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.     

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.     

Cretaceous and Eocene fossils of the rare extant genus Synneuron Lundstrom (Diptera: Canthyloscledidae): evidence of a true Pangean clade

The first two fossil species of the canthyloscelid genus Synneuron are described based on compression wings. Synneuron eomontana sp. nov. is described from the Middle Eocene Coal Creek Member of the Kishenehn Formation, in the USA, and Synneuron jelli sp. nov. is described from the Lower Cretaceous Koonwarra Fossil Bed of the Korumburra Group, in Australia. The wings are illustrated and compared to the extant species of the genus, to species of the three other recent genera of Canthyloscelidae and to an anisopodid. A phylogenetic analysis of the relationships between the species of Synneuron was performed. The Eocene fossil S. eomontana appears as sister of the pair of recent Holarctic species of the genus, while the Australian Cretaceous species S. jelli is sister of the clade with the species of Synneuron of the northern hemisphere. The sister group of Synneuron is the canthyloscelid clade (Hyperoscelis + Canthyloscelis), for which a middle Jurassic fossil is known. At the early Cretaceous, Gondwana was already separated from Laurasia and the disjunction between the species of Synneuron in Australia and the northern hemisphere clade of the genus suggest a true pangeic origin for the genus. The biology of the canthyloscelid larvae is shaped by its trophic specialization—xylosaprophagous. This suggests that the transition from the Pangean Jurassic gymnosperm-dominated forests to the late Cretaceous angiosperm-dominated forests may be related to the low recent diversity of Synneuron or of the canthyloscelids in the world—and maybe to the extinction of the genus in the southern hemisphere. This major turnover of the vegetation type along the Cretaceous may be also somehow related to the complete extinction of other groups of flies strictly associated with gymnosperms, as may be the case of the lower brachyceran family Zhangsolvidae. This speculation needs additional corroboration from other groups, that will become available with the combination of systematics, paleontology and biogeographical information of different early Cretaceous clades.     

A new tree fern stem,Heilongjiangcaulis keshanensis gen. et sp. nov., from the Cretaceous of the Songliao Basin,Northeast China: a representative of early Cyatheaceae

A new genus and species of Cretaceous Cyatheacean tree fern, Heilongjiangcaulis keshanensis gen. et sp. nov., is erected for several permineralized stems collected at the Keshan County in Songliao Basin, Heilongjiang Province, Northeast China. The new taxon is characterized by a dictyostelic, erect stem with dense multicellular scales and surrounded by persistent petiole bases and adventitious roots. The stem contains a central pith lacking medullary bundles, which is surrounded by a dictyostele, and the cortex externally. Each meristele of the dictyostelic ring is enclosed by a sclerenchyma sheath. The pith and cortex are parenchymatous. The proximal petiole bases present a frond trace composed of numerous meristeles, arranged in 1 abaxial and 2 adaxial arcs, with internally projecting bundles on the upper and lateral sides. The feature combination of the new genus is nearly identical to the anatomical structures of modern scaly genera of the Cyatheaceae apart from the absence of medullary bundles. It is interpreted as a primitive representative of early Cyatheaceae, that closely resembles the modern scaly genera, which suggests that in the Cretaceous, the tree ferns in this family were already in possession of most of the anatomical characteristics observed in extant taxa. The fossil records of the stems, petioles, and spores indicate that during the Jurassic and Cretaceous, the Northeastern region of Asia may have been one of the distribution centers of early Cyatheaceae.     

Tsuga shimokawaensis,a new species of permineralized conifer leaves from the Middle Miocene Shimokawa Group,Hokkaido, Japan

Silicified pinaceous leaves from the late Middle Miocene bed of Shimokawa Town, central Hokkaido, Japan are described asTsuga shimokawaensis sp. nov. Comparisons with leaves of extant species ofTsuga show that the new species is assigned to sect.Tsuga, resembling the extant speciesT. heterophylla in having usually one-cell-layered hypodermis, andT. sieboldii in having mesophyll cells between the resin canal and the hypodermis. From cladistic analyses we infer thatTsuga shimokawaensis is positioned the basal in sect.Tsuga and that sect.Tsuga comprises four species groups.     

The oldest species of the genus <Emphasis Type="Italic">Glaphyrus</Emphasis> Latr. (Coleoptera: Scarabaeoidea: Glaphyridae) from the Mesozoic of China

Glaphyrus ancestralis sp. nov. is described from the Yixian Formation (Upper Jurassic or Lower Cretaceous). The species is not only one of the earliest records of the family Glaphyridae but also the oldest representative of an extant genus of the family.     

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.     

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.     

BOTANICAL AFFINITIES OF SOME TRIPROJECTACITES FOSSIL POLLEN

During Late Cretaceous to Oligocene times, fossil pollen of the Triprojectacites group (also known as Aquilapolles or triprojectates), comprised a temporally and environmentally distinctive element of palynofloras in eastern Asia and western North America. Several species of this group serve as biostratigraphic index fossils for this interval. Using electron microscope and numerical analyses, primarily of North American triprojectate pollen, it is possible to recognize the presence of three distinct subgroups. One group, corresponding to the fossil genus Mancicorpus, has no morphologically close modern representative. Characters resembling those in Santalaceae (Santalales, Rosidae) occur in the second group, which is represented by a previously underscribed fossil triprojectate genus. The third group contains retipilate, isopolar pollen and strioreticulate, isopolar pollen. Forms exhibiting the latter morphology are commonly assigned to the genus Integricorpus, while the retipilate morphology characterizes another underscribed genus. This third group may have some phylogenetic connection to Apiaceae (Apiales, Asteridae). Pollen of some other extant families exhibits triprojectate features, although no close fossil representative can be presently identified. The triprojectate morphology is thus interpreted in modern and fossil forms as resulting from convergence rather than close phylogenetic relationships at the group level.     

Normapolles flowers of fagalean affinity from the Late Cretaceous of Portugal

 Exquisitely preserved, charcoalified fossil flowers with in situ pollen of the Normapolles-type from the Late Cretaceous of Portugal are described and a new genus and species of Fagales, Normanthus miraensis, are established. Floral organization and structure of floral organs were studied with scanning electron microscopy and microtome sections. Flowers are actinomorphic, epigynous, and pentamerous; the perianth is simple; stamens alternate with tepals; pollen is oblate and vestibulate; the exine is thick and the tectum is scabrate-microgranulate; the gynoecium is bicarpellate and unilocular; the fruits are probably one-seeded. Comparisons with extant taxa demonstrate that N. miraensis shares many similarities with Fagales and in particular with Betulaceae. However, it is not identical with any extant taxon and cannot be included in any extant family. The combination of characters found in the fossil flowers is congruent with wind-pollination syndromes present in many extant angiosperms and clearly indicates wind-pollination of N. miraensis. Received June 13, 2000 Accepted September 28, 2000     

The identification of fossil angiosperm pollen and its bearing on the time and place of the origin of angiosperms

Studies in the 1970's reporting the occurrence of fossil pollen types in the Cretaceous, coupled with surveys of extant pollen morphology of primitive flowering plants, laid the foundation for proposing a Lower Cretaceous origin of angiosperms. Over the last 30 years, morphological, ultrastructural, and ontogenetic studies of both extant and fossil pollen have provided an array of new characters, as well as greater resolution in defining character polarities. Moreover, a range of fossil pollen types exhibiting angiosperm characters occur in low frequency within Triassic and Jurassic sediments. The pollen data provide evidence of a pre-Cretaceous origin of angiosperms. Speciation and extinction rates were likely equal during the Triassic and Jurassic, resulting in the paucity of angiosperm pollen types from different geographic areas in the Atlantic – South American/African rift zone. It was not until the Lower Cretaceous that origination rates exceed extinction rates, resulting in the subsequent diversification of angiosperms and the origin of the eudicots.     

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).     

Species of the water-fern megaspore genus Molaspora from a Cenomanian deposit in western France: occurrence,sporoderm ultrastructure and evolutionary relationships

The partially reticulate sculpture of Molaspora aspera sp. nov., a marsileaceous megaspore from a Cenomanian deposit in western France, distinguishes it from other species of Molaspora. An acrolamella entirely surrounds and obscures a small tetrad scar, a feature that has been demonstrated hitherto within members of the genus only in M. fibrosa. It was also encountered for the first time in M. lobata, with which the new species is associated in the same French mesofossil assemblage. The ultrastructure of the sporoderm of M. aspera is similar to that of M. lobata, but differs particularly in that the inner epispore is markedly thicker and may also contain numerous large, homogeneous spherules or, alternatively, holes of comparable dimensions and only a few small spherules. It is possible that these are a response to some hostile bacterial or other activity when the developing sporoderm was partially permeable. The cavity replacing part of the epispore in one of the specimens, and in the specimen of M. lobata examined, may be a preservational feature or have served to increase buoyancy of the spore in water. Molaspora lobata is very similar to megaspores of fossil and extant Regnellidium, but M. aspera bears some resemblance to other members of extant Marsileaceae and certain species of Cretaceous Arcellites, although there are significant differences between them. This suggests that Molaspora is a heterogeneous taxon embracing megaspores produced by water ferns of more than one natural genus, of which only Regnellidium has survived to the present day.     

Early Cretaceous floral structures and in situ tricolpate‐striate pollen: New early eudicots from Portugal

Five new taxa with affinities to extant lineages that diverged at an early stage from the main line of eudicot evolution are established from the Early Cretaceous (late Aptian or early Albian) Vale de Agua locality, Portugal. Staminate flowers of Lusistemon striatus and pistillate flowers of Lusicarpus planatus are unisexual without rudiments of the opposite gender. They are linked by the association of an unusual pollen type found in situ in the stamens and adhering to the stigmatic surface. The staminate flower, Lusistemon striatus, is composed of six stamens subtended by small perianth parts. The arrangement of the stamens is difficult to ascertain, but their variable sizes suggests a spiral arrangement. Pollen found in situ is tricolpate and striate with densely‐spaced, sparsely diverging and anastomosing muri that are aligned more or less parallel to the polar axis. The muri have a conspicuous supratectal ornamentation of fine transverse ridges. The granular infratectal layer forms an indistinct internal reticulum. The foot layer is thin. Pollen is closely similar to dispersed grains from the Aptian of Egypt described as STRIOTRI‐SEGMUR. It also resembles pollen of the dispersed pollen genus Rutihesperipites, as well as some dispersed pollen assigned to Striatopollis. Pistillate flowers of Lusicarpus planatus consist of a bicarpellate, syncarpous gynoecium borne on a short stalk. The styles are bent outwards and expose the double‐crested stigmatic regions on their ventral sides. The only organ preserved besides the gynoecium is a lateral scale‐like organ at the base of the stalk. Pollen of the same type found in Lusistemon striatus occurs on the stigmatic surface of the carpels. Comparisons with extant taxa demonstrate that Lusistemon and Lusicarpus share many characters with early diverging groups of eudicots, in particular Buxaceae. In addition to the Lusistemon‐Lusicarpus flowers, the Vale de Agua samples also contain three other pistillate reproductive structures that may be related to early diverging lineages of eudicots. Silucarpus camptostylus has a bicarpellate and syncarpous gynoecium with two styles; Valecarpus petiolatus and Aguacarpus hirsutus have tricarpellate gynoecia that are distinguished from each other in the shape and extension of the stigma as well as other details.     

From near extinction to diversification by means of a shift in pollination mechanism in the gymnosperm relict Ephedra (Ephedraceae,Gnetales)

Pollination in gymnosperms is usually accomplished by means of wind, but some groups are insect‐pollinated. We show that wind and insect pollination occur in the morphologically uniform genus Ephedra (Gnetales). Based on field experiments over several years, we demonstrate distinct differences between two Ephedra species that grow in sympatry in Greece in pollen dispersal and clump formation, insect visitations and embryo formation when insects are denied access to cones. Ephedra distachya, nested in the core clade of Ephedra, is anemophilous, which is probably the prevailing state in Ephedra. Ephedra foeminea, sister to the remaining species of the genus, is entomophilous and pollinated by a range of diurnal and nocturnal insects. The generalist entomophilous system of E. foeminea, with distinct but infrequent insect visitations, is in many respects similar to that reported for Gnetum and Welwitschia and appears ancestral in Gnetales. The Ephedra lineage is well documented already from the Early Cretaceous, but the diversity declined dramatically during the Late Cretaceous, possibly to near extinction around the Cretaceous–Palaeogene boundary. The clade imbalance between insect‐ and wind‐pollinated lineages is larger than expected by chance and the shift in pollination mode may explain why Ephedra escaped extinction and began to diversify again.     

ULTRASTRUCTURE AND DEVELOPMENT OF MESOZOIC POLLEN: CLASSOPOLLIS

Because of its complexity one of the most unusual fossil pollen types is the genus Classopollis. Grains of this broadly defined Mesozoic taxon range from the late Triassic into the Cretaceous (Turonian), and include forms that are spherical with a subequatorial rimula. On the proximal pole is a trilete mark, and on the distal surface a thin area in the sporoderm termed the cryptopore. Ultrastructural studies of Classopollis have been completed on grains extracted from the pollen cone Classostrobus comptonensis collected from the Lower Cretaceous Wealden beds on the Isle of Wight, England. The sporoderm consists of clearly defined nexine and sexine components, with the mature nexine composed of approximately 20 electron dense lamellae, each about 10 nm thick. The sexine consists of four (S_1–4) easily recognizable layers, with the most prominent zone formed of coarse, inwardly-tapering elements. The S₂ layer is uniformly thickened, except in specialized areas (e.g., trilete, rimula, cryptopore) where it becomes thin. The remaining wall layers include spinules that ornament the surface and a uniform series of small lacunae associated with the spinule bases. The presence of orbicules and a complex system of membranes associated with the grains extracted from less mature cones provides an opportunity to trace some developmental stages in Classopollis sporoderm ontogeny, and to compare these stages with those of selected extant pollen types. The functional significance of the infrastructure in Classopollis pollen is discussed.     

Elatides sandaolingensis n. sp. (Cupressaceae sensu lato) — A new fossil conifer with cones from the Middle Jurassic of Xinjiang,northwestern China

Here we describe a new conifer fossil, Elatides sandaolingensis Z.X. Wang and B.N. Sun n. sp., from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. The materials consist of compressions represented by well-preserved leafy shoots, pollen cones, and seed cone. Leaves are characterized by long triangular shapes, with straight apex and entire margins, and two stomatal bands on the abaxial surface. Pollen cones are terminally disposed on the ultimate leafy shoots, borne singly or in clusters. Pollen sacs are long-oval shaped, with three pollen sacs fused together. Pollen grains are spherical and have small germinal papilla and few wrinkles. The seed cone is oblong, with more than 35 helically arranged bract-scale complexes, which are characterized by long-oval shape and triangular apex. Compared with the extant nine genera of Taxodiaceous Cupressaceae in the morphology of seed cone and pollen cones, the present fossil consistently shares many characteristics with the extant genus Cunninghamia, but differs in other aspects. After being compared with the reported fossil records of the fossil plants, the current species is found to be different from any known species; thus, the present fossil is referred to as a new species of Elatides. From the similarity between the present fossil and Cunninghamia, it can be inferred that there may be a genetic relationship between these two genera. Additionally, the new species has thin cuticles and slightly sunken stomata, which can provide evidence indicating that the climate of the Turpan-Hami Basin in Middle Jurassic might have been warm and humid. By studying the geological history and geographical distribution of Elatides, it can be inferred that this genus may have originated in Switzerland, and it was migrated from Xinjiang to North and South China through the Ural Mountains.     

Atoposaurid crocodyliforms from the Khorat Group of Thailand: first record of <Emphasis Type="Italic">Theriosuchus</Emphasis> from Southeast Asia

We describe a partial crocodilian skull from the Mesozoic non-marine sediments of the Khorat Plateau Sao Khua Formation (Berriasian-Barremian) in northeastern Thailand and assign it to Theriosuchus grandinaris sp. nov. An isolated dentary from the Phu Kradung Formation (latest Jurassic–Early Cretaceous) is also tentatively assigned to the genus Theriosuchus, and an isolated tooth from the Khok Kruat Formation (Aptian-Albian) may belong to this genus. The Thai fossils represent the first unambiguous evidence of presence of Theriosuchus outside Europe. Its occurrence in Thailand increases the known diversity of neosuchian crocodyliforms from Southeast Asia and suggests that Atoposauridae had a wide geographical distribution from the Late Jurassic to the Early Cretaceous.