Palaeobiogeography of late cretaceous belemnites of Europe

A North Temperate Realm, characterized by Belemnitellidae Pavlov, and a South Temperate Realm, characterized by Dimitobelidae WHITEHOUSE, existed throughout the Late Cretaceous, while Tethyan belemnites belonging to Belemnopseidae Naef existed only in the Cenomanian and disappeared afterwards. The North Temperate Realm may be subdivided into North European and North American Provinces. The latter province includes Greenland, Canada, the Western Interior Region of North America, and the Atlantic and Gulf coasts of North America. The belemnites from the North American Province, consisting of populations of the generaActinocamax Miller andBelemnitella d’Orbigny, are closely related to the belemnites of the North European Province and appear to have migrated from this province to North America via Greenland and Arctic Canada. The North European Province extends from Ireland to the Ural Mountains. Belemnites from this province belong to the following genera:Neohibolites Stolley,Parahibolites Stolley,Belemnocamax Crick,Actinocamax Miller,Belemnellocamax Naidin,Gonioteuthis Bayle,Belemnitella d’Orbigny,Belemnella Nowak, andFusiteuthis Kongiel. Two subprovinces within the North European Province have been recognized: the Central European and Central Russian Subprovinces. These subprovinces are well-defined in the late Coniacian-Early Campanian and are characterized by theGonioteuthis stock andBelemnitella stock, respectively. The two subprovinces are less distinct in other periods of the Late Cretaceous and may disappear completely.     

Large Shift in Symbiont Assemblage in the Invasive Red Turpentine Beetle

Changes in symbiont assemblages can affect the success and impact of invasive species, and may provide knowledge regarding the invasion histories of their vectors. Bark beetle symbioses are ideal systems to study changes in symbiont assemblages resulting from invasions. The red turpentine beetle (Dendroctonus valens) is a bark beetle species that recently invaded China from its native range in North America. It is associated with ophiostomatalean fungi in both locations, although the fungi have previously been well-surveyed only in China. We surveyed the ophiostomatalean fungi associated with D. valens in eastern and western North America, and identified the fungal species using multi-gene phylogenies. From the 307 collected isolates (147 in eastern North America and 160 in western North America), we identified 20 species: 11 in eastern North America and 13 in western North America. Four species were shared between eastern North America and western North America, one species (Ophiostoma floccosum) was shared between western North America and China, and three species (Grosmannia koreana, Leptographium procerum, and Ophiostoma abietinum) were shared between eastern North America and China. Ophiostoma floccosum and O. abietinum have worldwide distributions, and were rarely isolated from D. valens. However, G. koreana and L. procerum are primarily limited to Asia and North America respectively. Leptographium procerum, which is thought to be native to North America, represented >45% of the symbionts of D. valens in eastern North America and China, suggesting D. valens may have been introduced to China from eastern North America. These results are surprising, as previous population genetics studies on D. valens based on the cytochrome oxidase I gene have suggested that the insect was introduced into China from western North America.     

Putative source of the invasive Sirex noctilio fungal symbiont, Amylostereum areolatum, in the eastern United States and its association with native siricid woodwasps

Two genotypes of the fungal symbiont Amylostereum areolatum are associated with the invasive woodwasp Sirex noctilio first found in North America in 2004. S. noctilio is native to Europe but has been introduced to Australasia, South America and Africa where it has caused enormous losses in pine plantations. Based on nucleotide sequence data from the intergenic spacer region (IGS) of the nuclear ribosomal DNA, the A. areolatum genotypes found in North America are most similar to genotypes found in Europe, and not to genotypes from the southern hemisphere. Although two IGS strains of A. areolatum were found in North America it cannot be stated whether A. areolatum was introduced to North America from Europe once or twice based on our study. Genetic groupings formed by sequencing data were in most cases supported by vegetative compatibility groups (VCGs). Other siricid woodwasp species in the genus Sirex are native to North America. The North American native Sirex edwardsii emerging from the same tree as S. noctilio carried the same strain of A. areolatum as S. noctilio. The North American native Sirex sp. ‘nitidus’ collected outside the geographical range of S. noctilio carried a unique strain within A. areolatum. Our findings of A. areolatum in the native North American species, S. sp. ‘nitidus’, contrast with the previous view that A. areolatum was not present in North America before the accidental invasion of S. noctilio.     

On the origin of the North Atlantic redfish of the genus Sebastes (Scorpaenidae), as revealed by analysis of their parasite fauna

A comparative analysis of the parasite fauna of the Sebastes redfishes, which inhabit the North Atlantic and the North Pacific, was carried out. The relative youth of the host-parasite relationship in the North Atlantic Sebastes corroborates the hypothesis of their Pacific origin. The species attribution of the ancestor of the North Atlantic Sebastes is suggested.     

Des serpents (Reptilia,Squamata) de type nord-américain dans le Miocène Français. Évolution parallèle ou dispersion?

The French Miocene (Orleanian and Astaracian) yielded five new species of snakes referable to four genera from the Neogene or Recent of North America: Texasophis meini nov. sp., Paleoheterodon arcuatus nov. sp., Neonatrix europaea nov. sp., Neonatrix crassa nov. sp. (Colibridae) and Micrurus gallicus nov. sp. («Elapidae). It is suggested that Micrurus and the previously described boid snake Albaneryx from the French Miocene probably represent lineages that originated in North America and reached Europe by way of Asia. Paleoheterodon and Neonatrix could have originated in Asia and subsequently reached Europe on one hand and North America on the other. Texasophis may have originated in North America and followed the same route as Micrurus and Albaneryx, but an alternative hypothesis is that it originated in Asia and spread toward Europe and North America in the same manner as Paleoheterodon and Neonatrix.     

Intercontinental dispersal of Typha angustifolia and T. latifolia between Europe and North America has implications for Typha invasions

The full effects of biological invasions may be underestimated in many areas because of cryptogenic species, which are those that can be identified as neither native nor introduced. In North America, the cattails Typha latifolia, T. angustifolia, and their hybrid T. × glauca are increasingly aggressive invaders of wetlands. There is a widespread belief that T. latifolia is native to North America and T. angustifolia was introduced from Europe, although there has so far been little empirical support for the latter claim. We used microsatellite data and chloroplast DNA sequences to compare T. latifolia and T. angustifolia genotypes from eastern North America and Europe. In both species, our data revealed a high level of genetic similarity between North American and European populations that is indicative of relatively recent intercontinental dispersal. More specifically, the most likely scenario suggested by Approximate Bayesian Computation was an introduction of T. angustifolia from Europe to North America. We discuss the potential importance of our findings in the context of hybridization, novel genomes, and increasingly invasive behaviour in North American Typha spp.     

Evolutionary trends in Mio-Pliocene Leporinae, based onTrischizolagus (Mammalia, Lagomorpha)

New material ofTrischizolagus dumitrescuae from Moldova and Ukraine is described. The variation of p3 inTrischizolagus shows the gradual shift of morphotype frequencies from the ‘Hypolagus’ pattern in Turolian through the mixture of three patterns (including ‘Nekrolagus’ morphotype) in Early Ruscinian to the dominant ‘Alilepus’ pattern in the Late Ruscinian samples. These transformations took place parallel to that of the North AmericanNekrolagus. Probably North AmericanSylvilagus, Brachylagus, andRomerolagus had an North American origin fromNekrolagus, whereas Eurasiatic and AfricanOryctolagus, Caprolagus, Nesolagus, andPoelagus could have originated in the Old World fromTrischizolagus.     

Real-Time PCR Detection of Dogwood Anthracnose Fungus in Historical Herbarium Specimens from Asia

Cornus species (dogwoods) are popular ornamental trees and important understory plants in natural forests of northern hemisphere. Dogwood anthracnose, one of the major diseases affecting the native North American Cornus species, such as C. florida, is caused by the fungal pathogen Discula destructiva. The origin of this fungus is not known, but it is hypothesized that it was imported to North America with its host plants from Asia. In this study, a TaqMan real-time PCR assay was used to detect D. destructiva in dried herbarium and fresh Cornus samples. Several herbarium specimens from Japan and China were detected positive for D. destructiva, some of which were collected before the first report of the dogwood anthracnose in North America. Our findings further support that D. destructiva was introduced to North America from Asia where the fungus likely does not cause severe disease.     

Analysis of DNA sequences of six chloroplast and nuclear genes suggests incongruence, introgression, and incomplete lineage sorting in the evolution of Lespedeza (Fabaceae)

The genus Lespedeza (Fabaceae) consists of 40 species disjunctively distributed in East Asia and eastern North America. Phylogenetic relationships of all Lespedeza species and closely related genera were reconstructed using maximum parsimony, maximum likelihood, and Bayesian analyses of sequence data from five chloroplast (rpl16, rpl32-trnL, rps16-trnQ, trnL-F, and trnK/matK) and one nuclear (ITS) DNA regions. All analyses yielded consistent relationships among major lineages. Our results suggested that Campylotropis, Kummerowia, and Lespedeza are monophyletic, respectively. Lespedeza is resolved as sister to Kummerowia and these two together are further sister to Campylotropis. Neither of the two subgenera, subgen. Lespedeza and subgen. Macrolespedeza, in Lespedeza based on morphological characters, is recovered as monophyletic. Within Lespedeza, the North American clade is retrieved as sister to the Asian clade. The nuclear and chloroplast markers showed incongruent phylogenetic signals at shallow-level phylogeny, which may point to either introgression or incomplete lineage sorting in Lespedeza. The divergence times within Lespedeza and among related genera were estimated using Bayesian approach with BEAST. It is assumed that following the divergence between Kummerowia and Lespedeza in Asia in the late Miocene, the ancestor of Lespedeza diverged into the North American and the Asian lineages. The North American ancestor quickly migrated to North America through the Bering land bridge in the late Miocene. The North American and Asian lineages started to diversify almost simultaneously in the late Miocene but resulted in biased numbers of species in two continents.     

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.     

Phylogenetic relationships among New World Scrophularia L. (Scrophulariaceae): new insights inferred from DNA sequence data

The genus Scrophularia L. (Scrophulariaceae) comprises 200?C300 species, of which approximately 19 are distributed in North America and the Greater Antilles. To investigate phylogenetic and biogeographic relationships of the New World species, two intergenic spacers (trnQ-rps16 and psbA-trnH) of chloroplast DNA and nuclear ribosomal ITS were sequenced. Phylogenetic analyses revealed three distinct New World clades that correspond to their geographical distribution and are corroborated by morphological characters. Phylogenetic inference indicates the eastern American S. marilandica L. as sister to all Antillean species; for colonization of the Caribbean archipelago, a late Miocene dispersal event from the North American mainland is assumed. There is evidence for a hybrid origin of the most widespread North American species, S. lanceolata Pursh. The results further suggest that S. nodosa L. is sister to all New World and three Japanese species of Scrophularia. The latter form an Eastern Asian?CEastern North American (EA-ENA) disjunction with six New World species. We propose an eastern Asian origin for the New World taxa of Scrophularia. Divergence times estimated using a relaxed molecular clock model suggest one or more Miocene migration events from eastern Asia to the New World via the Bering Land Bridge followed by diversification in North America.     

Foliar pathogens of Populus angustifolia are consistent with a hypothesis of Beringian migration into North America

Populus angustifolia, the narrowleaf cottonwood, is considered one of two native species of Populus section Tacamahaca restricted to western North America. Efforts to construct a definitive phylogeny of Populus spp. are complicated by ancient hybridization, but some phylogenetic analyses suggest P. angustifolia is more closely related to Asian congeners than to Populus trichocarpa, the other species of Populus section Tacamahaca in western North America. Because hosts and their obligate symbionts can display parallel phylogeographic patterns, we evaluated the possibility of a Beringian migration into North America by an Asian ancestor of P. angustifolia by determining the distributions, host preferences, and, in some cases, closest phylogenetic relatives of fungal leaf pathogens of P. angustifolia. Phyllactinia populi, a common foliar pathogen on Populus spp. in Asia but unknown on P. trichocarpa, was found on P. angustifolia in multiple sites. Mycosphaerella angustifoliorum, also unknown on P. trichocarpa, was commonly collected on P. angustifolia. Conversely, many common foliar pathogens of P. trichocarpa in western North America were not found on P. angustifolia; only Melampsora×columbiana and Drepanopeziza populi were common to both Populus species. Phylogenetic analyses revealed that M. angustifoliorum was not part of the diversification of Mycosphaerella on Populus that includes all other Mycosphaerella species on Populus in North America: Mycosphaerella populicola, Mycosphaerella populorum, M. sp. 1, and M. sp. 2. The latter two undescribed species represent a newly discovered diversification of M. populorum in western North America. Overall, the leaf pathogen community of P. angustifolia is consistent with a Beringian migration into North America by the ancestor of P. angustifolia.     

A New Fishfly Species (Megaloptera: Corydalidae: Neohermes Banks) Discovered from North America by a Systematic Revision,with Phylogenetic and Biogeographic Implications

The taxonomy of Megaloptera from the Nearctic region is fairly well known and their faunal diversity has been largely surveyed, even in relatively remote regions. However, the evolutionary history of Nearctic Megaloptera is still poorly known with phylogenetic and biogeographic studies lacking. In this paper, we report a new fishfly species of the endemic North American genus Neohermes Banks, 1908, increasing the total number known of species to six. This new species (Neohermes inexpectatus sp. nov.) is currently known to occur only in California (USA) and is apparently confined to the Northern Coastal Range. The new species resembles the three Neohermes species from eastern North America based on the relatively small body size and the presence of female gonostyli 9. However, our phylogenetic analysis using adult morphological data recovered the new species as the sister species to the remaining Neohermes, which includes two species from western North America and three from eastern North America. According to the present interspecific phylogeny of Neohermes, with reconstructed ancestral areas, the initial divergence within the genus was found to take place in western North America, with a subsequent eastward dispersal. This likely lead to the modern distribution of Neohermes in eastern North America with the closure of the Mid-Continental Seaway, which separated western and eastern North America in the Mid-Late Cretaceous (100–80 MYA) and finally disappeared at the end of the Cretaceous (70 MYA). The uplift of the Cordilleran System probably accounted for the divergence between the eastern and two western Neohermes species.     

The species of Coleosporium (Pucciniales) on Solidago in North America

Species of Coleosporium (Pucciniales) are rust fungi that typically alternate between pines and angiosperms. In North America, species of Coleosporium often infect Solidago (goldenrods), although their taxonomy on these hosts is unresolved. Joseph. C. Arthur and George B. Cummins regarded these as a single species, Coleosporium solidaginis (fide Arthur) or C. asterum (fide Cummins), but later inoculation studies demonstrated the presence of more than one species, distinguishable by their aecial hosts. A more recent taxonomic study of Coleosporium found that specimens on Solidago identified as C. asterum in North America were not conspecific with the type, which is from Japan, prompting the present study. Herein, we conducted a systematic study on ca. 60 collections of Coleosporium infecting species of Asteraceae from North America using regions of ribosomal DNA and morphology of teliospores and basidia. Our data indicate at least three species of Coleosporium occur on Solidago in North America, C. solidaginis, C. montanum comb. nov., which is proposed for the taxon that has commonly been identified as C. asterum in North America, and C. delicatulum, all of which can be differentiated by morphology of their basidia. In addition, the challenges of marker selection for molecular barcoding of rust fungi is discussed.     

Integrated Fossil and Molecular Data Reveal the Biogeographic Diversification of the Eastern Asian-Eastern North American Disjunct Hickory Genus (Carya Nutt.)

The hickory genus (Carya) contains ca. 17 species distributed in subtropical and tropical regions of eastern Asia and subtropical to temperate regions of eastern North America. Previously, the phylogenetic relationships between eastern Asian and eastern North American species of Carya were not fully confirmed even with an extensive sampling, biogeographic and diversification patterns had thus never been investigated in a phylogenetic context. We sampled 17 species of Carya and 15 species representing all other genera of the Juglandaceae as outgroups, with eight nuclear and plastid loci to reconstruct the phylogeny of Carya. The phylogenetic positions of seven extinct genera of the Juglandaceae were inferred using morphological characters and the molecular phylogeny as a backbone constraint. Divergence times within Carya were estimated with relaxed Bayesian dating. Biogeographic analyses were performed in DIVA and LAGRANGE. Diversification rates were inferred by LASER and APE packages. Our results support two major clades within Carya, corresponding to the lineages of eastern Asia and eastern North America. The split between the two disjunct clades is estimated to be 21.58 (95% HPD 11.07-35.51) Ma. Genus-level DIVA and LAGRANGE analyses incorporating both extant and extinct genera of the Juglandaceae suggested that Carya originated in North America, and migrated to Eurasia during the early Tertiary via the North Atlantic land bridge. Fragmentation of the distribution caused by global cooling in the late Tertiary resulted in the current disjunction. The diversification rate of hickories in eastern North America appeared to be higher than that in eastern Asia, which is ascribed to greater ecological opportunities, key morphological innovations, and polyploidy.     

New adapiform primate of Old World affinities from the Devil's Graveyard Formation of Texas

Most adapiform primates from North America are members of an endemic radiation of notharctines. North American notharctines flourished during the Early and early Middle Eocene, with only two genera persisting into the late Middle Eocene. Here we describe a new genus of adapiform primate from the Devil’s Graveyard Formation of Texas. Mescalerolemur horneri, gen. et sp. nov., is known only from the late Middle Eocene (Uintan) Purple Bench locality. Phylogenetic analyses reveal that Mescalerolemur is more closely related to Eurasian and African adapiforms than to North American notharctines. In this respect, M. horneri is similar to its sister taxon Mahgarita stevensi from the late Duchesnean of the Devil’s Graveyard Formation. The presence of both genera in the Big Bend region of Texas after notharctines had become locally extinct provides further evidence of faunal interchange between North America and East Asia during the middle Eocene. The fact that Mescalerolemur and Mahgarita are both unknown outside of Texas also supports prior hypotheses that low-latitude faunal assemblages in North America demonstrate increased endemism by the late middle Eocene.     

A taxonomic study of campylopus schimperi and C. subulatus (Bryopsida: Dicranaceae) in North America

Campylopus schimperi is known in North America from Oaxaca, Colorado, British Columbia, Yukon, Alaska, Baffin Island, Gaspé Peninsula, and Newfoundland, and reported from Greenland. No specimens of the previously reported C. subulatus were found from North America and this species should be excluded from the North American bryophyte flora. A quantitative study and critical review of the characters distinguishing these two species indicate that leaf laminal cell shape and length, and costal anatomy are the most definitive taxonomic features. The distributional range of C. schimperi in North America suggests a close association with unglaciated areas and hypothesized glacial refugia.     

DomesticatedChenopodium of the Ozark Bluff Dwellers

Previous suggestions that prehistoric agriculturalists of the Ozark Bluff Dweller culture utilized a fully domesticated form ofChenopodium have been confirmed. Comparative examination of infructescence and fruit structure indicates that archaeological material is assignable toC. berlandieri ssp.nuttalliae, a product of Mexican agriculture. Large-fruited chenopod remains from other sites in eastern North America, often identified as those of wild species, may also belong to the Mexican domesticated form. A related, wild species of northeastern North America,C. bushianum, shows similarities to the Mexican weed-crop complex that may reflect prehistoric genetic interaction. This is the first documented report of domesticatedChenopodium from prehistoric North America.     

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.