Integrated conodont and radiolarian biostratigraphy of the upper Norian in Baoshan Block,Southwestern China

Two significant stratigraphical microfossils, conodonts and radiolarians, are usually used for the Upper Triassic chronostratigraphy. The Baoshan Block was located in eastern Tethys during the Late Triassic where the biostratigraphical data of Upper Triassic are still poorly known. We collected new samples from the Hongyan section (HY) for biostratigraphical study. This 24-m-thick section in Dabaozi Village, Baoshan City, is mainly composed of thin-layered limestones, sandstone and siltstone. The conodont fauna is referred to Sevatian 1 (late Norian), in which the species Mockina englandi, Mockina carinata and Mockina mosheri morphotype B are first recognized in the Baoshan Block, and thus eastern Tethys. The Norian radiolarian associations are first reported in the Baoshan Block, which correlate with the biozonation of North America and also that proposed for Central Japan. The radiolarian assemblages found in the analysed samples in HY section can be referred to the Sevatian Betraccium deweveri Zone. The Baoshan Block is a key area for conodont and radiolarian-based correlations between the Tethys, Japan and North American domains.     

Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera,Nymphalidae, Satyrinae)

Peña, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. (2010). Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae).—Zoologica Scripta, 39, 243–258. The diverse butterfly subtribe Euptychiina was thought to be restricted to the Americas. However, there is mounting evidence for the Oriental Palaeonympha opalina being part of Euptychiina and thus a disjunct distribution between it (in eastern Asia) and its sister taxon (in eastern North America). Such a disjunct distribution in both eastern Asia and eastern North America has never been reported for any butterfly taxon. We used 4447 bp of DNA sequences from one mitochondrial gene and four nuclear genes for 102 Euptychiina taxa to obtain a phylogenetic hypothesis of the subtribe, estimate dates of origin and diversification for major clades and perform a biogeographic analysis. Euptychiina originated 31 Ma in South America. Early Euptychiina dispersed from North to South America via the temporary connection known as GAARlandia during Eocene–Oligocene times. The current disjunct distribution of the Oriental Palaeonympha opalina is the result of a northbound dispersal of a lineage from South America into eastern Asia via North America. The common ancestor of Palaeonympha and its sister taxon Megisto inhabited the continuous forest belt across North Asia and North America, which was connected by Beringia. The closure of this connection caused the split between Palaeonympha and Megisto around 13 Ma and the severe extinctions in western North America because of the climatic changes of the Late Miocene (from 13.5 Ma onwards) resulted in the classic ‘eastern Asia and eastern North America’ disjunct distribution.     

The Phytogeography of the extinct angiosperm Nordenskioeldia (Trochodendraceae) and its response to climate changes

Elucidating the spatio-temporal distributions of terrestrial plants is a key for interpreting the origin of distribution patterns and the tempo of intercontinental disjunction. Nordenskioeldia was distributed in eastern Asia and North America from the Late Cretaceous to the Miocene. Its fossil record provides important information on former patterns of disjunction and dispersal in the Northern Hemisphere. New specimens from the Paleocene of China allow us to further extend the history of the group and provide the impetus to review its distribution in space and time. The comparative morphological survey on fossil Nordenskioeldia found in the Paleocene sediments in both eastern Asia and North America confirms that they belonged to the same morpho-species, which indicates a close floristic continuity between both continents due to land connection available during that time. The spatio-temporal distributions of Nordenskioeldia indicate that the taxon probably expanded eastward from eastern Asia into North America by the end of Early/Middle Maastrichtian, subsequently colonized Greenland, northeastern North America and Spitsbergen in the Early Paleocene, and finally became extinct in the Miocene. The fluctuations in its northern limits took place in response to climate changes: warming from the Paleocene to the Eocene, cooling during the Eocene–Oligocene and amelioration during the Late Oligocene–Mid-Miocene.     

Using ecological niche modelling to evaluate niche stability in deep time

Aim To investigate relative niche stability in species responses to various types of environmental pressure (biotic and abiotic) on geological time‐scales using the fossil record. Location The case study focuses on Late Ordovician articulate brachiopods of the Cincinnati Arch in eastern North America. Methods Species niches were modelled for a suite of fossil brachiopod species based on five environmental variables inferred from sedimentary parameters using GARP and Maxent . Niche stability was assessed by comparison of (1) the degree of overlap of species distribution models developed for a time‐slice and those generated by projecting niche models of the previous time‐slice onto environmental layers of a second time‐slice using GARP and Maxent , (2) Schoener’s D statistic, and (3) the similarity of the contribution of each environmental parameter within Maxent niche models between adjacent time‐slices. Results Late Ordovician brachiopod species conserved their niches with high fidelity during intervals of gradual environmental change but responded to inter‐basinal species invasions through niche evolution. Both native and invasive species exhibited similar levels of niche evolution in the invasion and post‐invasion intervals. Niche evolution was related mostly to decreased variance within the former ecological niche parameters rather than to shifts to new ecospace. Main conclusions Although the species examined exhibited morphological stasis during the study interval, high levels of niche conservatism were observed only during intervals of gradual environmental change. Rapid environmental change, notably inter‐basinal species invasions, resulted in high levels of niche evolution among the focal taxa. Both native and invasive species responded with similar levels of niche evolution during the invasion interval and subsequent environmental reorganization. The assumption of complete niche conservatism frequently employed in ecological niche modelling (ENM) analyses to forecast or hindcast species geographical distributions is more likely to be accurate for climate change studies than for invasive species analyses over geological time‐scales.     

Implications of early production in an invasive forest pest

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International importance of the eastern Chukchi Sea as a staging area for migrating king eiders

The evaluation of habitats used by arctic birds on migration is crucial for their conservation. We explored the importance of the eastern Chukchi Sea (ECS) as a staging area for king eiders (Somateria spectabilis) migrating between breeding areas in Siberia and western North America and wintering areas in the Bering Sea. We tracked 190 king eiders with satellite transmitters between 1997 and 2007. In late summer, 74% of satellite-tracked king eiders migrating south staged in the ECS for 13 ± 13 (SD) days between late June and early November. During spring migration, king eiders staged in the ECS between mid-April and early June for 21 ± 10 days. All instrumented birds migrating to breeding grounds in western North America (n = 62), and 6 of 11 males migrating to breeding grounds in Siberia, used this area for at least 1 week during spring migration. The importance of this staging area renders it possible that industrial development could adversely affect king eider populations in both Siberia and North America.     

The Middle‐Upper Tournaisian boundary event

The Middle‐Upper Tournaisian boundary broadly correlating with the Kinderhookian‐Osagean boundary in North America represents a level of a conspicuous lithological as well as biotic change. The decline of Chernyshinella and increased representation of eurythermal and agglutinated forms marks the end of Cherepet evolutionary cycle in calcareous foraminifers. In conodonts the decline of Siphonodella inhabiting the pelagic to hemipelagic dysphotic to aphotic environment accompanied by a widespread dispersal and diversification of nektobenthic gnathodids characterizes the end of the first Tournaisian conodont cycle. A major regression associated with important sedimentological changes marks the boundary between the middle and upper Tournaisian mega‐sequences. The above changes in fauna and lithology seem to be connected with climatic oscillations and accompanying oceanographic changes. They are a part of the broader Upper Devonian‐Lower Carboniferous cyclic scheme which appears to have been modified by a complicated interplay of cyclic and nonrecurring processes.     

Devonian Vertebrates From Colombia

Vertebrate remains are reported from the Emsian–Eifelian Floresta Formation and the Late Devonian (?Frasnian) Cuche Formation of north‐eastern Colombia. The material from the Floresta Formation is associated with a marine invertebrate fauna and includes an arthrodire and probably a rhenanid. Several vertebrate‐bearing localities are recorded from the Cuche Formation; vertebrates occur with plant remains and lingulid fragments. They include an acanthodian (Cheiracanthoides? sp.), a chondrichthyan (Antarctilamna? sp.), placoderms (Bothriolepis sp., Asterolepis? sp. and an undetermined groenlandaspidid or primitive brachythoracid arthrodire), a stegotrachelid actinopterygianand three sarcopterygians (a cosmine‐covered form tentatively referred to an osteolepidid, the porolepiform Holoptychiussp., and the rhizodontid Strepsodus? sp.). This assemblage suggests a Late Frasnian age and is surprisingly similar to Late Devonian vertebrate assemblages found in similar facies of Europe and North America, notwithstanding the presence of the Gondwanan chondrichthyan Antarctilamna?. key words: Vertebrata, Devonian, Colombia, South America, biostratigraphy, palaeobiogeography.     

The last diadectomorph sheds light on Late Palaeozoic tetrapod biogeography

Diadectomorpha is a clade of Late Palaeozoic vertebrates widely recognized as the sister group of crown-group Amniota and the first tetrapod lineage to evolve high-fibre herbivory. Despite their evolutionary importance, diadectomorphs are restricted stratigraphically and geographically, with all records being from the Upper Carboniferous and Lower Permian of North America and Germany. We describe a new diadectomorph, Alveusdectes fenestralis, based on a partial skull from the Upper Permian of China. The new species exhibits the derived mechanism for herbivory and is recovered phylogenetically as a deeply nested diadectid. Approximately 16 Myr younger than any other diadectomorph, Alveusdectes is the product of at least a 46 Myr ghost lineage. How much of this time was probably spent in Russia and/or central Asia will remain unclear until a specimen is described that subdivides this cryptic history, but the lineage assuredly crossed this region before entering the relatively isolated continent of North China. The discovery of Alveusdectes raises important questions regarding diadectomorph extinction dynamics including what, if any, ecological factors limited the diversity of this group in eastern Pangea. It also suggests that increased sampling in Asia will likely significantly affect our views of clade and faunal insularity leading up to the Permo-Triassic extinction.     

Ancient and modern colonization of North America by hemlock woolly adelgid,Adelges tsugae (Hemiptera: Adelgidae), an invasive insect from East Asia

Hemlock woolly adelgid, Adelges tsugae, is an invasive pest of hemlock trees (Tsuga) in eastern North America. We used 14 microsatellites and mitochondrial COI sequences to assess its worldwide genetic structure and reconstruct its colonization history. The resulting information about its life cycle, biogeography and host specialization could help predict invasion by insect herbivores. We identified eight endemic lineages of hemlock adelgids in central China, western China, Ulleung Island (South Korea), western North America, and two each in Taiwan and Japan, with the Japanese lineages specializing on different Tsuga species. Adelgid life cycles varied at local and continental scales with different sexual, obligately asexual and facultatively asexual lineages. Adelgids in western North America exhibited very high microsatellite heterozygosity, which suggests ancient asexuality. The earliest lineages diverged in Asia during Pleistocene glacial periods, as estimated using approximate Bayesian computation. Colonization of western North America was estimated to have occurred prior to the last glacial period by adelgids directly ancestral to those in southern Japan, perhaps carried by birds. The modern invasion from southern Japan to eastern North America caused an extreme genetic bottleneck with just two closely related clones detected throughout the introduced range. Both colonization events to North America involved host shifts to unrelated hemlock species. These results suggest that genetic diversity, host specialization and host phylogeny are not predictive of adelgid invasion. Monitoring non‐native sentinel host trees and focusing on invasion pathways might be more effective methods of preventing invasion than making predictions using species traits or evolutionary history.     

Variation in spring migration routes and breeding distribution of northern pintails Anas acuta that winter in Japan

In North America, spring migration routes and breeding distribution of northern pintails Anas acuta vary because some individuals opportunistically nest at mid‐latitudes in years when ephemeral prairie wetlands are available, whereas others regularly nest in arctic and sub‐arctic regions where wetland abundance is more constant. Less was known about migration routes and breeding distribution of pintails in East Asia. From 2007–2009 we marked 198 pintails on their wintering areas in Japan with satellite transmitters to: 1) document spring migration routes and summer distribution, 2) evaluate migratory connections and breeding season sympatry with North American pintails, and 3) determine if pintails used the same migration routes in fall as in spring. Most pintails (67%) migrated to the Kamchatka or Chukotka peninsulas in eastern Russia either directly from Japan or via Sakhalin Island, Russia. Remaining pintails primarily migrated to the Magadan region or Kolyma River Basin in eastern Russia via Sakhalin Island. The Chukotka Peninsula was the most common summer destination, with highest densities in the Anadyr Lowlands; a region also used by pintails that migrate from North America. One pintail migrated to St. Lawrence Island, Alaska, in spring and another briefly migrated to the western coast of Alaska in fall. Autumn migration routes generally mirrored spring migration although most pintails bypassed Sakhalin Island in fall. Compared to North American pintails, pintails that winter in Japan exhibited less variation in migration routes and breeding distribution, and nested at higher latitudes. In the Russian Far East there is no region with habitats comparable in extent to the ephemeral mid‐latitude wetlands of North America. Consequently, East Asian pintails mainly nest in arctic and sub‐arctic regions where annual consistency in wetlands promotes constancy in migration routes and breeding distribution. Breeding season sympatry between pintails from different continents results more from North American pintails migrating to eastern Russia than from Japanese pintails migrating to North America.     

Y‐chromosome evidence supports widespread signatures of three‐species Canis hybridization in eastern North America

There has been considerable discussion on the origin of the red wolf and eastern wolf and their evolution independent of the gray wolf. We analyzed mitochondrial DNA (mtDNA) and a Y‐chromosome intron sequence in combination with Y‐chromosome microsatellites from wolves and coyotes within the range of extensive wolf–coyote hybridization, that is, eastern North America. The detection of divergent Y‐chromosome haplotypes in the historic range of the eastern wolf is concordant with earlier mtDNA findings, and the absence of these haplotypes in western coyotes supports the existence of the North American evolved eastern wolf (Canis lycaon). Having haplotypes observed exclusively in eastern North America as a result of insufficient sampling in the historic range of the coyote or that these lineages subsequently went extinct in western geographies is unlikely given that eastern‐specific mtDNA and Y‐chromosome haplotypes represent lineages divergent from those observed in extant western coyotes. By combining Y‐chromosome and mtDNA distributional patterns, we identified hybrid genomes of eastern wolf, coyote, gray wolf, and potentially dog origin in Canis populations of central and eastern North America. The natural contemporary eastern Canis populations represent an important example of widespread introgression resulting in hybrid genomes across the original C. lycaon range that appears to be facilitated by the eastern wolf acting as a conduit for hybridization. Applying conventional taxonomic nomenclature and species‐based conservation initiatives, particularly in human‐modified landscapes, may be counterproductive to the effective management of these hybrids and fails to consider their evolutionary potential.     

Second occurrence of the dinosauriform ichnogenus Atreipus in the western United States,Upper Triassic Chinle Group of Eastern Utah

Abstract

A newly discovered track in the Chinle Group north of Moab, Utah, is attributable to the ichnogenus Atreipus, an ichnotaxon that is relatively common in eastern North America (Newark Supergroup) but very rare in the Late Triassic of the western part of the continent. This is only the second report of the genus from the Chinle Group. Atreipus has been attributed to a silesaurid dinosauriform, and dinosauriform taxa are relatively abundant by skeletal material in the Late Triassic of western North America, but track evidence in the same units is dominated by ichnotaxa attributed to dinosaurs. The rarity of Atreipus is currently an anomaly in the region.     

The insect trace fossil Tonganoxichnus from the middle Pennsylvanian of Indiana: Paleobiologic and paleoenvironmental implications

The ichnogenus Tonganoxichnus, produced by one or more monuran insect taxa, is now recorded from the Middle Pennsylvanian Mansfield Formation of Indiana. Tonganoxichnus is a resting trace that has three important implications. First, it represents a recurrent behavioral pattern in Upper Carboniferous to Lower Permian marginal marine environments of North America. Second, it provides finely resolved anatomical information for axial and appendicular body structures and behaviors that are difficult to determine from body‐fossil material alone. Third, integrated sedimentologic and ichnologic observations indicate that the Tonganoxichnus assemblage, inclusive of other ichnotaxa, is common in tidal rhythmites that were developed under freshwater conditions, probably in the innermost part of estuarine systems, close to or at the fluvioestuarine transition.     

Onshore expansion of benthic communities after the Late Devonian mass extinction

A detailed ichnological analysis of the Upper Devonian–Lower Mississippian Bakken Formation of sub‐surface Saskatchewan and the partially coeval Exshaw Formation of Alberta indicates the presence of an anomalous ichnofacies gradient. The distal Cruziana Ichnofacies, which in rocks of other ages is restricted to lower‐offshore facies, here ranges from this setting to the lower shoreface. No archetypal Cruziana Ichnofacies is present in these deposits. This pattern is interpreted as resulting from the differential effects of the Late Devonian mass extinction in shallow‐water ecosystems. The onshore expansion evidenced by ichnological data is consistent with the pattern displayed by the body‐fossil record, which indicates a re‐invasion of shallow‐water environments by the Palaeozoic evolutionary fauna during the Late Devonian and into the Early Carboniferous. The ichnofauna studied is overwhelmingly dominated by deposit feeders, with suspension feeders being notably absent, further underscoring the importance of trophic type as a selectivity trait during mass extinctions.     

Identification of glacial refugia in south-eastern North America by phylogeographical analyses of a forest understorey plant, Trillium cuneatum

Aim We examine several hypotheses emerging from biogeographical and fossil records regarding glacial refugia of a southern thermophilic plant species. Specifically, we investigated the glacial history and post‐glacial colonization of a forest understorey species, Trillium cuneatum. We focused on the following questions: (1) Did T. cuneatum survive the Last Glacial Maximum (LGM) in multiple refugia, and (if so) where were they located, and is the modern genetic structure congruent with the fossil record‐based reconstruction of refugia for mesic deciduous forests? (2) What are the post‐glacial colonization patterns in the present geographical range? Location South‐eastern North America. Methods We sampled 45 populations of T. cuneatum throughout its current range. We conducted phylogeographical analyses based on maternally inherited chloroplast DNA (cpDNA haplotypes) and used TCS software to reconstruct intraspecific phylogeny. Results We detected six cpDNA haplotypes, geographically highly structured into non‐overlapping areas. With one exception, none of the populations had mixed haplotype composition. TCS analysis resulted in two intraspecific cpDNA lineages, with one clade subdivided further by shallower diversification. Main conclusions Our investigation revealed that T. cuneatum survived the LGM in multiple refugia, belonging to two (western, eastern) genealogical lineages geographically structured across south‐eastern North America. The western clade is confined to the south‐western corner of T. cuneatum’s modern range along the Lower Mississippi Valley, where fossil records document a major refugium of mesic deciduous forest. For the eastern clade, modern patterns of cpDNA haplotype distribution suggest cryptic vicariance, in the form of forest contractions and subsequent expansions associated with Pleistocene glacial cycles, rather than simple southern survival and subsequent northward colonization. The north–south partitioning of cpDNA haplotypes was unexpected, suggesting that populations of this rather southern thermophilic species may have survived in more northern locations than initially expected based on LGM climate reconstruction, and that the Appalachian Mountains functioned as a barrier to the dispersal of propagules originating in more southern refugia. Furthermore, our results reveal south‐west to north‐east directionality in historical migration through the Valley and Ridge region of north‐west Georgia.     

A review of vital rates and cause‐specific mortality of elk Cervus elaphus populations in eastern North America

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Multiple Pleistocene refugia and Holocene range expansion of an abundant southwestern American desert plant species (Melampodium leucanthum,Asteraceae)

Pleistocene climatic fluctuations had major impacts on desert biota in southwestern North America. During cooler and wetter periods, drought‐adapted species were isolated into refugia, in contrast to expansion of their ranges during the massive aridification in the Holocene. Here, we use Melampodium leucanthum (Asteraceae), a species of the North American desert and semi‐desert regions, to investigate the impact of major aridification in southwestern North America on phylogeography and evolution in a widespread and abundant drought‐adapted plant species. The evidence for three separate Pleistocene refugia at different time levels suggests that this species responded to the Quaternary climatic oscillations in a cyclic manner. In the Holocene, once differentiated lineages came into secondary contact and intermixed, but these range expansions did not follow the eastwardly progressing aridification, but instead occurred independently out of separate Pleistocene refugia. As found in other desert biota, the Continental Divide has acted as a major migration barrier for M. leucanthum since the Pleistocene. Despite being geographically restricted to the eastern part of the species’ distribution, autotetraploids in M. leucanthum originated multiple times and do not form a genetically cohesive group.     

Rediscovery of Nebela ansata (Amoebozoa: Arcellinida) in eastern North America: biogeographical implications

Aim The question whether free‐living protists are generally cosmopolitan is currently a matter of debate. In this study we investigate the geographical distribution of a distinctive testate amoeba species, Nebela ansata, and use our data to assess the potential for highly restricted distribution patterns in some protist species. Location Global. Methods We analysed (1) 3400 testate amoeba publications from North America and other continents, (2) unpublished slides of the Penard Collection of the Natural History Museum, London, UK, and (3) 104 Sphagnum samples from eastern North America. Non‐metric multidimensional scaling (NMDS) was used to visualize the similarities in testate amoeba community composition among 1012 North American samples, including two communities that contained N. ansata. Results We rediscovered N. ansata at a site in New Jersey located close to its type locality, and in Nova Scotia. We also report the existence of an apparently unpublished museum specimen originally collected from New Jersey. Our extensive literature survey confirmed the presence of this species only in the temperate part of eastern North America. The NMDS revealed that communities with N. ansata were less similar to each other than to communities from other parts of North America, suggesting that favourable habitats for N. ansata occur in other Sphagnum‐dominated peatlands, a habitat type that has been extensively sampled in North America and elsewhere. Main conclusions These data provide an unusually convincing case of a free‐living microorganism with a very limited distribution range in the temperate part of eastern North America. The remarkably restricted distribution of N. ansata highlights the extent of our ignorance about the natural history of free‐living microorganisms, and raises questions about the lack of attention to microbial diversity in conservation biology.     

Biogeography and evolution of the Holarctic zooplankton genus Leptodora (Crustacea: Branchiopoda: Haplopoda)

Aim To reconstruct the phylogeographic history of the Holarctic carnivorous genus Leptodora (Crustacea: Cladocera: Haplopoda). Location We studied the DNA of between one and five specimens each from 28 populations distributed across the Holarctic, but with emphasis on Eurasia. Methods We sequenced a mitochondrial (cytochrome c oxidase subunit I) and a nuclear (elongation factor‐1α) gene, and combined this molecular information with geological and palaeoclimatological data. Haplotype networks and phylogenetic trees were constructed using a Bayesian and maximum likelihood approach. A molecular clock was applied. Results Leptodora consists of three clades (Leptodora kindtii in Europe, Leptodora richardi in China and Japan, and Leptodora sp. in North America), with insular subclades in Japan and in the eastern Mediterranean. The North American clade was not studied in detail. Leptodora richardi is the more thermophilic of the three. It extends from the Tropic of Cancer in the south to the Heilong Basin in the north. The western European L. kindtii is more cold‐water adapted than the eastern Mediterranean subclade. ‘West European’ and ‘Chinese’ clades are broadly separated by a hybrid zone in Siberia and European Russia as far west as the Volga. These hybrids have the mitochondrial DNA of L. kindtii, the nuclear DNA of L. richardi and the low‐temperature preference of L. kindtii, and may have formed as recently as the Holocene hypsithermal. A pure L. kindtii population in the Upper Irtysh catchment, east of the Dzungarian Gates, has been sequestered in endorheic Lake Wulungu, Xinjiang, since the mid‐Pleistocene. Main conclusions Application of a molecular clock places the most recent common ancestor of the North American, East Asian and European populations in the mid‐Miocene. The North American taxon is still living in isolation, while the Eurasian taxa, separated by the Alpine folding, made contact again in the Pleistocene, when the cold‐stenothermic L. kindtii repeatedly moved eastwards across Siberia and back. The population in Xinjiang is a relict of an early wave coming from western Europe: it crossed the Dzungarian Gates during a humid mid‐Pleistocene event, probably corresponding to the Apsheron transgression in the Caspian Basin. Later aridity isolated it there, and it started accumulating private haplotypes. The Holocene Euro‐Siberian hybrid zone may eventually engulf all European populations.