Embryology and systematic position ofLiparis (Orchidaceae)

The embryology ofLiparis paradoxa andL. rostrata has been studied. The young anther wall consists of an epidermis, endothecium, three middle layers and secretory tapetum with uninucleate cells. In the mature anther, two or three sub-epidermal layers develop fibrous thickenings. The anther wall development corresponds to the massive type. Simultaneous cytokinesis results in decussate, isobilateral, linear, T-shaped and tetrahedral pollen tetrads. The pollinia are compact and at anthesis the pollen grains are 2-celled. Ovules are anatropous, bitegmic and tenuinucellate. Both the integuments are dermal in origin and 2-layered. Development of female gametophyte is of the monosporic, 8-nucleate type. Double fertilization occurs. The primary endosperm nucleus degenerates. The mature embryo is organized from the derivatives of tiers 1, 1, m, and n. Its development conforms to a variation of the Onagrad type. The seed coat is formed entirely by the outer layer of the outer integument. There are three sterile and three fertile valves in the ovary. In the prefertilization stages valves consist of parenchymatous cells. After fertilization, the sterile valves become sclerenchymatous whereas the fertile valves remain parenchymatous. The embryological characters support the disputed systematic position ofLiparis within subtribeLiparidinae ofEpidendreae.     

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.     

The genusCarex (Cyperaceae) in the Flora Iranica area

88 species and 20 subspecies ofCarex, altogether 94 taxa, occur in the area ofRechinger's Flora Iranica. Of these, 48 taxa, have an Euro-Siberian, 11 an Irano-Turanian and 35 a Central Asiatic distribution, the last including five species with connections to E. and tropical SE. Asia. The Saharo-Sindian element is missing. Endemic taxa are found in all groups. Eight taxa, endemic to the NW. Himalayas occur in the E. part of the area. The Euro-Siberian taxa are concentrated in the NW. of the area, but have connections to the east; similarly, Central-Asiatic taxa are concentrated in E. Afghanistan and N. Pakistan, but have connections to the west. Some Euro-Siberian taxa are widely distributed in the area, but avoid the most arid regions. The Irano-Turanian taxa are few and extend from Egypt through the area of Flora Iranica to Kazakhstan SSR and Central China.Dedicated to Hofrat Univ.-Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

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.     

Genetic variation and hybridization in SwedishSchoenus (Cyperaceae)

Schoenus ferrugineus andS. nigricans have restricted distributions in Sweden and are almost exclusively confined to calcareous fen habitats. AtS. nigricans sites,S. ferrugineus is usually also present, and hybrids are frequently found. In this report, I used allozymes to estimate the amount of gene flow between the two species, and to compare the partitioning of genetic diversity in each of them. Thirteen loci were analysed at eight different enzyme systems. Seven loci were variable between or within the species. The two species had completely different alleles at two of the seven variable loci, whereas there was overlap at five loci. In all, 22 different alleles were found. Six of these alleles were confined toS. nigricans, and five alleles were confined toS. ferrugineus. Nei's genetic identity was 0.55.—InS. ferrugineus, three loci (23%) were polymorphic, and the average number of alleles per polymorphic locus was 2.0 (each polymorphic locus had two alleles). InS. nigricans, three loci (23%) were polymorphic, and the average number of alleles per polymorphic locus was 2.3.—The proportion of genetic diversity due to variation among sites (G _ST) was fairly similar in the two species, mean over loci = 0.12 inS. ferrugineus and 0.15 inS. nigricans. However, the proportion of genetic diversity due to variation among individuals within sites (G _IS) differed markedly between the two species, mean over loci = 0.54 inS. ferrugineus and 0.17 inS. nigricans. Accordingly, there was a much higher individual heterozygosity inS. nigricans than inS. ferrugineus. — Most hybrids were interpreted as F₁ hybrids. However, a small proportion, 0.5–1.6 %, were F_n hybrids or back-crosses.—On the Swedish mainland, all former occurrences ofS. nigricans are extinct, but viable hybrids are still present at a few sites in southernmost Sweden.     

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.     

Embryology ofCrocus thomasii (Iridaceae)

The embryology ofCrocus thomasii is described. Male meiosis is of simultaneous type, and gives rise to starchy microspores which develop into lipoid pollen grains; these are two-celled and show a spinulate acolpate, abaculate exine lacking apertures. The tapetum is glandular and its cells become bi- or sometimes multinucleate. The ovule is anatropous and bitegmic; the inner integument forms the micropyle. Megasporogenesis is heteropolar with starch accumulation in the functional chalazal megaspore. Embryo sac development conforms to thePolygonum type. The endosperm development is nuclear. The embryo develops according to the Caryophyllad type. In the ripe seed it is differentiated and enveloped by a starchy cellular endosperm. The embryological characters observed strongly favour a close relation betweenC. thomasii andC. sativus.     

The systematic position of theHolarrheninae (Apocynaceae)

The genusHolarrhena, described byRobert Brown in 1811, has had a problematic taxonomic history, in part due to a suite of characters that does not conform with accepted concepts within theApocynaceae. In a number of important taxonomic charactersHolarrhena is typical of subfam.Apocynoideae. But due to the relatively unspecialized structure of the anthers most recent authors have placedHolarrheng, together withCarruthersia andSpirolobium, as the subtribeHolarrheninae in subfam.Plumerioideae. For the present investigation the floral structure and pollen morphology ofHolarrhena, Carruthersia andSpirolobium were analyzed. From the chemical literature reports of the occurrence of steroidal alkaloids in thePlumerioideae were evaluated. Our results indicate that the three genera belong to subfam.Apocynoideae in the tribeNerieae, but that the Holarrheninae is an unnatural group, and that the three genera should be accommodated individually within the tribe.     

Relationships among species inCarex sect.Phyllostachys (Cyperaceae) based on allozyme divergence

Allozyme variation was examined inCarex sect.Phyllostachys (Cyperaceae) to provide insight into phylogenetic relationships hypothesized in an earlier study and to determine the degree of genetic differentiation within and between taxa. Genetic identity values are concordant with the morphological differences found between species. The lowest values are found between species with the greatest morphological dissimilarity. Conversely, the highest values are associated with species pairs distinguished by relatively few morphological differences. Conspecific populations possess high genetic identities, although interpopulation differentiation has characterized the evolutionary history of some species. Geographic patterning is also evident within species, with geographically proximate populations often having the highest identity values. Phylogenetic trees produced using different cladistic methods were poorly supported and varied in their depiction of relationships among species. One cladogram produced using presence/absence allelic data is more or less congruent with a topology recovered from an earlier analysis utilizing molecular and morphological data. The wide- and narrow-scaled clades are maintained as are the sister species pairsC. backii/C. saximontana, C. basiantha/C. superata, andC. jamesii/C. juniperorum. Contrary to the finding of our previous study, however,C. willdenowii is aligned withC. jamesii/C. juniperorum.     

The impact of species-specific traits and phylogenetic relatedness on allozyme diversity inCarex sect.Phyllostachys (Cyperaceae)

Allozyme variation was examined inCarex sect.Phyllostachys (Cyperaceae) to study the effects of species-specific traits and phylogenetic relatedness on genetic structure. In contrast to the findings of similar studies, genetic variability in thePhyllostachys is poorly correlated with geographic range and putative differences in breeding systems (as inferred from morphology). This suggests that other patterns of evolution, colonization, and gene flow characterize the species found in this section. Fixation indices are negative for all populations suggesting that mechanisms such as disassortative mating and selection are maintaining heterozygous excess within populations. Closely related taxa often exhibit different genetic variability statistics. In some instances, however, clades (e.g.C. jamesii andC. juniperorum) display very similar levels of genetic variability despite marked differences in species-specific traits. Recent speciation coupled with the ability to maintain historical levels of variation within populations may be factors accounting for this phenomenon. Contrary to similar studies, species restricted to known glacial refugia have lower genetic diversity than those species that underwent mass migrations in response to deglaciation. Narrowly endemic species were found to partition their genetic diversity within, as opposed to between populations. The opposite trend was evident in wider ranging congeners.     

Agmatoploidy inCarex laevigata (Cyperaceae). Fusion and fission of chromosomes as the mechanism of cytogenetic evolution in Iberian populations

In this paper cytogenetic studies on 64 specimens from 20 Iberian populations ofCarex laevigata (Cyperaceae) are presented. Chromosome behaviour in meiosis suggests that the different chromosome numbers obtained (ranging from 2n = 69 to 2n = 80) were distributed according to an increasing geographic gradient of chromosome fission along the North South direction. Four relatively stable areas were also delimited according to chromosome numbers displayed by this species, i.e. 2n = c. 72, c. 74, c. 76, and 78. The meiotic behaviour ofCarex ×deserta (C. laevigata ×C. binervis) was also studied.     

The embryology ofStegnospermataceae,with a discussion on its status,affinities and systematic position

The embryology ofStegnosperma halimifolium andS. watsonii has been studied in detail. The tapetum is of the secretory type and its cells become multinucleate. Simultaneous cytokinesis in the pollen mother cells follows meiosis. The ripe pollen grains are 3-celled. The ovule is crassinucellate, bitegmic and amphitropous, with the micropyle formed by the inner integument alone. The female archesporium is one celled, and the parietal tissue 3–5 layered. The embryo sac development conforms to thePolygonum type. A central strand, 6 or 7 cells thick, differentiates inside the nucellus and extends from the base of the embryo sac to the chalazal region. The endosperm is nuclear. The embryogeny conforms to the Caryophyllad type. The seed coat is formed by the outer epidermis of the outer integument and the inner epidermis of the inner integument. Based on this evidence and other data, the status of the genus as an independent family,Stegnospermataceae (Stegnospermaceae) is confirmed. Apparently, it forms a connecting link betweenPhytolaccaceae andCaryophyllaceae.     

Embryogeny in someCypereae

The paper presents a comparative account of embryo development inPycreus pumilus Nees.,Cyperus alternifolius Willd.,Mariscus paniceus Vahl. andRemirea maritima Abul., belonging to the tribeCypereae. Embryogeny follows the Juncus variation of the Onagrad type or conforms to Period I, Series A₂, Megarchetype IV. InPycreus andRemirea the precocious differentiation of dermatogen initials is followed by that of plerome and periblem elements. Mature embryos ofPycreus, Mariscus, andRemirea are identical withCyperus. Embryogeny and structure of mature embryo support the fusion of these taxa withCyperus. Embryogeny in SomeCyperaceae 1.     

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.     

火炭母的一个新异名及其系统位置初探

该文将铺地火炭母(Polygonum chinense L. var. procumbens Z. E. Zhao et J. R. Zhao)处理为火炭母原变种(Persicaria chinensis var. chinensis)的异名,并讨论了火炭母在命名上的问题,及其系统学位置。     

Embryology ofCordia sebestena (Boraginaceae)

Sporogeneses and gametogeneses inCordia sebestena are described. Anther wall develops according to the Dicotyledonous type. Pollen grains are three-celled at the time of dehiscence. The occurrence of enlarged, precociously developing, densely cytoplasmic cells toward the proximal end of the anther and between adjacent pollen sacs is a unique feature in this species. The possible functions of these cells are discussed. A placental obturator develops simultaneously with the ovules. The dual origin of parietal tissue (from nucellar epidermis and the archesporial initial) is an unusual feature in this species. Megagametophyte development is monosporic or (occasionally) bisporic. Megaspore tetrads are linear or T-shaped. Double dyads and tetrads are frequently observed.—Maintainance of subfamily status forCordioideae is supported.     

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.