Some evidence for infrageneric classification in Ziziphora L. (Lamiaceae:Mentheae)

The genus Ziziphora L. (Lamiaceae) consists of annual herbs, except Ziziphora clinopodioides, and is represented by six taxa in the flora of Turkey: Z. clinopodioides, Z. capitata, Z. persica, Z. tenuior, Z. taurica subsp. taurica and Z. taurica subsp. cleonioides. They are strong aromatic herbs and important medicinal plants. This study has been conducted to determine the palynological and karyological features of these taxa. The pollen morphology and exine structure of Ziziphora species were investigated by light microscope and scanning electron microscope. Pollen grains were determined as hexacolpate, medium in size and a circular or elliptic amb. The shape of pollen grains varies from oblate to prolate spheroidal. Ornamentation types of pollen grains are bireticulate, microreticulate or perforate. Somatic chromosome number has been determined and counted as 2n = 16, 18. Results indicate that the pollen characters and chromosome numbers of the genus are valuable for taxonomic applications and may be useful for infrageneric classification.     

A new taxon in the genusMoehringia (Caryophyllaceae)

The new taxonMoehringia intricata subsp.giennensis, from the calcareous mountains of the south-eastern Iberian Peninsula is described. A comparative morphological study with regard to the most closely related taxa, contributes information of leaf anatomy, seed and pollen morphology, ecology and distribution.     

Pollen morphology and phylogenetic analysis of Eperua Aublet (Detarieae: Caesalpinioideae: Leguminosae)

The diverse pollen morphology of fourteen species of the genus Eperua is described and illustrated using light, scanning electron and transmission electron microscopy. Six pollen types are described and a key for their identification is provided. A cladistic analysis was carried out using macromorphological and palynological characters to form a hypothesis of relationships between taxa. The pollen morphology is discussed with regard to systematic relationships, function and phylogenetic significance of certain pollen morphological structures within the genus.     

POLLEN MORPHOLOGY AND SYSTEMATICS OF SIPHONOGLOSSA SENSU LATO (ACANTHACEAE)

Pollen morphology of 15 species of Siphonoglossa and of two closely related groups was investigated. Two tribal-specific pollen types are found within Siphonoglossa sensu lato suggesting that the genus is artificial, composed of taxa belonging in several genera among two tribes (subtribes sensu Bremekamp) of Acanthaceae. Five taxa currently included in an informal subgeneric category of Siphonoglossa have tricolporate, prolate pollen (termed Type I) that is characteristic of Odontonemeae (= Odontoneminae, Justicieae). Pollen of the remaining taxa, belonging in two formal sections of the genus, are mostly 2-porate, bilateral (Type II) with a sexine sculpturing characteristic of Justicieae (= Justiciinae). Pollen of section Siphonoglossa is rather uniform, 2-porate, bilateral with lolongate pores, and seem to delimit a natural group. Taxa of section Pentaloba have a more heterogeneous pollen morphology, mostly 2-porate, bilateral with lalongate pores. Controversial aspects of the interpretation of pollen morphology in Justiciinae are presented and their relevance to this study are examined. Hypothetical trends in the evolution of pollen of Justiciinae are discussed and the application of pollen morphology to taxonomy of the genus is presented, including a recommendation for narrowing the generic concept of Siphonoglossa to the taxa of the type section.     

Pollen morphology of the family Cistaceae in Egypt and its systematic significance

The pollen morphology of 11 species (including two subspecies and two varieties) belonging to two genera (Helianthemum and Fumana) of the family Cistaceae in Egypt was studied using light and scanning electron microscopy.Pollen grains of the studied taxa were found to be radially symmetrical and tricolporate.Pollen size,shape,apertures,and exine ornamentation characteristics were valuable parameters among the studied taxa.The largest pollen size was recorded in H.salicifolium and the smallest one observed in H.kahiricum subsp,schweinfurthii.Pollen shape in the Egyptian taxa varied from (sub-)prolate to prolate spheroidal,but F.arabica is different in having sub-oblate grains.The pollen data confirm that H.lippii and H.sessiliflorum are very closely related species.Pollen sculpture was useful in distinguishing between H.vesicarium var.vesicarium and H.vesicarium var.ciliatum.Three main pollen types of exine ornamentation were recognized:retipilate; reticulate to verrucate; and striate.Based on palynological data,a key for the studied taxa is suggested.     

The metareticulate pollen morphology of Alternanthera Forssk. (Gomphrenoideae,Amaranthaceae) and its taxonomic implications

Alternanthera (Amaranthaceae) is a diverse genus largely restricted to the American Tropics that belongs to the alternantheroid clade containing C₄ and C₃–C₄ intermediate species. This research focuses on the study of pollen characters by studying 13 species, representatives of the two major clades and subclades of Alternanthera. General palynological comparisons were conducted with light microscopy (LM), scanning electron microscopy (SEM) and with confocal laser scanning microscopy (CLSM) for exine ultrastructure. Twenty-five characters were measured and described for Alternanthera and among these, 14 pollen characters were used to discriminate pollen groups using cluster analysis and canonical analysis of principal coordinates (CAP). Pollen form and ornamentation, pores number, spines length, number of ektexinous bodies and nanospines on the ektexinous bodies on pore membranes, arrangement of nanopores and spines on structural elements, and metareticula form were taxonomically important and therefore used to construct the first palynological key to the alternantheroid clade species. Our study indicates that the seemingly subtle morphological variation of pollen is useful for recognising three main pollen types within Alternanthera. The much needed palynological terminology for describing the mesoporium in the metareticulate pollen of Amaranthaceae is provided.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCanina G. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.     

Taxonomic importance of seed morphology in Gentiana (Gentianaceae)

The seeds of Gentiana show high diversity in size, shape and surface pattern. Until now, seeds of only a limited number of taxa have been studied in detail and their utility in the subgeneric classification of Gentiana has not been evaluated. In this study we investigated seed micromorphology of the poorly known sections Pneumonanthe, Frigidae and Isomeria. In order to evaluate the relevance of seed sculpturing for taxonomy, we selected qualitative characters of seed shape and testa ornamentation of all documented sections and performed cluster analyses based on presence and absence of coded characters. Several new scanning electron microscopy pictures of seeds provided a number of valuable characters for the subgeneric differentiation of Gentiana. The results of the cluster analyses of seed characters generally support the sectional classification given by former authors. However, the European Gentiana asclepiadea does not group together with other taxa of section Pneumonanthe, but clusters near to the strictly European section Gentiana. Gentiana froelichii, endemic to the south‐east Alps and treated traditionally within section Frigidae, clusters together with taxa of the European section Ciminalis. Our results reveal that seed micromorphology supports the reassignment of some taxa incertae sedis within Gentiana. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 101–115.     

Pollen morphology of the genus Iris L. (Iridaceae) from Croatia and surrounding area: taxonomic and phylogenetic implications

In Croatia and the surrounding area, the genus Iris is represented with about 20 Alpine-Dinaric, Mediterranean and Pannonian taxa from the subgenera Iris and Limniris. We researched pollen morphology of all taxa by using scanning electron microscopy. All pollen grains are sulcate with few palynological features, which could have taxonomic importance on different classification levels: shape of dry pollen grains; shape, outline and approximate size of hydrated pollen grains, approximate size of the sulcus, and ornamentation of the sulcus membrane and of the exine. At least four pollen types were recognised and taxonomically delimited to the series level: two characterise the subgenus Iris, section Iris (series Elatae and Pumilae), and two the subgenus Limniris, section Limniris (series Laevigatae, Sibiricae and Spuriae). Taxonomic implications of pollen morphology of their critical groups and taxa have been evaluated. Possible phylogenetic implications of pollen morphology of the genus Iris on the Southern European area were suggested: the subgenus Iris is more advanced than the subgenus Limniris; I. graminea seems to be the most primitive member of the subgenus Limniris on the Southern European territory; and pollen morphology of the population of the subspecies I. sibirica subsp. erirrhiza from the Mountain Bjelolasica could represent a link between the subgenera Limniris and Iris in the territory of Southern Europe. The possible pathway of evolution of the genus Iris on the territory of Southern Europe was suggested: from the subgenus Limniris, through the “linked taxon” I. sibirica subsp. erirrhiza from the Bjelolasica Mountain, to the subgenus Iris, series Pumilae, and finally to the series Elatae.     

Pollen morphology of endemic species of the Gonospermum Less., Lugoa DC. and Tanacetum L. complex (Asteraceae: Anthemideae) in the Canary Islands (Spain), and its taxonomical implications

The pollen of nine endemic taxa of the Gonospermum Less., Lugoa DC. and Tanacetum L. complex and of T. parthenium L. (introduced species from the Canary Islands) was studied using light and scanning electron microscopy with the aim of extending the knowledge of their morphology and to investigate whether pollen morphology may help clarifying the controversial taxonomy of these taxa. Twenty six populations in the Canary Islands (Spain) were studied and the following parameters were measured: polar axis (P), equatorial diameter (E), polar exine thickness (Ex1), length (L) and thickness (A) of the spines in the meridional optical section, exine thickness at the centre of the mesocolpium in the interapertural area (Ex2) and distance between spines (De) in the equatorial optical section. The studies show that the pollen of the three genera is very similar regarding shape, apertures, and ornamentation, but may be distinguished by size and exine thickness.     

Pollen morphology and localization of Ni in some Ni-hyperaccumulator taxa of Alyssum L. (Brassicaceae)

Pollen morphology of seven Alyssum L. taxa growing on serpentine soils in different places in the European Mediterranean macrobioclimate territory were studied, described and compared. Cluster analysis was performed to show similarity between species and their populations. The shape of the pollen grains varies among the species and among the grains within the same anther. The pollen grains are 3-colpate, prolate, with long and narrow colpi reaching the poles. The ornamentation of the exine varies from micro-reticulate to reticulate between the species. Pollen sterility/fertility was also calculated. The highest percentage of sterile pollen (73.76%) was calculated for Alyssum murale subsp. murale and the lowest (9.54%) for A. bertolonii subsp. bertolonii. All species are representatives of sect. Odontarrhena (C.A. Meyer) Koch well known as Ni-hyperaccumulators. Nickel and other elements present in pollen and stamen were studied by inductively coupled plasma-mass spectrometry. The stamen parts of all species were micromorphologically analyzed by scanning electron microscopy coupled to an energy-dispersive X-ray probe. Accumulation of Ni was detected in the stamens of all studied species and rarely in the pollen grains. The distribution patterns of Ni were similar among the species examined.     

Taxonomy and cytogeography of Cardamine raphanifolia and C. gallaecica (Brassicaceae) in the Iberian Peninsula

For a long time it was considered that Cardamine raphanifolia is widely distributed with several infraspecific taxa in the Mediterranean region. Recent studies showed that C. raphanifolia is restricted to the northern Iberian Peninsula. Cardamine gallaecica from NW Spain was regarded to be closely related, but was originally regarded as C. amara, and later described as C. raphanifolia subsp. gallaecica. In this study, we validate the treatment of C. raphanifolia and C. gallaecica as distinct species, and show their clear differentiation based on morphometric and molecular AFLP analyses. Moreover, both species are clearly distinct from C. amara subsp. pyrenaea, which has an overlapping distributional range with C. raphanifolia. Hexaploid (2n = 48) and octoploid cytotypes (2n = 64) of C. raphanifolia, and tetraploid (2n = 32) and hexaploid (2n = 48) cytotypes of C. gallaecica were found. Due to their weak morphological and genetic differentiation, these polyploid races are not classified as separate taxa.     

Comparative pollen morphology of Turkish species of <Emphasis Type="Italic">Petrorhagia</Emphasis> (Caryophyllaceae) and its systematic implications

Pollen morphology of 10 taxa from Turkey, four of them endemic, belonging to the genus Petrorhagia (Caryophyllaceae), P. alpina subsp. alpina, P. alpina subsp. olympica, P. cretica, P. dubia, P. hispidula, P. lycica, P. pamphylica, P. peroninii, P. prolifera and P. saxifraga, has been investigated using light (LM) and scanning electron (SEM) microscopy. Pollen morphology differences among these taxa have been determined. The pollen type of investigated taxa is polyporate (12–22-porate), sphaeroidal, the exine exhibits a tectate structure and a microechinate ornamentation. Petrorhagia alpina subsp. alpina has the smallest pollen grain diameter (18.57 μm) and P. dubia the largest one (37.80 μm). The number of pores ranges from 12 (in P. dubia and P. saxifraga) to 22 (in P. alpina subsp. alpina and P. alpina subsp. olympica and P. hispidula) with a minimum pore diameter of 2.37 μm in P. alpina subsp. alpina and a maximum pore diameter of 4.23 μm in P. peroninii. The exine thickness ranges from 1.73 μm in P. saxifraga to 3.78 μm in P. pamphylica. In this study, the systematic implications are discussed in the light of palynological results.     

Palynological characters of Glycyrrhiza,Glycyrrhizopsis, and Meristotropis (Leguminosae), with special reference to their taxonomic significance

Abstract The pollen morphology of 11 species of the genus Glycyrrhiza L. with one from each of the genera Glycyrrhizopsis Boiss. & Bal. and Meristotropis Fisch. & C. A. Mey. was investigated by scanning electron microscopy. In pollen morphology, the main differences between Glycyrrhizopsis and Glycyrrhiza are: Glycyrrhizopsis—pollen grains 36.63 × 40.42 μm in size, oblate spheroidal in shape; and Glycyrrhiza—pollen grains 24.47–33.18 × 23.82–31.83 μm in size, prolate spheroidal in shape. Glycyrrhizopsis and Glycyrrhiza should be recognized as two distinct genera based on palynological and morphological characters. Meristotropis and Glycyrrhiza are similar in many important palynological and morphological characters, suggesting that the two should be merged. In Glycyrrhiza, two types of pollen grains, 3‐lobed‐circular or subtriangular in polar view, are found in different species, in accordance with morphological differences in the two groups, shedding light on the classification and evolution of the genus.     

Geographical Distribution of Diploid and Tetraploid Cytotypes of Thymus sect. Mastichina (Lamiaceae) in the Iberian Peninsula, Genome Size and Evolutionary Implications

The occurrence of diploid and tetraploid cytotypes in Thymus section Mastichina (Lamiaceae) was investigated by chromosome counting and flow cytometric measurements of DNA content. To cover the entire native distribution of the section across the Iberian Peninsula, plant material was sampled from 29 populations, representing all three taxa (Thymus albicans, T. mastichina subsp. mastichina and T. mastichina subsp. donyanae). We provide first estimates of genome size for taxa of this section. Analyses revealed the existence of new cytotypes in T. mastichina subsp. mastichina (2n?=?2x?=?28, 30) and confirmed the existence of previous ones (2n?=?4x?=?56, 58, 60). We also confirmed the presence of exclusively diploid cytotypes (2n?=?2x?=?30) in the southwestern Iberian endemics T. albicans and T. mastichina subsp. donyanae. We conclude that the southwestern Iberian Peninsula acted as a Pleistocene glacial refugium that may have permitted speciation processes within this section. Several hypotheses concerning these processes and the origin and distribution of the studied taxa are discussed.     

Ecogeographical differentiation patterns inAdonis sect.Consiligo (Ranunculaceae)

Adonis sect.Consiligo is split into three subsections based on morphology, growth pattern, pollen and fruit morphology. For all species distribution maps are provided and discussed with regard to their ecology and climatic requirements. These data reveal additional characters that support the classification of the taxa. Most species of the section grow in continental Eurasiatic steppes and mountain steppe habitats. Only the species of subsect.Amurenses grow in Far Eastern temperate forests and prefer a more oceanic climate. The taxonomic position of these strongly divergent species is supported by the distributional and ecological data. The evolution ofAdonis sect.Consiligo occurred under strong climatic control. During evolution the taxa could only slightly widen their ecological amplitude and, thus, ecology remained rather stable except that of the subsect.Amurenses. Ecological and distributional data can be used to validate taxonomy and phylogeny and reveal additional informations.     

Micromorphological observations on leaf and pollen of Capparis L. sect. Capparis (Capparaceae)

Sect. Capparis is represented by a single species, Capparis spinosa L., divided into several intraspecific taxa showing plesiomorphic features and disjunct distributions in the Old World. Leaf surface and pollen features were investigated in the whole group by SEM and light microscope observations. The section is characterized by simple hairs, a reticulate to undulate cuticle, anomocytic stomata surrounded by a peristomal rim, and trizonocolporate, prolate pollen grains. The characteristics of the indumentum appear constant, while the studied taxa are fairly differentiated with respect to cuticular patterns and dimensions of the stomata, and show slight differences in pollen size and exine surface. This micromorphological evidence, coupled with other phenotypic features, supports the placement of this section at the base of the genus Capparis in the paleotropical area. Considering the striking geographic disjunction and symplesiomorphies of the group, its biogeographical and systematic aspects are also discussed.     

Palynological investigations on some Hypericum taxa (Hypericaceae) growing naturally in Turkey

This study examined the detailed pollen morphological structures of sixteen Hypericum taxa (four endemic, twelve non endemic) including eight sections showing the natural distribution in Turkey: H. sect. Ascyreia (H. calycinum L.), H. sect. Heterophyllum (H. heterophyllum Vent.), H. Sect. Taeniocarpium (Hypericum confertum Choisy subsp. confertum, H. venustum Fenzl, H. linaroides Bosse), H. sect. Drosocarpium (H. montbretii Spach, H. bithynicum Boiss.), H. sect. Crossophyllum (H. adenotrichum Spach, H. orientale L.), H. sect. Olympia (H. olympicum L. subsp. olympicum), H. sect. Origanifolia (H. origanifolium Willd., H. avicularifolium Jaub and Spach subsp. depilatum (Freyn and Bornm.) Robson var. depilatum, H. avicularifolium Jaub. and Spach subsp. byzantinum) and H. sect. Hypericum (H. tetrapterum Fries, H. perforatum L., H. triquetrifolium Tura). These taxa were studied under light microscope and scanning electron microscope for the first time. Of the taxa studied, H. tetrapterum has the smallest pollen grains (on average 15.85–17.20 × 15.45–16.05 μm); and H. olympicum subsp. olympicum the largest grains (on average 22.90–23.10 × 21.40–22.10 μm). The pollen grains of 15 taxa are subprolate and one taxon is prolate-spheroidal. The regular pollen grains of all 16 taxa are 3-zonocolporate. Ornamentation is microreticulate; lumina regularly spaced in eight taxa, tectum perforatum; tectal perforations regularly spaced in seven taxa and tectum perforatum; and tectal perforations grouped together in one taxon. Endoaperture is cruciform porus, with short lateral and meridional extensions in ten taxa, cruciform porus, transversally elongated, with very small lateral extensions in one taxon and lalongate colpus in five taxa. Basic pollen types are ten taxa in type X, five taxa in type IV and one taxon in type II.     

The evolution of pollen heteromorphism inViola: A phylogenetic approach

Pollen heteromorphism, defined here as the production within all flowers of a plant of several pollen morphs differing in aperture number, is common in angiosperms. We have focused on the evolution of pollen heteromorphism in the genusViola, in which about 1/3 of the species are pollen-heteromorphic. We have studied the distribution of pollen heteromorphism in the genusViola using a molecular phylogeny based on ITS sequence data. We show that pollen heteromorphism has evolved independently at least six times inViola. A comparative analysis shows that the occurrence of pollen heteromorphism is correlated with sporophytic polyploidy in all sections of the genus apart from sectionMelanium. This section differs from all other sections on several aspects such as flower morphology, absence of cleistogamous flowers, and a high proportion of heteromorphic species. We discuss the possible adaptiveness of pollen heteromorphism in this section.     

A study of Sarcocornia A.J. Scott (Chenopodiaceae) from Western Mediterranean Europe

The latest publications on Sarcocornia taxonomy and phylogeny recognize six taxa in this genus on the Iberian Peninsula: S. perennis, S. fruticosa, S. alpini, S. alpini subsp. carinata, S. hispanica, and S. pruinosa. The present study represents a comprehensive revision of the different taxa in the Sarcocornia genus present in Western Mediterranean Europe by means of morphological, micromorphological and phylogenetic internal transcribed spacer (ITS) analysis. Morphological and micromorphological data were studied from Sarcocornia samples from 113 populations in coastal salt marshes and inland salt pans in Portugal, France, Spain and Italy. Sixteen new ITS sequences were obtained from Mediterranean Sarcocornia species and analysed together with previous reported data. Published karyological, ecological and biogeographical data from Western Mediterranean Europe were also reviewed. The results indicate the presence of a new species, S. lagascae, found growing in coastal Mediterranean areas of the Iberian Peninsula. The species S. fruticosa was found to be absent from the Iberian territories.