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.     

Aspects of evolutionary differentiation of theHamamelidaceae and the LowerHamamelididae

New investigations on the flower and fruit structure of extantHamamelidaceae and other LowerHamamelididae together with new finds of fossil flowers and seeds from the Upper and Lower Cretaceous provide the outline of an increasingly more differentiated picture of the early evolution of the subclass. Three patterns of valvate anther dehiscence are recognized in the subfamilyHamamelidoideae (and the subclassHamamelididae). The basic (plesiomorphic) type within theHamamelididae has 2 valves per theca. The type with 1 valve but 2 pollen sacs per theca is both consistent and exclusive for the 5 southern genera of theHamamelidaceae. They seem to be the remnants of a homogeneous group that originated before the Upper Cretaceous. This is supported by fossil hamamelidaceous flowers from the Upper Cretaceous that have thecae with 1 valve. Since several-seededHamamelidaceae predate one-seeded forms in the fossil seed record (in Europe) and the systematic structure of the one-seeded group is relatively more homogeneous, several-seeded groups are considered to be more ancient. Several parallel evolutionary trends are recognized within theHamamelidaceae as well as within the LowerHamamelididae: anther dehiscence with 2 valves per theca 1 slit or 1 valve; pollen sacs per theca 2 1; pollen tricolpate polyforate; exine coarsely reticulate finely reticulate; loss of perianth (tepals or petals and sepals) and concomitant loss of fixed number of floral organs; differentiation of exposed nectaries.     

Fossil flowers and pollen ofLauraceae from the Upper Cretaceous of New Jersey

A fossil trimerous flower from the Turonian (ca. 90 MYBP, Upper Cretaceous) of New Jersey is described as a new genus in the familyLauraceae. The fossil flower is charcoalified and preserved in exceptional detail. This fossil specimen is particularly remarkable in that several pollen grains have been preserved; pollen grains ofLauraceae generally have very thin exine and are rarely preserved in the fossil record. Although the specimen is incomplete and lacks anthers, there are sufficient structural details preserved to permit an assignment to theLauraceae, as well as comparisons with the tribePerseeae. This new genus provides an important addition to our knowledge of systematic and structural diversity in CretaceousLauraceae.     

Pollen-ovule ratios in some Neotropical Araceae and their putative significance

The correlation between pollen-ovule (P/O) ratio and breeding system has generally been analysed with respect either to pollination efficiency, or in terms of sex allocation theory. Pollen/ovule ratios were measured in nine species of Araceae belonging to two genera with bisexual flowers (Anaphyllopsis, Monstera) and three genera with unisexual flowers (Dieffenbachia, Philodendron, Montrichardia). The family Araceae with its unique inflorescence morphology allows the analysis of variations of the P/O ratio with respect to two basal morpho-functional pollination units: the flower or the inflorescence. We found a relationship between the value of the P/O ratio and the breeding system that is partially different from Cruden's results (1977). Some facultative xenogamous species have a higher P/O than the obligatory xenogamous species. A link was found between the P/O and the type of inflorescence, the floral cycle, and the mode of growth.     

Structure of the inflorescence and flower ofPetalonyx linearis (Loasaceae)

Fossil chloranthoid androecia,Chloranthistemon endressii gen. et spec. nov. are described from the Upper Cretaceous (Upper Santonian or Lower Campanian) of Scania, southern Sweden. They are three-lobed and dorsiventrally flattened with all pollen sacs borne laterally and inclined toward the presumed adaxial surface. The central lobe bears two pairs of pollen sacs, the lateral lobes a single pair each. The morphology, anatomy and valvate dehiscence of the fossil androecia is very similar to that seen in extant species ofChloranthus andSarcandra, but the in situ pollen differs from that of all extantChloranthaceae in being spiraperturate. A single chloranthoid androecium from the Lower Cretaceous (Upper Albian) of Maryland, North America has a more generalized structure thanChloranthistemon endressii. It consists of three stamens that are fused at the base, and each stamen bears two pairs of oppositely positioned pollen sacs. Combined with anatomical information from recentChloranthus the Lower Cretaceous specimen suggests that the androecium in the living genus has arisen by fusion and other modifications of three separate stamens each with a normal complement of four pollen sacs. The structure of both the Upper and Lower Cretaceous androecia suggest that these fossilChloranthaceae were insectpollinated. Macrofossil evidence combined with information from dispersed pollen indicates that theChloranthaceae diversified early in angiosperm fossil history and were an important component of Mid-Cretaceous plant communities.     

A new Monstera (Araceae: Monsteroideae) from Brazil

A new species ofMonstera (Araceae),M. praetermissa, is described, illustrated, and compared to its most similar relatives,M. xanthospatha andM. obliqua. The new species is so far only known from Brazil, but has a broad distribution there, ranging from the south to the northeast of the country.     

Pollen morphology ofLactoridaceae

The pollen wall ultrastructure of the primitive AngiospermLactoris fernandeziana Phil. (Lactoridaceae) is described. The monosulcate aperture, granular wall structure and sacci (all primitive features) suggest placement of this family in theMagnoliales. Pollen ofLactoris is compared to fossil dispersed pollen from the Lower and lower-Upper Cretaceous. The fossil pollen shares characteristics which are restricted to theLactoridaceae suggesting that this family was present during the early Cretaceous.     

Normapolles plants: a prominent component of the Cretaceous rosid diversification

The Normapolles complex, characterised by its oblate and triaperturate pollen, constitutes an important and diverse element of many Late Cretaceous and Early Cainozoic floras of the Northern Hemisphere. Based on the dispersed pollen record alone it has been difficult to assess systematic affinities, but relationships with Fagales have been proposed. Over the past twenty years several exquisitely preserved Late Cretaceous reproductive structures with Normapolles type pollen in situ have been described. In this study we provide a summary and new information of these floral structures. Further, a new genus, Dahlgrenianthus, is described from the Late Cretaceous of southern Sweden. The genus includes the type species Dahlgrenianthus suecicus, a number of reproductive structures referred to Dahlgrenianthus sp., and Dahlgrenianthus trigonus (Knobloch et Mai) comb. nov. from the Maastrichtian flora of Walbeck, Germany. Dahlgrenianthus comprises small flowers with pentamerous perianth and androecium and a tricarpellate gynoecium. It is distinguished from all other Normapolles floral structures in its hypogynous floral organisation. All Normapolles floral structures described so far are thought to be related to various members of the core Fagales, but the group is obviously not monophyletic. The stratigraphic range of the Normapolles taxa and other fagalean fossils strongly suggests that all major fagalean lineages were present by the Cenomanian or earlier.     

The mangrove palm Nypa in the geologic past of the New World

The paleontological history of Nypa, known today as the mangrovepalm, is traced through geological time back to the Late Cretaceous. Emphasis is laid on the New World occurrences, especially in NorthAmerica, where Nypa is known from fossil fruits and pollen. In SouthAmerica, the stratigraphic range of this palm extends from theMaastrichtian to the late Eocene, whereas in North America, Nypa isrestricted to only the Eocene. Nypa occurs as pollen all along theAmerican Gulf Coast from the early Eocene (Ypresian) to the late Eocene(Priabonian), while fruit records come from the early and middle Eoceneof Maryland and Texas, respectively. The floristics of these Eocenemangroves, including possible mangrove associates, and the developmentof mangrove vegetation in the neotropics through the Tertiary andQuaternary, are discussed. New paleobotanical evidence from a middleEocene faunal and floral assemblage in Texas, the Casa Blanca flora (LaredoFormation, Claiborne Group), which contains fossil Nypa pollen andfruits, is described in detail.     

Fossil Araceae from a Paleocene neotropical rainforest in Colombia

Both the fossil record and molecular data support a long evolutionary history for the Araceae. Although the family is diverse in tropical America today, most araceous fossils, however, have been recorded from middle and high latitudes. Here, we report fossil leaves of Araceae from the middle-late Paleocene of northern Colombia, and review fossil araceous pollen grains from the same interval. Two of the fossil leaf species are placed in the new fossil morphogenus Petrocardium Herrera, Jaramillo, Dilcher, Wing et Gomez-N gen. nov.; these fossils are very similar in leaf morphology to extant Anthurium; however, their relationship to the genus is still unresolved. A third fossil leaf type from Cerrejón is recognized as a species of the extant genus Montrichardia, the first fossil record for this genus. These fossils inhabited a coastal rainforest ～60-58 million years ago with broadly similar habitat preferences to modern Araceae.     

Mesofossils with platanaceous affinity from the Dakota Formation (Cretaceous) in Kansas, USA

The Platanaceae holds a basal position in the phylogeny of eudicots and therefore is of great interest to angiosperm systematists. The fossil record of the family is found in strata ranging from the Cretaceous to Recent in America, Europe and Asia. The research on the Platanaceae in the Dakota Formation can be traced back to 19th century; however, mesofossils of reproductive organs of the Platanaceae were never reported in the Midwest of North America before. This paper reports several specimens of Friisicarpus (Platanaceae) from the Dakota Formation in Kansas, USA. It complements the existing fossil records, and provides more information on reproductive biology of the family. The comparison with similar fossils from eastern North America and Europe provides some hints on biostratigraphy of the Cretaceous.     

Allonia decandra: Floral remains of the tribeHamamelideae (Hamamelidaceae) from Campanian strata of southeastern USA

A single flower, detached anthers with in situ pollen grains, and isolated seeds from Campanian strata (Upper Cretaceous) of Georgia, southeastern USA, document the presence of plants assignable toHamamelidaceae in the Upper Cretaceous. The fossil flower is actinomorphic, pentacyclic and pentamerous. Irregular sepals are preserved as lobes of the floral cup, and petals are narrow, with parallel margins. The androecium has two whorls of functional stamens. Anthers are tetrasporangiate, dehisce through two valves, and have strongly elongate connective protrusions which converge over the center of the flower. The organizational and architectural features of the fossil document its affinity within subtribeLoropetalinae (Hamamelideae, Hamamelidoideae). Cladistic phylogenetic analyses using parsimony were conducted to explore the relationships between the fossil flower and extant genera of the tribeHamamelideae. The strict consensus of the four most parsimonious trees showsHamamelideae andLoropetalinae as well-supported monophyletic taxa. The fossil flower is clearly included within theLoropetalinae, and is placed as sister taxon to the southeastern Asian genusMaingaya. The occurrence of fossils assignable toLoropetalinae during the Campanian documents the existence ofHamamelidaceae with a level of floral organization and character evolution equivalent to that of extant genera, early in the evolutionary history of the family.     

Origin and divergence of Afro-Indian Picrodendraceae: linking pollen morphology,dispersal modes,fossil records,molecular dating and paleogeography

The pantropical Picrodendraceae produce mostly spheroidal to slightly oblate, echinate pollen grains equipped with narrow circular to elliptic pori that can be hard to identify to family level in both extant and fossil material using light microscopy only. Fossil pollen of the family have been described from the Paleogene of America, Antarctica, Australia, New Zealand, and Europe, but until now none have been reported from Afro-India. Extant pollen described here include representatives from all recent Picrodendraceae genera naturally occurring in Africa and/or Madagascar and south India and selected closely related tropical American taxa. Our analyses, using combined light microscopy and scanning electron microscopy, show that pollen of the Afro-Indian genera encompass three morphological types: Type 1, comprising only Hyaenanche; Type 2, including Aristogeitonia, Mischodon, Oldfieldia and Voatamalo; Type 3, comprising the remaining two genera, Androstachys and Stachyandra. Based on the pollen morphology presented here it is evident that some previous light microscopic accounts of spherical and echinate fossil pollen affiliated with Arecaceae, Asteraceae, Malvaceae, and Myristicaceae from the African continent could belong to Picrodendraceae. The pollen morphology of Picrodendraceae, fossil pollen records, a dated intra-familial phylogeny, seed dispersal modes, and the regional Late Cretaceous to early Cenozoic paleogeography, together suggest the family originated in the Americas and dispersed from southern America across Antarctica and into Australasia. A second dispersal route is believed to have occurred from the Americas into continental Africa via the North Atlantic Land Bridge and Europe.     

Fossil spores and pollen grains of cretaceous (Upper campanian) from Sakhalin, Russia

This paper shows fossil spores and pollen grains from Cretaceous (Upper Campanian) of Sakhalin, Russia, with scanning electron microscopy. A total of 520 palynomorph assemblages consisting of 25% spores of pteridophytes and bryophytes, 4.5% of ephedroid pollen grains, 6.5% of coniferous pollen grains, and 64% of angiospermous pollen grains were recovered in the present study. 5 genera of pteridophytes, 4 genera of gymnosperms, and 18 genera of angiosperms are described in the present study. The frequent and representative genera from the stratum areEphedripites, Liliacidites, Clavatipollenites, Tricolpites, Aquilapollenites, andAzonia. A new genus,Sciadopitipollenites, that is comparable with extantSciadopitys is proposed in the present study. Polycolpate pollen with the same exine sculpture ofClavatipollenites suggests a generic differentiation in the Chloranthaceae during the Cretaceous age. The diverse spores and pollen paleoflora shown in the present study suggests a wide diversification of angiosperms in the Upper Campanian at the eastern side of Laurasia (Aquilapollenites province).     

The Phytogeography of the extinct angiosperm Nordenskioeldia (Trochodendraceae) and its response to climate changes

Elucidating the spatio-temporal distributions of terrestrial plants is a key for interpreting the origin of distribution patterns and the tempo of intercontinental disjunction. Nordenskioeldia was distributed in eastern Asia and North America from the Late Cretaceous to the Miocene. Its fossil record provides important information on former patterns of disjunction and dispersal in the Northern Hemisphere. New specimens from the Paleocene of China allow us to further extend the history of the group and provide the impetus to review its distribution in space and time. The comparative morphological survey on fossil Nordenskioeldia found in the Paleocene sediments in both eastern Asia and North America confirms that they belonged to the same morpho-species, which indicates a close floristic continuity between both continents due to land connection available during that time. The spatio-temporal distributions of Nordenskioeldia indicate that the taxon probably expanded eastward from eastern Asia into North America by the end of Early/Middle Maastrichtian, subsequently colonized Greenland, northeastern North America and Spitsbergen in the Early Paleocene, and finally became extinct in the Miocene. The fluctuations in its northern limits took place in response to climate changes: warming from the Paleocene to the Eocene, cooling during the Eocene–Oligocene and amelioration during the Late Oligocene–Mid-Miocene.     

Upper Cretaceous sulcate pollen from the Timerdyakh Formation,Vilui Basin (Siberia)

One inaperturate and 16 monosulcate pollen types are described from the latest Campanian to earliest Maastrichtian sediments of the Vilui basin, Siberia, using both light and scanning electron microscopy, and assigned systematically when possible to modern families or subfamilies. Despite their scant occurrence, the pollen grains show considerable diversity. Two new genera have been erected. Lasioideaecidites with two species, Lasioideaecidites hessei and Lasioideaecidites bogneri, represents the earliest record of the subfamily Lasioideae (Araceae). Aristolochiacidites with one species, Aristolochiacidites viluiensis, is assigned to the subfamily Aristolochioideae (Aristolochiaceae) and represents the first fossil pollen record of the family. A new species of Liliacidites, Liliacidites goldblattii, is closely similar to pollen of Isophysioideae (Iridaceae) and/or Doryanthaceae and is the first fossil evidence of the clade that includes these closely related families. A further nine new fossil species are described (Clavatipollenites timerdyakhensis, Liliacidites palaeofritillaria, Retimonocolpites longosucatus, R. microreticulatus, R. microrugulatus, R. lysichitonoides, R. chapmaniae, Monosulcites parvus and Arecipites tyungensis) and assigned to Chloranthaceae, Araceae (Orontioideae, ?Pothoideae), Liliaceae, ?Hypoxidaceae and Arecaceae. The Hydatellaceae (Nymphaeales) may also be present in the palynoflora of the Vilui Basin, represented by pollen similar to M. rivularis Braman from the Santonian to Campanian (Upper Cretaceous) Milk River Formation, Canada.     

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.     

Mesozoic Araucariaceae: Morphology and systematic relationships

The Southern Hemisphere conifer family Araucariaceae has a very restricted present day distribution, but was more widespread in the past. The genusAraucaria is represented by good fossil material in both hemispheres as early as the Jurassic, whileAgathis is only known from the Southern Hemisphere beginning in the Cretaceous. Cuticle studies of extant araucarians have enabled accurate comparisons of fossil leaves to living taxa.Araucaria SectionBunya is represented by cones of several types in the Jurassic. In addition to these remains, a suite of araucarian cones showing affinities to several sections of the genusAraucaria have been described from England, Japan and North America. Evidence that fossil araucarian cones may have produced seeds with hypogeal germination is discussed in light of recent work on germination of extant bunya seedlings and the discovery of new fossil shoots from the Jurassic Morrison Formation of Utah.     

Phylogeny of Schisandraceae based on morphological data: evidence from modern plants and the fossil record

Schisandraceae are traditionally subdivided in two genera, Schisandra and Kadsura, based on differences in the organisation of the floral receptacle, the carpels, and the presence or absence of a ``pseudostigma'. Recently, phylogenetic analyses utilizing ITS sequence data and morphological data resulted in incongruent tree topologies, with the morphological trees suggesting monophyly of the two genera, whereas ITS trees did not resolve Schisandra and Kadsura as monophyletic clades. In the present paper we study seed morphology and leaf epidermal features of 22 species of Schisandraceae in order to provide additional data for a morphological data matrix. Seed morphological characters are highly homoplastic and do not yield further evidence for monophyly of the two genera. Instead, a number of characters appear to support sister group relationships between taxa within the genera, such as, for instance, for K. coccinea and K. scandens, both of which have large seeds along with a multi-layered mesotesta. Considering leaf epidermal characteristics, species of Kadsura were found to be consistently amphistomatic, whereas species of Schisandra are always hypostomatic. Phylogenetic analysis using the extended data matrix resulted in weakly supported Kadsura and Schisandra clades with five and four synapomorphies indicating monophyly of Kadsura and Schisandra, respectively. Fossils ascribed to Schisandraceae date back to the Late Cretaceous. These are tri-and hexacolpate pollen types displaying a combination of features found in modern Schisandraceae and partly also in Illiciaceae. Leaf remains from this period are poorly preserved and difficult to ascribe to Schisandraceae because of the lack of synapomorphies for the family. In the Early Cainozoic, leaf and seed remains from North America and Europe unambiguously belong to the family. Seeds from the Eocene of North America show some similarities to the modern Schisandra glabra from North America, while fossils from Europe show more similarities to modern Asian species.     

The genusBuxus (Buxaceae): Aspects of its differentiation in space and time

Trends of pollen grain aperture evolution and exine characters as well as characters of leaf venation, petiole and axial vascularization are briefly described and related to geographical distribution and classification ofBuxus. A review of fossil records is given. Three major taxonomic groups can be delimitated within the genus, and aspects of their relationships and chorogenesis are presented. The level of differentiation, the pattern of distribution and the fossil record speak in favour of an ancient origin of the genus.