal a great diversity of epicalyx and calyx, often relevant to fruit dispersal. The present contribution deals with theScabiosa group of genera, the core of theScabioseae tribe. Most of its taxa develop a diaphragma from a meristem on the inside of the epicalyx. This diaphragma, together with the lower part of the epicalyx encloses the fruit proper, whereas the upper parts form a so-called “epi-diaphragma” (ed) and a ± hyaline corona. Differences of the epicalyx with respect to the size and position of the ed, elaboration of the corona, origin of pits (=foveoles) and other morphological and anatomical specializations can be demonstrated. Together with palynological and karyological data these new facts support an improved concept of relationships and systematics for the taxa studied:Scabiosa sect.Scabiosa and sect.Cyrtostemma are closely related and should be united to form the genusScabiosa s. str.;Pycnocomon can be maintained as an independent genus, sister toScabiosa sect.Trochocephalus which then has to be treated as a genus,Lomelosia. In contrast, the following genera have to be included inLomelosia:Tremastelma asLomelosia sect.Callistemma, andScabiosiopsis as part ofLomelosia sect.Lomelosia. Pseudoscabiosa deviates in so many features that it has to be excluded from the redefinedScabioseae s. str.

Effects of elevated CO2 on flowering phenology and nectar production were studied in five important nectar plants of calcareous grasslands, i.e. Lotus corniculatus, Trifolium pratense, Betonica officinalis, Scabiosa columbaria and Centaurea jacea. Glasshouse experiments showed that flowering probability was significantly enhanced in C. jacea. B. officinalis flowered carlier and L. corniculatus produced more flowers under elevated CO2. In contrast, the number of flowers decreased in T. pratense. The amount of nectar produced per flower was not affected in the investigated legumes (T. pratense, L. corniculatus), but was significantly reduced in the other forbs. Elevated CO2 did not significantly affect nectar sugar concentration and sugar composition. However, S. columbaria and C. jacea produced significantly less total sugar per flower under elevated CO2. The nectar amino acid concentration remained unaffected in all investigated plant species whereas the total of amino acids produced per flower was significantly reduced in all non-legumes. In addition, the amino acid composition changed significantly in all investigated species except for C. jacea. The observed effects are unexpected and are a potential threat to flower visitors such as most butterflies which have no alternative food resources except nectar. Changes in nectar production due to elevated CO2 could also generally have detrimental effects on the interactions of flowers and their pollinators.

Hispano-Lusitanian plants traditionally identified as Scabiosa semipapposa Salzm. ex DC. are recognized with a new name, Scabiosa galianoi Devesa, Ortega-Olivencia & J. LOpez, after it was found that they differ clearlyfrom the Moroccan plants on which De Candolle's description was based.

The microdistribution of five butterfly species through their flying season was analyzed in a mosaic-like habitat, brought about by secondary succession. In order to explain the patterns observed, activity patterns and the use and distribution of nectar sources were determined. Emphasis was laid on the changing allocation of visits to flower species and changing abundances of flowers during the season. The use of nectar sources was basically limited to three flower species, Centaureaa scabiosa, C. bracteata and Serratula tinctoria. As a consequence, niche breadth values were generally low and niche overlaps generally high. Some butterflies changed their patterns of flower visits during the season and therefore reduced niche overlap with the other butterfly species. The microdistribution of Melanargia galathea, Lysandra coridon, don, Ochlodes venatus and Lictoria achilleae was strongly influenced by the distribution of their preferred nectar sources as well as by areas generally rich in flowers. Changing flower preferences of Melanargia galathea and Lysandra coridon males during the course of the season were also expressed by changes in the correlations between the distribution of these butterflies and their nectar plants. The distribution of nectar sources was not found to be of importance for Coenonympha arcania, a species which rarely visited flowers

A phytochemical and morphological comparative study of two taxa of the Scabiosa L. genus has been carried out. The chemical investigation concerns the isolation and identification of six flavonoids from Sc. tenuis and seven flavonoids from Sc. argentea. The morphological study deals with ten traits of both species (on a total of 213 specimens). The relation between the chemical pattern and phylogenesis is discussed. The results of morphometric study are connected with their geographical distribution; Figures of the latter are also given. Correlation of phytochemical and morphological results is attempted. Es wurde eine vergleichende phytochemische und morphologische Studie von zwei Arten der Gattung Scabiosa L. durchgefuhrt. Die chemische Untersuchung umfaste Isolierung und Bestimmung von sechs Flavonoiden von Sc. tenuis und sieben Flavonoiden von Sc. argentea. Die morphologische Studie umfaste zehn Eigenschaften beider Arten (von insgesamt 231 Proben). Das Verhaltnis zwischen chemischem Typ und Phylogenese wird diskutiert. Die Ergebnisse der morphometrischen Studie hangen mit ihrer geographischen Verbreitung zusammen. Die geographische Verbreitung wird in zwei Abbildungen dokumentiert. Es wird versucht, die phytochemischen und morphologischen Resultate miteinander zu vergleichen.

Zymograms of glucosephosphate isomerase (GPI) and phosphogluconate dehydrogenase (PGD) revealed three isozymes for each enzyme in the plant species Scabiosa columbaria. Intergenic heterodimers are formed between the polypeptides coded by Gpi-1 and Gpi-2 and between those coded by Pgd-1 and Pgd-2, indicating that a GPI and a PGD locus have been duplicated in the past. The ancestral genes assort independently with their duplicated gene, suggesting that the duplications have originated from a process of translocation. Linkage was found only between Gpi-1 and Pgd-2 and between Gpi-2 and Pgd-1, suggesting that the duplicated loci were located on the same translocated chromosomal segment. Both duplications are present in all other examined species of Scabiosa and in Cephalaria and Knautia, two other genera of the Dipsacaceae. The genera Succisa and Dipsacus, also belonging to the Dipsacaceae, do not show Gpi-1 activity, making Gpi-2 and Pgd-1 the most likely ancestral genes. In Succisa, the isozymes of Gpi-1 and Gpi-2 either overlap or Gpi-1 has been silenced. The combined results suggest that a chromosomal segment containing Gpi-2 and Pgd-1 has been translocated before the divergence of Scabiosa, Cephalaria, Knautia, and Succisa.

We provide information on the typification of ten species names described by Roberto de Visiani and Josif Pančić in Plantae Serbicae Rariores aut Novae—Decas II, published in 1865, and one published by Pančić in Flora of the Principality of Serbia (1874). Ten lectotypes are designated here. The name Scabiosa achaeta Visiani & Pančić is synonymised with Scabiosa fumarioides Visiani & Pančić. Differential characters distinguishing Allium serbicum Visiani & Pančić and the similar A. pallens L. are given.

Phylogenetic relationships among 17 taxa of Dipsacaceae were inferred from nucleotide sequence variation in both the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA and the chloroplast trnL (UAA) intron sequences. The combined phylogenetic analysis, carried out by using two taxa from Valerianaceae as an outgroup yielded a single most parsimonious tree, in which Dipsacaceae are divided into two major clades: one including Lomelosia and Pycnocomon, both in a sister group relationship with a clade containing Pterocephalus, Scabiosa and Sixalix; the other including Pseudoscabiosa, Succisa and Succisella is sister group to Knautia, Pterocephalidium, Dipsacus and Cephalaria. The results obtained here greatly differ from previous ones based on classical morphology, but are congruent with recent findings on epicalyx differentiation and with pollen characters. In particular, our results would confirm on molecular grounds the recently restricted circumscription for Scabioseae proposed by other authors. Our phylogenetic hypothesis indicates that adaptations to seed dispersal have been a very strong driving force in Dipsacaceae evolution, with similar selective pressures causing the onset of similar epicalyx shapes and dispersal modes in a parallel fashion in various taxa. For this reason, the gross morphology of the involucel is deceptive in inferring relationships.

The amount of genetic variation within a population is, among other things, related to population size. In small populations loss of genetic variation due to high levels of genetic drift and inbreeding may result in decline of individual fitness and increase the chance of population extinction. This chain of processes is known as genetic erosion. In this study we tested the genetic erosion hypothesis by investigating the relation between morphological variation and population size in two perennial, outbreeding plant species, Salvia pratensis and Scabiosa columbaria. To relate phenotypic variation to genetic variation the experiments were performed under common environmental conditions. For both species a positive correlation was observed between the amount of phenotypic variation and population size (Salvia r=0.915; Scabiosa r=0.703). Part of this variation is likely to have a genetic base, although maternal effects were present in the seedling and juvenile life stages. Differences between populations could in both species be attributed to parameters related to fitness, i.e. growth rate in Salvia and reproductive effort in Scabiosa. Discriminant functions reflecting these parameters did not however discriminate between large and small populations.Results are discussed in relation to the common environment approach and to electrophoretic results obtained earlier (Van Treuren et al. 1991).

SUMMARY (1) Seeds of thirteen herbaceous species from chalk grassland in The Netherlands were buried in their natural habitat. Samples were exhumed at regular intervals over 3 years. Mortality and germination in situ were recorded. The surviving seeds were tested for germination in light and darkness and with different temperature regimes. (2) Seed mortality was low, except in Gentianella germanica and Origanum vulgare. In Carlina vulgaris, Leontodon hispidus and Scabiosa columbaria disappearance of seeds was mainly caused by germination in the soil. (3) Freshly harvested seeds of Arenaria serpyllifolia and Linum catharticum had an absolute light requirement for germination which was retained during burial. Burial induced a light requirement for germination in Origanum vulgare. (4) All species that germinated showed some degree of primary dormancy and seasonal change in secondary dormancy. This was most pronounced in shortlived species. (5) The seasonal change in secondary dormancy of surviving seeds was most evident either from germinability in light (Arenaria, Daucus carota and Linum), in darkness ( Carlina, Leontodon, Plantago lanceolata and Scabiosa), or both (Centaurea scabiosa). The response of the seeds to temperature also varied seasonally. (6) The amplitude of seasonal change in dormancy tended to decrease during the 3-year of burial in some species (Arenaria, Linum and Daucus). (7) Rhinanthus minor and R. alectorolophus seeds only germinated in late winter and were dormant during the remainder of the year.

Biosystematics of some species of Centaurea, sect. Acrocentron Cass.—Morphometrical investigations on herbarium specimens, and serological as electrophoretic studies on seed proteins, were carried out on Centaurea rupestris, C. scabiosa, C. dichroantha and C. ceratophylla (in this case only morphometrical and serological studies were possible). The morphometrical and electrophoretic results make us think that C. rupestris, C. dichroantha and C. scabiosa are species taxonomically almost equidistant. C. ceratophylla is distinct but very similar to C. rupestris, making it correct to consider the two taxa as subspecies. Serologically the four species appear very homogeneous: it seems appropriate to include them in a single section. C. dichroantha shows some peculiar morphological and biochemical characters, non-intermediate between C. rupestris and C. scabiosa, therefore we would exclude a recent genesis through its hybrid derivation from the crossing of these species. On the other hand, if C. dichro...

The purpose of this study is to analyse, by means of Scanning Electron Microscopy (SEM), the validity of the typification of the pollen of Centaurea L. s.s., carried out by Wagenitz (1955) with light microscopy and based on the exine structure and sculpturing. The pollen of six species have been analyzed: one species for every type of pollen present in Italy: C. sempervirens L., C. alpina L., C. scabiosa L., C. alba L., C. montana L., C. cyanus L. The validity of the pollen typification suggested by Wagenitz has been confirmed also at the ultramicroscopic level. The ultrastructure and the sculturing of the sporoderm are described in detail and some discrepancies, mainly due to the different potentialities offered by the two methods, are pointed out.

Chromosome numbers for 28 Iranian populations ofDipsacaceae, corresponding to 14 taxa, are presented. New are those forScabiosa aff.olivieri var.pinnatisecta, S. persica, Pterocephalus canus, P. kurdicus, Dipsacus strigosus, Cephalaria dichaetophora, C. hirsuta, C. microcephala, C. subindivisa andC. aff.sublanata. Except forC. syriaca (x = 5) andC. dichaetophora (x = 7 : new basic chromosome number forCephalaria), all species examined are characterized by the basic chromosome number (x = 9). Instances of aneusomaty, B-chromosomes, aneuploidy, dysploidy, and polyploidy have also been found.

Critical notes on the flora of Italy. VI. Final miscellaneous remarks. – The present paper includes a number of changes of rank and nomen-clatural transfers in the genera Jovibarba, Saxifraga, Buglossoides, Stachys, Scabiosa, Helichrysum, Anthemis, Leucanthemopsis, Centaurea, Avenula some of them proposed by other Authors (B. Anzalone, W. Greuter, P. Marchi). Achillea lucana Pign. from the Southern Apennines near Potenza is described as a new species.

The chromosome numbers of several Mediterranean and Central European species from the genusCentaurea L. have been studied. The material was collected in natural habitats, exceptC. thracica andC. maxima. The numbers forC. cineraria (2n = 36; tetraploid!),C. subtilis (2n = 22),C. sonchifolia (2n = 44),C. sphaerocephala (2n = 44),C. napifolia (2n = 22),C. achaia (2n = 22),C. thracica (2n = 18), andC. maxima (2n = 28) are reported for the first time. For the following species earlier counts are confirmed:C. cineraria (diploid, 2n = 18),C. maculosa (2n = 18),C. rhenana (2n = 18),C. triumfettii (2n = 22 + 0−1 B; first report of accessory chromosome for this species),C. scabiosa (2n = 20 + 0−3 B),C. crassifolia (2n = 30). B-chromosomes are reported for 3 populations ofCentaurea scabiosa from Berlin; they lack in 2 populations from Bayern and Hessen.

Trematocarpus Kutzing in southern Africa is found to comprise four taxa in three species: newly described is T. fragilis (Ag.) De Toni var. divaricatus Simons, var. nov. Two of the species, T. flabellatus (J. Ag.) De Toni and T. affinis (J. Ag.) De Toni are restored to independent specific rank after being regarded as synonyms of T. scutellatus (Her.) Searles. It is shown that Sphaerococcus (Chondrus) scutellatus Her., the basionym of T. scutellatus refers to a species of Gigartina, therefore a new combination Gigartina scutellata (Her.) Simons is made for this species which was formerly known as G. fastigiata J. Ag. and G. scabiosa (Kiitz.) Papenf. Trematocarpus elongatus Kiitz. is relegated to synonymy under T. fragilis var. divaricatus .

Ausgehend von der Forderung nach einer umfassenden, neben Morphologie und Anatomie auch Wuchsrhythmik und Standortbindung berucksichtigenden Wuchsformenanalyse werden einige Wuchsformenreihen mediterran-mitteleuropaischer Angiospermen-Taxa behandelt. Bei Carlina, Scabiosa und Digitalis werden die Beziehungen von kanarischen oder sudmediterranen Kandelaberstrauchern zu submediterran-mitteleuropaischen Pleiokormstauden einerseits und zu Annuellen andererseits dargestellt. Besondere Beachtung findet dabei die Differenzierung des Pleiokorm (Scabiosa, Digitalis) und der Radikation (Carlina). Die verschiedene Ausdehnung des verholzten Teiles innerhalb der Sprossysteme bei mediterranen und submediterranen Strauchern, Halb- und Zwergstrauchern sowie bei mitteleuropaischen mehr oder weniger krautigen und geophytischen Vertretern wird am Beispiel von Arten der Gattungen Thymus und Teucrium erortert. Auserdem werden die Wuchsformen einiger geophytischer Stauden der mitteleuropaischen Waldvegetation aus der Gattung Euphorbia mit den Wuchsformen ihrer mediterranen Verwandten verglichen. For the understanding of the growth forms are to consider not only the morphological and anatomical characters but also the growth rhythm and the relations of plants to their habitat. Representing the growth forms of Carlina, Scabiosa, and Digitalis the relations between Canarian or South-Mediterranean candelabrum shrubs and Submediterranean-Middle-European pleiocorm perennials or annuals are demonstrated. Especially the differentiations of the pleiocorm (Scabiosa, Digitalis) and the radication (Carlina) are examined. Further there are discussed the different extension and intensity of the shoot lignification of Mediterranean shrubs, semishrubs, and dwarf shrubs on the one hand and of Middle-European more or less herbaceous and geophytic plants on the other hand with regard to several species of Thymus and Teucrium. Finally the growth forms of some geophytic Middle-European and some related Mediterranean species of Euphorbia are compared.

On some hybrid plants gathered in the March-Aprile region. — The A. reports the occurring in the March-Aprile of some interspecific or intergeneric hybrids not recorded in this region till now; they are: Polygonum × condensatum F. Schultz (= P. mite Schrank × P. persicaria L.), Centaurea rupestris L. × C. scabiosa L., × Danthosieglingia breviaristata (Beck ex Vierh.) Domin (= Danthonia provincialis DC. × Sieglingia decumbens (L.) Bernh.), and Orchis × dietrichiana Bogenh. (= Orchis tridentata Scop. × O. ustulata L.). Particulars are given about the characters of the specimens gathered, and about the localities where they were found and the distribution of the different taxa in the region and in Italy. Special attention is given to × Danthosieglingia breviaristata as it is a little known taxon, almost not dealt with in publications (at least as a hybrid) as regards the Italian Peninsula.

Darwin (1877) attempted to understand the significance and evolution of unisexuality in plants. In addition to dioecy, he recognized gynodioecy, in which separate female and hermaphrodite individuals are found, androdioecy, where there are males and hermaphrodites, and several other conditions. Most of the gynodioecious species studied by Darwin were members of the Labiatae, with one or a few species in other groups, such as the genera Scabiosa, Plantago and Cirsium (then called Cnicus). These species rarely showed any tendency to be dioecious. Since especially the Labiatae contain many gynodioecious and many hermaphroditic species but few dioecious ones, Lewis (1942) concluded that there was little evidence for an evolutionary connection between gynodioecy and dioecy. Such a conclusion is in accord with most of the then available evidence, but recent studies have shown that there are groups of related species of which some are gynodioecious and others are dioecious or nearly so, e.g. in Pimelea (Burrows, 1960; Ross, 1970b) and in New Zealand Umbelliferae (Lloyd, 1973). In one section of the genus Fuchsia three species are morphologically gynodioecious but functionally subdioecious (Arroyo and Raven, 1975). Since these groups have hermaphrodite relatives it seems that gynodioecy first evolved from hermaphroditism and then gave rise to near dioecy by gradual reduction of seed fertility of hermaphrodites. Consequently we may now recognize two evolutionary types of gynodioecy, namely a stable or classical type which rarely gives rise to dioecy, and an unstable type which appears to do so frequently. A model of the evolution of dioecy with male heterogamety through the development of complete linkage between a recessive gene m for male sterility, and a dominant gene F for female sterility, was presented by Lewis (1942). Males had genotype MF/mf and females mf/mf, but if the dominance of the two mutations was reversed, females would be heterogametic. Studies of gene substitutions in populations under Lewis' model with male heterogamety showed that dioecy could evolve, but only with large selection differentials (Ross and Weir, 1976). This result, together with the evidence that subdioecy evolved from gynodioecy by gradual reduction of seed set on hermaphrodites, suggested that a new hypothesis for the evolution of dioecy was required. Ross and Weir (1976) proposed a multilocus hypothesis with several recessive genes for male sterility and dominant or nondominant genes for partial female sterility. Valdeyron (1972) also proposed a multilocus control of female sterility in a subdioecious species, and Westergaard (1958) has inferred the presence of two male-sterility genes in Melandrium. The present paper has two main aims. First it aims to test Ross and Weir's hypothesis by studying a recessive gene for male sterility and a nonrecessive gene for partial female sterility. It is shown that there are interactions such that even without linkage there may be more females in equilibrium populations in the presence of partially female-sterile hermaphrodites ("subhermaphrodites") than in their absence, and vice versa. Linkage results in increased proportions of females and subhermaphrodites, and decreased proportions of hermaphrodites. It is held that additional mutations would behave in the same way. making the evolution of sub

(1) Competition for pollinators and avoidance of interspecific pollen transfer have been proposed as selective factors influencing flowering time and floral specializations. Their importance in a community of chalk grassland plants was studied. (2) The flowering periods of the important insect-pollinated herbs were determined. The most prominent flowering species were Helianthemum nummularium, Hippocrepis comosa and Onobrychis viciifolia during June and early July, and Centaurea nigra, C. scabiosa, Knautia arvensis and Scabiosa columbaria from late July to September. (3) Bumblebees, particularly Bombus lapidarius, were the most abundant visitors to most of these plant species; Apis mellifera, some Lepidoptera and Syrphidae were of lesser importance. (4) Some insect species differed in the flowers they visited most frequently. (5) In the early-flowering groups of plants there was some specialization for different pollinators and divergences in flowering time, suggesting that competition for pollinators or avoidance of interspecific pollen transfer has exerted selection pressure in the past. (6) In the late-summer group all species overlapped considerably in flowering time and their insect visitors, which suggests that there has not been any strong selection against an overlap. There appeared to be a saturation of insect visitors in late summer, with particularly large numbers of Bombus lapidarius.

l. The basic temperatures for growth of Ageratum houstonianum 'Capri', Calendula officinalis 'Dania', Scabiosa atropurpurea 'Imperial' and Tagetes patula var. nana 'Naughty Marietta' were estimated to be 8, 2, 4, and 60C respectively. The plants were raised under constant climatic conditions at temperatures of 3, 6, 9, 10, 12, 15, 18, and 21C. 2. The plants tended to be tallest at intermediate temperatures (15 and 18'C). At 210C the elongation growth was partially inhibited, especially in Scabiosa. At the highest temperatures Tagetes and Calendula reached first a maximum total length and then decreased somewhat again possibly because of decrease in turgor pressure after flowering. Internodal length was found to be positively correlated with temperature. 3. For Ageratum, Calendula and Tagetes the

The respiration for Ageratum houstonianum 'Capri', Calendula officinalis 'Dania', Scabiosa atropurpurea 'Imperial', and Tagetes patula var. nana 'Naughty Marietta' was examined manometrically in multithermostat at temperatures from 0 to 30'C. Within this temperature range the respiration increased more than linearly with the temperature, but slowest for Scabiosa. The regression equations y = a + bt2 gave a satisfactory expression for the respiration (see Figs. 2a-d). The respiration intensity at the various temperatures varied considerably for the cultivars. At 0C higher respiration was found for Calendula and Scabiosa than for Tagetes and Ageratum; at 30'C the respiration was highest for Tagetes and Calendula. The relative respiration with 10C chosen as a standard equal to 1.00 was for Calendula similar to values for spruce and pea reported by Dahl and Mork (Tab. 3). Above 15'C the deviations from these values were considerable for Ageratum and Tagetes. Still greater deviations from spruce and peas were found for Scabiosa. Correlations between growth and temperature data for the cultivars in the second growth phase were compared with correlations between growth data and calculated respiration sums according to the regression equations in the same growth period. By experiments in constant climate the elongations were also correlated with respiration equivalents calculated for corresponding periods as the other data. Higher correlation coefficients were in constant climate usually found for elongation correlated with one of the respiration measures than with the temperature itself. It suggests that respiration was one of the most important growth limiting factors under these conditions. In field experiments, however, highest correlation between growth data and calculated respiration sums was usually obtained only after exclusion of the experimental plots with highest temperatures.