Die pollenmorphologie der afrikanischen buxus- und notobuxus-arten (buxaceae) und ihre systematische bedeutung

Pollen grains of 8 African species of Buxus and Notobuxus were examined by light and scanning electron microscopy. Three pollen types are recognized based on the exine sculpture and the evolutionary trends of the pollen characters are discussed. Pollen morphology shows the existence of three major systematic groups. These results support the previous classification of an African section Probuxus with two subsections and speak in favor of the generic rank of the Notobuxus species. Together with anatomical and taxonomical data pollen morphology and its evolutionary trends lend support to the suggestions that the centre of origin of the genus Buxus might have been in Africa, while secondary centres of evolution have developed in the neotropical and Eastasian region.     

Normapolles flowers of fagalean affinity from the Late Cretaceous of Portugal

 Exquisitely preserved, charcoalified fossil flowers with in situ pollen of the Normapolles-type from the Late Cretaceous of Portugal are described and a new genus and species of Fagales, Normanthus miraensis, are established. Floral organization and structure of floral organs were studied with scanning electron microscopy and microtome sections. Flowers are actinomorphic, epigynous, and pentamerous; the perianth is simple; stamens alternate with tepals; pollen is oblate and vestibulate; the exine is thick and the tectum is scabrate-microgranulate; the gynoecium is bicarpellate and unilocular; the fruits are probably one-seeded. Comparisons with extant taxa demonstrate that N. miraensis shares many similarities with Fagales and in particular with Betulaceae. However, it is not identical with any extant taxon and cannot be included in any extant family. The combination of characters found in the fossil flowers is congruent with wind-pollination syndromes present in many extant angiosperms and clearly indicates wind-pollination of N. miraensis. Received June 13, 2000 Accepted September 28, 2000     

Elucidating the unusual floral features of Swartzia dipetala (Fabaceae)

In this study, we evaluated the floral ontogeny of Swartzia dipetala, which has peculiar floral features compared with other legumes, such as an entire calyx in the floral bud, a corolla with one or two petals, a dimorphic and polyandrous androecium and a bicarpellate gynoecium. We provide new information on the function of pollen in both stamen morphs and whether both carpels of a flower are able to form fruit. Floral buds, flowers and fruits were processed for observation under light, scanning and transmission electron microscopy and for quantitative analyses. The entire calyx results from the initiation, elongation and fusion of three sepal primordia. A unique petal primordium (or rarely two) is produced on the adaxial side of a ring meristem, which is formed after the initiation of the calyx. The polyandrous and dimorphic androecium also originates from the activity of the ring meristem. It produces three larger stamen primordia on the abaxial side and numerous smaller stamen primordia on the adaxial side. These two types of stamens bear morphologically similar ripening pollen grains. However, prior to the dehiscence of thecae and presentation of pollen in the anther, only the pollen grains of the larger stamens contain amyloplasts. Two carpel primordia are initiated as distinct protuberances, alternating with the larger stamens, in a slightly inner position in the floral meristem, constituting the bicarpellate gynoecium. Both carpels are able to form fruit, although only one fruit is generally produced in a flower. The increase in gynoecium merism probably results in an increase in the surface deposition of pollen grains and consequently in the chance of pollination. This is the first study to thoroughly investigate organogenesis and the ability of the carpel to form fruit in a bicarpellate flower from a member of Fabaceae, in addition to the pollen ultrastructure in the heteromorphic stamens associated with the ‘division of labour’ sensu Darwin. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 303–320.     

Neighbourhood interactions and environmental factors influencing old‐pasture succession in the Central Pyrenees

Abstract. The shrub Buxus sempervirens and the trees Abies alba and Fagus sylvatica have recently recolonized old‐pastures in the Central Pyrenees. We mapped all live and dead individuals (> 1.3 m tall) in a large forest plot in Ordesa Valley to examine the importance of density‐dependent processes during recolonization. Biotic interactions were inferred from changes in horizontal structure and the influences of neighbours on tree survival. Buxus differentially influenced establishment and survival of tree species, thereby controlling future canopy composition and spatial structure. The rapidly invading Abies formed denser patches on elevated sites less occupied by Buxus, whereas Fagus preferentially established within shrubs. Abies reached densities which led to intense intraspecific competition and high mortality rates among saplings. Self‐thinning in Abies led to smaller numbers of regularly spaced survivors, and greater relative dominance of Fagus. Disregarding intraspecific effects and abiotic environment, Abies survival was significantly lower under Buxus shrubs, which suggests that the spatial location and abundance of Abies was constrained by the location of Buxus. Fagus survival was not related to Buxus density, but remained significantly lower in denser Abies patches. The higher mortality of Fagus in denser Abies patches, and the resulting spatial segregation of the species, reflects asymmetric interspecific competition. Inhibition from Buxus shrubs and intraspecific competition prevent invading Abies from dominating and may therefore help in maintaining a mixed Abies‐Fagus stand.     

Unreduced diploid nuclei in Cupressus dupreziana A. Camus pollen

Size and DNA content of pollen of Cupressus dupreziana A. Camus, a highly endangered Mediterranean conifer, were assessed by cytomorphological observations and flow cytometric analyses and then compared to C. sempervirens L. pollen. Mature C. dupreziana pollen was composed of two uninucleated types of pollen grains differing in size. Around 35% of the grains exhibited a size similar to C. sempervirens pollen, while 65% exhibited a larger diameter. However only one peak of fluorescence was detected by flow cytometry. DNA content of C. dupreziana pollen was twice the DNA content of C. sempervirens pollen. Comparison of DNA contents of mature and germinating pollen revealed that mature pollen of both species were arrested in the G2 phase. Comparison with the DNA content of somatic tissue (2C) provided evidence for the production of unreduced pollen in C. dupreziana. This unexpected feature in gymnosperms is discussed in terms of reproductive strategy of this species. Received: 11 December 1999 / Accepted: 22 December 1999     

Variation and similarities in pollen features in some basal angiosperms, with some taxonomic implications

Species within three families of basal angiosperms (Trimeniaceae, Winteraceae, Monimiaceae) illustrate differences and similarities in pollen within a species, between species and between genera. Trimenia papuana (Trimeniaceae) has dimorphic pollen (inaperturate, polyforate), each confined to different individual plants. Other species have either disulculate or polyforate pollen. Evolution seems to be from disulculate to inaperturate to polyforate. Present-day Winteraceae have pollen in permanent tetrads except four species of Zygogynum with monads. Why? Did such monads appear as fossils before tetrads in Winteraceae? Molecular studies of Takhtajania perrieri indicate it is basal but its unique bicarpellate unilocular gynoecium seems derived. Although Hedycarya arborea and Kibaropsis caledonica have near-identical permanent pollen tetrads, many other features are very different. Hedycarya species have permanent tetrads or inaperturate monads with spinulose, `starry' or other sculpturing, and it is suggested this and recent molecular data indicate further studies are needed to determine generic limits.     

Identifying the ecological and societal consequences of a decline in <Emphasis Type="Italic">Buxus</Emphasis> forests in Europe and the Caucasus

The potential impact of new invasive tree pests and diseases is usually quantified in economic terms. The ecological and social impacts are less often assessed. Using a comprehensive literature review we assess the potential ecological and social impact of two non-native invasive species (the box tree moth, Cydalima perspectalis and the fungus Calonectria pseudonaviculata) that threaten the survival of box tree, Buxus spp. in forests in Europe and the Caucasus. A total of 132 fungi, 12 chromista (algae), 98 invertebrate and 44 lichens were found to use Buxus spp. Of these, 43 fungi, 3 chromista and 18 invertebrate species have only been recorded on Buxus spp., suggesting that these species are obligate on Buxus spp. and are most at risk from in the loss of Buxus spp. due to these invasive pest and disease species. Buxus spp. was shown to be important for soil stability and water quality but there was no information on other ecosystem functions provided by Buxus spp. Buxus was found to be of considerable historical cultural importance but there was very limited information on current social values and uses. Buxus trees, wood and leaves are associated with different folklore and sacred rites which are still particularly important in the Caucasus. While we could not find any assessment of the economic value of Buxus forests the biodiversity, cultural and social values of Buxus identified here indicate that its loss could have major indirect and non-market economic effects. This work highlights the importance of studying the ecological and societal implications of biological invasions.     

Relating postglacial relict plants and Holocene vegetation dynamics in the Balearic Islands through field surveys,pollen analysis and GIS modeling

Abstract

An inventory of the current Balearic plants occurring in mesophyllous environments, with climatic conditions similar to those in the first half of the Holocene, was compiled. The presence of these floristic elements, extending mainly into the postglacial time, has been related to pollen records of different localities in the Balearic Islands. GIS techniques have been applied in order to map the past and the present-day vegetation suitability of some of these mesophyllous species using pollen surface samples, forest inventory data and Digital Climatic Models. We integrated the information provided by field survey, pollen records and suitability mapping in order to discuss the vegetation dynamics of different habitats. We focused on the nature and the possible causes of the vegetation landscape change that occurred approximately 5000 – 6000 uncal. years BP, when Fagus, Corylus and Buxus formations were replaced by xeric-type formations with Olea dominance.     

Early Cretaceous floral structures and in situ tricolpate‐striate pollen: New early eudicots from Portugal

Five new taxa with affinities to extant lineages that diverged at an early stage from the main line of eudicot evolution are established from the Early Cretaceous (late Aptian or early Albian) Vale de Agua locality, Portugal. Staminate flowers of Lusistemon striatus and pistillate flowers of Lusicarpus planatus are unisexual without rudiments of the opposite gender. They are linked by the association of an unusual pollen type found in situ in the stamens and adhering to the stigmatic surface. The staminate flower, Lusistemon striatus, is composed of six stamens subtended by small perianth parts. The arrangement of the stamens is difficult to ascertain, but their variable sizes suggests a spiral arrangement. Pollen found in situ is tricolpate and striate with densely‐spaced, sparsely diverging and anastomosing muri that are aligned more or less parallel to the polar axis. The muri have a conspicuous supratectal ornamentation of fine transverse ridges. The granular infratectal layer forms an indistinct internal reticulum. The foot layer is thin. Pollen is closely similar to dispersed grains from the Aptian of Egypt described as STRIOTRI‐SEGMUR. It also resembles pollen of the dispersed pollen genus Rutihesperipites, as well as some dispersed pollen assigned to Striatopollis. Pistillate flowers of Lusicarpus planatus consist of a bicarpellate, syncarpous gynoecium borne on a short stalk. The styles are bent outwards and expose the double‐crested stigmatic regions on their ventral sides. The only organ preserved besides the gynoecium is a lateral scale‐like organ at the base of the stalk. Pollen of the same type found in Lusistemon striatus occurs on the stigmatic surface of the carpels. Comparisons with extant taxa demonstrate that Lusistemon and Lusicarpus share many characters with early diverging groups of eudicots, in particular Buxaceae. In addition to the Lusistemon‐Lusicarpus flowers, the Vale de Agua samples also contain three other pistillate reproductive structures that may be related to early diverging lineages of eudicots. Silucarpus camptostylus has a bicarpellate and syncarpous gynoecium with two styles; Valecarpus petiolatus and Aguacarpus hirsutus have tricarpellate gynoecia that are distinguished from each other in the shape and extension of the stigma as well as other details.     

Jasminum honghoense sp. nov. (Oleaceae), from the dry‐hot Yuanjiang valley in Yunnan,China

Jasminum honghoense (Oleaceae), a new species from the dry‐hot Yuanjiang valley in southern central Yunnan, China, is described and illustrated. Differences between J. honghoense and two morphologically similar species are presented. Moreover, the pollen morphology of J. honghoense is reported.     

Chromosomal location of a pollen fertility-restoring gene, Rf, for CMS in Japanese bunching onion (Allium fistulosum L.) possessing the cytoplasm of A. galanthum Kar. et Kir. revealed by genomic in situ hybridization

In a previous study, we developed cytoplasmic male sterile lines of Allium fistulosum possessing the cytoplasm of A. galanthum, a wild species, by continuous backcrossing. Furthermore, we reported the presence of a pollen fertility-restoring gene (Rf) for cytoplasmic male sterility (CMS) in A. fistulosum from segregation of pollen fertility of backcross progenies. In the present study, genomic in situ hybridization (GISH), using genomic DNA of A. galanthum as the probe DNA and that of A. fistulosum as the blocking DNA, was applied to F₁ hybrids between both species and backcross progenies to determine the chromosomal location of the Rf locus. By means of GISH, eight chromosomes from A. galanthum were clearly discriminated from those of A. fistulosum in the F₁ hybrids, and chromosome substitution process through continuous backcrossing was visualized. Interestingly, the chromosome region from A. galanthum, specific to male fertile plants, was detected in one chromosome of BC₄ to BC₇ generations. Based on the karyotype analysis of the male fertile plants, the chromosome was identified as the 5F chromosome. Our results confirm that the Rf locus is located on the 5F chromosome of the male fertile plants. This is the first report that identified the chromosomal location of the pollen fertility-restoring gene in A. fistulosum.     

Studies in Montia L. and Claytonia L. and Allied Genera

The pollen morphology of several genera in Portulacaceae is described. Particular attention has been paid to the genera of the subfamily Montioideae, as a stage of continued monographical studies. Among genera especially dealt with are Claytonia, Montia, Crunocallis, Naiocrene, Neopaxia, Mona, Maxia, Limnalsine, and Montiastrum. In the taxonomical treatment of these genera the pollen morphology has proved to afford many important additional characters.

The pollen grains of Claytonia are distinguished from those of the remainder in being 3-colpate. The grains of the Claytonia-type have many similarities with those of Lewisia, a genus of the subfamily Portulacoideae. The other genera of Montioideae have pantocolpate pollen grains. Among these genera several different pollen types are distinguished, chiefly with regard to the sexine structures and the aperture membranes. The Montiastrum-type is especially interesting, with tholate grains, a particular pollen type not met with in any other genus in the family. The pollen morphology of some genera in the Portulacoideae is also treated. In some species in Calandrina and Talinum pantotreme pollen grains are observed with apertures transitional between pori and colpi. The apertures of the pantotreme grains are arranged in characteristic patterns.

Particular attention has been given to the variation of the pollen morphological characters. This variation has been examined with regard to the differences between different populations of the same species as well as between different species. The greatest variation has been observed in the shape and size of the grains. The structure and sculpture and thickness of the sexine and the aperture membranes are less variable. Some polyploid taxa are connected with the occurrence of pollen grains with divergent and varying aperture numbers.

In a survey of the genera the taxonomical results of the investigation are presented with particular regard to the pollen morphology. The new genus, Maxia Ö. Nilss., is described. One new species, Montia clara Ö. Nilss., is described and some new combinations are made.

Pollen morphological diagnoses are given for 46 different taxa. The aperture conditions for 96 different species are presented.     

SYSTEMATIC SIGNIFICANCE OF POLLEN NUCLEAR NUMBER IN EUPHORBIACEAE,TRIBE EUPHORBIEAE

Pollen nuclear number is determined in 139 species of 5 genera in the Euphorbieae, subtribe Euphorbiinae. The 111 new determinations are tabulated along with previous reports, and the results indicate that the distribution of binucleate (II) and trinucleate (III) pollen is strongly associated with the taxonomic groupings within the Euphorbieae. Although binucleate pollen is probably primitive within the tribe Euphorbieae, as suggested by the nuclear condition in Neoguillauminia, the situation in Euphorbia still requires further elucidation. Within Euphorbia, the morphologically most primitive species studied have III pollen despite the fact that II pollen is presumably the original condition for the subtribe Euphorbiinae. In Euphorbia, II pollen only is reported from nine sections and III pollen only from ten sections, while in four sections (Esula, Goniostema, Aphyllis, and Deuterocalli) both II and III pollen have been found. The New World species of Euphorbia nearly all have III pollen, whereas the vast majority of the African succulents have II pollen. The genera of New World origin, Chamaesyce and Pedilanthus, have III pollen, while the African genera Monadenium and Synadenium have II pollen. Independent derivations of III pollen from II pollen appear to have occurred in sections Goniostema, Aphyllis, and Deuterocalli (all of subg. Euphorbia). There is no evidence that reversals from III to II pollen have occurred.     

Aperture Orientation in Laurelia Pollen (Atherospermataceae Syn. Subfamily Atherospermoideae of Monimiaceae)

The development of the encircling aperture of pollen grains or the New Zealand species Laurelia novae-zelandiae is described by means of light and scanning electron microscopy. Some observations are also reported from pollen development and structure in the South American species Laurelia sempervirens, mature pollen of which can be distinguished from that of L. novae-zelandiae. The aperture begins to develop while pollen of L. novaezelandiae is still in tetrads, and passes through the distal and proximal poles of each grain. The widest parts of the mature aperture are at and near what are morphologically the distal and proximal poles of each grain. No such meridionosulcate type of pollen was recorded in a recent survey of pollen aperture types of primitive angiosperms and contradicts a recent report that pollen in the Atherospermataceae had equatorially aligned apertures.     

Comparative palynology and seed morphology in annual pansies (Viola sect. Melanium,Violaceae): implications for species delimitation

Abstract

This integrated study provides new insights into pollen and seed morphology and pollen heteromorphism of four closely related annual taxa of Viola sect. Melanium. The plant material, both fresh and dried, was collected in Italy and studied using light and scanning electron microscope. Palynological data for V. hymettia together with a detailed comparative analysis of seed morphology and micromorphology of the four species are reported for the first time. Results of this work highlight some pollen and seed features as useful diagnostic characters. The pollen size proves to be of diagnostic value to easily separate V. kitaibeliana, having the smallest pollen grains, from the others, especially from V. arvensis with the largest ones. Exine ornamentation is microreticulate, showing no relevant differences among species. We can partially confirm the diagnostic value of the prevailing pollen morph as it can be useful only for V. arvensis (five-aperturate) versus V. tricolor and V. hymettia (four-aperturate). The macro- and micromorphology of seeds provide additional useful distinguishing characters. Particularly, seed size was found to be a good delimitating character, especially to distinguish V. kitaibeliana (with the smallest seeds) from V. arvensis, and easy to be measured with no need of particular equipment.     

Airborne‐pollen pool and mating pattern in a hybrid zone between Pinus pumila and P. parviflora var. pentaphylla

The reproductive isolation barriers and the mating patterns among Pinus pumila, P. parviflora var. pentaphylla and their hybrids were examined by flowering phenology and genetic assays of three life stages: airborne‐pollen grains, adults and seeds, in a hybrid zone on Mount Apoi, Hokkaido, Japan. Chloroplast DNA composition of the airborne‐pollen was determined by single‐pollen polymerase chain reaction. Mating patterns were analysed by estimating the molecular hybrid index of the seed parent, their seed embryos and pollen parents. The observation of flowering phenology showed that the flowering of P. pumila precedes that of P. parviflora var. pentaphylla by about 6 to 10 days within the same altitudinal ranges. Although this prezygotic isolation barrier is effective, the genetic assay of airborne‐pollen showed that the two pine species, particularly P. pumila, still have chances to form F₁ hybrid seeds. Both parental species showed a strong assortative mating pattern; F₁ seeds were found in only 1.4% of seeds from P. pumila mother trees and not at all in P. parviflora var. pentaphylla. The assortative mating was concluded as the combined result of flowering time differentiation and cross‐incompatibility. In contrast to the parental species, hybrids were fertilized evenly by the two parental species and themselves. The breakdown of prezygotic barriers (intermediate flowering phenology) and cross‐incompatibility may account for the unselective mating. It is suggested that introgression is ongoing on Mount Apoi through backcrossing between hybrids and parental species, despite strong isolation barriers between the parental species.     

Plant male germ line transformation

A new method to produce transgenic plants - ma le ge rm li ne tr ansformation (MAGELITR) is reported. Unicellular tobacco microspores were isolated from excised anthers, and DNA carrying two marker genes was transferred biolistically. The bombarded microspores were matured in vitro for 6 days, and the mature pollen was used for in vivo pollination. Seeds were recovered and putative transformants were selected on the basis of their antibiotic resistance. Five kanamycin-resistant plants were chosen for further analysis, four contained the first construct, one the second. Parallel experiments with bicellular immature pollen did not produce any transgenic plants. A detailed DNA blot and expression analysis confirmed the transgenic nature of the five plants, and a genetic analysis showed that the transgenes are transmitted to subsequent generations. MAGELITR is a fast, regeneration-independent method, not prone to chimerism and somaclonal variation which should be genotype-independent and may be applicable in a wide range of species once in vitro maturation of pollen is established.     

Saussurea pseudograminea sp. nov. (Asteraceae) from the Qinghai–Tibetan plateau,China

Morphological characteristics, as well as ultrastructure of pollen grains, chromosome numbers and karyotype analysis have been used to establish a new species of Saussurea from the Qinghai–Tibetan plateau. The new species, Saussurea pseudograminea Y. F. Wang, G. Z. Du et Y. S. Lian is easily distinguished from the similar S. graminea Dunn by having 2–3 capitula, involucre 0.7–1.2 cm in diameter, smaller pollen grains, pollen surface with larger and denser spines, achenes 4.0–5.5 mm long, 32 chromosomes, and a karyotype formula 2n = 2x = 32 = 18m + 10sm + 4st, whereas S. graminea has solitary capitula, involucre 1.2–2.0 cm in diameter, larger pollen grains, pollen surface with smaller and sparser spines, achenes 3–4 mm long, 28 chromosomes and a karyotype formula 2n = 2x = 28 = 6m + 20sm + 2st. The new species is distributed in Dianzhangou, Awanchang, and Gamaliang mountain regions of Maqu county in Gansu province.     

Myosotis wumengensis sp. nov. (Boraginaceae) from central Yunnan,southwest China

Myosotis wumengensis, a distinct new species endemic to Yunnan Province, China, is described and illustrated. The new species differs from all Myosotis species in China by having exserted anthers and a long corolla tube. Phylogenetic analysis confirms the generic position of the new species. In addition, the pollen morphology of the new species is described.