A CONTRIBUTION TO THE EARLY TERTIARY FOSSIL RECORD OF THE SALVINIACEAE

Megaspores and massulae of Azolla, Azollopsis, and Salvinia are recorded from the Paleocene and Eocene of Montana and the Dakotas. Three new species of Azolla are described and two species of Azolla are redescribed. The columella of the floating apparatus of the megaspore apparatuses of Azolla appears to be a phylogenetic localization of the perispore around the megaspore. A new section of Azolla (section Kremastospora) is created for species whose megaspore apparatuses have many floats and whose massulae have hooked (anchor-shaped) glochidia. Salvinia preauriculata is based only on leaf remains from the Eocene of North America. Megaspores and massulae which probably are conspecific with the leaf remains are described. The megaspores and massulae of the fossil species are much smaller than those of the living species, S. auriculata, with which S. preauriculata has been compared. The similar size of the megaspores and massulae of the fossil species suggests a less specialized condition than in living species, where these two structures differ greatly in size. Azollopsis tomentosa, previously known from the late Cretaceous, has been found in the Eocene.     

EM-studies on Salviniaceae Megaspores from the Middle Miocene Fasterholt Flora,Denmark

Reproductive organs of Salvinia and Azolla were found together with other small plant remains in the Middle Miocene Fasterholt flora, Denmark. They were examined by the use of SEM and TEM; this was done with the three species of Azolla for the first time. Salvinia is represented by one, or possibly two, species. The megaspore of S. cerebrata has a characteristic surface sculpture of irregular, rounded ridges. Some specimens differ from the typical S. cerebrata by having strongly developed verrucae or clavae. They are described as Salvinia sp., but it is possible that they represent a variety of S. cerebrata. Azolla is represented by three species. A. nikitinii and A. ventricosa have nine floats in two tiers; massulae have not been found. A. tuganensis has three floats in one tier, massulae have anchor-shaped glochidia.     

The role of Hydropteris pinnata gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns

Large segments of intact plants that represent a heterosporous fern have been discovered within an aquatic plant community from the Late Cretaceous St. Mary River Formation near Cardston in southern Alberta, Canada. Branching rhizomes of Hydropteris pinnata gen. et sp. nov. are 1–2 mm wide. They produce fronds at intervals of 2–12 mm and bear numerous elongated roots. Fronds, up to approximately 6 cm long, are pinnate with subopposite to alternate pinnae that exhibit anastomosing venation. Large, multisoral sporocarps occur at the junctures of the rhizome and frond rachides. Both microsporangiate massulae and megaspore complexes occur within each sporocarp. Megaspore complexes are assignable to the sporae dispersae genus Parazolla Hall. Microspores are trilete, smooth-walled, and are embedded in episporal material of the massulae. A numerical cladistic analysis indicates that the heterosporous aquatic ferns are monophyletic, and not as closely related to either schizaeaceous or hymenophyllaceous ferns as they are to some other filicaleans. Systematic revisions are proposed to reflect newly recognized cladistic relationships within the heterosporous clade, and character originations in the evolution of heterosporous aquatic ferns are evaluated. Hydropteridaceae fam. nov. is proposed, and included with Salviniaceae and Azollaceae in the Hydropteridineae subord. nov., and the Hydropteridales Willdenow.     

Palynological evidence of Azolla nilotica Dec. in recent Holocene of the eastern Nile Delta and palaeoenvironment

Megaspores, microspores and massulae of the free-floating fern, Azolla nilotica, were found in Late Holocene sediments obtained by coring in the eastern Nile Delta. Nowadays, the nearest station for this fern is southern Sudan. The determination of the species is based on spiny projections on the megaspore body and on the verrucate microspores. Palynological studies reveal that the habitat of the fern consisted of extensive papyrus marshes, now disappeared. Several causes for the disappearance of the fern from the Nile Delta are proposed amongst which the most probable is human influence which has completely modified the vegetation and the hydrology.     

Taxonomic re-evaluation of the Azolla genus in Portugal

ABSTRACT

The Portuguese and the European Flora refer to the presence of two or three Azolla species in Portugal: A. filiculoides Lam., A. caroliniana Willd. and/or Azolla mexicana Presl., the latter included in the last edition of Flora Europaea. In the present work, the taxonomy of Azolla species is reviewed using the two most important characters that can distinguish between these two/three species: papillae in the dorsal leaf lobe and perine architecture of the megaspore apparatus. Other characteristics, such as the hyaline border cells of the dorsal leaf lobes and the number of glochidia septa in microsporangium massulae, are also used. All the Azolla specimens, collected from several locations in Portugal, were identified as Azolla filiculoides Lam. This identification disagrees with previous published reports on Azolla taxonomy in Portugal as well as with herbarium identification.     

Ontogeny of the Calliandra – massulae (Mimosaceae: Ingeae), and the associated viscin body

Polyads, called also massulae, are common in the Mimosaceae. Within this family, only in Calliandra (s. l.) polyads show a remarkable morphological variation: 16-grained lens-shaped massulae in the neotropical Zapoteca (syn. Calliandra p.p.), and highly asymmetric eight-grained massulae in Calliandra (s. str.). As a further specialisation the massulae in Calliandra (s. str.) possess a sticky appendage called a viscin body. The form and ontogeny of this unique two-dimensional construction of the massula is of particular interest. In Calliandra (s. str.) the octad is normally the developmental product of the initiation stage, beginning with one pollen mother cell (PMC). In total, Calliandra (s. str.) possesses eight PMCs in each anther.The dicotyledons are characterised by the simultaneous type of cytokinesis, but wall formation in Calliandra (s. str.) during meiotic division is successive, as common in monocotyledons. The first wall divides the PMC diagonally, and during the tetrad period, two clear variants of tetrads are found. One shows the typical rhomboidal arrangement and the other forms two congruent symmetrical halves. The latter probably results from early fixed, quite inflexible walls. The cells of tetrad do not separate, and four successive mitotic divisions give rise to the eight-celled massula. The direction of growth of the PMC and the massula, the sequence of division, and the position of the walls all show a strict polarity. The normal number of eight cells in each massula is sometimes exceeded without loss of viability.Calliandra (s. str.) lacks a developed “Pollenkitt”, and its asymmetrical massula is combined with the viscin body, which is involved in pollination. This viscin body is located on a specialised pollen grain at the top of the massula. With the viscin body, Calliandra (s. str.) massulae have a mechanism for immediate and persistent attachment to different surfaces. SEM is used to understand the mechanisms of attachment in detail. Furthermore, the relation between pollinators and the massula shape is investigated. Massula transport and the role of the viscin body in pollen–stigma interaction are discussed as well. The stigma of Calliandra (s. str.) is papillate and in the receptive stage, it is characterised by acquisition of a copious exudate.Zapoteca (syn. Calliandra p.p.) massulae could be distinguished histochemically from those of Calliandra (s. str.) by the main components of the “Pollenkitt”. Histochemical and physical properties of the viscin body suggest a waxy nature, including lipids in combination with unsaturated fatty acids. Proteins are present in less amounts and polysaccharides and polyglucans are absent. These results are discussed in relation to analogous phenomena in other families.The term “apex cell” is introduced to indicate the specialisation of the tip-cell of the massula by ontogeny and polarity.     

The Micro-and Mega-Sporogenesis and the Formation of Male and Female Gametophytes in Gastrodiadata Blume

The present paper deals with the morphogenesis of the micromegaspore and the formation of male and female gametophytes in Gastrodia elata Blume. The embryo sac is originated from the chalazal megaspore of a triad. In the mature embryo sac, the egg apparatus and the single polar nucleus are derived from the four microphylar nuclei. These two chalazal once are disappered before the maturation of the embryo sac. It is a simplified form of the Polygonum type. The origin and the formation of the male gametophyte in Gastrodia elata are similar to that of the Orchidaceous in general. The tetrads of each anther are adhered and become into massulae. The extent range of the adherent are gcnerally from hundreds to more or less one thousand. A preliminary observation are revealed by the writer under the fluorescence microscope relating to the variation of the components of their cell walls during the different stage of the sporogenesis as well as the formation of the male gametophyte. In this papre, the signifieancc of the existence of pollinia substanee arounding the massulae as well as the temporal occurance callour of the generative cell have been disseussed.     

Some structures in Azolla megaspores,and an anomalous form

On the assumption that the “float” apparatus of Azolla arose by the “capture” and retention of separate floats developed as massulae round abortive megaspores, a model is proposed, accounting hypothetically for the separate origin of three-floated, nine-floated, fifteen-floated and twenty-four-floated species in different types of cytological behaviour during origin of the functional megaspore. It is considered irrelevant to ask whether nine-floated arose from three-floated or vice versa. Each type arose de novo and possibly more than once, out of the initial cytological situation which is constant for all species.An abnormal form of Azolla imbricata is described, which in terms of the model appears to represent a reversion to a Salvinia-like ancestral form, by loss of linked genes responsible for the Azolla “syndrome”.Anatomical and morphological studies on five modern and two Paleocene species are used to illustrate adaptation of perine as float-retaining mechanisms. A new structure, the manicula, produced from the perispore, is described in A. teschiana and A. velus.     

The pollinium ofLoroglossum hircinum (Orchidaceae) between pollination and pollen tube emission

The structure of the massulae composing the pollinium ofLoroglossum hircinum was studied before pollination and 12 and 24 hours afterwards. The grains are grouped in tetrads closely packed in massulae. The exine is only present on the outside of the massulae. The intine consists of two layers: a compact layer surrounding the pollen grain and a looser layer surrounding the pollen grain and a looser layer surrounding the tetrad. Twelve hours after pollination, pollen volume and the space between the tetrads increase due to vacuolization. Twenty-four hours after pollination, pollen volume and tetrad spacing are higher due to vacuolization and some grains have emitted pollen tubes. Pollen growth due to vacuole formation, and the absence of common walls between adjacent tetrads lead to crumbling of the massulae. The mature pollen grain does not have apertures: the site of pollen tube emission is determined after pollination. The first grains to germinate are those in the centre of the massula. The vegetative cell nucleus is the first to enter the pollen tube; the generative cell elongates and undergoes the second haploid mitosis shortly after entering the pollen tube.     

STUDIES ON FOSSIL AZOLLA: AZOLLA MONTANA,A CRETACEOUS MEGASPORE WITH MANY SMALL FLOATS

Azolla montana is a new species from the late Cretaceous; it has a megaspore apparatus with a single large and conspicuous columella which resembles the floats of the megaspore apparatus of many extant species. The columella differs from true floats in being pilose, like the perispore of the megaspore body. The 10–20 small true floats are appressed to the columella, and are difficult to distinguish from it. It is suggested that the floats were derived, phyletically, by segmentation of the columella; the columella represents the primitive “float” of the megaspore apparatus.     

STUDIES ON FOSSIL AZOLLA: PRIMITIVE TYPES OF MEGASPORES AND MASSULAE FROM THE CRETACEOUS

Fossil Salviniaceae are described from the Claggett Shale and Judith River Formation, late Cretaceous (Campanian stage) of Montana. A new genus, Parazolla, from the Claggett Shale, has megaspores in which the swimming apparatus is composed of a number of elongate floats attached to the megaspore body and invested by coiled hairs. The floats separate at maturity. Massulae (bearing microspores) have simple hair-like glochidia, many of which are knobbed at their tips. Glochidia tend to resemble the perisporial hairs of the megaspore body. This resemblance provides fossil evidence of the homology of these two hair-like structures among living species of Azolla. In Azolla simplex from the Judith River Formation the megaspore has a single cap-like so-called columellate float. Massulae, which have anchor-shaped glochidia, are associated with these megaspores. A. simplex is the oldest species of Azolla and Parazolla the oldest member of the Salviniaceae so far found.     

EMBRYOLOGICAL STUDIES IN THE FAMILY SAXIFRAGACEAE (S.L.). I. DEVELOPMENT OF THE OVULE AND EMBRYO SAC IN SAXIFRAGA FORTUNEI VAR. PARTITA (MAKINO) NAKAI

The development of the ovule, megaspore and megagametophyte in Saxifraga fortunei var. partita (Makino) Nakai was observed. The ovule is anatropous, bitegmic, and crassinucellate. Both integuments originate from the epidermis. The archesporium is considered to be multicellular. The primary sporogenous cell functions as the megaspore mother cell which forms a T-shaped tetrad. The chalazal member of the megaspore tetrad is functional and develops into a Polygonum-type embryo sac. In the pyriform synergids the filiform apparatus is observed, but any hook or indentations could not be recognized. The antipodal cells are detectable until the Helobial endosperm undergoes several nuclear divisions. Secondary multiplication of the nuclei or the cells of the antipodals could not be observed.     

A contribution to the embryology of Desmodium motorium (Houtt.) Merr. (= D. gyrans (L.f.) DC.) (Fabaceae)

Abstract

The anthers are tetrasporangiate. The anther wall comprises epidermis, fibrous endothecium, middle layer and tapetal layer. The tapetum is of the Glandular type and its cells remain uninucleate. Meiosis in pollen mother cells is normal and simultaneous cytokinesis leads to the formation of tetrahedral and decussate microspore tetrads. The pollen grains are shed at 2-celled stage. The ovule is campylotropous, bitegmic and crassinucellate. Meiosis in megaspore mother cell results in the formation of linear or occasionally T-shaped megaspore tetrad. The chalazal megaspore develops into Monosporic Polygonum type of embryo sac. Endosperm development is of the Nuclear type.     

珍稀濒危植物巴东木莲胚胎学研究

对巴东木莲(Manglietia patungensis Hu)的花发育以及胚胎发育过程进行了系统研究。巴东木莲花顶生,花器官头年年底开始分化到第二年3月分化出花被、雌雄蕊群直至6月发育成熟。雌蕊成熟时胚珠倒生,双珠被,厚珠心,大孢子四分体线形排列,合点端发育成功能大孢子,珠孔端的3个退化,大孢子为单孢子发生型,胚囊发育方式属蓼型;雄蕊花药外侧壁玫瑰红色,内侧有4个白色花粉囊,绒毡层有1层多核细胞,小孢子四分体排列方式多为左右对称形和交叉形,四面体形,偶为T字形和线形,成熟花粉粒为二细胞型。在巴东木莲花发育和大、小孢子发生以及雌、雄配子体形成过程中未见异常现象,因此笔者认为该物种的花器官发育以及雌、雄配子体发育并不构成导致该物种濒危的因素。     

ACHLAMYDOCARPON VARIUS COMB. NOV.: MORPHOLOGY AND REPRODUCTIVE BIOLOGY

Structurally preserved arborescent lycopsid fructifications are described from Pennsylvanian age strata in eastern Kentucky and southern Illinois. Achlamydocarpon varius comb. nov. is the name proposed for these cones and previously reported isolated megasporophylls described as Lepidostrobophyllum varius. The specimens range up to 3.5 cm long and represent cone apices. Megasporophylls are spirally arranged and attached to the cone axis at an acute angle. Megasporangia are large with a wall two cell layers thick. Each sporangium contains one large, presumably functional megaspore, and three smaller, abortive megaspores. Functional and abortive spores possess a prominent apical tuft (massa) that covers up to one-third of the proximal surface of the spore. Sporoderm ultrastructure is detailed together with a comparison of morphologically similar sporae dispersae megaspores. The possible function of the megaspore massa is discussed as it relates to the reproductive biology of the cone.     

Viscin Threads in Zeuxine Strateumatica (Orchidaceae)

In the pollinium of Zeuxine strateumatica (L.) Schlecht., the massulae are held together by viscin threads. These threads are elastic in nature and seem to lack lignin and sporopollenin.     

Microtubular configurations during meiosis and megasporogenesis in Gasteria verrucosa and Chamaenerion angustifolium

Summary In Gasteria and Chamaenerion, microtubular configurations were visualized immunocytochemically during meiosis and megasporogenesis in order to study their relationship to cell development, meiotic divisions and selection of the functional megaspore. In Chamaenerion, the intensity of the fluorescence found in megaspores was weaker than that found in Gasteria. Both plants exhibited concentrations of microtubules around the meiocyte nuclei during pachytene-diplotene. Preprophase bands were not observed. In Chamaenerion, cytoplasmic microtubules radiating from meiocyte nuclei were found at late prophase, the dyad stage and in the functional megaspore; in Gasteria, they were observed only at the dyad stage and in the functional megaspore. During the second meiotic division of Gasteria, dividing cells and their nuclei exhibited differences in volumes. Also, the two microtubular spindles of the dyad cells had different widths. Fluorescence indicating the presence of the cytoskeleton diminished during maturation of the large functional megaspores in both plants, whereas in the three degenerating smaller megaspores, fluorescence intensity persisted. Our conclusion is that only an indirect relationship exists between the organization of the microtubular cytoskeleton and selection of the functional megaspore.     

MEGASPORE WALL ULTRASTRUCTURE IN ISOETES

In Selaginella, megaspore wall ultrastructure (unit morphology and arrangement) is correlated with taxonomic position. In some Pennsylvanian lycopods there is a correlation with dispersal strategy. This study was designed to assess any correlation between habitat preference and wall ultrastructure in Isoetes. Except for a few minor structural correlations, wall ultrastructure appears to be constant in terrestrial, amphibious, and aquatic species of Isoetes. Several distinct sizes of megaspores occur in some megasporangia and correspond to 1) full-size megaspores with siliceous coatings, 2) small megaspores with siliceous coatings, and 3) full-size megaspores without siliceous coatings. The uniformity in wall ultrastructure within modem Isoetes together with the lack of uniformity of fossil isoetalean megaspore wall ultrastructure suggest that the modem species of Isoetes are closely related.     

Megaspore development in Selaginella. I. “Wicks”, their presence,ultrastructure, and presumed function

Summary Structures have been found in the locular space between the tapetal cells and megaspores in Selaginella argentea and S. kraussiana that enter the megaspore wall and extend to the plasma membrane of the megaspore cytoplasm. We have called these structures wicks. Unless special fixation procedures are used wicks are either very poorly preserved or not apparent. Wicks appear to be routes for the transport of materials from the tapetum to developing megaspores. The entry of the wicks into the megaspore wall and their passage throughout the wall implies that the megaspore wall of Selaginella is a three-dimensional mesh-work of inter-connecting spaces. Wicks have several macromolecular-sized subunits, and the results of our histochemical reactions indicated the presence of glycoprotein and/or mucopolysaccharide. X-ray microanalysis of the S. convoluta exospore showed that silicon is present in rod-shaped structures between units of the exospore in mature megaspores. Because of the size and form of the structures between the exospore units we consider that they are remnants of wicks stabilized by silicon.Present address:Cátedra de Palinologia, Museo de La Plata, Paseo del Bosque s/nro., 1900 La Plata, Argentina.     

Polarity and competition between megaspores in the ovule ofOenothera hybrids

New data on the development of polarity in the ovules during megasporogenesis and early stages of embryo sac development inOenothera-hybrids are presented. It is confirmed that allOe. hookeri-hybrids show a strong tendency to form heteropolar tetrads, with the micropylar megaspore developing into an embryo sac. This preference is seen in the delay of the second meiotic division on the chalazal side, the absence of callose in the lateral wall of the micropylar megaspore, and the accumulation of starch in this megaspore. However, homopolar tetrads, chalazal preference, and ovules with two developing embryo sacs are also observed with considerable frequency. Quantitative data on the frequency of the different developmental types are compared with earlier genetic results about competition in the haplophase. There is sufficiently good agreement to support the hypothesis ofRenner that there is a correlation between the developmental processes in the megaspore tetrad and the genetic phenomena of competition in the haplophase.