The Miocene floras of Iceland and their significance for late Cainozoic North Atlantic biogeography

A large number of plant macrofossils from several Middle to Upper Miocene localities from Iceland have been studied. The fossil material includes four ferns and fern allies, seven conifers, and about 40 species of flowering plants. Betula islandica and Salix gruberi are described as new species. Coniferous twigs previously ascribed to the genus Sequoia are shown to belong to Cryptomeria based on macro‐morphological and epidermal features. Fossil plants from Iceland are compared with coeval fossil taxa from Europe and North America and with living plants. The main finding is that the Miocene flora of Iceland belongs to a widespread Neogene northern hemispheric floral type including plants whose representatives are restricted to East Asia, North America and to western Eurasia at the present time. Previously inferred conspicuous similarities to North American modern equivalents appear to be misleading. The type of vegetation in four plant‐bearing sedimentary formations from the late Mid Miocene to Late Miocene, the 12 Ma Brjánslækur‐Seljá Formation, the 10 Ma Tröllatunga‐Gautshamar Formation, the 9–8 Ma Skarðsströnd‐Mókollsdalur Formation, and the 7–6 Ma Hreðavatn‐Stafholt Formation, corresponds to a humid temperate broadleaved (deciduous)–coniferous mixed forest dominated by Betulaceae, Fagaceae and Acer. Changes in species composition in the sedimentary formations reflect a shift from warm temperate (Cfa climate) to cool temperate (Cfb climate) conditions from the late Mid Miocene to the latest Miocene. This shift was connected to repeated phases of extinction and colonization. Specifically, one set of thermophilic taxa including Magnolia, Liriodendron, Sassafras and Comptonia went extinct between 12 and 10 Ma, and appears to have been replaced by another set of thermophilic taxa in the 10 Ma formation (Juglandaceae aff. Pterocarya/Cyclocarya, Rhododendron ponticum type). The 9–8 and 7–6 Ma formations are characterized by taxa that migrated to Iceland from Europe, such as Fagus gussonii, Betula cristata and Pterocarya fraxinifolia type. Although there is convincing evidence that plants colonized Iceland both from North America and Europe until 12 Ma, migration in the younger formations (9–8, 7–6 Ma) is suggested to have occurred mainly from Europe. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149 , 369–417.     

Floristic turnover in Iceland from 15 to 6 Ma – extracting biogeographical signals from fossil floral assemblages

Aim This study aims to document the floristic changes that occurred in Iceland between 15 and 6 Ma and to establish the dispersal mechanisms for the plant taxa encountered. Using changing patterns of dispersal, two factors controlling floristic changes are tested. Possible factors are (1) climate change, and (2) the changing biogeography of Iceland over the time interval studied; that is, the presence or absence of a Miocene North Atlantic Land Bridge. Location The North Atlantic. Methods Species lists of fossil plants from Iceland in the time period 15 to 6 Ma were compiled using published data and new data. Closest living analogues were used to establish dispersal properties for the fossil taxa. Dispersal mechanisms of fossil plants were then used to reconstruct how Iceland was colonized during various periods. Results Miocene floras of Iceland (15–6 Ma) show relatively high floristic turnover from the oldest floras towards the youngest; and few taxa from the oldest floras persist in the younger floras. The frequencies of the various dispersal mechanisms seen in the 15‐Ma floras are quite different from those recorded in the 6‐Ma floras, and there is a gradual change in the prevailing mode of dispersal from short‐distance anemochory and dyschory to long‐distance anemochory. Two mechanisms can be used to explain changing floral composition: (1) climate change, and (2) the interaction between the dispersal mechanisms of plants and the increasing isolation of proto‐Iceland during the Miocene. Main conclusions Dispersal mechanisms can be used to extract palaeogeographic signals from fossil floras. The composition of floras and dispersal mechanisms indicate that Iceland was connected both to Greenland and to Europe in the early Middle Miocene, allowing transcontinental migration. The change in prevalence of dispersal modes from 15 to 6 Ma appears to reflect the break‐up of a land bridge and the increasing isolation of Iceland after 12 Ma. Concurrent gradual cooling and isolation caused changes in species composition. Specifically, the widening of the North Atlantic Ocean prevented taxa with limited dispersal capability from colonizing Iceland, while climate cooling led to the extinction of thermophilous taxa.     

Phylogeny, biogeography, and molecular dating of cornelian cherries (Cornus, Cornaceae): tracking Tertiary plant migration

Data from four DNA regions (rbcL, matK, 26S rDNA, and ITS) as well as extant and fossil morphology were used to reconstruct the phylogeny and biogeographic history of an intercontinentally disjunct plant group, the cornelian cherries of Cornus (dogwoods). The study tests previous hypotheses on the relative roles of two Tertiary land bridges, the North Atlantic land bridge (NALB) and the Bering land bridge (BLB), in plant migration across continents. Three approaches, the Bayesian, nonparametric rate smoothing (NPRS), and penalized likelihood (PL) methods, were employed to estimate the times of geographic isolations of species. Dispersal and vicariance analysis (DIVA) was performed to infer the sequence and directionality of biogeographic pathways. Results of phylogenetic analyses suggest that among the six living species, C. sessilis from western North America represents the oldest lineage, followed by C. volkensii from Africa. The four Eurasian species form a clade consisting of two sister pairs, C. mas-C. officinalis and C. chinensis-C. eydeana. Results of DIVA and data from fossils and molecular dating indicate that the cornelian cherry subgroup arose in Europe as early as the Paleocene. Fossils confirm that the group was present in North America by the late Paleocene, consistent with the DIVA predictions that, by the end of the Eocene, it had diversified into several species and expanded its distribution to North America via the NALB and to Africa via the last direct connection between Eurasia and Africa prior to the Miocene, or via long-distance dispersal. The cornelian cherries in eastern Asia appear to be derived from two independent dispersal events from Europe. These events are inferred to have occurred during the Oligocene and Miocene. This study supports the hypothesis that the NALB served as an important land bridge connecting the North American and European floras, as well as connecting American and African floras via Europe during the early Tertiary.     

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.     

Spread and potential host range of the invasive oak lace bug [Corythucha arcuata (Say, 1832) – Heteroptera: Tingidae] in Eurasia

1. The North American oak lace bug feeds on leaves of ‘white oaks” in its native range. In Europe, it was first discovered in northern Italy in 2000. In recent years, it has subsequently spread rapidly and population outbreaks have been observed in several European countries. In the present study, we summarize the steps of its expansion.
2. To predict its potential host range, we checked 48 oak species in 20 sentinel gardens in seven countries between 2013 and 2018.
3. In total, 27 oak species were recorded as suitable hosts; 13 of them are globally new ones, 23 out of the 29 in section Quercus (～ white oaks, an intrageneric taxonomic unit within genus Quercus), including Asian oaks, native to Japan, Korea and China, and four out of five in section Cerris (another intrageneric unit of the same genus), were accepted as hosts. None of the species in section Lobatae (red oaks) or in the Ilex group was accepted.
4. Host records were also collected in forest stands of 10 countries. We found 11 oak species that were infested. Outbreak populations were most commonly found on Quercus robur, Quercus frainetto, Quercus petraea and Quercus cerris, comprising widespread and outstandingly important oaks species in Europe.
5. Based on our findings, we conclude that suitable hosts for oak lace bug are present in most of Europe and Asia. This means that a lack of hosts will likely not restrict further range expansion.

     

Phylogenetic placement and biogeography of the North American species of Valerianella (Valerianaceae: Dipsacales) based on chloroplast and nuclear DNA

Although Valerianaceae is a relatively small group of angiosperms (ca. 350 species), sample sizes of previous phylogenetic studies have been limited and taxonomic sampling has been usually geographically biased to species from Europe and/or South America. One group that has never been included in any analyses to date is the North American representatives of Valerianella. In this study I sampled 17 additional accessions from Valerianaceae, including six of the 12 described North American Valerianella species and four additional samples of European Valerianella species. Phylogenies based on parsimony and Bayesian methods show strong support for placing the North American Valerianella nested within the European species. These analyses also found Fedia to be nested within Valerianella, making Valerianella paraphyletic, a result consistent with several previously published molecular phylogenies. Divergence times estimated from the molecular data using recently proposed Bayesian 'relaxed' clock methods, suggest Valerianella was in North America by the middle Miocene (ca. 16-14 million years ago). These dates would suggest the North American species of Valerianella arrived in the New World just prior to the North Atlantic land bridge (NALB) no longer being a viable conduit for migration. Given the large amount of uncertainty in the divergence time estimates, dispersal across the North Atlantic may not be the only viable alternative to explain the current distribution of Valerianella. However, biogeographic analyses using an explicit model based approach, strongly favor the NALB as the route of dispersal for Valerianella.     

The Oldest Cranium of <Emphasis Type="Italic">Sinomastodon</Emphasis> (Proboscidea,Gomphotheriidae), Discovered in the Uppermost Miocene of Southwestern China: Implications for the Origin and Migration of This Taxon

The origin of the Old World brevirostrine gomphotheriid taxon Sinomastodon has been debated intensively. The discovery of the oldest known Sinomastodon cranium, reported herein, supports its endemic origin and contradicts the prevalent theory of its North America origin. The new cranium was discovered from the Shuitangba locality, southwestern China, and is dated at about 6.5–6.0 Ma, corresponding to the latest Miocene. The new specimen shows distinct characters from the other species of Sinomastodon and was therefore named Sinomastodon praeintermedius, sp. nov. Newly discovered, isolated Sinomastodon-like teeth from the upper Miocene to the lower Pleistocene of southwestern China and Southeast Asia indicate a long evolution of Sinomastodon endemically. Remains of this species are frequently accompanied by those of stegodontid species. These two groups may have had a similar migration route, invading northern China and Japan during the latest Miocene, and retreating or becoming extinct from the Palearctic realm by the end of the Pliocene. The migrations of proboscideans may have been sparked by major paleoenviromental changes, i.e., the strengthened summer monsoon beginning in the late Miocene (~7–8 Ma) and global cooling due to the expansion of ice sheets from the middle Pliocene to the early Pleistocene. The new finding reveals a close relationship of the early Pliocene fauna of northern China and the latest Miocene fauna of southwestern China, and thus provides novel insight into the origin and components of Pliocene fauna in northern China.     

Plant survival in Iceland during periods of glaciation?

Aim The paper addresses the classical question of possible plant survival in Iceland during the last glacial period in the light of a palaeobotanical record from northern Iceland, spanning the period 11,300–9000 BP , including the Younger Dryas stadial. We review the Late Cenozoic fossil plant record, the past debate on glacial plant refugia in Iceland, and the evidence for ice-free areas during the Weichselian. Location The investigated lake sediment record comes from Lake Torfadalsvatn, which is situated in the northwestern part of the Skagi peninsula in northern Iceland. Methods The sediment chronology was constructed from the cccurrence of the Vedde Ash and the Saksunarvatn ash, two well-dated Icelandic tephras, together with the results from five AMS and conventional radiocarbon dates performed on bulk sediment samples. The vegetational reconstruction was based on detailed pollen analysis of the sediment sequence. Results The pollen analysis revealed that many of the taxa present in the area prior to the Younger Dryas stadial continued to produce pollen during that cold event. The more or less immediate reappearance of a few other pollen taxa at the Younger Dryas-Preboreal boundary suggests that these plants also survived, even if they did not produce sufficient pollen to be recorded during the Younger Dryas stadial. Main conclusions We conclude that the relatively high plant diversity found in high Arctic areas and present-day nunataks in Iceland and Greenland, together with the fact that many plant species were able to survive the Younger Dryas stadial on the Skagi peninsula, suggest that species with high tolerance for climate fluctuations also survived the whole Weichselian in Iceland. This conclusion is supported by recent palaeoclimatic data from ice-cores and deep-sea sediments, indicating that Icelandic climate during the last glacial was only occasionally slightly colder than during the Younger Dryas stadial.     

The Tertiary history of the northern temperate element in the northern Latin American biota

The time of origin of cool-to-cold-temperate plants of northern affinities in the Latin American biota is unsettled. Two models have been proposed-a Paleogene origin from a once widespread temperate rain forest, and a Neogene origin by introductions from the north which is best supported by new evidence. Fourteen palynofloras of Tertiary age are now available from Mexico and Central America, in addition to numerous others from the southeastern United States and northern South America. Pollen of cool-temperate plants occurs in the Eocene of southeastern United States, but not in northern Mexico, central Panama, or northern South America. In the Miocene this pollen is sparse in deposits from Mexico and Guatemala, rare in Panama, and absent from northern South America. In the Pliocene pollen representing a diverse northern temperate element of ten genera is present in the Pliocene of southeastern Veracruz, Mexico, five in northeastern Guatemala, and two (Myrica, Salix) first appear in northern South America; Alnus and Quercus are added in the Pleistocene. This north-to-south and early-to-late pattern is consistent with the appearance of highlands in southern Central America and northern South America in the Neogene, closure of the isthmian marine portal between 3.5 and 2.5 Ma (million years ago), and the late Cenozoic cooling trend evident in the O/O-based paleotemperature curve.     

Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest (Greece)

The Lesvos Petrified Forest (western Lesvos, Greece) has long been famous for its plant fossils. Recently, one proboscidean (from the Gavathas locality) and seven micromammalian species (from the Lapsarna locality) were described; these were the first animals to be found in the Early Miocene subtropical forest. For the first time, a fauna of gastropods and ectothermic vertebrates from the Lapsarna locality is now available. This fauna derives from lacustrine sediments under the pyroclastic material that contains the petrified plants. Based on fragmented mollusc remains, isolated fish pharyngeal teeth and utricular otoliths (lapilli), fragmented amphibian vertebrae and a tooth-bearing element, and reptile fragmented dentaries, teeth, osteoderms and vertebrae, the presence of eight freshwater and three terrestrial gastropod species, three freshwater cyprinid species, and two amphibian and five reptile taxa has been confirmed. Stratigraphical and radiometric data suggest an age older than 18.4 ± 0.5 Ma (latest Early Miocene), in good agreement with the faunal composition. This paper is the first report of the concurrent presence of three cyprinid fish species in a Greek Early Miocene locality, as well as the first documentation of an Early Miocene proteid amphibian in southeastern Europe. The present findings represent one of the best-documented Early Miocene gastropod and fish faunas in the Aegean/southern Balkans, thus adding to our knowledge of Early Miocene amphibians and reptiles from that region and providing valuable information on the local subtropical ecosystem.     

POLLEN MORPHOLOGY AND PLANT TAXONOMY OF WHITE OAKS IN EASTERN NORTH AMERICA

An evaluation of possible approaches to fossil oak pollen identification utilized scanning electron microscopy to examine exine-surface features of 171 collections, representing 16 Quercus subgenus Lepidobalanus species and varieties of eastern North America. Twenty qualitative pollen morphological characters were defined and tabulated for each of 217 pollen grains. The data were subjected to cluster analysis and cluster diagrams were compared with published white oak taxonomy. Pollen morphology and plant taxonomy compared well in series of the subgenus Lepidobalanus due primarily to consistency of character presence and absence within species and varieties. Pollen morphology of white oaks appears to reflect plant systematics above the species level. Use of routine SEM analysis to identify series of white oaks among fossil pollen grains likely will yield valid results.     

Selection and dispersal in a multispecies oak hybrid zone

The four western North American red oak species (Quercus wislizeni, Q. parvula, Q. agrifolia, and Q. kelloggii) are known to produce hybrid products in all interspecific combinations. However, it is unknown whether hybrids are transitory resulting from interspecific gene flow or whether they are maintained through extrinsic selection. Here, we examine cryptic hybrid structure in Q. wislizeni through a broad region including contact and isolation from three other western North American red oaks using amplified fragment length polymorphism molecular markers. All four species were simultaneously detected in the genetic background of individuals morphologically assigned to Q. wislizeni, although the contribution of Q. kelloggii was minor. In some cases, introgression was detected well outside the region of sympatry with one of the parental species. Molecular structure at the individual level indicated this was due to long-distance pollen dispersal and not to local extinction of parental species. Species admixture proportions were correlated with climatic variables and greater proportions of Q. agrifolia and Q. parvula were present in the genetic background of Q. wislizeni in sites with cooler and more humid summers, corresponding with habitat preferences of the parental species. Partial Mantel tests indicated that climate was more important than distance from pollen source in this association. Despite high levels of introgression, species integrity was maintained in some populations in close proximity to the other species, providing further support to environmental selection in determining population genetic structure. Thus, the contribution of species mixtures to population genetic structure varies across the landscape according to availability of pollen, but more importantly to varying environmental selection pressures that produce a complex pattern of hybrid and pure gene pools.     

Anatomical and developmental study of petrified Quercus (Fagaceae) fruits from the Middle Miocene, Yakima Canyon, Washington, USA

The first reported petrified acorns to show internal anatomical structure are here described from Middle Miocene (～15.6 million years old) chert of the Columbia River Basalt Group in Yakima Canyon, Washington. Quercus hiholensis Borgardt et Pigg sp. nov. is described from anatomical and morphological fruit features, as well as a little recognized anatomical feature, the umbilical complex. Acorns, each comprising a nut and its cupule, are up to 15.3 mm long and 18.8 mm wide with helically arranged, imbricate, tuberculate cupule scales. They show basal aborted ovules, short styles, broad stigmas, and lack grooves in their cotyledons. These characters and the developmental pattern seen in these fossil acorns demonstrate that Q. hiholensis conforms to genus Quercus (Fagaceae), subgenus Quercus, section Quercus (the white oaks). The correspondence of Q. hiholensis to the modern section Quercus reveals that the derived floral and fruit characters that distinguish section Quercus within the genus had evolved by the Middle Miocene.     

First report of rodents from the late Hemphillian (late Miocene) Zwiebel Channel and a revised late Neogene biostratigraphy/biochronology of the Sand Draw area of Nebraska

A new late Hemphillian (late Miocene) rodent assemblage is reported from Zwiebel Channel, a channel cut into underlying Ash Hollow Miocene sediments along Sand Draw, Brown County, Nebraska. This locality extends the temporal range of rodent history in the Sand Draw area. A new biostratigraphic hypothesis proposes that previously described assemblages with Ogmodontomys are older than those with Ophiomys, as is the case in the Meade Basin of southwestern Kansas. Consequently, two Pliocene temporal zones are recognised. Based on a phylogenetic analysis of Ophiomys, rodent biostratigraphy, and paleomagnetic profiles, Sand Draw assemblages with Ogmodontomys are considered to have been deposited about 3.0–2.8 Ma, while those with Ophiomys were laid down between about 2.8–2.5 Ma. The 1.6 Ma date previously suggested for Ophiomys parvus from Froman Ferry, Idaho is probably too young; it is more likely that O. parvus became extinct in Idaho prior to the North American Microtus immigration event at about 2.0 Ma, inhabiting the Snake River basin until around 2.2 Ma.     

Subglacial refugia in Iceland enabled groundwater amphipods to survive glaciations

An endemic family of subterranean amphipods (Crustacea) has recently been discovered in Iceland, in addition to a new species of a previously known amphipod family. These findings are remarkable because Iceland was covered by glaciers from about 2.6 million BP to about 10,000 BP and is isolated on the mid-Atlantic Ridge, far from the North American and European continents. We argue that there were subglacial refugia in Iceland during the Quaternary glacial period. The presence of subterranean amphipods, belonging to an old group with its present distribution mainly in North America and the Eurasian continent, indicates past contact of subterranean fresh waters of Iceland and the North Atlantic continent. The amphipods currently found in Iceland may have been present in Greenland at 40 Ma, when the precursor of Iceland drifted together with the hot spot (Iceland plume) from Greenland.     

The biogeography of aquatic macrophytes in North America since the Last Glacial Maximum

Aim To document the post‐glacial migration of the major aquatic macrophytes of North America. Location North America north of Mexico. Methods Aquatic macrophyte pollen were extracted from the North American Pollen Database. The modern pollen distribution was mapped and related to the climate to document the geographical and climatic constraints on these taxa. The fossil pollen were mapped at 2‐ka intervals for the past 21 ka. Results Numerous genera were present in ice‐free Alaska during the Last Glacial Maximum, and south of the Laurentide Ice Sheet in the southeast. Those taxa with the widest modern climatic ranges migrated rapidly into ice‐marginal areas, first in the west and then in the east of North America. Subsequent changes in the range and abundance were smaller. Main conclusions There were four migration routes of aquatic macrophytes during the late‐glacial and post‐glacial periods: a southward migration from Alaska between 14–13 and ka, a northern migration in the west at the same time into the ice‐free Cordilleran region, and movements east and west of Appalachia as early as 19 ka for some taxa into the lower Mississippi and into the upper Mississippi and Great Lakes by 11 ka. As the Laurentide ice sheet wasted, aquatic taxa with the broadest contemporary temperature tolerances rapidly occupied ice‐marginal environments.     

Genetic evidence for hybridization in red oaks (Quercus sect. Lobatae, Fagaceae)

? Premise of the study: Hybridization is pervasive in many plant taxa, with consequences for species taxonomy, local adaptation, and management. Oaks (Quercus spp.) are thought to hybridize readily yet retain distinct traits, drawing into question the biological species concept for such taxa, but the true extent of gene flow is controversial. Genetic data are beginning to shed new light on this issue, but red oaks (section Lobatae), an important component of North American forests, have largely been neglected. Moreover, gene flow estimates may be sensitive to the choice of life stage, marker type, or genetic structure statistic. ? Methods: We coupled genetic structure data with parentage analyses for two mixed-species stands in North Carolina. Genetic structure analyses of adults (including F(ST), R(ST), G'(ST), and structure) reflect long-term patterns of gene flow, while the percentage of seedlings with parents of two different species reflect current levels of gene flow. ? Key results: Genetic structure analyses revealed low differentiation in microsatellite allele frequencies between co-occurring species, suggesting past gene flow. However, methods differed in their sensitivity to differentiation, indicating a need for caution when drawing conclusions from a single method. Parentage analyses identifed >20% of seedlings as potential hybrids. The species examined exhibit distinct morphologies, suggesting selection against intermediate phenotypes. ? Conclusions: Our results suggest that hybridization between co-occurring red oaks occurs, but that selection may limit introgression, especially at functional loci. However, by providing a source of genetic variation, hybridization could influence the response of oaks and other hybridizing taxa to environmental change.