FOSSIL LEAVES OF DENDROPANAX FROM EOCENE SEDIMENTS OF SOUTHEASTERN NORTH AMERICA

As part of a reinvestigation of the Eocene floras from southeastern North America, a leaf form similar to some that were formerly placed in the genus Aradia has been identified as Dendropanax. Studies of the fine venation and cuticular characters have been important in this identification. This is the first report of leaf material of the genus Dendropanax in the fossil record.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: RHAMNUS MARGINATUS (RHAMNACEAE) REEXAMINED

Leaf compressions, previously assigned to Rhamnus marginatus Lesquereux, were collected from the Middle Eocene Claiborne Formation of western Kentucky and Tennessee. The leaf architecture and cuticular features of over 40 compressions were carefully examined and compared to those of many extant species of Rhamnaceae and related families as well as fossil specimens previously assigned to this taxon. This leaf type appears to belong to the Rhamnaceae, however, it conforms more closely to species of several genera in the tribe Zizypheae than to those of Rhamnus or other genera in the tribe Rhamneae. Confident assignment to any specific genus within this complex of genera cannot be made on the basis of leaf characteristics alone and would require discovery and analysis of additional vegetative and reproductive organs. Because this fossil leaf form cannot be confidently assigned to any modern genus and earlier classifications appear to be improper, this leaf type has been reassigned to the taxon Berhamniphyllum claibornense gen. et sp. nov. The transfer of this leaf form at the tribe level reaffirms the need for close examination of taxonomic determinations made by early workers.     

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.     

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.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF SOUTHEASTERN NORTH AMERICA: PHILODENDRON LEAF REMAINS

Large leaves, new to the fossil record, from the Claiborne Formation in western Tennessee have been collected and analyzed. Careful analyses of venation and cuticular anatomy indicate that these fossil leaves contain specific characters found in modern species of the genus Philodendron subgenus Meconostigma. Features of venation were taxonomically more useful than other features. Reports of fossils of the Araceae are scarce and those few fossils assigned to the family frequently have been inadequately studied. This report establishes a reliable and carefully documented occurrence of the Araceae in the fossil record, and provides information concerning the differentiation of the family in time and the distributions of the genus Philodendron.     

The Phylogenetic Affinities of Nothofagus (Nothofagaceae) Leaf Fossils based on Combined Molecular and Morphological Data

The phylogenetic placements of leaf fossils of Nothofagus (Nothofagaceae) were determined using parsimony analyses of molecular and morphological data for extant species combined with morphological data for fossils. Placement was possible for only seven of the 30 or so described fossil species of Nothofagus because only these had sufficiently good preservation of both cuticular and leaf architectural characters. In combined analyses of morphology and molecular data, leaf cuticular characters showed little homoplasy. In contrast, many architectural characters, including some leaf margin and venation characters, showed high homoplasy, making it difficult or impossible to accurately determine the phylogenetic affinities of impression fossils of this genus.     

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.     

COMPARATIVE PALEOBIOCHEMISTRY OF SOME FOSSIL AND EXTANT FAGACEAE

A Fagus-like leaf fossil (cuticular compression) with an attached fruit, differing from any known Fagus species (fossil or extant) or other fagoid taxa, has been discovered from the Miocene Clarkia Lake deposits of northern Idaho. Because of its unusual morphology (especially the fruit) the fossil taxon has been described as a new genus and species, Pseudofagus idahoensis Smiley and Huggins. The successful previous use of paleobiochemistry in studies of fossil taxa from the Miocene Succor Creek Flora of Oregon suggested that chemical data might help clarify the taxonomic affinities of Pseudofagus. Indeed, examination of the chemistry of the fossil, Pseudofagus idahoensis, and comparison with extant Fagus species and related fagoid genera indicate that: 1) based on steroid chemistry, Pseudofagus idahonesis does belong in the Fagaceae; 2) like all extant species of Fagus, the fossil lacks the tannin component, ellagic acid, which separates it from other extant fagoid genera, and 3) its simple flavonoid pigment profile places it closest to the extant North American Fagus grandifolia or the European/Eurasian Fagus sylvatica. However, the exclusive presence of an isorhamnetin (3'-methoxyquercetin) 3-0-glycoside, onocerane, and 5α-cholestane imparts a species-specific chemical character to Pseudofagus idahoensis, which also sets it apart from extant species of Fagus. While the chemistry does not decide the taxonomic level to be accorded to the fossil, it certainly supports, along with morphology and anatomy, the distinctness of Pseudofagus and its proposed relationships within the Fagaceae.     

Approaches to the identification of angiosperm leaf remains

During the past 125 years the history of early angiosperms, interpreted through the fossil leaf record has been largely an exercise in paleofloristic studies, ignoring evolution. Imprecise identifications of ancient leaves “matched” to extant genera and families have been used as the basis for reconstructions of paleocommunities and paleoclimates. However, as the result of careful morphological studies of leaf form, venation and cuticular features new insights into the evolution of angiosperms are now available. In this paper considerations are given to the usefulness and shortcomings of leaf form, venation and cuticular analysis as diagnostic tools of plant identification. Many techniques for the study of the morphology of modern and fossil leaves are included in this paper as well as tables outlining features of leaf venation and the epidermis. Careful morphological studies of leaf form (such as the venation and epidermal characters emphasized in this paper) will provide better understanding of the relationships of living angiosperms and transform the fossil leaf record into useful data that can be used to study the evolution of the angiosperms.     

A new species of Zelkova (Ulmaceae,Ulmoideae) with leaves and fruits from the Oligocene of South China and its biogeographical implications

A new fossil species of Zelkova is described from the Oligocene Ningming Formation of Guangxi, South China. Zelkova ningmingensis sp. nov. is characterized by leaves with craspedodromous venation pattern and drupaceous fruit type. The new species possesses elliptical to ovate leaves, bearing 7–12 pairs of secondary and simple toothed margin, as well as epidermal cells with straight or rounded anticlinal walls. The species is compared with extant and other fossil species hitherto reported of the genus. It is most similar to the living Zelkova schneideriana in the leaf gross morphology and epidermal characters, which may be suggested to be the ancestral type of Zelkova schneideriana. The discovery of Zelkova ningmingensis sp. nov. in Guangxi indicates that Zelkova has already existed in southern China as early as the Oligocene. In combination with Zelkova material from the Oligocene of Europe, it can be inferred that Eurasian Zelkova had begun to diversify by at least the Oligocene. Because China is the biodiversity centre of modern Zelkova, the fossil herein provides new insights into Zelkova biogeography.     

Defining the gigantopterid concept: a reinvestigation of Gigantopteris (Megalopteris) nicotianaefolia Schenck and its taxonomic implications

Reinvestigation of the first described gigantopterid plant, Gigantopteris (Megalopteris) nicotianaefolia Schenk ex Potonié, 1902 from the Permian of China, has been conducted using the original specimens documented in 1883. The application of new techniques has permitted a comprehensive review of their morphology. This study represents the first photographic record of Schenk's specimens and provides a standardised terminology to describe them. This species of Palaeozoic leaf megaphyll is characterised by its pinnate venation, with tertiary veins that anastomose, and give rise to branches that may be of the same or higher order. These anastomosed branches form a reticulum of irregular polygonal meshes, within which higher order veins anastomose and again form meshes. On examination of Schenk's original specimens only one, MB.Pb.2002/989, can be assigned to Gigantopteris nicotianaefolia, and this is herein designated the lectotype. The remaining specimens described by Schenk are reassigned to cf. G. nicotianaefolia, cf. Gigantopteris sp., Gigantonoclea sp. or cf. Gigantonoclea sp. This re‐analysis of Schenk's specimens has led to an emended diagnosis of both the genus Gigantopteris and the species G. nicotianaefolia. Current evidence suggests that Gigantopteris nicotianaefolia is restricted to the southern floral regions of Cathaysia, as typically are other species of this genus. Fertile specimens of Gigantopteris are unknown, and the status of these plants as a natural group relies upon a combination of their complex foliar physiognomy, stratigraphic range, and geographical distribution. This study indicates that the genera GigantopterisGigantonoclea form the basis of the gigantopterid concept. These two genera share characters of leaf morphology including megaphylls with eucamptodromous venation across which extends a continuous lamina, and higher order veins, third order or above, that arise from the secondary veins and anastomose to form complex meshes. These genera are distinguished in that the venation of Gigantopteris is far more complex than that of Gigantonoclea, with a greater number and complexity of vein orders, the penultimate and ultimate of which form meshes within meshes, and of these the finest may terminate in blind endings.     

A new species of Eretmophyllum (Ginkgoales) from the Middle Jurassic of Turpan-Hami Basin,Xinjiang, China

A few specimens previously assigned to Eretmophyllum, based on the gross leaf morphology, lack cuticular structures. In this paper, we describe a new species of Eretmophyllum from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China, based on fossil leaves preserved as compressions, and characterized by paddle shapes with conspicuous petioles, retuse apices, entire margins and two veins dichotomously branched from near the base. Leaf amphistomatic, the cuticles are composed of stomatal and non-stomatal bands. Based on a detailed comparison of the gross morphology and cuticular structures with previously reported fossils, Eretmophyllum hamiensis n. sp. is erected. The presence of Eretmophyllum indicates that Middle Jurassic climate in the Turpan-Hami Basin was likely warm and humid. On the basis of the geological history and geographical distributions of Eretmophyllum, we conclude that the genus probably originated in Europe and migrated from Siberia to northwestern China during the Middle Jurassic.     

THE SYSTEMATIC POSITION OF EOGINKGOITES

Eoginkgoites is a fan-shaped, imparipinnate leaf with a short rachis and long petiole. It was first described from the Upper Triassic Newark Group of Pennsylvania by Bock who assigned the fossil to the Ginkgoales. The fossil has also been found in the Upper Triassic Chinle Formation in Utah and Arizona and in the Newark Group in North Carolina. Investigation of the well-preserved specimens found in the Chinle Formation shows that the leaf has anastomosing venation, a marginal vein and paracytic (syndetocheilic) stomata. These characters indicate that the leaf is bennettitalean and Eoginkgoites is reassigned to the Bennettitales although its shape is perplexing. Eoginkgoites may be an important index fossil to the lower Upper Triassic (middle Carnian) rocks of North America.     

THE PLATYCARYA PERPLEX AND THE EVOLUTION OF THE JUGLANDACEAE

We report on the leaves, fruits, inflorescences, and pollen of two fossil species in the genus Platycarya. The association of these dispersed organs has been established by their repeated co-occurrence at a large number of localities, and for two of the organs (fruit and pistillate inflorescence, and pollen and staminate inflorescence) by apparent organic attachment of compression fossils. Each of the two species can be distinguished by characteristics of all the known megafossil organs. We also review the fossil record of dispersed platycaryoid fruits and inflorescences, recognizing three additional species of Platycarya and two of Hooleya. Two of the fossil Platycarya species are morphologically very different from the living Platycarya strobilacea Sieb. et Zucc., but they show the diagnostic features of the genus. Hooleya is a generalized member of the Platycaryeae that is probably close to the ancestry of Platycarya. The two Platycarya species known from multiple organs provide a remarkable example of mosaic evolution in which fertile and foliar structures have attained different levels of morphological specialization. The leaves, often considered the most plastic of plant organs, retain several features that are otherwise seen only in the Engelhardieae. These similarities in leaf architecture between the fossil Platycarya species and Engelhardieae are advanced features for the Juglandaceae, and thus indicate a sister-group relationship between the two lines. In contrast to the leaves, the fruits, inflorescences, and pollen of the fossil Platycarya species are almost as specialized as those of the extant P. strobilacea and bear little resemblance to the same structures in other genera of the family. The morphology, taphonomy, sedimentary setting, and geographic and stratigraphic distribution of three of the fossil platycaryoid species suggest that they were wind-dispersed, early successional plants that grew in thickets. This habit is retained by Platycarya strobilacea and is typical of many of the amentiferae (e.g. Myricaceae, Betulaceae). The r-selected life-history pattern of the Platycarya line may well have contributed to its low diversity through geologic time.     

Bennettitalean leaf cuticle fragments (here Anomozamites and Pterophyllum) can be used interchangeably in stomatal frequency‐based palaeo‐CO2 reconstructions

Abstract: Bennettites are an abundant and frequently well‐preserved component of many Mesozoic fossil floras, often playing an important ecological role in flood plain vegetation communities. During a recent study focusing on stomatal indices of Triassic–Jurassic fossil plants, it became evident that the leaf fragments of two bennettite genera Anomozamites Schimper (1870) emend. Harris (1969) and Pterophyllum Brongniart (1825) display a significant overlap of leaf shape as well as cuticular characters. Owing to the preference of recognition of single taxa (ideally species) for the stomatal method, we use a database of 70 leaf fragments of Anomozamites and Pterophyllum compressions from five isotaphonomic Late Triassic sedimentary beds of Astartekløft in East Greenland to test whether leaf and cuticle fragments of the two genera can be separated using a range of quantitative and qualitative morphological and statistical analyses. None of the observed characters – including stomatal frequencies – could be applied to separate the fragments of the two genera into well‐defined groups. Our results therefore indicate that fragmented material and dispersed cuticles cannot be utilized to distinguish between Anomozamites or Pterophyllum at the genus level, but that instead these cuticle fragments may be used interchangeably as stomatal proxies. Classification of fossil leaves into either of these genera is thus only possible given adequate preservation of macro‐morphology and is not possible based solely on cuticle morphology. We suggest that this large inter‐ and intra‐generic morphological variation in both leaf and cuticle traits within Anomozamites and Pterophyllum may be related to the bennettites’ role as understory plants, experiencing a range of micro‐environmental conditions, perhaps depending mainly on sun exposure. Based on the results obtained in this study, we conclude that Anomozamites and Pterophyllum cuticle fragments can be employed interchangeably in palaeo [CO₂] reconstructions based on the stomatal method, thus potentially annexing a plethora of bennettitalean fossil plant material as CO₂ proxies, including dispersed cuticles.     

A RECORD OF PAULOWNIA IN THE TERTIARY OF NORTH AMERICA

Smiley , Charles J. (Macalester Coll., St. Paul, Minn.) A record of Paulownia in the Tertiary of North America. Amer. Jour. Bot. 48(2): 175–179. Illus. 1961.—Paulownia, an eastern Asiatic genus in the family Scrophulariaceae, has been identified from fossil leaves in the Ellensburg flora of Washington. Age of these fossils is late Miocene (Barstovian) to early Pliocene (Clarendonian). They are nearly identical to leaves of the living P. tomentosa (Thunberg) Steudel, a deciduous tree now growing in temperate parts of China and one that has escaped from cultivation in eastern United States. Identification is based on comparative leaf morphology. Modern and fossil Paulownia leaves may be distinguished from those of such other genera as Buettneria, Catalpa, and Cercis by secondary and tertiary venation. Central Washington environment during this time was well suited to the growth and reproduction of Paulownia, judging from such fossil associates as Liquidambar, Nyssa, Passiflora, Persea, Rhododendron, Ulmus, and Zelkova. Deciduous habit, present temperate environment, possible early Tertiary record in Alaska, later Tertiary records in middle latitudes of North America and Europe, and present occurrence in China indicate that Paulownia was once widely distributed over the northern hemisphere. Secular climatic trends toward cooler conditions since the Eocene appear to have resulted in the southward migration of Paulownia as a member of the Arcto-Tertiary Geoflora. Subsequent extinctions in Europe and North America may have resulted from Pleistocene glacial climates, and from barriers to further migrations to more southern latitudes.     

POLLEN MORPHOLOGY AND THE RELATIONSHIPS OF CIRCAEASTER,OF KINGDONIA,AND OF SARGENTODOXA TO THE RANUNCULALES

The pollen of three monotypic genera, Circaeaster, Kingdonia, and Sargentodoxa has been examined by light and scanning electron microscopy and in the case of the last genus, also by transmission electron microscopy. The type of tectum found in Circaeaster and Kingdonia, derivations of a compound layer of striae, has a restricted distribution in the Order Ranunculales. Of 64 genera examined in this order only six had species with a similar tectum. They include Achlys, Epimedium, Jeffersonia, and Vancouveria of the Berberidaceae s.l., the controversial Hydrastis, and Trollius of the Ranunculaceae. Circaeaster and Kingdonia have been considered as related since both have rare and primitive vegetative characteristics, the most notable being open dichotomous leaf venation. They are probably best treated as a ditypic family, Circaeasteraceae. The pollen of Sargentodoxa, especially the structure of the exine, closely resembles that of the Lardizabalaceae. However, the fruits of Sargentodoxa have been considered to be distinct from those of the Lardizabalaceae, suggesting that it be treated as a separate, but closely allied, family.     

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.     

A STUDY OF LEAF COMPRESSIONS OF KNIGHTIOPHYLLUM FROM EOCENE DEPOSITS OF SOUTHEASTERN NORTH AMERICA

A study was made of the gross morphological characters and cuticular features of several fossil leaves of Knightiophyllum wilcoxianum Berry. These fossil leaves were collected from Eocene deposits exposed in three clay pits in western Tennessee. This investigation refutes the relationship of Knightiophyllum wilcoxianum to the modern genus Knightia and the family Proteaceae as suggested by E. W. Berry.