Cenozoic migration of a desert plant lineage across the North Atlantic.

Previous paleobotanical work concluded that Paleogene elements of the sclerophyllous subhumid vegetation of western Eurasia and western North America were endemic to these disjunct regions, suggesting that the southern areas of the Holarctic flora were isolated at that time. Consequently, molecular studies invoked either parallel adaptation to dry climates from related ancestors, or long-distance dispersal in explaining disjunctions between the two regions, dismissing the contemporaneous migration of dry-adapted lineages via land bridges as unlikely. We report Vauquelinia (Rosaceae), currently endemic to western North America, in Cenozoic strata of western Eurasia. Revision of North American fossils previously assigned to Vauquelinia confirmed a single fossil-species of Vauquelinia and one of its close relative Kageneckia. We established taxonomic relationships of fossil-taxa using diagnostic character combinations shared with modern species and constructed a time-calibrated phylogeny. The fossil record suggests that Vauquelinia, currently endemic to arid and subdesert environments, originated under seasonally arid climates in the Eocene of western North America and subsequently crossed the Paleogene North Atlantic land bridge (NALB) to Europe. This pattern is replicated by other sclerophyllous, dry-adapted and warmth-loving plants, suggesting that several of these taxa potentially crossed the North Atlantic via the NALB during Eocene times.

Urticaceae leaves with stinging trichomes were already present in latest early Eocene Okanogan Highlands, British Columbia, Canada.

PREMISE: Paleontologists use tooth form to assess diets of fossil mammals. Plants would also be expected to adapt their morphology to respond to herbivory. Fossil nettle leaves with definitive stinging trichomes (tribe Urticeae, family Urticaceae) are described from the early Eocene upland lacustrine floras of the Okanogan Highlands, British Columbia, Canada. This is the first report of stinging trichomes in the fossil record. Their occurrence in western North America at a time of major large herbivorous mammal radiation suggests they acted, as they do today, as a deterrent for mammal herbivory. METHODS: Fossil leaf compressions and extant leaves were photographed with standard methods. Focus-shift stacking was used to layer photos of the fossil leaves. RESULTS: Urticaceous fossil leaves from the Okanogan Highlands greatly resemble their modern relatives in leaf morphology and particularly in both stinging and nonstinging trichomes. Nettles are common components of the flora of the Volcanoes National Park in Rwanda. This region is used as a modern analogue for the Okanogan Highlands, based on comparable elevation, equable conditions that host both similar floras and large folivores. CONCLUSIONS: Nettles in tribe Urticeae (Urticaceae) producing leaves with stinging and nonstinging trichomes were already present in the early Eocene of western North America at a pivotal time during the early radiation of modern mammalian herbivore groups. They offer tantalizing evidence of a selective response that plants may have developed to protect themselves from the evolving mammalian herbivores of that time.

New evidence of the reproductive organs of Glossopteris based on permineralized fossils from Queensland, Australia. II: pollen-bearing organ Ediea gen. nov.

Ediea homevalensis H. Nishida, Kudo, Pigg & Rigby gen. et sp. nov. is proposed for permineralized pollen-bearing structures from the Late Permian Homevale Station locality of the Bowen Basin, Queensland, Australia. The taxon represents unisexual fertile shoots bearing helically arranged leaves on a central axis. The more apical leaves are fertile microsporophylls bearing a pair of multi-branched stalks on their adaxial surfaces that each supports a cluster of terminally borne pollen sacs. Proximal to the fertile leaves there are several rows of sterile scale-like leaves. The pollen sacs (microsporangia) have thickened and dark, striate walls that are typical of the Arberiella type found in most pollen organs presumed to be of glossopterid affinity. An examination of pollen organs at several developmental stages, including those containing in situ pollen of the Protohaploxypinus type, provides the basis for a detailed analysis of these types of structures, which bear similarities to both compression/impression Eretmonia-type glossopterid microsporangiate organs and permineralized Eretmonia macloughlinii from Antarctica. These fossils demonstrate that at least some Late Permian pollen organs were simple microsporophyll-bearing shoot systems and not borne directly on Glossopteris leaves.

Rolled liverwort mats explain major Prototaxites features: Response to commentaries.

In volume 97(2) of the American Journal of Botany (pp. 268-275), we published an article entitled "Structural, physiological, and stable carbon isotopic evidence that the enigmatic Paleozoic fossil Prototaxites formed from rolled liverwort mats". Here, we respond to a letter and a commentary on our article in the present issue, welcoming this opportunity to continue the scientific dialogue about an issue that has long been stimulating and controversial. For the reader's benefit, we first briefly describe the recent scholarly context of our article.

Structural, physiological, and stable carbon isotopic evidence that the enigmatic Paleozoic fossil Prototaxites formed from rolled liverwort mats.

New structural, nutritional, and stable carbon isotope data may resolve a long-standing mystery-the biological affinities of the fossil Prototaxites, the largest organism on land during the Late Silurian to Late Devonian (420-370 Ma). The tree trunk-shaped specimens, of varying dimensions but consistent tubular anatomy, first formed prior to vascular plant dominance. Hence, Prototaxites has been proposed to represent giant algae, fungi, or lichens, despite incompatible biochemical and anatomical observations. Our comparative analyses instead indicate that Prototaxites formed from partially degraded, wind-, gravity-, or water-rolled mats of mixotrophic liverworts having fungal and cyanobacterial associates, much like the modern liverwort genus Marchantia. We propose that the fossil body is largely derived from abundant, highly degradation-resistant, tubular rhizoids of marchantioid liverworts, intermixed with tubular microbial elements. Our concept explains previously puzzling fossil features and is consistent with evidence for liverworts and microbial associates in Ordovician-Devonian deposits, extensive ancient and modern marchantioid mats, and modern associations of liverworts with cyanobacteria and diverse types of fungi. Our interpretation indicates that liverworts were important components of Devonian ecosystems, that some macrofossils and microfossils previously attributed to "nematophytes" actually represent remains of ancient liverworts, and that mixotrophy and microbial associations were features of early land plants.

Fruits of icacinaceae (tribe iodeae) from the late paleocene of Western north america.

The Icacinaceae occur pantropically today, but are well represented by fossil fruits of the warm Early Middle Eocene, when tropical plants that currently occupy low latitudes were more widely distributed in higher latitudes. Members of this family are first known in the Late Cretaceous; however, fossil fruits of tribe Iodeae are quite rare before the Eocene. In this paper we describe the first formally recognized Late Paleocene icacinaceous taxa from western North America. We name two new species of Icacinicarya based on anatomically preserved fruits and establish a new genus, Icacinicaryites, for impressions with a strong similarity to Icacinicarya that lack anatomical preservation. These new records from the Almont/Beicegel Creek flora in North Dakota and several localities in Wyoming, Colorado, and Montana complement records known from the Early Eocene of England and document an increased diversity of Iodeae and related forms in the Paleogene of western North America.

New evidence of reproductive organs of Glossopteris based on permineralized fossils from Queensland, Australia. I. Ovulate organ Homevaleia gen. nov.

This study describes Homevaleia gouldii H. Nishida, Pigg, Kudo et Rigby gen. et sp. nov., an ovule-bearing glossopterid organ, based on a combination of recently collected permineralized specimens from the Late Permian Homevale Station locality in the Bowen Basin of Queensland, Australia, and on previously studied material from the 1977 Gould and Delevoryas study. Homevaleia, which resembles the compression-impression genus Dictyopteridium, is an inrolled megasporophyll with a distinct keel that bears numerous (over 70) stalked ovules on its adaxial surface. Ovules are small, oval, with an elaborate mesh-like structure that is developed from the outermost integumentary layers. Specimens interpreted as representing different developmental stages show there is an apparent interrelationship between megagametophyte development and the opening of the surrounding fertile structure for pollination. Together, new information provided by this material enables better understanding of glossopterid reproductive structure and its function in one distinctive form.

Comparative infructescence morphology in Altingia (Altingiaceae) and discordance between morphological and molecular phylogenies.

Altingia (Altingiaceae) is a tropical to subtropical Asian genus of lowland trees for which 5-15 species have been recognized. Morphological diversity, particularly of the mature infructescence, has been poorly known, especially for species with relatively localized and narrow distributions, and our understanding of Altingia has lagged behind that of its close temperate relative Liquidambar (sweet gum). In this contribution, mature infructescence structure, at the levels of anatomy, morphology, and micromorphology, and some distinctive inflorescence features, are described for five recognized species of Altingia, some for the first time. In the phylogenetic framework of both morphology and molecules, characters of Altingia contrast with those of Liquidambar and suggest that character evolution within Altingiaceae is at least partly related to geographic and climatic distribution. Differences in rates of evolution and morphological convergence suggest complex patterns of diversification in Altingiaceae at several different phylogenetic levels: (1) at the deep nodes, characters of the stem lineage fossil Microaltingia persist into crown group Altingiaceae, morphological stasis; (2) at the generic level, convergence within both Liquidambar and Altingia toward their respective habitats; (3) at the infrageneric level, morphological divergence in species diversification within Altingia, in response to diverse habitats of the eastern Asian subtropics; and (4) within the intercontinental disjunct species pair L. orientalis-L. styraciflua, morphological stasis.

Utility of high resolution x-ray computed tomography (HRXCT) for paleobotanical studies: an example using London Clay fruits and seeds.

High resolution x-ray computed tomography (HRXCT) was used to image pyritized fossil fruits from the Lower Eocene London Clay flora to test the utility of this technique for paleobotanical application. The combination of carbon-pyrite preservation and void spaces between fruit and seed layers within fossils provides differences in density and composition that enable excellent imaging. Fossil fruits of Palaeorhodomyrtus subangulata (Bowerbank) Reid & Chandler (Myrtaceae) were investigated in situ within their silicone fluid conservation medium, which protects these unstable fossils from oxygen and humidity. HRXCT recovers taxonomically informative anatomical and morphological detail and provides a means of nondestructive examination of delicate type materials and other important specimens. These results suggest that HRXCT will be applicable to a broad spectrum of pyritized fossils to record structural details in inherently unstable materials.

Paleoactaea gen. nov. (Ranunculaceae) fruits from the Paleogene of North Dakota and the London Clay.

Paleoactea nagelii Pigg & DeVore gen. et sp. nov. is described for a small, ovoid ranunculaceous fossil fruit from the Late Paleocene Almont and Beicegel Creek floras of North Dakota, USA. Fruits are 5-7 mm wide, 4.5-6 mm high, 10-13 mm long, and bilaterally symmetrical, containing 10-17 seeds attached on the upper margin in 2-3 rows. A distinctive honeycomb pattern is formed where adjacent seeds with prominent palisade outer cell layers abut. Seeds are flattened, ovoid, and triangular. To the inside of the palisade cells, the seed coat has a region of isodiametric cells that become more tangentially elongate toward the center. The embryo cavity is replaced by an opaline cast. This fruit bears a striking resemblance to extant Actaea, the baneberry (Ranunculaceae), an herbaceous spring wildflower of North Temperate regions. A second species, Paleoactaea bowerbanki (Reid & Chandler) Pigg & DeVore nov. comb., is recognized from the Early Eocene London Clay flora, based on a single fruit. This fruit shares most of the organization and structure of P. nagelii but is larger and has a thicker pericarp. This study documents a rare Paleocene occurrence of a member of the buttercup family, a family that is today primarily herbaceous, and demonstrates a North Atlantic connection for an Actaea-like genus in the Paleogene.

Comparative infructescence morphology in Liquidambar (Altingiaceae) and its evolutionary significance.

The sweet gum genus Liquidambar (Altingiaceae) has two species in eastern Asia, one in eastern North America, and one in western Asia. Mature infructescences are studied to provide anatomical, morphological, and micromorphological details, some of which are newly recognized. Homology is suggested between extrafloral spinose processes of L. formosana and L. acalycina, braid-like ornamentation of L. styraciflua, and broad intercarpellate areas of L. orientalis. Morphology, position, number, and the presence of similar structures in the closely related Hamamelidaceae s.s. support their derivation from sterile flowers. Morphological cladistic analysis using 43 characters supports the monophyly of Liquidambar with Altingia as its sister. The matK analysis contrastingly places Altingia sister to the L. acalycina-L. formosana clade, rendering Liquidambar paraphyletic. Discordance between morphological and matK data sets may result from both different rates of morphological evolution and convergence. Several similarities between Altingia and L. acalycina are symplesiomorphic in the morphological cladistic analysis. Microaltingia apocarpela, from the Cretaceous of eastern North America, documents the earliest known fossil divergence within Altingiaceae. The Miocene Liquidambar changii of western North America is sister to a clade of extant Liquidambar species. Consideration of this fossil evidence reveals complex intercontinental biogeographic disjunctions in Altingiaceae.

Shirleya grahamae gen. et sp. nov. (Lythraceae), Lagerstroemia-like fruits from the middle Miocene Yakima Canyon flora, central Washington State, USA.

Shirleya grahamae Pigg & DeVore gen. et sp. nov. (Lythraceae) is established for silicified fruits from the middle Miocene Yakima Canyon of central Washington State, USA. The capsules are 10 mm long × 11.5-12.5 mm wide, enclosed in a persistent floral tube and contain 5-7 locules. They are loculicidally dehiscent, fracturing into fragments and leaving the central axis free. Placentation is axile. Five to seven mature seeds are tightly packed per locule, often with several smaller seeds. Seeds are winged, anatropous, and narrowly attached subapically to the central axis, curving basally and radially within the fruit. They are up to ∼4.6 mm long × 1.9 mm wide, with a small, triangular embryo cavity and a prominent distal wing. The inflated wing is filled with a bilobed parenchymatous pad of tissue with a central cavity. Shirleya grahamae is assigned to the Lythraceae, and is most similar to Lagerstroemia, based on the synapomorphies of distally winged seeds and revolute cotyledons. Shirleya differs from Lagerstroemia in seed arrangement, and pericarp and wing anatomy. This study provides the first anatomical information about a Miocene Lagerstroemia-like fruit and documents further diversity of the Lythraceae in the Neogene of northwestern North America.

Zooidogamy in the Late Permian genus Glossopteris.

We describe details of anatomically preserved fossil glossopterid ovules from the Late Permian of Queensland, Australia, that contain several pollen tubes at various stages of releasing flagellated sperm. Each sperm is approximately 12.7 microm long and 13.9 microm wide, with a conspicuous spiral structure comprised of a series of dots that resemble the position of basal bodies of flagella aligned along the multilayered structure (MLS). This configuration is similar to the helically arranged flagella in the sperm of cycads, Ginkgo, and many pteridophytes. However, the motile gametes of Glossopteris are considerably smaller than those of Ginkgo and cycads, and more similar in size, number of basal bodies, and number of gyres in their helix to pteridophyte forms. Glossopteris thus shares the intermediate stage of motile male gamete formation and apparently that of haustorial pollen tubes with cycads and Ginkgo.

Anatomically preserved Liquidambar (Altingiaceae) from the middle Miocene of Yakima Canyon, Washington state, USA, and its biogeographic implications.

Liquidambar changii Pigg, Ickert-Bond & Wen sp. nov. (Altingiaceae) is established for anatomically preserved, middle Miocene infructescences from Yakima Canyon, Washington, USA. Specimens are spherical, ∼2.5 cm in diameter, and have ∼25-30 tightly packed, bilocular fruits per head. Fruits are 3.4-4.7 mm wide × 2.6-3.5 mm long and wedge shaped, fused at the base, and free distally. Each locule contains 1-2 mature, elongate seeds proximally and 5-9 aborted seeds of more irregular shape distally. Mature seeds are 1.5 mm long × 1.2 mm wide, elongate, and triangular transversely, with a slight flange. Seeds have a seed coat for which three zones can be well defined, a uniseriate outer palisade layer, a middle region of isodiametric cells comprising most of the integument, and a uniseriate inner layer of tangentially elongate cells lining the embryo cavity. Liquidambar changii is most similar to the eastern Asian L. acalycina H.-T. Chang on features of infructescence, fruit, and seed morphology and quite unlike the North American L. styraciflua L. and other species. Such a close relationship between these two species supports a Beringian biogeographic track between eastern Asia and western North America during the Miocene. Previous phylogenetic and allozyme analysis of modern Liquidambar demonstrates a close relationship between North American-western Asian taxa and suggests a North Atlantic biogeographic track in the middle Miocene. Together, these biogeographic tracks underscore the complexity of the biogeographic history of the Altingiaceae in the Northern Hemisphere throughout the Neogene.

ANATOMICALLY PRESERVED GLOSSOPTERIS STEMS WITH ATTACHED LEAVES FROM THE CENTRAL TRANSANTARCTIC MOUNTAINS, ANTARCTICA.

Stems and buds of Glossopteris skaarensis Pigg and buds of G. schopfii Pigg from the Permian Skaar Ridge locality in the central Transantarctic Mountains, Antarctica demonstrate the first anatomically preserved glossopterids known with stem/leaf attachment. Stems of G. skaarensis are 1-12 mm in diameter ( = 3.1 mm) with a broad pith, poorly defined primary xylem, and a zone of secondary xylem up to 6 mm thick. Pycnoxylic wood conforming to Araucarioxylon Kraus is composed of tracheids with uni- to biseriate oval to hexagonal bordered pits on radial walls, uniseriate rays one to a few cells high, and cupressoid to taxodioid cross-field pitting. Stems have a narrow zone of secondary phloem, aerenchymatous cortex with scattered sclereids, and sometimes a narrow periderm. Two wedge-shaped leaf traces each bifurcate to form four strands in the base of each petiole. Small axillary branches are vascularized by double branch traces that fuse at the margin of the main axis. Buds of G. skaarensis have leaves with narrow lateral laminae and a thickened midrib containing a wide lacuna, delicate vascular strands, and a prominent hypodermis. In contrast, buds of G. schopfii have uniformly thick leaves with prominent, circular vascular bundle sheaths. These anatomical details are used to reconstruct individual types of glossopterid plants, providing new information toward understanding the ecology and evolution of this important group of Permian seed plants.