Embryology ofEriocaulon setaceum (Eriocaulaceae)

Eriocaulon setaceum can be characterized by: young microsporangium wall with epidermis, endothecium (with fibrous thickenings), and glandular tapetum (uninucleate cells); pollen grains 3-celled, spiraperturate; embryo sac development according to the Polygonum type and with antipodal cyst; endosperm nuclear; embryo small, with incipient differentiation into cotyledonary and epicotylary loci; seed coat mainly from the inner layers of the integuments; pericarp 2-layered and membranous. Embryologically, theEriocaulaceae are nearer to theXyridaceae than to otherFarinosae. Their elevation to the rank of an order,Eriocaulales, therefore appears justified.     

Seed structure and systematic position ofHampea nutricia (Malvaceae)

The mature seeds ofHampea nutricia are glabrous, ovoid, arillate and dark tan in colour. Longitudinal streaks on the seed surface correspond to the underlying integumentary vascular strands. Testa and tegmen are derived from the outer and inner integuments, respectively. The outer epidermis of the tegmen forms a palisade-like macrosclereid layer, the inner epidermis a fringe layer. The endosperm is single-layered and also fills the space between the two cotyledons. The embryo is nearly straight, gland-dotted; it has asymmetrical and folded cotyledons, and gossypol ducts. Systematic position ofHampea is discussed and its placement inMalvaceae is supported.     

Embryology ofKyllinga monocephala (Cyperaceae) and its systematic position

Aspects of the life history ofKyllinga monocephala are described. Anther wall development corresponds to the Monocot type. The endothecium shows spiral thickenings. The tapetum is glandular and has uninucleate cells. Ubisch granules are present. Mature pollen grains (pseudomonads) are 3-celled at maturity. Ovules are bitegmic, crassinucellate and develop a funicular obturator. The embryo development conforms to theJuncus-variation of the Onagrad type. Endosperm, seed coat and pericarp are described.     

The systematic position ofAsplenium cardiophyllum (Aspleniaceae)

Asplenium cardiophyllum is a morphologically unusual species with simple leaves and anastomosing venation, and is often placed in the segregate genusBoniniella. To determine its systematic position, character comparisons were made of vascular anatomy, raphides in leaf epidermis, chromosome number and perispore of this species and those ofAsplenium sect.Hymenasplenium. Asplenium cardiophyllum conforms with sect.Hymenasplenium in its dorsiventral dictyostele, the presence of raphides, a chromosome number of 2n=156 (x=39), and lophate peristore with spinulate projections on the lumina. We therefore propose to includeA. cardiophyllum in that section. Dedicated to the memory of the late Professor Kunio Mitui.     

Embryology ofMalaxis saprophyta,with comments on the systematic position ofMalaxis (Orchidaceae)

InMalaxis saprophyta, anther wall development corresponds to the Monocotyledonous type. The uninucleate tapetum is of secretory type and the endothecium develops U- and V-shaped thickenings on the inner tangential and radial walls. Cytokinesis is simultaneous; tetrahedral, isobilateral and T-shaped tetrads are formed which are compactly aggregated in pollinia. At anthesis the microspore tetrads are 2-celled. The ovule is anatropous, bitegmic and both integuments are dermal in origin. A single hypodermal cell develops directly into a megaspore mother cell. Embryo sac development is predominantly monosporic and less often bisporic. Irrespective of the type of development, the mature embryo sac is 6-nucleate. Although double fertilization occurs, the primary endosperm nucleus degenerates. Embryogeny is of the Onagrad type. The mature embryo lacks differentiation into cotyledon, plumule and radicle. The reticulate seed coat is formed entirely by the outer layer of outer integument. There are three sterile and three fertile valves in the ovary. Although initially parenchymatous, the entire three sterile valves in the ovary and the upper half of the three fertile valves become sclerified after fertilization. The embryological characters support the disputed systematic position ofMalaxis within subtribeMalaxidinae ofEpidendreae.     

Floral anatomy ofHypseocharis (Oxalidaceae) with a discussion on its systematic position

The floral anatomy of threeHypseocharis spp. has been studied. The genus resemblesOxalidaceae as well asMonsonia andSarcocaulon of theGeraniaceae. As it is closer toGeraniaceae than toOxalidaceae, it perhaps serves as a connecting link between them.     

On the morphological position of Paepalanthus subgenus Psilandra (Eriocaulaceae)

Paepalanthus subgen. Psilandra comprises two species known only from the type collections. The aim of this study is to evaluate their unique floral morphology within a given phylogenetic context and its implications for taxonomy. Flowers of both species were analyzed, described, and photographed using a digital microscope. Taxonomically relevant characters were mapped on previous molecular phylogenies. The flowers of both species fit the Paepalanthoideae pattern and are quite similar to each other. The petals of the pistillate flowers, although free, show evidence of previous fusion in the middle. The free condition is interpreted to be secondary, probably because of the handling of the specimens or to the current developmental stage. Character mapping indicates that both species share floral features with Syngonanthus but not with Paepalanthus. Therefore, based on the median fusion of the pistillate flower petals, absence of floral bracts, petals of the pistillate flower being shorter than the sepals, simple stigmas in the commissural position, the pedicellate pistillate flower, and the glabrous petals and sepals, we transfer P. subgen. Psilandra to synonymy of Syngonanthus sect. Syngonanthus and combine both species in this genus.     

Female reproductive organization and systematic position ofPlumariella yoshikawae Okamura (Ceramiaceae,Rhodophyta)

A detailed account of female reproductive features ofPlumariella yoshikawae is presented.Plumariella yoshikawae is recognized as morphologically similar toDelesseriopsis elegans in several features, particularly in the sequence of branch initiation, the presence of gland cells on the abaxial sides of the basal segments of the lateral branches, having carpogonial branches on the basal segments of unmodified lateral branches, and the maturation of the carposporophyte at well below the thallus apices. These features indicate thatPlumariella yoshikawae is best removed from the Ptiloteae and is correctly placed in the tribe Delesseriopsideae. The generaPlumariella andBalliella are retained along withDelesseriopsis within the tribe Delesseriopsideae.     

On the taxonomic position ofSilene thebana (Caryophyllaceae)

Silene thebana Orph. exBoiss. is transferred as a subspecies toS. fabaria (L.)Sm., resulting in the new combinationS. fabaria (L.)Sm. subsp.thebana (Orph. exBoiss.)Melzh. The chromosome count of 2n = 24 is recorded for the first time for this taxon.Dedicated to Prof. DrK. H. Rechinger on the occasion of his 80th birthday, 16th October 1986.     

The systematic significance of surface features of theBalanophora tuber (Balanophoraceae)

The surface ofBalanophora tubers consists of a nonepidermal layer made up of two distinctive types of cells, armature cells and stellate wart cells. Both cell types are provided with a heavy wall, and are dead at maturity. Stellate warts in the three species investigated,B. elongata, B. fungosa, andB. hansenii, séem to be uniform in appearance, but armature cells are extremely distinctive for each species. They are present in large, agglomerate masses in the first, singly or in very small clusters in the second, and as completely free, individual, acicular cells in the third species. Such differences are believed to be significant systematically, and the separation ofB. hansenii is thus probably justified. Notwithstanding superficial similarities, stellate warts do not seem to be comparable to the lenticels of other plants.     

Evolution and systematic relationships in theTriticeae (Poaceae)

The evolution and taxonomic relationships in theTriticeae are discussed with the view to highlight aspects of this agronomically important group of plants, which may be of interest to molecular biology. Some of these aspects are addressed in more detail in adjoining papers in which specific genomic loci have been examined at the DNA sequence or isozyme level. Aspects discussed include the systematics and geographic distribution of theTriticeae species, isozyme and chromosome pairing studies on some of the species as well as more recent developments in DNA analyses. A survey of the systematics of theTriticeae indicated that the genomic system ofLöve is probably the most useful starting point for interpreting molecular data even though the system has many problems from a taxonomic point of view. The geographical distribution ofTriticeae species, using both published and unpublished data, suggested that information of this type taken together with the theory of continental drift provides a broad time-span for considering data from DNA sequence studies. The significance, and modes of analyses, of isozyme studies were assessed because they often provide valuable characters in determining relationships between species. The main character underlyingLöve's andDewey's analyses of theTriticeae, namely chromosome pairing, is discussed with particular reference to isozyme studies to show that in some cases, such as species ofHordeum sensu lato, consistent relationships are obtained. Finally, new developments in understanding chromosome structure are considered in relation to the above variables in the taxonomy and evolution of theTriticeae.     

The taxonomy and morphological and anatomical differentiation of populations ofLeiothrix crassifolia (Eriocaulaceae)

The morphological and anatomical leaf and scape differentiation ofLeiothrix crassifolia is analysed and classified into seven types (I, II, III, IV, V, VI, VII). The species has a relatively wide distribution in the campo rupestre of Minas Gerais in Brazil. Variation in the isolated populations on different mountain ranges is complex, does not follow a clear geographical pattern, and defies taxonomic classification.L. nubigena andL. obtusifolia are included inL. crassifolia.     

A new infrageneric classification and phylogenetic trends in the genusRhododendron (Ericaceae)

Recent studies have improved the infrageneric classification ofRhododendron, including my own investigations on flavonoids and anthocyanins as chemosystematic markers. From a synoptical comparison of morphological, anatomical and phytochemical characters a new system for the genus is proposed. Phylogenetic character progressions and relationships among subgenera, sections and subsections are discussed and illustrated. Key positions for subg.Candidastrum between chori subgenerumRhododendron andNomazalea, and for subg.Choniastrum between chori subgenerumHymenanthes andNomazalea are suggested.     

Biosystematics of two sympatric species ofEucharis (Amaryllidaceae)

Eucharis candida andE. formosa are two often sympatric species of bulbous geophytes restricted to neotropical rain forest understory. The species are most common in eastern Ecuador, and are the only two east Ecuadorean species of the genus found north of the Pastaza valley. Data from phenetic, karyotypic, and preliminary isozyme electrophoretic analyses of both species are represented. The species are distinguishable phenetically and karyologically, but isozyme-based relationships are more complex. Phenetic resolution of the isozyme phenotypes supports recognition of two species in Ecuador. A Peruvian isolate ofE. formosa, though not morphologically distinct, shows both allozyme and chromosomal divergence from Ecuadorean populations. Cladistic relationships based on overall allozyme data do not support species distinction, but a novel electrophoretic phenotype for glutathione reductase is shared only by individuals ofE. candida. An apparent geographic component within the monophyletic groups resolved in the cladogram suggests that some degree of gene flow between these two species has been maintained without the complete loss of morpholgoical species identity. This may have been mediated either by artificial population structures due to a probable long history of cultivation, or via Pleistocene refugia effects. Both species may have originated in eastern Ecuador from a common ancestral population which has since radiated outward, perhaps several times.     

A search for the phylogenetic position of the seven-son flower (Heptacodium, Dipsacales): Combining molecular and morphological evidence

A first report on the problematic phylogenetic position ofHeptacodium (2 spp.; China) using molecular data from chloroplast DNA is presented. Amplification of ORF2280 homolog region was executed in a number of representative taxa in order to determine ifHeptacodium shows similar structural rearrangements as other Dipsacales. DNA sequences ofndhF were generated to clarify the phylogenetic position ofHeptacodium among Caprifoliaceae (s.l.). Six outgroup taxa and fifteen representatives of Dipsacales were sampled and more than 2100 basepairs ofndhF sequence were used in a cladistic analysis. Parsimony analysis produced two shortest trees and showedHeptacodium as sister to all members of Caprifoliaceae (s.str.), although weakly supported. Additionally, trees were constructed withndhF data supplemented with availablerbcL sequences and a morphological data set. Results of all analyses support an unresolved basal position forHeptacodium among Caprifoliaceae (s.l.), which in part explains the difficulty experienced previously in classifying the genus.     

Sporoderm characters ofTetrameranthus duckei (Annonaceae) and their systematic implications

Ultrastructural and micromorphological investigations of the sporoderm ofTetrameranthus duckei pollen confirm the interpretation of this genus as being isolated within theAnnonaceae.     

Taxonomic and morphological studies inHypecoum sect.Hypecoum (Papaveraceae)

Hypecoum sect.Hypecoum comprises 8 species, one with 2 additional subspecies, in the Mediterranean area and the Middle East. Five of these are recognized for the first time, viz.Hypecoum procumbens L. subsp.fragrantissimum Å. E. Dahl,H. procumbens L. subsp.atropunctatum Å. E. Dahl,H. trullatum Å. E. Dahl,H. angustilobum Å. E. Dahl andH. torulosum Å. E. Dahl. H. dimidiatum Delile andH. pseudograndiflorum Petr., which have generally been included inH. imberbe Sm., are reestablished as distinct species. Morphology and variation patterns are described and discussed with special reference to mating systems. Self-incompatibility is dominating butH. procumbens subsp.procumbens andH. torulosum are self-compatible. The broad variation ranges of the former taxon in traits presumably related to mating system (petal, anther and stigma sizes as well as pollenovule ratios) indicate varying rates of outcrossing. SEM micrographs are given of petal and stigma surfaces and of pollen grains. Chromosome counts are presented for seven taxa. All are diploid with 2n = 16.     

Aspects of floral structure and phenology in the genusConophytum (Mesembryanthemaceae)

The complex floral structure in the southern African genusConophytum (Mesembryanthemaceae; 77 spp.) is described in detail and assigned to three basic floral types, two of which can be divided into two subtypes. Correlations between structural features and phenological patterns are demonstrated and discussed in the family context as well as in relation to the systematic subdivision of the genus.     

Studies on systematic embryology inScilla (Hyacinthaceae)

Scilla persica and 5 species of the so-calledS. hohenackeri group, namely,S. furseorum, S. puschkinioides, S. vvedenskyi, S. hohenackeri, andS. greilhuberi, have been investigated embryologically with special reference to embryo sac and endosperm development.Polygonum-type embryo sac development was stated inS. puschkinioides andS. greilhuberi. 8-nucleate, normally structured embryo sacs, which could not be specified further due to sparse availability of the material, were stated inS. furseorum, S. vvedenskyi, andS. hohenackeri. InS. persica the embryo sac develops according to the bisporicAllium-type. In most species endosperm development was stated to be nuclear, exceptS. hohenackeri, where the type could not be specified. Other traits of possible taxonomic significance are the number of layers in the outer integument, which is mostly 4, or 5–6 inS. furseorum, and the occurrence of polyploid versus haploid and early degenerating antipodal nuclei, the latter occurring only inS. persica andS. furseorum. These embryological characters may be useful for assessing taxonomic relationship of the present species with other allied groups withinScilla, in particular, theS. siberica alliance,S. messeniaca, and theS. bifolia alliance. TheAllium-type embryo sac, which occurs inS. persica, is also characteristic for theS. siberica alliance, and may be a common derived character. Lack of antipodal polyploidization, as characteristic forS. persica andS. furseorum, occurs also in theS. siberica alliance, and is perhaps another common derived trait indicating phylogenetic relationship. Nuclear endosperm development is more frequent in spring-flowering squills than helobial development, which has previously been stated inS. messeniaca, some species of theS. siberica alliance, and inS. litardierei. While helobial endosperm may be primitive forHyacinthaceae in general, it may, by reversal, also occur as a derived character, at least in some species of theS. siberica alliance.