Phylogenetics and biogeography of eastern Asian-North American disjunct genus Pachysandra （Buxaceae） inferred from nucleotide sequences

Pachysandra is an eastern Asian-North American disjtunct genus with three species, two in eastern Asia （Pachysandra axillaris and Pachysandra terminalis） and one in eastern North America （Pachysandra procurnbens）. Although morphological and cytological studies suggest a close affinity of Pprocumbens with P axillaris, molecular data from nuclear and chloroplast DNA regions have provided conflicting signals. In this study, we tested previous phylogenetic hypotheses using sequences of nuclear ribosomal DNA internal transcribed spacers and chloroplast ndhF gene from multiple individuals of each of the three species. We also estimated the time of divergence between eastem Asia and eastern North America. Our results support the morphological and cytological conclusion that P procumbens is more closely related to P axillaris than to P terminalis. The estimated time of divergence of P axillaris and P procumbens was 14.6±5.5 mya, consistent with estimates from many other eastern Asian-North American disjunct genera. The migration of Pachysandra populations from eastern Asia to North America might have occurred by way of the North Atlantic land bridge.     

Phylogenetic analysis of Toxicodendron （Anacardiaceae） and its biogeographic implications on the evolution of north temperate and tropical intercontinental disjunctions

Toxicodendron is a genus in the Rhus complex of Anacardiaceae with a disjunct distribution between eastern Asia and North America, extending to southeastern Asia and the neotropics. Nuclear （internal transcribed spacer, external transcribed spacer, and NIA-i3） and chloroplast （ndhF and trnL-F） sequences were used to construct phylogenetic relationships of Toxicodendron. Phylogenetic analysis of these data strongly support Toxieodendron as a monophyletic group distinct from other genera of the Rhus complex, and the phylogeny does not fully corroborate classification at the sectional level. Two temperate disjunct lineages were detected, one from section Toxicodendron and the other between the eastern North American Toxicodendron vernix and the eastern Asian Toxicodendron vernieifluum. Their divergence times were estimated to be 13.46 （7.95-19.42） and 7.53 （2.76-12.86） mya, respectively. The disjunction between section Griffithii （taxa from warm temperate to tropical Asia） and Toxieodendron striatum （from the neotropics） was supported and their divergence time was estimated to be 20.84 （11.1 6-30.52） mya in the early Miocene. Our biogeographic results and the paleontological data support the Bering land bridge as the most likely route to explain the temperate disjunctions, yet the tropical disjunction in Toxicodendron seems to be best explained by the North Atlantic land bridge hypothesis.     

Segregation of F-2 interspecific hybrid growth performance and wing color patterns relative to parental species in the Papilio machaon species group （Lepidoptera： Papilionidae）

A strange Papilio female specimen with a prominent yellow wing band type very much like P. brevicauda （and also seen in P. machaon, and P. zelicaon） was captured in the forested mountains of southwest Vermont, USA. Despite its brevicauda-like appearance, it had mt-DNA of P. polyxenes. In eastern North America the P. brevicauda （Maritime Provinces of Canada） is currently believed to reflect previous （ancient） hybridization between ancestral P. machaon and P. polyxenes. It is uncertain whether the brevicauda-like wing trait features in the odd Vermont female may be a result of some introgression of P. machaon or P. brevicauda with P. polyxenes in eastern North America as seen also in the Ozark Mountains with P. joanae. Nonetheless, the significance of wing trait analyses in relation to hybrid zones with P. polyxenes remains relevant. The segregation of wing color patterns and hostplant use abilities （efficiencies and growth rates） were examined for laboratory hand-paired F-2 hybrids of P. zelicaon ~ Papilio polyxenes relative to the parental species and primary hybrids. The black adult color pattern characterizing P. polyxenes is presumed to be inherited as a simple autosomal dominant. These hybrids were examined to determine： （i） how F-1 primary hybrid offspring perform relative to the parental types in survival, growth rates, and growth efficiencies; and （ii） whether these larval traits and the adult color patterns might be linked with wing morphology in the F-2 hybrids. While the F-1 hybrids grew as well as the P. polyxenes parent （P. zelicaon was significantly less efficient and slower in growth than either of these）, we could not convincingly determine whether the growth performances of the F-2 segregants were linked with the parental wing patterns.     

Evolution of the Madrean-Tethyan disjunctions and the North and South American amphitropical disjunctions in plants

The present paper reviews advances in the study of two major intercontinental disjunct biogeographic patterns： （i） between Eurasian and western North American deserts with the Mediterranean climate （the Madrean- Tethyan disjunctions）; and （ii） between the temperate regions of North and South America （the amphitropical disjunctions）. Both disjunct patterns have multiple times of origin. The amphitropical disjunctions have largely resulted from long-distance dispersal, primarily from the Miocene to the Holocene, with available data indicating that most lineages dispersed from North to South America. Results of recent studies on the Mediterranean disjuncts between the deserts of Eurasia and western North America support the multiple modes of origin and are mostly consistent with hypotheses of long-distance dispersal and the North Atlantic migration. Axelrod＇s Madrean-Tethyan hypothesis, which implies vicariance between the two regions in the early Tertiary, has been favored by a few studies. The Beringian migration corridor for semiarid taxa is also supported in some cases.     

Molecular Epidemiological Investigation of Infectious Hypodermal and Hematopoietic Necrosis Virus and Taura Syndrome Virus in Penaeus Vannamei Cultured in China

The Infectious hypodermal and hematopoietic necrosis virus(IHHNV) and Taura syndrome virus(TSV) are two important shrimp viruses in cultured shrimp in America.These two viruses were transmitted to China at the beginning of the 21st century.In this study,214 shrimp samples of Penaeus vannamei were collected from seven different areas of China and tested by PCR for IHHNV and TSV infection.The results showed that there were a high prevalence of IHHNV(65.42%) and low prevalence of TSV(3.27%) in the tested samples.Several samples were found to be co-infected with these two viruses.A 3 kb fragment of 7 positive IHHNV samples and a structure protein region(ORF2) of three TSV positive samples were amplified and sequenced.The sequence comparison indicated that both IHHNV and TSV sequenced in China have a low genetic variations compared with the prototype IHHNV and TSV from Hawaii.Phylogenetic analysis showed that TSV isolates were clustered into two groups,Asia and America group,which was genetically correlated to geographic distribution.     

桂西北典型喀斯特区生态服务价值的环境响应及其空间尺度特征

Study on ecosystem service values (ESVs) is the bridge of understanding ecosystem and economic decision-making. To investigate the response of ESVs to ambient environment and their spatial scales is an urgent work in the process of ecological restoration and sustainable development in southwest China. Based on the previous research results, the remote images, and weather data of 31 years (1975-2005), the response of ESVs to ambient environment and their spatial scales in a typical karst area of northwest Guangxi, China were evaluated with the method of empirical mode decomposition (EMD) in this paper. The results showed that ESVs had obvious variations with changes in elevation, rocky types and soil types, and had corresponding characteristics of spatial scales. The ESVs in middle and low elevation areas were higher, but they decreased due to human activities such as deforestation. The ESVs in peak-cluster depression areas were lower, and they would increase as influenced by policies such as returning farmland to forestland. The ESVs were influenced by rocky types, showing lower values but an increasing trend in the typical karst regions while higher values but a decreasing trend in non-karst regions. The average ESVs in the west part of the study region, which is not only the main nature reserve for rare wildlife species in Guangxi, but also one of the best preserved natural vegetation regions in China, were more than 15,000 RMB Yuan ha-1,. Comparatively, the ESVs were less than 10,000 Yuan ha-1 in the middle part of the region attributed to low vegetation coverage, serious peak-cluster depression and karst rocky desertification. After about 20 years, that is from 1985 to 2005, the ecosystem conditions had been improved and the ESVs had increased in the middle and eastern part of this study region. On the contrary, in most western part, which was dominated by subtropical evergreen broad-leaved forest, they had been worsened, and the balance values of ESVs were negative. As for the relationships of ESVs with rocky types and soil types, lime soil and red soil were the two main soil types that contributed to ESVs (The total contribution was above 60%, and it is 63.77%,64.37%,64.56%,64.91% respectively in the four years of 1985, 1990, 2000 and 2005.). The variance contribution of intrinsic mode function (IMF) and trend (R) showed that there were obvious special sales for ESVs in this study area, and the main spatial scales were about 2.7 km, 5.5 km, and 11.6 km. The variance contribution ratio was 12.29%, 11.26%, 11.49% respectively, and the trend (R) was high (17.74%). In conclusion, this study indicated that ecosystem conditions in the typical karst area were improved owing to the application of rocky desertification control policies, such as ecological migration and returning farmland to forestland. Besides, the main spatial scales of ESVs in the study area were the interactions of terrain, physiognomy, land use / land cover and human activities.     

会议报道：北京生物入侵国际研讨会: 2004年6月6日－11日

 Scientists and policy-makers are now well aware of the ecological and societal problems associated with the introduction of species by humans across natural barriers to dispersal. Some introduced species spread abundantly in their new habitats and have negative effects on existing species, both agricultural and natural. Such introduced, invasive species can alter fire, nutrient, and hydrological regimes and have large economic costs. In many parts of the world, this awareness has come rather late: commerce and travel have already carried large numbers of plants, animals, and microbes between many regions. In contrast, China's relative self-remove from global trade during much of the second half of the 20th century may have earned it a partial reprieve from invasions from abroad and reduced the number of invasions from China to elsewhere. If so, there could be a new wave of species invasions between China and North America impending. China's trade with the U.S. increased by 460% from 1992 to 2002, according to the U.S. Department of Commerce. This may be the critical moment to apply the lessons learned from invasions across other oceans and forestall new invasions across the Pacific. We need to match growth in trade with growth in the science and policy needed to prevent and control introductions. To this end, some 100 scientists and representatives of governmental and non-governmental organisations from eight countries gathered on June 6-11 for the Beijing International Symposium on Biological Invasions, subtitled "Species Exchanges Between Eastern Asia and North America: Threats to Environment and Economy". The Institute of Botany of the Chinese Academy of Sciences and the Sino-Ecologists Club Overseas organized the meeting, under the leadership of Xinguo Han and Ming Dong for the Institute and Shili Miao for the Club. The National Science Foundation of China, the U.S. Geological Survey, and the Asian Ecology Section of the Ecological Society of America provided additional support. Shili Miao and Jianhui Huang coordinated the program and logistics, and the Beijing Botanical Garden provided the venue, appropriately stocked with introduced plants from around the world, presumably non-invasive ones. There were two keynote addresses. Harold Mooney (Stanford, USA) highlighted the need for scientists to develop quantitative tools that managers can use to analyze the extent and the impacts of invasions. Richard Mack (Washington State, USA) noted that the phylogenetic, latitudinal, and climatic similarities between China and the U.S. may predispose introductions from one to the other to bloom into invasions. The 37 other talks and the posters focused on current patterns of invasion in eastern Asia, the theory of invasions, techniques to manage invasions, and policies to prevent them.     

THE SKULL OF HAPALODECTES (?ACREODI, MAMMALIA), WITH NOTES ON SOME CHINESE PALEOCENE MESONYCHIDS

<正> Hapalodectes has long been considered as a small mesonychid specialized toward a carnivorous and piscivorous mode of life. Its rare fossil record was known from the early Eocene of North America and middle and upper Eocene of Asia. Knowledge of this genus is still restricted to two species based mainly on some fragmentary materials.A part of the right mandible of the type species, H. leptognathus, was first described by Osborn and Wortman (1892), but was referred by them to Dissacus, and renamed as Hapalodectes by Matthew (1909). Szalay and Gould (1966) erected a new subfamily, Hapalodectinae, and then all known material were restudied by the former author (Szalay, 1969). No additional material of the genus has been reported since then.The newly discovered specimens described in this paper are the first, and so far the only, cranial evidence of Hapalodectes. The fossils were collected from early Eocene (Ling-cha Formation), Hengdong, Hunan, during two field seasons of 1982-1983. The purpose of the present paper is to place on record the morphology of this well-preserved cranial material and to discuss briefly its significance, it is hoped that the information about the cranial morphology of Hapalodectes will increase our understanding of its relationship with Mesonychidae.We are indebted to Drs. Everett H. Lindsay and Chow Minchen for correcting English summary. We would like to express our gratitude to Dr. Malcolm C. McKenna for his valuable comments and correcting English summary and the facilitating the casts of the type species of Hapalodectes at the American Museum of Natural History. Ms. Ann Bleefield and Meng jin help us to get the casts, Zhai R. j. and Qi T. lend us unpublished material for the comparison, all the photographs are taken by Wang J. f. and figures are drawn by shen W. L., we thank all of them for their kind help.     

Teleconnection between the early immigration of brown planthopper (Nilaparvata lugens St?l) and ENSO indices: implication for its medium- and long-term forecast

Teleconnection between the early immigration of brown planthopper (BPH) and El Nino/Southern Oscillation (ENSO) indices from January of two years previously to the current June was investigated to make long-term forecast. The teleconnection results were as follows: ENSO indices which were significantly correlated with the early immigration of BPH were primarily sea surface temperature anomalies (SSTA) in N3, N4 and N3.4 regions, accounting for 71.8% of the total. Significant ENSO indices from two years and one year before the immigration events had a proportion of about 84%, while those in the current year only accounted for 16.7%. There was significantly negative correlation between the early immigration of BPH and SSTA in each Nino region from two years before to the previous spring, whereas there was significantly positive correlation between these two factors during the period from the previous winter to the current spring. The significant correlation between the early immigration of BPH and SSTA in each Nino region in the last summer and autumn did not show any obvious tendencies. The relationship between the southern oscillation index (SOI) and the early immigration of BPH was opposite to that between the immigration and SSTA in each Nino region. The above mentioned significant ENSO indices were used as key factors to build forecasting models for the early immigration of BPH by step-wise multiple linear regression analysis. Finally, 12 integrated forecasting models were obtained, which could make predictions 3–27 months ahead and had a predictive accuracy of 88.9%.     

Naturally occurring Trichogramma species in olive farms in Egypt

A survey of two-year studies (2001-2003) was carried out in two olive groves sited at two representative olive growing areas, namely Paradise Park (arid area) and Burg E1-Arab farm (semi-arid area) to monitor the frequency of endemic Trichogramma species on olive moth (Prays oleae) and jasmine moth (Palpita unionalis). The suspended host bait traps were found to be a more practical and effective tool for collecting Trichogramma wasps than the attached ones. Four naturally occurring Trichogramma species were collected for the first time in Egypt from the olive groves, where releases have never been conducted. T.bourarachae was collected exclusively from Burg El-Arab farm. It seems that this wasp species adapts well to the semi-arid area. Three species, namely T. cordubensis, T. nr.pretiosum and T. cacociae were isolated from Paradise Park farm. All of these wasps were also bred from naturally parasitized host eggs during favorable and even at unfavorable temperature conditions of June-August. However, these endemic species did not occur naturally in sufficient numbers to keep the pest populations from reaching damaging levels.The excessive usage of insecticides and the oophagous predators (e.g., ants and lacewing larvae) are some factors that affect the performance of Trichogramma wasps in olive farms.The presence of warm weather wasp strains suggests the existence of well-adapted wasp species or swains which may be appropriate candidates for the control of target pests in olive groves. Additional study is required to determine the best “habitat-specific” species/strains of Trichogramma for augmentative release of naturally occurring wasps and to incorporate them into integrated pest management programs. Efforts should be made to conserve these endemic species from oophagous predators, hot weather and insecticides.The olive and jasmine moth-larvae and pupae found under tree canopies were bred and emerged parasitoids were listed.     

Phylogeny and Biogeography of Thuja L. (Cupressaceae), an Eastern Asian and North American Disjunct Genus

In order to develop better insights into biogeographic patterns of eastern Asian and North American disjunct plant genera, sequences of nuclear ribosomal DNA internal transcribed spacer (nr DNA ITS) region were used to estimate interspecific relationships of Thuja L. (Cupressaceae) and infer its biogeography based on the phylogeny. According to the phylogenetic analysis, two clades were recognized. The first clade included Thuja plicata D. Don (western North America) and T. koraiensis Nakai (northeastern Asia), and the second one contained T. occidentalis (Gord.) Carr. (Japan). The ancestral area of Thuja was inferred to be eastern Asia, and two dispersal events were responsible for the modern distribution of Thuja in North America. Both the North Atlantic land bridge and Bering land bridge were possible routes for the migration of ancestral populations to North America.     

Phylogeny and Phytogeography of Eupatorium (Eupatorieae, Asteraceae): Insights from Sequence Data of the nrDNA ITS Regions and cpDNA RFLP

Eupatorium were examined by sequencing the internal transcribed spacers (ITS) of nuclear ribosomal DNA and restriction site analysis of chloroplast DNA. Molecular data provided strong evidence that (1) this genus originated in North America, (2) the genus diverged into three morphological species groups, Eutrochium, Traganthes and Uncasia in North America, and (3) one of the North American Uncasia lineages migrated into temperate Europe and eastern Asia over the Bering land bridge. The estimated divergence times support a late Miocene to early Pliocene migration from North America to Eurasia via the Bering land bridge. A European species was sister to all of the eastern Asian species examined. The disjunct distribution pattern of the genus Eupatorium is incongruent with the classical Arcto-Tertiary geoflora concept. Received 13 September 1999/ Accepted in revised form 4 January 2000     

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.     

Phylogenetics and biogeography of eastern Asian–North American disjunct genus Pachysandra (Buxaceae) inferred from nucleotide sequences

Abstract Pachysandra is an eastern Asian–North American disjunct genus with three species, two in eastern Asia (Pachysandra axillaris and Pachysandra terminalis) and one in eastern North America (Pachysandra procumbens). Although morphological and cytological studies suggest a close affinity of P. procumbens with P. axillaris, molecular data from nuclear and chloroplast DNA regions have provided conflicting signals. In this study, we tested previous phylogenetic hypotheses using sequences of nuclear ribosomal DNA internal transcribed spacers and chloroplast ndhF gene from multiple individuals of each of the three species. We also estimated the time of divergence between eastern Asia and eastern North America. Our results support the morphological and cytological conclusion that P. procumbens is more closely related to P. axillaris than to P. terminalis. The estimated time of divergence of P. axillaris and P. procumbens was 14.6±5.5 mya, consistent with estimates from many other eastern Asian–North American disjunct genera. The migration of Pachysandra populations from eastern Asia to North America might have occurred by way of the North Atlantic land bridge.     

Phylogenomics,biogeography, and evolution of morphology and ecological niche of the eastern Asian–eastern North American Nyssa (Nyssaceae)

Nyssa (Nyssaceae, Cornales) represents a classical example of the well‐known eastern Asian–eastern North American floristic disjunction. The genus consists of three species in eastern Asia, four species in eastern North America, and one species in Central America. Species of the genus are ecologically important trees in eastern North American and eastern Asian forests. The distribution of living species and a rich fossil record of the genus make it an excellent model for understanding the origin and evolution of the eastern Asian–eastern North American floristic disjunction. However, despite the small number of species, relationships within the genus have remained unclear and have not been elucidated using a molecular approach. Here, we integrate data from 48 nuclear genes, fossils, morphology, and ecological niche to resolve species relationships, elucidate its biogeographical history, and investigate the evolution of morphology and ecological niches, aiming at a better understanding of the well‐known EA–ENA floristic disjunction. Results showed that the Central American (CAM) Nyssa talamancana was sister to the remaining species, which were divided among three, rapidly diversified subclades. Estimated divergence times and biogeographical history suggested that Nyssa had an ancestral range in Eurasia and western North America in the late Paleocene. The rapid diversification occurred in the early Eocene, followed by multiple dispersals between and within the Erasian and North American continents. The genus experienced two major episodes of extinction in the early Oligocene and end of Neogene, respectively. The Central American N. talamancana represents a relic lineage of the boreotropical flora in the Paleocene/Eocene boundary that once diversified in western North America. The results supported the importance of both the North Atlantic land bridge and the Bering land bridge (BLB) for the Paleogene dispersals of Nyssa and the Neogene dispersals, respectively, as well as the role of Central America as refugia of the Paleogene flora. The total‐evidence‐based dated phylogeny suggested that the pattern of macroevolution of Nyssa coincided with paleoclimatic changes. We found a number of evolutionary changes in morphology (including wood anatomy and leaf traits) and ecological niches (precipitation and temperature) between the EA–ENA disjunct, supporting the ecological selection driving trait evolutions after geographic isolation. We also demonstrated challenges in phylogenomic studies of lineages with rapid diversification histories. The concatenation of gene data can lead to inference of strongly supported relationships incongruent with the species tree. However, conflicts in gene genealogies did not seem to impose a strong effect on divergence time dating in our case. Furthermore, we demonstrated that rapid diversification events may not be recovered in the divergence time dating analysis using BEAST if critical fossil constraints of the relevant nodes are not available. Our study provides an example of complex bidirectional exchanges of plants between Eurasia and North America in the Paleogene, but “out of Asia” migrations in the Neogene, to explain the present disjunct distribution of Nyssa in EA and ENA.     

<Emphasis Type="Italic">Altingioxylon hainanensis</Emphasis> sp. nov.: earliest fossil wood record of the family Altingiaceae in Eastern Asia and its implications for historical biogeography

A new species, Altingioxylon hainanensis, is described from the Eocene Changchang Formation of the Changchang Basin on Hainan Island, South China. It is the first record of a fossil wood assigned to Altingiaceae found in China, and the most ancient evidence of wood for this family in eastern Asia. The new species is similar to A. rhodoleioides, known since the Miocene in India and Java Island, and to Altingia hisauchii from the Miocene to Pliocene of Japan. The close resemblance between these species and Liquidambar sp., known from the Middle Miocene of western North America, provides additional evidence for the migration of their ancestors from Asia to North America across the Bering land bridge during the Miocene. Distinctions in ray sizes between the eastern Asian specimens and their contemporaries from Europe to Kazakhstan is suggested as a result of the divergence between the large eastern Asian clade and the North American–west Asian clade within Altingiaceae during the Eocene–Oligocene. The presence of crystals in ray cells may be considered an ancestral condition that persists in the eastern Asian lineages up to the extant Altingia and Semiliquidambar, but which was lost in other Altingiaceae in the course of evolution.     

Molecular phylogenetics and biogeography of the eastern Asian–eastern North American disjunct Mitchella and its close relative Damnacanthus (Rubiaceae,Mitchelleae)

Mitchella is a small genus of the Rubiaceae with only two species. It is the only herbaceous semishrub of the family showing a disjunct distribution in eastern Asia and eastern North America, extending to Central America. Its phylogeny and biogeographical diversification remain poorly understood. In this study, we conducted phylogenetic and biogeographical analyses for Mitchella and its close relative Damnacanthus based on sequences of the nuclear internal transcribed spacer (ITS) and four plastid markers (rbcL, atpB‐rbcL, rps16 and trnL‐F). Mitchella is monophyletic, consisting of an eastern Asian M. undulata clade and a New World M. repens clade. Our results also support Michella as the closest relative to the eastern Asian Damnacanthus. The divergence time between the two intercontinental disjunct Mitchella species was dated to 7.73 Mya, with a 95% highest posterior density (HPD) of 3.14?12.53 Mya, using the Bayesian relaxed clock estimation. Ancestral area reconstructions suggest that the genus originated in eastern Asia. The semishrub Mitchella appears to have arisen from its woody ancestor in eastern Asia and then migrated to North America via the Bering land bridge in the late Miocene. © 2013 The Linnean Society of London     

Morphological and molecular comparisons ofCampsis grandiflora andC. radicans (Bignoniaceae), an eastern Asian and eastern North American vicariad species pair

Morphological and molecular comparisons were made forCampsis grandiflora (Thunb.)K. Schumann (Bignoniaceae) from eastern Asia andC. radicans (L.)Seemann from eastern North America. Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases. The cpDNA divergence between the two vicariad species was 2.44%, which is the highest reported among North Temperate disjunct taxa and one of the highest reported for infrageneric taxa. Detailed morphological comparisons also suggest a high level of divergence. Cluster analyses based on 22 morphological characters and 39 OTUs revealed two distinct groups corresponding with the two species. The average taxonomic distance between the two species was 1.806. Analyses of variance (ANOVA) revealed that 12 of the 18 quantitative characters differed significantly ( 0.01) betweenC. grandiflora andC. radicans. Divergence time based on cpDNA data was estimated as 24.4 million years. The Bering land bridge hypothesis was favored over the North Atlantic land bridge hypothesis based on the estimated divergence time and the geological history of the North Temperate region. The high levels of morphological and cpDNA divergence are not consistent with morphological stasis, which has been proposed as a common mode of evolution for North Temperate disjunct taxa.     

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.