Sclerosperma fossils from the late Oligocene of Chilga,north-western Ethiopia

The palm family, Arecaceae, is notoriously depauperate in Africa today, and its evolutionary, paleobiogeographic, and extinction history there are not well documented by fossils. In this article we report the pollen of two new extinct species of the small genus, Sclerosperma (Arecoideae), from a late Oligocene (27–28 Ma) stratum exposed along the Guang River in Chilga Wereda of north-western Ethiopia. The pollen are triporate, and the two taxa can be distinguished from each other and from modern species using a combination of light and scanning electron microscopy, which reveals variations in the finer details of their reticulate to perforate exine sculpture. We also report a palm leaf fragment from a stratum higher in the same section that is in the Arecoideae subfamily, and most likely belongs to Sclerosperma. The implications of these discoveries for the evolutionary history of this clade of African arecoid palms is that their diversification was well underway by the middle to late Oligocene, and they were much more widespread in Africa at that time than they are now, limited to West and Central Africa. Sclerosperma exhibits ecological conservatism, as today it occurs primarily in swamps and flooded forests, and the sedimentology of the Guang River deposits at Chilga indicate a heterogeneous landscape with a high water table. The matrix containing the fossil pollen is lignite, which itself indicates standing water, and a variety of plant macrofossils from higher in the section have been interpreted as representing moist tropical forest or seasonally inundated forest communities.     

Triaperturate pollen in the monocotyledons: configurations and conjectures

Triaperturate pollen are known in at least twenty seven genera of monocotyledons. Differences between aperture type and polarity indicate that the development of three apertures has occurred a number of times. Mode of cytokinesis during microsporogenesis is compared with differences in aperture configuration, to assess the extent to which this appears to influence aperture arrangement. Triapertury in monocot pollen tends to fall into one or another of three situations: 1) it is the normal state, 2) it is fairly common, but pollen with more or less apertures also occur in the taxon or sample, 3) it is a rare, or abnormal state for pollen which usually has less than three apertures. The various forms of triaperturate pollen are described, as well as monosulcate pollen of the orchid genera Cypripedium and Paphiopedilum, often misinterpreted as tri-sulcate, and the unusual extended trichotomosulcate pollen of Agrostocrinum (Hemerocallidaceae). Monosulcy, trichotomosulcy, and zonasulcy, with unusual and rare exceptions of zonasulcy in the eudicots, are aperture states shared exclusively with the basal dicots. Furthermore, to some extent all have links with the triaperturate condition in monocots and basal dicotyledons. This is discussed, as well as the association of tripory with polypory in monocots and basal dicots. The fossil pollen record is considered.This paper is dedicated to Klaus Kubitzki in recognition, not only for his extensive contribution to systematic botany, but also for his firm belief that pollen characteristics contribute to a better understanding of plant systematics and evolution.     

APERTURE EVOLUTION IN THE GENUS PTYCHOPETALUM BENTH. (OLACACEAE)

The genus Ptychopetulum (Olacaceae), disjunct between South America (2 spp.) and Africa (2 spp. West Africa; I sp. Rhodesia), exhibits predominantly diploporate pollen among the West African species and predominantly triporate pollen among the South American taxa. An analysis of aperture evolution suggests two possible schemes of aperture evolution for the genus. Scheme I avers the independent origin of both triporate and diploporate apertures from a basic brevicolpate condition while scheme II proposes a basic diploporate condition in which the diploporate apertures have migrated to the equator, fused to form brevicolpi and have ultimately become reduced to pores. Pollen evidence in support of both schemes is discussed. The distribution of aperture types within the genus also suggests West Africa as the original site of aperture evolution and South America as the second area of aperture development with types traceable to West African stock.     

Microsporogenesis in Monocotyledons

This paper critically reviews the distribution of microsporogenesistypes in relation to recent concepts in monocot systematics.Two basic types of microsporogenesis are generally recognized:successive and simultaneous, although intermediates occur. Theseare characterized by differences in tetrad morphology, generallytetragonal or tetrahedral, although other forms occur, particularlyassociated with successive division. Successive microsporogenesisis predominant in monocotyledons, although the simultaneoustype characterizes the ‘lower’ Asparagales. Simultaneousmicrosporogenesis also occurs inJaponolirion and Petrosavia(unplaced taxa), some Araceae, Aponogeton, Thalassia andTofieldia(Alismatales), Dioscorea, Stenomeris and Tacca (Dioscoreales),and some Commelinanae: Arecaceae (Arecales), and Cyperaceae,Juncaceae and Thurniaceae (Poales). Simultaneous microsporogenesisis of phylogenetic significance within some of these groups,for example, Asparagales, Dioscoreales and Poales. An intermediatetype is recorded in Stemonaceae (Pandanales), Commelinaceae(Commelinales) and in Eriocaulaceae and Flagellariaceae (Poales).There is little direct relationship between microsporogenesistype and pollen aperture type in monocots (except for trichotomosulcateand pantoporate apertures), although trichotomosulcate aperturesin monocot pollen, and equatorial tricolpate and tricolporateapertures in eudicot pollen, are all related to simultaneousmicrosporogenesis. Copyright 1999 Annals of Botany Company Microsporogenesis, monocotyledons, pollen apertures, phylogeny, tetrads, simultaneous, successive, systematics.     

Aperture Orientation in Laurelia Pollen (Atherospermataceae Syn. Subfamily Atherospermoideae of Monimiaceae)

The development of the encircling aperture of pollen grains or the New Zealand species Laurelia novae-zelandiae is described by means of light and scanning electron microscopy. Some observations are also reported from pollen development and structure in the South American species Laurelia sempervirens, mature pollen of which can be distinguished from that of L. novae-zelandiae. The aperture begins to develop while pollen of L. novaezelandiae is still in tetrads, and passes through the distal and proximal poles of each grain. The widest parts of the mature aperture are at and near what are morphologically the distal and proximal poles of each grain. No such meridionosulcate type of pollen was recorded in a recent survey of pollen aperture types of primitive angiosperms and contradicts a recent report that pollen in the Atherospermataceae had equatorially aligned apertures.     

A survey of species assigned to the fossil pollen genus Anacolosidites

Anacolosidites Cookson & Pike, a fossil pollen genus recorded since the Campanian, is peculiar in its morphology – six‐porate with three apertures on each hemisphere, located away from the equator, and with the distal and proximal apertures positioned over each other. Representatives of this fossil genus are widely considered to represent extant Olacaceae from tribe Anacoloseae. Olacaceae is an exclusively tropical angiosperm family with a pantropical distribution; consequently the fossils are often used to suggest a tropical climate and in addition are frequently used as a stratigraphic marker. Fossil species assigned to Anacolosidites are quite variable and may not all represent Olacaceae, in which case they may not indicate tropical climate.

The present study is a morphological survey of fossil pollen assigned to Anacolosidites; it identifies the published reports of the fossil species that probably represent positive occurrences of Olacaceae pollen in the fossil record. Within Olacaceae, Anacolosidites‐type pollen is usually compared with pollen of genera in tribe Anacoloseae, in particular: Cathedra, Anacolosa and Phanerodiscus, but never with the pollen of Ptychopetalum, a genus from tribe Olaceae with closely similar pollen to the other three genera, but with a reticulate tectum and very small circular apertures located near the equator. Nevertheless, the records of reticulate Anacolosidites species which have been excluded from the emended diagnosis are unlikely to be related to Ptychopetalum. The earliest accepted record of Anacolosidites is from the Late Cretaceous (Maastrichtian) of Germany. However, most Late Cretaceous records, and later Russian and Chinese occurrences referred to Anacolosidites, consist mainly of Normapolles‐type pollen, whereas many of the Cenozoic records assigned to Anacolosidites have a much clearer affinity with the pollen of Anacolosa, Cathedra and Phanerodiscus (tribe Anacoloseae). The newly emended genus Anacolosidites may be used as a stratigraphic marker for tropical or megathermal climatic conditions.     

New Striate Pollen from the Pennsylvanian of Kansas

A new genus of striate saccate pollen, Caelopollis is described from the Virgilian (Pennsylvanian) of Kansas. A striate distal leptoma serves to distinguish the genus and four new species of striate pollen. One new species of Vittatina, V. moorei, is described that also possesses an unusual type of distal ornamentation.     

Pollen morphology of Strobilanthes Blume (Acanthaceae) in China and its taxonomic implications

The pollen morphology of 37 Chinese species of Strobilanthes Blume (Acanthaceae), a large and taxonomically problematic genus, was investigated by LM and SEM. Two distinct pollen shape classes, ellipsoid and spheroidal, recognised by earlier workers were found. Eight pollen types could be distinguished on the basis of variation in ornamentation, and the number, distribution and type of apertures. Five of these fall within twenty two pollen types reported from Strobilanthes in southern India and Sri Lanka. Three new pollen types are also described, further extending the palynological diversity of the genus. From the results presented, the characters of palynological diversity and the related taxonomic implications are discussed.     

Unique tetrads of <Emphasis Type="Italic">Epilobium luteum</Emphasis> (Onagraceae: Onagreae) pollen from Alaska

Aperture morphology of tetrad pollen of Epilobium luteum (Onagraceae: Epilobieae) from three Alaskan collections is highly variable. The first collection appears to lack apertures altogether and is presumed to consist of immature pollen gains in a genus known to achieve mature size before the apertures become distinctly protruding. A second collection has tetrads with 3- and 4-apertured grains, the apertures in the latter are often irregularly spaced and not in apposition with the apertures of neighboring members. The third collection consists of the more typical 3-apertured members that characterize the majority of Epilobium pollen grains. In all of these collections individual pollen grains (monads) are interspersed among the tetrads. The variations in the number of apertures emphasize the importance of having a comprehensive understanding of the stage of development of the pollen (taxon) examined when describing pollen collections. In the first collection this would mean the recognition that in Onagraceae apertures occur in the later stages of microspore ontogeny. In the latter two collections a thorough background of the literature of the pollen morphology on this largest Onagraceae taxon is useful for the understanding of the significance of a range of aperture numbers on Epilobium pollen grains.     

Arecaceae potential for production of monofloral bee pollen

Bees often collect pollen from plants that are near their colonies or apiaries. Arecaceae are characterised by having species that flower throughout the year and produce large quantities of pollen. Pollen grains of Arecaceae have two main forms, namely monosulcate and trichotomosulcate. The study area is located in the coastal region of the state of Sergipe, Brazil, where large amounts of bee pollen are produced by Apis mellifera. This study aims to evaluate how Arecaceae contribute to the production of monofloral bee pollen. Samples of bee pollen loads were collected each month for two years (totalling 24 samples), acetolysed, and mounted on permanent slides. At least 500 grains of pollen were counted for each sample, the data was analysed, and figures were made using the Tilia software. The pollen grains of Cocos nucifera were present in 23 samples (96%), and concentrations of this pollen type varied between 1.1 and 46.8% (but were commonly above 20%). The concentration of Arecaceae pollen grains is the result of the large-scale production of coconuts along the coastline. However, the presence of other pollen types, such as Mimosa arenosa, Mimosa pudica, Mikania and Myrcia, compliments the diets of bees in the region. The results show that even when in low frequency, Cocos nucifera pollen grains represent a large proportion of pollen content compared with other pollen types, because of the large size of Arecaceae pollen grains.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A CATKIN WITH JUGLANDACEOUS AFFINITIES

Three specimens of one type of fossil catkin from the Middle Eocene of Tennessee are excellently preserved and have been investigated morphologically. The flowers on these catkins are subtended by elongate, three-lobed bracts, are exclusively staminate, and have three conspicuous, obovate, perianth parts that bear large peltate scales. The stamens are well preserved and contain triporate pollen grains that are equivalent to the dispersed pollen genus Momipites. Floral morphology, cuticular features, and pollen indicate close affinities with the extant genera Engelhardia, Oreomunnea, and Alfaroa of the Juglandaceae; but because the fossil catkins are distinct and are a dispersed plant organ, they are placed in a new form genus: Eokachyra. These fossil flowers represent a rare opportunity to correlate the micro- and macrofossil record and to compare the relative rates of evolution of these features. The fossil catkins also demonstrate that much structural information may be gained from the study of fossil angiosperm flowers. The similarities between the staminate flowers of the fossil catkins and the staminate flowers of Engelhardia, Oreomunnea, and Alfaroa confirm the idea that this complex has had a long evolutionary history and suggest that the pollination system of certain extant genera was well developed during Middle Eocene times.     

Pollen morphology of Astragalus section Hymenostegis (Fabaceae) and evaluation of its systematic implications

Astragalus is with nearly 3000 described species the largest genus of flowering plants. So far analyses of pollen characters have only been conducted for a few species of the groups within the genus. Here we analyse pollen grains of 22 species representative for Astragalus section Hymenostegis using scanning electron microscopy. We found the basic shape of the pollen grains to be oblate-spheroidal and apertures to be tricolpate as for other eudicots. The sculpturing pattern of the exine is micro-reticulate. Pollen grains show low morphological variation among different species of this section, but differences occur between sections of the genus. We conclude that the vast morphological differentiation that occurred during the rapid radiation of section Hymenostegis was not accompanied by comparable differentiation in pollen morphology.     

Formation and function of a new pollen aperture pattern in angiosperms: The proximal sulcus of Tillandsia leiboldiana (Bromeliaceae)

Pollen grains are generally surrounded by an extremely resistant wall interrupted in places by apertures that play a key role in reproduction; pollen tube growth is initiated at these sites. The shift from a proximal to distal aperture location is a striking innovation in seed plant reproduction. Reversals to proximal aperture position have only very rarely been described in angiosperms. The genus Tillandsia belongs to the Bromeliaceae family, and its aperture pattern has been described as distal monosulcate, the most widespread aperture patterns recorded in monocots and basal angiosperms. Here we report developmental and functional elements to demonstrate that the sulcate aperture in Tillandsia leiboldiana is not distal as previously described but proximal. Postmeitotic tetrad observation indicates unambiguously the proximal position of the sulcus, and in vitro germination of pollen grains confirms that the aperture is functional. This is the first report of a sulcate proximal aperture with proximal germination. The observation of microsporogenesis reveals specific features in the patterns of callose thickenings in postmeiotic tetrads.     

Studies in Montia L. and Claytonia L. and Allied Genera

The pollen morphology of several genera in Portulacaceae is described. Particular attention has been paid to the genera of the subfamily Montioideae, as a stage of continued monographical studies. Among genera especially dealt with are Claytonia, Montia, Crunocallis, Naiocrene, Neopaxia, Mona, Maxia, Limnalsine, and Montiastrum. In the taxonomical treatment of these genera the pollen morphology has proved to afford many important additional characters.

The pollen grains of Claytonia are distinguished from those of the remainder in being 3-colpate. The grains of the Claytonia-type have many similarities with those of Lewisia, a genus of the subfamily Portulacoideae. The other genera of Montioideae have pantocolpate pollen grains. Among these genera several different pollen types are distinguished, chiefly with regard to the sexine structures and the aperture membranes. The Montiastrum-type is especially interesting, with tholate grains, a particular pollen type not met with in any other genus in the family. The pollen morphology of some genera in the Portulacoideae is also treated. In some species in Calandrina and Talinum pantotreme pollen grains are observed with apertures transitional between pori and colpi. The apertures of the pantotreme grains are arranged in characteristic patterns.

Particular attention has been given to the variation of the pollen morphological characters. This variation has been examined with regard to the differences between different populations of the same species as well as between different species. The greatest variation has been observed in the shape and size of the grains. The structure and sculpture and thickness of the sexine and the aperture membranes are less variable. Some polyploid taxa are connected with the occurrence of pollen grains with divergent and varying aperture numbers.

In a survey of the genera the taxonomical results of the investigation are presented with particular regard to the pollen morphology. The new genus, Maxia Ö. Nilss., is described. One new species, Montia clara Ö. Nilss., is described and some new combinations are made.

Pollen morphological diagnoses are given for 46 different taxa. The aperture conditions for 96 different species are presented.     

Pollen wall development in Vigna vexillata I. Characterization of wall layers

Light and electron microscope observations characterized the layers that comprise Vigna vexillata L. pollen walls, and identified the timing of their development. Exine sculpturings form an unusually coarse ektexinous reticulum. The structure of the ektexine is granular; this differs from the columellate/tectate type of structure typical of most angiosperm pollen. The ektexine overlies a homogeneous-to-lamellar, electron-dense endexine, which in turn surrounds a thick, microfibrillar intine. Pollen grains are triporate and operculate, with Zwischenkörper and thickened intine underlying the apertures. The ektexine forms during the tetrad period of microspore development, the endexine and Zwischenkörper during the free microspore stage, and the intine during the bicelled (pollen) stage. Coarsely reticulate exine sculpturings and the granular structure of the patterned exine wall of the pollen grains are features that make this species suitable for detailed studies of pollen wall pattern formation.     

CHLORANTHUS-LIKE STAMENS FROM THE UPPER CRETACEOUS OF NEW JERSEY

Fossil angiospermous stamens with in situ pollen from the Turonian (ca. 90 million years before present, Late Cretaceous) of New Jersey are described and assigned to the Chloranthaceae. The fossil stamens, which are three-parted and bear two bisporangiate thecae on the central lobe and one bisporangiate theca on each lateral lobe, are indistinguishable from stamens of several extant species of Chloranthus. The pollen is spheroidal, 13–18 μm in diameter, with a reticulate exine and apparently elongate/elliptical apertures. The pollen is similar to that in extant Chloranthus in grain size, shape, exine sculpture, and aperture structure. Like pollen of some extant species of Chloranthus, aperture number in the fossil pollen appears to be variable. Because fossil pistillate chloranthoid reproductive structures have not been found at this locality it is unknown whether the fossil stamens described here were borne on the side of the ovary, as in extant Chloranthus, or in another arrangement. The three-parted stamen of Chloranthus is unique in angiosperms and there has been considerable debate concerning the origin and evolutionary significance of the structure. Uncertainty as to whether the three-parted stamen represents a synapomorphy for the genus or a retained plesiomorphy in angiosperms is the primary reason why these fossil stamens are not assigned to the extant genus Chloranthus.     

Ultrastructure and diversity of recent and fossil zona-aperturate pollen grains

A survey of recent and fossil zona-aperturate pollen grains within the angiosperms was undertaken. Zona-aperturate pollen (with complete ring-like apertures) has been evolved independently in several genera of basal angiosperm families (Nymphaeaceae, Eupomatiaceae, Annonaceae, Monimiaceae, Atherospermataceae) and monocots (Araceae, Iridaceae, Laxmanniaceae, Arecaceae, and Rapateaceae). The special case of the eudicot Limnanthes (Limnanthaceae) is discussed. New examples of recent and fossil fully zonate pollen are presented; a noteworthy new example are Scrophulariaceae, with Pedicularis; our fossil examples show affinities either to Nymphaeaceae, or Araceae, or Iridaceae, or cannot be related to an extant taxon. The zona-aperturate pollen grains differ in ornamentation and stratification mostly in the respect of the aperture areas, but sometimes also outside the aperture. The question of polarity is settled only for some taxa because of the frequent lack of tetrad observations. A ring-like aperture may divide the pollen grains in symmetric halves, or if running asymmetrically, divides the pollen grain in two more or less unequal halves; the presence of clearly unequal halves gives a strong argument to assume an equatorial ring in such cases where tetrad configurations were not available. The zona-aperturate condition may be a functional benefit in wet environments: while in dry pollen the apertural ring is completely closed, in wet or very moist realm it expands greatly, and the pollen tube can be formed quickly and everywhere in this area.     

Beiträge zur Pollenmorphologie und Systematik der Convolvulaceen-Gattungen Maripa und Mouroucoa

Suggested by a previous communication on the occurence of two types of apertures among the pollen grains of Maripa s. l. (Convolvulaceae), 16 species of this genus were investigated palynologically. 3-zonocolpare grains mixed with 4–6-pantocolpate and 6–15-pantocolpate grains were found. The structure of the exine (infratectale bacula with or without supratectale processes) reveals further differences. The combination of these characters in the genus Maripa offers the separation into four types of pollen grains. The genus Mouroucoa is represented by its own type. These facts stress the necessity of the generical separation of the two genera. Consideration of pollen morphology yields some new aspects for subdivision of the genus Maripa.     

A monograph of the betel nut palms (Areca: Arecaceae) of East Malesia

The palm genus Areca is widespread in tropical Asia and includes the economically important betel nut palm, A. catechu. The genus has three centres of high species diversity: the Sunda Region, the Philippines and East Malesia (to the east of Wallace's line). The taxonomy of the genus in East Malesia has been neglected. Prior to this study, 19 species were accepted for this area, all but one endemic, but their limits and differences were not understood. Here, we provide a taxonomic monograph of East Malesian Areca spp., based on an extensive study of the genus in herbaria and in the field. We recognize six species of Areca in East Malesia, including the widespread cultivated A. catechu. Five wild species are accepted, namely A. macrocalyx, A. mandacanii, A. novohibernica, A. oxycarpa and A. vestiaria. We place 12 of the previously accepted species into synonymy, and provide additional new synonymy in A. catechu. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 147–173.