The embryology and relationships ofPenaeaceae (Myrtales)

Two species ofPenaeaceae (Penaea mucronata andSaltera sarcocolla), a unique South African family ofMyrtales, were investigated embryologically.Penaeaceae clearly agrees with otherMyrtales in its basic embryological characteristics, and further is characterized by its highly specialized features: ephemeral endothecium, 16-nucleatePenaea-type embryo sac, and unique ovular form. A wider range of affinities of families includingPenaeaceae, Oliniaceae, Rhynchocalycaceae, Alzateaceae, andCrypteroniaceae sensu stricto, as well as a possible common divergence from an ancestral line leading toLythraceae and/orMelastomataceae, are discussed on embryological and other grounds.     

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.     

The embryology ofEpipogium roseum (Orchidaceae)

Development of pollen and female gametophyte inEpipogium roseum (D. Don)Lindl. has been investigated. The embryo sac conforms to the Apinagia type. The taxonomic position ofEpipogium within the family is discussed.     

Studies in the embryology of the diandrous orchidCypripedium cordigerum (Cypripedieae,Orchidaceae)

The anther wall layers ofCypripedium cordigerum are six to eight. The glandular tapetum is 2- or 3-layered and its cells are uninucleate. Simultaneous cytokinesis results in decussate, isobilateral and tetrahedral pollen tetrads. Ripe pollen grains are 2-celled. The mature ovules are anatropous, bitegmic and tenuinucellate. Both the integuments are dermal in origin and 2-layered. The inner integument alone forms the micropyle. The female gametophyte is 6-nucleate and bisporic. The reduction of nuclei is due to the strike phenomenon. Double fertilization occurs. The primary endosperm nucleus divides to form two free endosperm nuclei. The mature embryo is undifferentiated. The cells ca, m and n contribute to the embryo. The suspensor is single-celled. 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 these valves consist of parenchymatous cells with starch and raphides. After fertilization, the sterile valves develop sclerotic cells whereas the fertile valves remain parenchymatous. The pericarp structure and embryological features support the retention of tribeCypripedieae within theOrchidaceae.     

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.     

The embryology of angiosperms: Its broad application to the systematic and evolutionary study

Embryology allows one to work with a wide array of characters (more than 50 in general) for each taxon of angiosperms. This paper, while providing a brief review of recent studies on Myrtales and associated families by me and my co-workers, discusses evidence for the general utility of embryological characters for the study of plant systematics. In particular, evidence is given that characters of seed coat anatomy may be best applied to the study of specific and sectional (and even familial) relationships, those of seed appendages as well as of integumentary morphology and histogenesis to the study of generic relationships, and other major characters to the study of familial relationships. Embryology thus provides many features that are complex and, when properly applied along with evidence from other sources, offers good indications of relationships at various taxonomic level, from the ordinal to the specific level. Despite its evident systematic value and increasing need, however, information on embryological characters is still lacking for a majority of genera, and even at the family level, data is lacking or insufficiently available for more than 30% of families. Recipient of the Botanical Society Award for Young Scientists, 1987.     

Studies in the embryology ofHeliantheae (Compositae)

The embryology ofLagascea mollis, Zinnia angustifolia andGalinsoga parviflora has been studied. The anther archesporium is hypodermal and consists of a single row of 6–8 cells, there are two layers below the epidermis of the anther and a periplasmodial tapetum. Ripe pollen grains are tricolpate and 3-celled. The ovary contains a single ovule, but in a few cases ofGalinsoga parviflora two ovules have been found. The female archesporium is unicellular, but sometimes more than one archesporial cell occurs inLagascea mollis andZinnia angustifolia. The embryo sac development is of the Polygonum type, the synergids are hooked, antipodal cells show great variation. The endosperm development is cellular inGalinsoga parviflora and peripheral layer persists in the mature seed. The embryo development conforms to the Senecio variation of the Asterad type. The pericarp structure and cmbryological features support the disputed systematic position ofLagascea withinHeliantheae. Part of a Ph.D. thesis, accepted at Andhra University.     

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.     

The grass megagametophyte and its possible phylogenetic implications

The structure of the grasses megagametophyte is considered to be characteristic enough as to deserve a particular place in the megagametophyte typology. Furthermore, it is compared with those of other Monocotyledonous families to point out embryological affinities.Both are members of the Carrera del Investigador (Conicet, Argentina).     

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.     

Floral structure and relationships ofHortonia (Monimiaceae)

The flowers ofHortonia angustifolia were investigated for their phyllotaxis, morphology, anatomy and development of the perianth, androecium and gynoecium. Certain features were also studied inH. ovalifolia. Characters so far overlooked further support the isolated and intermediate position of the genus between theAtherospermataceae andMonimiaceae s. str. and its archaic position among theLaurales. Dedicated to Professor Dr.W. Leinfellner on the occasion of his 70th birthday.     

Intergeneric relationships ofLolium,Festuca, andVulpia (Poaceae) and their phylogeny

Morphological and seed protein analyses of 26 species of the generaLolium, Festuca andVulpia confirmed their close systematic affinities. Six inflorescence characters readily differentiatedFestuca fromLolium. Protein similarities betweenFestuca of sect.Bovinae and cross-pollinated species ofLolium, coupled with cytogenetic and crossability data, substantiate that they should be united into one genus.Vulpia had phenetic similarities with sect.Scariosae, Montanae andOvinae ofFestuca. Lolium, Festuca, andVulpia are most likely derived from a common ancestral form which was close toFestuca pratensis andLolium perenne.     

The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae)

TOBE, H. & PENG, C.-I, 1990. The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae). We present the first report on the embryology of Bretschneidera , the only genus of Bretschneideraceae (which are one of 15 glucosinolate-producing families), to clarify its relationships. Embryologically Bretschneidera is characterized by the following features: ovule campylotropous, bitegmic and crassinucellate; outer integument thick, multiplicative and vascularized; embryo sac formation of the Allium type; seed exalbuminous; seed coat 'exotestal' with a palisade of columellar, thick-walled exotestal cells; mesotesta thick with the inner half aerenchymatous. These features suggest that Bretschneidera is distinct from any of the taxonomically related families but resembles both Hippocastanaceae and Sapindaceae (Sapindales) more closely than Moringaceae or Capparaceae (Capparales) which have been considered alternative allies, supporting most of the modern taxonomic treatments that place Bretschneidera as a separate family in Sapindales.     

Leaf flavonoid chemistry and the relationships of theLactoridaceae

Leaves of the monotypic angiosperm familyLactoridaceae exhibit flavonoid constituents consisting of six 3-0-diglycosides of the flavonols kaempferol and isorhamnetin. The presence of flavonols is concordant with the placement ofLactoridaceae among the archaic or primitive flowering plants. Flavonoid chemistry is less informative on the relationships of the family within the primitive dicots. The presence of isorhamnetin suggests closer affinities with families in theLaurales, particularly theGomortegaceae andMonimiaceae. Phenetic and cladistic analyses of morphological features place theLactoridaceae near several families in theMagnoliales.     

Species relationships inFestuca (sect.Ovinae,Poaceae)

Morphological and seed protein studies of selected species ofFestuca were performed to elucidate the relationships between species of sect.Ovinae and their affinity with other fescues.Festuca rubra andF. heterophylla (extravaginale group) had higher phenetic affinity with taxa of sect.Scariosae, Montanae andBovinae than with members of their own section. 5 species of the intravaginale group (F. ovina, F. capillata, F. valesiaca, F. sulcata, F. trachyphylla) were fairly uniform and well defined by protein and morphological data. This supports current systematic thinking and does not agree withHackel's (1882) classification of this group.     

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.     

Embryology and relationships ofGyrocarpus andHernandia (Hernandiaceae)

Two representative genera of Hernandiaceae,Gyrocarpus andHernandia, were investigated embryologically to contribute to a better understanding of their respective evolutionary position. Comparisons with other lauralean families using Chloranthaceae or Annonaceae (as a representative of Magnoliales) as an outgroup of Laurales (if present, plus other related taxa) support a lauraceous affinity for the two genera because of the presence of ramified raphal vascular bundles at the chalaza (a synapomorphy), but do not provide evidence for the separation of Hernandiaceae from Lauraceae.Hernandia rather shares with Lauraceae two apomorphies (i.e., the seed pachychalazy and the ruminate seed) which may be homoplasies judged from results of cladistic and molecular studies published elsewhere.Hernandia is greatly divergent from an ancestral line common withGyrocarpus and is even diversified within the genus. Based on evidence from embryology as well as from other sources, it seems best to accept two separate subfamilies in Hernandiaceae as usually have been accepted: one is a derived subfamily Hernandioideae, and the other a less specialized Gyrocarpoideae.     

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.