Inflorescence bracts of fossil and extant Tilia in North America,Europe, and Asia: patterns of morphologic divergence and biogeographic history

The enlarged inflorescence bract diagnostic of extant Tilia has an extensive Tertiary fossil record in the Northern Hemisphere. Diversity of bract morphology, and the extent of adnation between peduncle and bract, is reviewed for fossil and extant species of Tilia. An extinct type of bract with an orbicular outline and palmate venation is documented by the fossil species Tilia circularis (Chaney) comb. nov. from the early Oligocene of Oregon and is designated Type A. Living species of the genus have elongate bracts with predominately pinnate venation that are borne in two basic configurations: Type B, with the peduncle fused only to the extreme base of the bract lamina, as in extant Tilia endochrysea Hand.-Mzt. of southern China; and Type C with the peduncle fused medially along the basal one-third of the bract lamina, as in most extant species. Bracts of Type B were widely distributed in the Tertiary of western North America (late Eocene to Miocene) and Europe (early Miocene to Pliocene), while those of Type C are known in the fossil condition only from the middle and late Tertiary of Asia and Pliocene of Europe. The bracts of T. circularis, like those of type B, are borne on relatively long stalks and have the peduncle fused only at the extreme base. The fossil record supports recognition of the following characters as apomorphic in Tilia bract evolution: bracts sessile, peduncle adnate to the upper surface of the bract, and pinnate bract venation.     

Exploring the Formation of a Disjunctive Pattern between Eastern Asia and North America Based on Fossil Evidence from Thuja (Cupressaceae)

Thuja, a genus of Cupressaceae comprising five extant species, presently occurs in both East Asia (3 species) and North America (2 species) and has a long fossil record from Paleocene to Pleistocene in the Northern Hemisphere. Two distinct hypotheses have been proposed to account for the origin and present distribution of this genus. Here we recognize and describe T. sutchuenensis Franch., a new fossil Thuja from the late Pliocene sediments of Zhangcun, Shanxi, North China, based on detailed comparisons with all living species and other fossil ones, integrate the global fossil records of this genus plotted in a set of paleomaps from different time intervals, which show that Thuja probably first appeared at high latitudes of North America in or before the Paleocene. This genus reached Greenland in the Paleocene, then arrived in eastern Asia in the Miocene via the land connection between East Asia and western North America. In the late Pliocene, it migrated into the interior of China. With the Quaternary cooling and drying, Thuja gradually retreated southwards to form today’s disjunctive distribution between East Asia and North America.     

Microsatellite DNA and behavioural studies provide evidence of host-mediated speciation in Myzus persicae (Hemiptera: Aphididae)

The fossil record of mammals records a major interchange of northern and southern faunas in the New World, upon closure of the Panamanian isthmus approximately 3 Mya, termed the Great American Biotic Interchange (GABI). Due to their poor preservation in the fossil record, the degree of participation of birds in this interchange remains largely unknown. A phylogeny for wrens of the genus Campylorhynchus (Aves: Passeriformes) was reconstructed using DNA sequences from the mitochondrial control region and cytochrome b gene. This phylogeny, in combination with biogeographical inference and molecular clock methods, allows estimates of the importance of Late Pliocene interchange to the history of the group. Biogeographical reconstructions and divergence date estimates suggest that the genus began diversification in North America prior to closure of the Panamanian isthmus, consistent with a hypothesized North American origin for the family Troglodytidae. These reconstructions are consistent with pre-GABI dispersal of at most a single Campylorhynchus lineage into South America, with subsequent dispersal of additional lineages, probably across the fully formed isthmus. Increased sampling of avian taxa with widespread New World distributions will continue to clarify the timing and direction of continental interchange.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 687–702.     

EVOLUTION OF HORNS IN UNGULATES: ECOLOGY AND PALEOECOLOGY

(1) The savanna ungulate faunas of the North American Miocene were broadly similar to those of present-day East Africa in terms of overall morphological and taxonomic diversity. However, the predominant ungulates of the African faunas are bovids, which possess bony horns that are primitively sexually dimorphic in their occurrence. The predominant ungulates of the North American Tertiary were equids, camelids and oreodonts, which all lacked horns. A limited number of horned ruminants were present, but these were largely Miocene immigrants from Eurasia. Horns were also absent from the large-bodied herbivores in the endemic faunas of South America and Australia. (2) The absence of horns in equids and tylopod artiodactyls is unlikely to be due to genetic insufficiency. Bony horns were present in brontotheres, which were closely related to equids, and in protoceratids, which were closely related to camelids. Nasal horns were present in one oreodont genus. (3) Studies on living ungulates show that a strong correlation exists between habitat type, feeding behaviour, social behaviour and morphology. It is possible to use the morphological remains of extinct ungulates to reconstruct the types of feeding and social behaviour, and to use the distribution of morphologies and body sizes in a community of mammals, in conjunction with geological and paleobotanical evidence, to reconstruct the type of habitat. (4) The importance of the post-Eocene climatic changes to the history of mammalian evolution is stressed. Continents at higher latitudes have become increasingly seasonal in terms of temperature and rainfall since the equable global conditions of the early Tertiary. Savanna mosaic were the predominant biome in North America by the early Miocene, and in Eurasia by the middle Miocene. Living temperate-latitude species of ungulates may not be a reliable guide for the assessment of the interrelationship between behaviour and morphology in an evolutionary perspective, as their behaviour may have been recently adapted to a habitat type that has only been in existence since the Pleistocene. (5) The primitive condition in eupecorans and protoceratids is the absence of horns, with the presence of large sabre-like canines in the males. The first horned members of these divisions had horns in the males only. Small present-day antelope, where horns may also be present in the females of the species, are probably secondarily small. (6) Horns were acquired independently in ruminant artiodactyls at least three times, and a maximum number of seven times is not unlikely. In each case, horns first appeared at a critical body weight of about 18 kg, and in correlation with a change in habitat from closed to open woodland. (7) Horns in living ruminants are associated with territorial defence by males holding exclusive feeding and reproductive territories in woodland habitats. Such behaviour in present-day antelope is correlated with a body size of greater than 15 kg and a folivorous diet. It is argued that horns evolved in ruminant artiodactyls on the adoption of this type of territorial behaviour once the critical combination of body size, diet and habitat type had been attained in their evolution from small, essentially frugivorous, forest-dwelling animals. (8) Perissodactyls never evolved sexually dimorphic bony horns of the type seen in ruminant artiodactyls. This is because their foraging and digestive strategies necessitate a larger daily intake of food. In a woodland habitat they were never able to adopt a feeding area small enough to make exclusive territory maintenance an economical proposition. Territory holding in male perissodactyls is seen, but under the opposite conditions of habitat to territorial behaviour in ruminant artiodactyls. (9) Study of the morphology and paleoecology of oreodonts suggests that they were woodland herd-forming browsers with exclusively folivorous diets. They probably had some forestomach fermentation, but did not chew the cud. Similar studies of Tertiary camelids suggest that they were predominantly selective browsers eating herbage at a low level in open country and formed mixed-sex feeding groups. These combinations of feeding and social behaviour suggest a more open structure of the mid-Tertiary habitat in North America than in Eurasia. (10) Studies of the behaviour and morphology of living members of the Ruminantia, and of the morphology and paleoecology of their fossil ancestors, suggest that they were primitively tree browsers living in closed woodland habitats. Such habitats were abundant in the Old World, but in limited supply in North America during the Oligocene, where the protoceratids were the only ungulates to parallel the eupecoran type of feeding and social behaviour. South America appears to have had an even more open habitat in the Oligocene than North America, and no parallel to the eupecorans was seen amongst the indigenous ungulates. The radiation of the Bovidae into open grassy habitats in the Pliocene may have been dependent on the immigration of grazing equids into the Old World. (11) I conclude that there was a difference in habitat structure between North America and the Old World during the Tertiary. The food resources in North America were more widely dispersed, and this may have been the result of the trees being more widely spaced. A possible causal mechanism for this was the stable land mass of the North American continent during the Tertiary, resulting in a more continental climate, with a more severe effect of the post-Eocene seasonality on the vegetation. The faunal record of the two continents also implied a greater density of trees in the Old World. (12) Thus most endemic North American ruminants did not evolve horns because, at the critical combination of body size and diet seen in the evolution of horns in the Old World ruminants, the dispersal of the food resources within the vegetation was too great for an effective home range to be maintained as an exclusive territory. (13) Attention is drawn to the dangers of constructing evolutionary stories about living animals without primary reference to the fossil record to see if the hypotheses are upheld, and of assuming that fossil animal communities can be made to fit models of existing communities.     

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.     

鹅观草属的几个新组合

本文报道了禾本科鹅观草属的三个种级新组合和四个变种级新组合。即大丛鹅观草Roegneria magnicaespis (D.F.Cui)L.B.Cai;新疆鹅观草Roegneria sinkiangensis(D.F.Cui)L.B.Cai;阿尔泰鹅观草Roegneria altaica(D.F.Cui)L.B.Cai;短芒鹅观草Roegneria glaberrima var.breviarista (D.F.Cui)L.B.Cai;林缘鹅观草Roegneria mutabilis var.nemoralis (D.F.Cui)L.B.Cai;多花鹅观草Roegneria abolinii var.pluriflora (D.F.Cui)L.B.Cai和曲芒鹅观草Roegneria tschimganica var.glabrispicula (D.F.Cui)L.B.Cai。     

A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.     

Phylogeny and biogeography of the flowering plant genus Styrax (Styracaceae) based on chloroplast DNA restriction sites and DNA sequences of the internal transcribed spacer region

Phylogenetic relationships within the flowering plant genus Styrax were investigated with DNA sequence data from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA) and with chloroplast DNA restriction site data from the genes trnK, rpoC1, and rpoC2. The data sets from each genome were analyzed separately and in combination with parsimony methods. The results strongly support the monophyly of each of the four series of the genus but provide little phylogenetic resolution among them. Reticulate evolution may at least partly explain discordance between the molecular phylogenetic estimates and a prior morphological estimate within series Cyrta. The historical biogeography of the genus was inferred with unweighted parsimony character optimization of trees recovered from a combined ITS and morphological data set, after a series of combinability tests for data set congruence was conducted. The results are consistent with the fossil record in supporting a Eurasian origin of Styrax. The nested phylogenetic position of the South American members of the genus within those from southern North America and Eurasia suggests that the boreotropics hypothesis best explains the amphi-Pacific tropical disjunct distribution occurring within section Valvatae. The pattern of relationship recovered among the species of section Styrax ((western North America + western Eurasia) (eastern North America + eastern Eurasia)) is rare among north-temperate Tertiary forest relicts. The monophyly of the group of species from western North America and western Eurasia provides qualified support for the Madrean-Tethyan hypothesis, which posits a Tertiary floristic connection among the semiarid regions in which these taxa occur. A single vicariance event between eastern Asia and eastern North America accounts for the pattern of relationship among intercontinental disjuncts in series Cyrta.     

New fossil fruits of Carya (Juglandaceae) from the latest Miocene to earliest Pliocene in Tennessee,eastern United States

Fossil fruits including nuts and associated husk valves of a new species of Carya (Juglandaceae) are described from the latest Miocene to earliest Pliocene in northeastern Tennessee, eastern United States. The husk valves are elliptic, 1.2–4.5 mm thick, with a convex exterior face and a concave interior face; the nuts are globose to ovoid in shape, smooth and longitudinally ribbed on exterior surface, with a short protruding apex and a slightly 4-angled base; inner ribs, lacunae and primary septa are well-developed, while secondary septa are absent or weakly developed. The combination of these carpological characteristics clearly shows a close resemblance to the genus Carya in Juglandaceae. Detailed comparisons of carpological morphology and anatomy indicate that the present fossil taxon is different from both living and most other fossil species of the genus, and therefore warrants the designation of a new fossil species, Carya tennesseensis Huang et al., sp. nov. Carya tennesseensis displays a carpological similarity to C. ventricosa from the late Oligocene to early Pliocene in Europe, suggesting a potential species exchange of the genus between Europe and southeastern North America during the late Neogene. The new fossil species represents one of the few fruit fossil species of Carya from its modern distribution range in southeastern North America. It provides crucial information for better understanding the rapid diversification of the genus from the late Miocene to early Pliocene, and the origin and establishment of today's Carya biodiversity in this region.     

Neogene Corylopsis seeds from eastern Tennessee

A new fossil species of Corylopsis (Hamamelidaceae), C. grisea Quirk & Hermsen sp. nov, based on seeds from the early Pliocene Gray Fossil Site (GFS), eastern Tennessee, USA, is described. The assignment of the seeds to Hamamelidaceae, subfamily Hamamelidoideae, is based on the overall size of the seeds, smooth testa, lack of a seed wing, and the presence of a terminal hilar scar. The assignment to the genus Corylopsis is based on seed size as well as the presence of a hilar facet, in addition to the hilar scar. Although Corylopsis persists only in East Asia today, its fossil record indicates that the genus was widespread across the Northern Hemisphere in the past. Prior to its discovery at GFS, Corylopsis was only known from the Paleogene in North America. The presence of C. grisea at GFS extends the fossil record of Corylopsis in North America to the Neogene and reinforces the interpretation of GFS as a forested refugium that provided a relatively moist, equable environment where subtropical to warm temperate plants could persist during a time of cooling and drying in the continental interior of North America. Its presence provides additional evidence for the biogeographic connection between the GFS paleoflora and the modern flora of eastern Asia.     

Multilocus molecular systematics and evolution in time and space of Calligrapha (Coleoptera: Chrysomelidae,Chrysomelinae)

Calligrapha (Coleoptera: Chrysomelidae) is a genus with species present in most of the American continent, from the Arctic polar circle to the Pampas in Argentina. In its current concept, the genus comprises some 80 species, but the diagnosis of the genus is problematic, based on a combination of potentially symplesiomorphic character states. In this study, we investigate the largest taxonomic sample of Calligrapha diversity to date (43 species) using a phylogenetic perspective based on more than 6000 molecular characters from eight genes (four mitochondrial and four nuclear) for a systematic evaluation of the genus. The analyses also include thirteen species in the closely related Zygospila (currently a subgenus of Zygogramma) to assist the systematic delimitation of Calligrapha. Partitioned and total evidence phylogenetic trees were additionally used for molecular clock analyses and dating based on standard mtDNA evolutionary rates, and for likelihood‐based inference of ancestral areas. Calligrapha and Zygospila are reciprocally paraphyletic, and our interpretation of taxonomic stability merges both taxa into a larger genus Calligrapha which plausibly originated in the dry steppes of southern North America in the Late Miocene. The genus includes a minimum of five strongly supported lineages which initially diversified in the Pliocene, fully congruent with expectations from morphology, but of uncertain mutual relationships. Only two of these lineages dispersed to South America: the group of C. polyspila right at the time of the final closure of the Isthmus of Panama in the Early Pliocene and the group of C. argus only in recent times, well in the Pleistocene. The most species‐rich lineage of Calligrapha, associated to trees and shrubs typical of riverine and lacustrine environments (as opposed to herbaceous steppe plants, generally Malvaceae and Asteraceae, for most other groups) diversified and spread in North America in the Late Pliocene. The ecological shift to a stable habitat spreading in the continent due to climate change is hypothesized as one possible explanation for the evolutionary success of this group.     

壳斗科的地质历史及其系统学和植物地理学意义

在收集整理现有壳斗科化石资料的基础上,讨论了壳斗科及其各属的起源时间、地史分布和地史 演替过程以及这些化石资料在系统学和植物地理学上的意义。白垩纪尚无壳斗科可靠的大化石记录, 微化石需要进一步研究才能确定亲缘关系以及古新世壳斗科已经分化出两个类群。从以上这些事实推 论壳斗科起源于白垩纪晚期,而壳斗科现代各属出现的时间应不晚于古新世。最早发现的壳斗科化石和现代栗亚科和水青冈亚科在形态结构上非常相似,这一事实表明,壳斗科分为两个亚科的观点更接近客观事实。在水青冈亚科中,三棱栎类的化石最早出现;在栎属中,青冈亚属更接近祖先类群;在地史中全缘栎类较具齿栎类出现早,粗齿的落叶栎类出现最晚。三棱栎属、栲属和石栎属的化石在老第三纪出现于北美和欧洲的事实说明,北美、欧洲和东亚在老第三纪时有一个相通的壳斗科植物区系。南美的三棱栎是通过北美进入南美的。中国横断山、欧洲地中海沿岸和北美西北部有一类形态特征相似、亲缘关系相近的硬叶栎类,它们之间有相同的地质演替历史,它们现代分布边界可能就是古地中海的边界。美洲的栎类有两个来源,常绿硬叶栎类是通过古地中海沿岸而经北美-欧洲陆桥到达的,落叶栎类则是在中新世以后通过白令海峡到达的。     

Fossils of the Fagaceae and their implications in systematics and biogeography

The fossil history of the Fagaceae from China and its systematic and biogeographic implications are discussed based on revisionary studies of the fossil records. No creditable macrofossil record of the Fagaceae exists in the Cretaceous deposits and all the Cretaceous microfossil reports remain equivocal and require further study. The Paleocene fossils show the appearance and diversification of the two groups corresponding to the subfamilies Fagoideae and Castaneoideae sensu Nixon. By the Eocene, all modern genera had been present. The oldest fagaceous fossils represent subfamily Fagoideae with affinities to the extant genus Trigonobalanus. The leaf fossil genus Berryophyllum, with affinities to Quercus subg. Cyclobalanopsis, has been documented by the early Eocene and might have occurred earlier than other fossils assignable to Quercus. The appearance of evergreen sclerophyllous Ouercus with entire leaves might have occurred earlier than those with toothed leaves. Deciduous, urticoid-leaved oak fossils (Quercus subg. Quercus sect. Quercus) had not appeared until the Miocene. Fossil equivalents of Trigonobalanus, Castanopsis and Lithocarpus had occurred in Europe and North America by the early Tertiary, suggesting that continuous distributions were achieved via the northern hemisphere land bridges. Three groups of evergreen sclerophyllous oaks of apparent close phylogenetic relationships occurred in the Hengduan mountains, the Mediterranean area and northwestern North America. Their fossil forms have become dominant elements of those vegetation zones since the Miocene. A shared fossil history indicates a possible biogeographic boundary formed by the ancient Mediterranean. The evidence suggests that the oaks might arrive in North America during two distinct geologic periods: evergreen sclerophyllous entire-leaved oaks appeared by the Early Tertiary, whereas thedeciduous oaks with urticoid leaves appeared in the Late Tertiary.     

An extinct juglandaceous fruit from the Upper Palaeocene of southern England

A new genus of juglandaceous winged fruit, is described from the Reading Beds (Upper Palaeocene) of southern England. It comprises one of the earlier macrofossil records of the Juglandaceae, and is the earliest from the European Tertiary. The fruit represents an extinct genus related to the extant tribe Engelhardieae, but excluded from it by its simple unlobed bract. Cladistic analysis shows Casholdia to display generalized engelhardioid fruit morphology. It lacks the tri-lobed bract diagnostic of the Engelhardieae, and predates the first occurrence of such bracts in the fossil record. Casholdia adds to the mounting evidence indicating an early Palaeogene radiation of the Juglandaceae.     

MOLECULAR DIVERGENCE BETWEEN ASIAN AND NORTH AMERICAN SPECIES OF LIRIODENDRON (MAGNOLIACEAE) WITH IMPLICATIONS FOR INTERPRETATION OF FOSSIL FLORAS

Botanists have long been aware of the floristic similarities between eastern Asia and eastern North America. Most who have considered this classic disjunction pattern have suggested that it arose through range disruption of a flora that was once more widely distributed in the Northern Hemisphere. There is less agreement on the timing of this process, with suggestions ranging from the Paleocene to the Neogene. In this study, molecular markers from two different plant genomes were used to assess the degree of genetic divergence between the two interfertile, morphologically similar species of the genus Liriodendron, i.e., L. tulipifera and L. chinense. Resulting molecular divergence estimates were translated into approximate dates of separation, independent of evidence from the fossil record. Allozyme data (Nei's genetic identity = 0.434) suggested a divergence time of 10–16 million years before present, whereas sequence divergence in the plastid genomes (1.24%) led to an estimate of approximately 11–14 million years before present. A review of the paleobotanical literature indicated that the fossil floras that included, or might have included Liriodendron could not have survived in Beringia after the late Miocene and the onset of southward-migrating Arctic air masses on the North American continent. This interpretation suggests a minimum time of separation of approximately 13 million years before present. Thus, both molecular data sets and the paleobotanical evidence concur in suggesting a divergence time of 10–16 million years before present. Interspecific compatibility and relative morphological stasis must have, therefore, persisted from at least the late Miocene. We emphasize the need for similar studies in other genera, especially those that have both a reasonable Tertiary fossil history and extant species in mesic temperate refugia in Asia, Europe, and western as well as eastern North America.     

ORIGINS AND RELATIONSHIPS OF TROPICAL NORTH AMERICA IN THE CONTEXT OF THE BOREOTROPICS HYPOTHESIS

The boreotropics hypothesis postulates a preferential tropical biotic interchange between North America and Eurasia during the early Tertiary that was directed by Eocene thermal maxima and the close proximity of these two continental plates. This preferential interchange occurred at a time when South America was geologically and biotically isolated. A prediction of this hypothesis posits that a taxon with a present-day center of diversity in tropical North America, and with an early Tertiary fossil record from any region there, has a high probability of having sister-group relatives in the Paleotropics and derived relatives in South America. We propose a test of this prediction with phylogenetic studies of two pantropical taxa of Leguminosae that have early Tertiary North American fossil records. Our findings are consistent with the boreotropics hypothesis, and additional evidence suggests that many tropical elements in North America could be descendants of northern tropical progenitors. Ramifications of this hypothesis include the importance of integrating the fossil record with cladistic biogeographic studies, theoretical bases for recognizing tropical taxa with such disjunct distributions as Mexico and Madagascar, identification of taxa that may be most useful for testing vicariance models of Caribbean biogeography, and integrating the study of disjunct distributions in temperate regions of the northern hemisphere with those in the neo- and paleotropics.     

Fossil bananas (Musaceae): Ensete oregonense sp. nov. from the Eocene of western North America and its phytogeographic significance

Fossil seeds of Ensete, a genus presently native to Asia and Africa, have been recovered from the middle Eocene of Oregon, confirming the presence of Musaceae in the North American Tertiary. The seed of Ensete oregonense sp. nov. is operculate, with a well-defined micropylar collar, a pronounced chalazal chamber, and a wide hilar cavity. A survey of seed morphology in extant Zingiberales provides characters for distinguishing Musaceae from other families of the order, furnishes criteria for distinguishing the three extant genera of Musaceae (Musa, Ensete and Musella), and facilitates critical assessment of fossil seed remains. “Musa” cardiosperma Jain from the Cretaceous/Tertiary boundary Deccan Series of India is excluded from Musaceae (although retained in Zingiberales) on the basis of fruit and seed characters, including the lack of laticifers and absence of a chalazal chamber. We reexamined the musaceous seeds from Colombia that previously were described as Tertiary fossils (Musa enseteformis Berry, 1925) and now believe that they are recent, nonfossil remains, evidently from Ensete ventricosum, which is grown in the region where the specimens were originally obtained. In addition, a reputed fossil banana fruit from the Cretaceous of Colombia was reexamined and determined to be a concretion of nonbiological origin. Ensete oregonense is significant therefore, as the first unequivocal fossil record of Ensete and of Musaceae. Although the Musaceae are currently native only to the Old World tropics, this discovery establishes that the family was present in North America about 43 million years ago.     

The historical biogeography of sturgeons (Osteichthyes: Acipenseridae): a synthesis of phylogenetics, palaeontology and palaeogeography

Abstract. The historical biogeography of sturgeons is explored using information from palaeogeography, palaeontology and phylogenetic interrelationships. The integration of information from these diverse sources indicates that sturgeons reached a wide Laurasian distribution in the Cretaceous and Tertiary by freshwater and coastal dispersal routes across land connections and along newly forming continental margins. The fossil record also suggests a considerable degree of morphological stasis and also supports an estuarine habit, and perhaps diadromy, as an old and conserved life history trait. While a ‘centre of origin’ for sturgeons remains elusive, phylogenetic relationships indicate that diversification appears to have been associated with fragmentation of biota, and of landmasses and basins, by late Tertiary geological and climatic phenomena, such as orogeny and unequal glaciation over North America, the desiccation of central Asia and alteration of its drainages, and the formation of discrete Ponto-Caspian basins by the fragmentation of the Paratethys. Amphi-oceanic distributions of certain species (Acipenser medirostris Ayres) and sister taxa (e.g. A. oxyrhynchus Mitchill and A. sturio L.) are explained by coastal dispersal and subsequent vicariance by geological (sea-floor spreading and development of new continental margins) and climatic (Pliocene cooling) changes during the Tertiary. An hypothesis is developed for the relationships of the North American sturgeons and their potential relationships with the Siberian sturgeon A. baeri. Late Tertiary climatic and geological phenomena are hypothesized as mediators of vicariance and subsequent diversification of these acipenserids. It appears that although acipenserids are a geologically old group, the historical biogeography of surviving lineages is best explained by more recent geological and climatic changes.