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.     

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.     

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.     

Paleoazolla,a new heterosporous fern from the Upper Cretaceous of Argentina

Megaspore and microspore massulae of Paleoazolla patagonica gen. et sp. nov. are described from the Upper Cretaceous of La Colonia Formation, Chubut Province, Argentina. The new fern possesses megaspore complexes with three to four glochidiate floats attached directly to the megaspore; a columella-like structure appears to be absent. The megaspore wall consists of a two-layered exine that is smooth to irregularly perforate, a two-layered perine with a spongy, densely packed endoperine, and a loosely organized exoperine. Infrafilosum hairs cover the exoperine. Microspore massulae are irregular in size and shape and bear multibarbed glochidia that generally have anchor-shaped tips. A comparison with other azollaceous and salviniaceous genera, particularly Azolla, is provided, together with a discussion of some evolutionary trends within the family.     

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.     

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.     

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.     

POROSTROBUS NATHORSTII SP. NOV.: A NEW SPECIES OF LYCOPSID CONE FROM THE EARLY PENNSYLVANIAN OF ILLINOIS

Thirty-one specimens of a small megasporangiate lycopsid cone referable to the genus Porostrobus Nathorst and abundant associated dispersed megaspores have been collected from Early Pennsylvanian strata in the Allied Stone Company quarry, Milan, Illinois. Based on other elements in the flora, the deposit is considered to be part of the Morrowan Caseyville Formation and probably of Namurian age. This is the first reported occurrence of Porostrobus in North America and the cones are recognized as a new species, P. nathorstii. The environment of deposition indicates that the cones may have been transported from the parent plant prior to preservation. Cones are preserved as coalified compressions measuring 15–36 mm long by 2.5–7 mm wide, and are characterized by an apical tuft of leaves up to 20 mm long. Sporophylls are spirally arranged on a narrow cone axis, lack a heel or keel, and have a long distal lamina. Sporangia contain a single functional megaspore tetrad. Mature megaspores are 750–1, 150 μm in diameter, have prominent trilete sutures raised to form a gula, and have numerous branched hairs confined to an equatorial band. Megaspores correspond to the dispersed form Setosisporites praetextus (Zerndt) Potonie and Kremp. Porostrobus nathorstii is the only species of the genus described to date that is monosporangiate.     

ARIADNAESPORITES AND GLOMERISPORITES IN THE LATE CRETACEOUS: ANCESTRAL SALVINIACEAE

Glomerisporites and Ariadnaesporites are two genera based on megaspores and microspores from the late Cretaceous. Each has megaspores with characteristics shared with other late Cretaceous Salviniaceae, including numerous floats, an acrolamella, and a complex perispore. Microspores are borne singly, however, not in characteristic salviniaceous massulae. Microspores of Glomerisporites have a pseudovacuolate basal region and a single spore in an apical neck. Microspores of Ariadnaesporites are structurally like the megaspores but of smaller size. Ariadnaesporites varius occurs in the Cenomanian Stage (early late Cretaceous) and is the oldest presumed member of the Salviniaceae.     

Ultrastructure and affinity of Lower Carboniferous megaspores from the Moscow Basin, Russia

Ten megaspore species isolated from Moscow Basin lignites of Lower Carboniferous (Viséan) age have been studied by scanning and transmission electron microscopy (SEM and TEM). These species belong to seven megaspore genera: Lagenicula, Sublagenicula, Crassilagenicula, Setosisporites, Zonalesporites, Caudatosporites, and Cystosporites. Megaspores of the genus Caudatosporites have only been described previously from the Duckmantian (Westphalian B); a new species is duly erected. The ultrastructure of megaspore walls from the genera Crassilagenicula and Zonalesporites has not been previously described. This study also places them in context with other contemporaneous megaspores. The study shows that during the Viséan, in the Moscow Basin, megaspores expressed a similar wall ultrastructure despite large differences in external appearance. The genus Crassilagenicula may represent a group of megaspores from plants that had evolved from those bearing gulate megaspores here typified by Lagenicula acuminata, Setosisporites brevispinosus, and Sublagenicula hirsutoida. Zonalesporites brasserti also appears to show affinities to this group, and may be representative of a plant species in a transitional state between the Lagenicula bearing lycopsids and those more isoetalean in nature.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

A TETRAHEDRAL MEGASPORE ARRANGEMENT IN A SEED FERN OVULE OF PENNSYLVANIAN AGE

A tetrahedral arrangement of one functional and three aborted megaspores has been found in the fossil seed fern ovule Conostoma anglo-germanicum. This is the first conclusive report of a Pennsylvanian age ovule with such a primitive megaspore arrangement, suggesting that this feature was more widespread and persistent among Paleozoic pteridosperms than previously assumed.     

Paleobotanical remains from the Paleocene–lower Eocene Vagadkhol Formation, western India and their paleoclimatic and phytogeographic implications

A small assemblage of macro- and micro floral remains comprising fossil leaf impressions, silicified wood, spores, and pollen grains is reported from the Paleocene–lower Eocene Vagadkhol Formation (=Olpad Formation) exposed around Vagadkhol village in the Bharuch District of Gujarat, western India. The fossil leaves are represented by five genera and six species, namely, Polyalthia palaeosimiarum (Annonaceae), Acronychia siwalica (Rutaceae), Terminalia palaeocatapa and T. panandhroensis (Combretaceae), Lagerstroemia patelii (Lythraceae), and a new species, Gardenia vagadkholia (Rubiaceae). The lone fossil wood has been attributed to a new species, Schleicheroxylon bharuchense (Sapindaceae). The palynological assemblage, consisting of pollen grains and spores, comprises eleven taxa with more or less equal representation of pteridophytes, gymnosperms, and angiosperms. Angiospermous pollen grains include a new species Palmidites magnus. Spores are mostly pteridophytic but some fungal spores were also recovered. All the fossil species have been identified in the extant genera. The present day distribution of modern taxa comparable to the fossil assemblage recorded from the Vagadkhol area mostly indicate terrestrial lowland environment. Low frequency of pollen of two highland temperate taxa (Pinaceae) in the assemblage suggests that they may have been transported from a distant source. The wood and leaf taxa in the fossil assemblage are suggestive of tropical moist or wet forest with some deciduousness during the Paleocene–early Eocene. The presence of many fungal taxa further suggests the prevalence of enough humidity at the time of sedimentation.     

PINUS WOLFEI,A NEW PETRIFIED CONE FROM THE EOCENE OF WASHINGTON

A silicified cone from the Late Eocene of Washington is described as a new fossil species of Pinus. The cone was probably 9–10 cm long and 3–5 cm at its widest diam in the living condition and is peculiar in having abundant resin canals in the secondary xylem of the axis arranged in three concentric rings near the cone base. The bract of the fossil is also unusual in having resin canals of distinctly unequal sizes and a vascular strand that is adaxially concave. In the absence of external features of the scale tips, these anatomical conditions along with the construction of the outer cortex of the axis of thick-walled cells suggest closest affinity of the new species with the subsections Contortae, Oocarpae, and Sylvestres of the section Pinus, subgenus Pinus.     

DIVERSITY AND HOST SPECIFICITY OF AZOLLA CYANOBIONTS1

A unique, hereditary symbiosis exists between the water fern Azolla and cyanobacteria that reside within a cavity in the dorsal leaf‐lobe of the plant. This association has been studied extensively, and questions have frequently been raised regarding the number and diversity of cyanobionts (cyanobacterial symbionts) among the different Azolla strains and species. In this work, denaturating gradient gel electrophoresis (DGGE) and a clone library based on the 16S rRNA gene were used to study the genetic diversity and host specificity of the cyanobionts in 35 Azolla strains covering a wide taxonomic and geographic range. DNA was extracted directly from the cyanobacterial packets, isolated after enzymatic digestion of the Azolla leaves. Our results indicated the existence of different cyanobiont strains among Azolla species, and diversity within a single Azolla species, independent of the geographic origin of the host. Furthermore, the cyanobiont exhibited host‐species specificity and showed most divergence between the two sections of genus Azolla, Azolla and Rhizosperma. These findings are in agreement with the recent redefinition of the taxon Azolla cristata within the section Azolla. With regard to the taxonomic status of the cyanobiont, the genus Anabaena of the Nostocaceae family was identified as the closest relative by this work.     

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.     

MORPHOLOGICAL STUDIES IN ALETRIS. I. DEVELOPMENT OF THE OVULE,MEGASPORES AND MEGAGAMETOPHYTE OF A. AUREA AND THEIR CONNECTION WITH THE SYSTEMATICS OF THE GENUS

Browne , Edward T., Jr . (U. of Kentucky, Lexington.) Morphological studies in Aletris. I. Development of the ovule, megaspores and megagametophyte of A. aurea and their connection with the systematics of the genus. Amer. Jour. Bot. 48(2): 143–147. Illus. 1961.—Development in a North American species of this variously classified genus has shown great similarity with the development in several genera of Hutchinson's Liliaceae-Narthecieae: Pleea, Tofieldia, Nanhecium and ∗∗∗Metaparthecium. The ovules are anatropous, bitegmic, crassinucellate and arranged in 4 rows in each locule of the tricarpellate pistil. There is a hypostase and an obturator. The primary archesporial cell is hypodermal. This undergoes a division to form a wall cell and the megaspore parent cell (MPC). The megaspores usually have a linear arrangement although occasionally a T-shaped tetrad may be formed. Most frequently the chalazal megaspore functions, but rarely the one adjacent to it may enlarge instead. Megagametophyte development is of the Polygonum type. A characteristic narrowed chalazal constriction is formed during the development of the megagametophyte. It is recommended on the basis of this information that Aletris be classified with the genera of the Liliaceae-Narthecieae.     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

FRUITS AND SEEDS OF TRIBE GORDONIEAE (THEACEAE) FROM THE EOCENE OF NORTH AMERICA

A new genus and species Gordoniopsis polysperma and two new species of Gordonia (Gordonieae, Camellioideae, Theaceae) are described based on fossil fruit and seed remains. These specimens are part of a large flora consisting of various plant organs from the middle Eocene Claiborne Formation in western Kentucky and Tennessee. Gordoniopsis is a five-valved loculicidally dehiscent capsule similar to capsules of Gordonia but differing in having unwinged seeds and a greater number of seeds per locule. The two Gordonia species are among the earliest unequivocal records of the genus and two of only four fossil Gordonia species known with in situ seeds. Two extinct genera, Gordoniopsis and Andrewsiocarpon, and the extant genus Gordonia in the tribe Gordonieae are known from the middle Eocene Claiborne flora, suggesting an early radiation within the tribe. Based on a survey of Recent fruits and seeds we concur with Keng's proposal to merge Laplacea with Gordonia.     

LIRIODENDRON (MAGNOLIACEAE) FROM THE MIOCENE CLARKIA FLORA OF IDAHO

Miocene Liriodendron carpels, whole fruiting structures and leaves from Clarkia and Oviatt Creek sites in northern Idaho are preserved as imprints and compressed fossils in soft lacustrine clays. The isolated carpels are indistinguishable from those described as L. hesperia Berry from the Spokane Latah flora. Fruit aggregates from the type Clarkia and Oviatt Creek localities and leaves from three Clarkia sites are considered to be within the range of variation of the single species L. hesperia. Comparisons were made regarding leaf architecture, lower leaf epidermal structures, leaf flavonoid and steroid analysis, morphological features of receptacles and carpels, and the venation pattern of carpels of the fossil material to the two extant species, L. tulipifera L. (native to southeastern United States) and L. chinense Sarg. (native to southeastern Asia). Leaf architecture features analyzed by standard statistical and canonical tests and marginal venation patterns near the base of leaves suggest that L. hesperia is more similar to L. tulipifera, whereas the size dimensions of lower epidermal cells and the common presence of two sterane compounds imply that L. hesperia is more similar to L. chinense. The fossil species, however, is a distinct taxon indicated by statistical discriminant and canonical tests, leaf base shape, often smaller epidermal cell dimensions, and the shape of round receptacle carpel scars. Both the fossil and the two living Liriodendron species are associated with comparable mixed mesophytic floras.