Phylogenetic relationships and biogeography of the Japanese clawed salamander, Onychodactylus japonicus (Amphibia: Caudata: Hynobiidae), and its congener inferred from the mitochondrial cytochrome b gene

Using the mitochondrial cytochrome b gene, we investigated phylogenetic relationships between and within the Japanese clawed salamander, Onychodactylus japonicus, and its close continental relative O. fischeri. Monophyly of O. japonicus was well supported, and O. japonicus was clearly distinguished from O. fischeri. However, O. fischeri comprises genetically distinct populations from Russia, NE China, and Korea that do not form a monophyletic group. Within O. japonicus, four major clades were clearly recognized: Clade I from northern Tohoku district, Clade II from southern Tohoku district and the Tsukuba Mountains, Clade III from southwestern Honshu, and Clade IV from Kinki and Chugoku districts in Honshu and from Shikoku. Although genetic distances among these clades were large (5.5-9.6%), relationships among the clades were unresolved. All clades except Clade I contained two or three distinct subclades. In several localities in Kinki and Chugoku, Clades III and IV were sympatric. The estimated divergence times and available geohistorical data suggest that O. japonicus began to differentiate in the Upper Late Miocene and that the pattern of genetic differentiation of this species has been affected strongly by climate changes and geohistorical events such as volcanic activity and mountain formation. Our results suggest that both O. fischeri and O. japonicus comprise multiple cryptic species.     

‘Out‐of‐Africa’ dispersal of tropical floras during the Miocene climatic optimum: evidence from Uvaria (Annonaceae)

Aim African–Asian disjunctions are common in palaeotropical taxa, and are typically explained by reference to three competing hypotheses: (1) ‘rafting’ on the Indian tectonic plate, enabling Africa‐to‐Asia dispersal; (2) migration via Eocene boreotropical forests; and (3) transoceanic long‐distance dispersal. These hypotheses are tested using Uvaria (Annonaceae), which is distributed in tropical regions of Africa, Asia and Australasia. Recent phylogenetic reconstructions of the genus show a clear correlation with geographical provenance, indicating a probable origin in Africa and subsequent dispersal to Asia and then Australasia. Ancestral areas and migration routes are inferred and compared with estimates of divergence times in order to distinguish between the prevailing dispersal hypotheses. Location Palaeotropics. Methods Divergence times in Uvaria are estimated by analysing the sequences of four DNA regions (matK, psbA–trnH spacer, rbcL and trnL–F) from 59 Uvaria species and 77 outgroup species, using a Bayesian uncorrelated lognormal (UCLD) relaxed molecular clock. The ancestral area of Uvaria and subsequent dispersal routes are inferred using statistical dispersal–vicariance analysis (s‐diva ). Results Uvaria is estimated to have originated in continental Africa 31.6 Ma [95% highest posterior density (HPD): 38.4–25.1 Ma] between the Middle Eocene and Late Oligocene. Two main migration events during the Miocene are identified: dispersal into Madagascar around 17.0 Ma (95% HPD: 22.3–12.3 Ma); and dispersal into Asia between 21.4 Ma (95% HPD: 26.7–16.7 Ma) and 16.1 Ma (95% HPD: 20.1–12.1 Ma). Main conclusions Uvaria fruits are widely reported to be consumed by primates, and are therefore unlikely candidates for successful long‐distance transoceanic dispersal. The other biogeographical hypotheses, involving rafting on the Indian tectonic plate, and dispersal via the European boreotropical forests associated with the Eocene thermal maximum, can be discounted due to incongruence with the divergence time estimates. An alternative scenario is suggested, involving dispersal across Arabia and central Asia via the tropical forests that developed during the late Middle Miocene thermal maximum (17–15 Ma), associated with the ‘out‐of‐Africa’ dispersal of primates. The probable route and mechanism of overland dispersal between Africa and Asia for tropical plant groups during the Miocene climatic optimum are clarified based on the Uvaria data.     

The Evolution and Paleobiogeography of Flying Squirrels (Sciuridae, Pteromyini) in Response to Global Environmental Change

Flying squirrels are strictly arboreal squirrels adopting a special gliding form of locomotion. This group of animals has a long history that has mirrored the vicissitude of forests. The discrepancy in the distribution between fossils and extant species indicates a mysterious evolution history requiring further exploration. This study compiles the worldwide fossils of Pteromyini to the species level in order to reproduce the spatiotemporal distribution pattern of flying squirrels and deduce the ancestral distribution according to dispersal-vicariance analysis of a phylogeny of the extant species. In addition, we reconstruct the paleoenvironmental background and find that flying squirrels probably originated in the Oligocene–Miocene transition from Europe and immediately dispersed to Asia and North America. Influenced by glaciation, CO₂ reduction, geologic movements and the Paratethys retreat, the Northern Hemisphere underwent climate deterioration and grassland expansion during the late Miocene, and thus the diversity of Pteromyini dramatically decreased. The uplift of the Tibet Plateau in addition to the strengthened Asian monsoons intensified the aridity in central Asia, but brought sufficient water to the densely forested regions of South and Southeast Asia. These forests are likely both refugia and diversification center for flying squirrels during glacial periods in the Quarternary. The subsequent connection and separation events among these heterogeneous habitats has probably been a driving force in the speciation of flying squirrels. Based on this work, we predict a bleak future for the flying squirrels, one which is closely associated with the fate of forests in Asia.     

The Biogeographic South-North Divide of Polygonatum (Asparagaceae Tribe Polygonateae) within Eastern Asia and Its Recent Dispersals in the Northern Hemisphere

Eastern Asia (EA) is a key region for the diversification of flowering plants in the Northern Hemisphere, but few studies have focused on the biogeographic history within EA in the context of the other northern continents. Polygonatum is an important medicinal genus widely distributed in the Northern Hemisphere with its highest species richness in EA, and it represents an excellent model for studying the evolution of biogeographic patterns in this region. Divergence time estimation was used to examine the biogeographic history of Polygonatum based on nuclear ITS and four plastid sequences (rbcL, matK, psbA–trnH and trnC–petN) from 30 Polygonatum species and 35 outgroup taxa. The ancestral area of Polygonatum and subsequent dispersal routes were inferred using Bayes-Lagrange. Polygonatum was estimated to have originated in southern EA during the middle Miocene (14.34–13.57 Ma) with subsequent south-to-north expansion in the late Miocene. Multiple intercontinental dispersal events were inferred between EA and Europe or North America, and all of them have occurred recently in the late Miocene to Pliocene. The separation of Polygonatum into the south and north lineages and their subsequent diversifications in the late Miocene supports the existence of a biogeographic divide between the northern and southern parts of EA that also coincides with the retreat and redevelopment of the arid zone in EA in the Neogene. Our results demonstrate the complexity of biogeographic history of Polygonatum in the Northern Hemisphere including early vicariance followed by frequent and recent dispersals in the Neogene.     

Die Cypriniden (Teleostei: Cypriniformes) des oberoligozänen Maares von Enspel nebst Bemerkungen zur Phylogenie und Biogeographie der Phoxininae

Palaeorutilus enspelensis (Böhme 1996), the only fish species from the Upper Oligocene maar lake Enspel (Westerwald), is described and its phylogenetical position within the subfamily Phoxininae is discussed. The cladistic analysis of osteological and lepidological features indicates that the genusPalaeorutilus Gaudant 1988 is a member of the Chub Clade. Furthermore, including of the recent Eurasian taxaLagowskiella andEupallasella in the analysis, the Chub Clade can be divided into two monophyletic Clades: the EurasianPhoxinus Clade (including the holarcticPhoxinus) and the eastern North American Creek Chub Clade which includes the fossil genusPalaeorutilus as its basal sister taxon. These phylogenetical investigation as well as the recent and fossil distribution pattern of the Phoxininae support the following biogeographical hypothesis: The settlement of cyprinids in North America took place in several waves and different directions. The separation of Western Clade and Chub Clade did not occur in North America but in Eocene times in Asia. Members of the Creek Chub Clade migrated from Europe to eastern North America via a transatlantic route prior to the Middle Miocene.     

Did geckos ride the Palawan raft to the Philippines?

Aim We examine the genetic diversity within the lizard genus Gekko in the Philippine islands to understand the role of geography and geological history in shaping species diversity in this group. We test multiple biogeographical hypotheses of species relationships, including the recently proposed Palawan Ark Hypothesis. Location Southeast Asia and the Philippines. Methods Samples of all island endemic and widespread Philippine Gekko species were collected and sequenced for one mitochondrial gene (NADH dehydrogenase subunit 2) and one nuclear gene (phosducin). We used maximum likelihood and Bayesian phylogenetic methods to derive the phylogeny. Divergence time analyses were used to estimate the time tree of Philippine Gekko in order to test biogeographical predictions of species relationships. The phylogenetic trees from the posterior distribution of the Bayesian analyses were used for testing biogeographical hypotheses. Haplotype networks were created for the widespread species Gekko mindorensis to explore genetic variation within recently divergent clades. Results Both maximum likelihood and Bayesian phylogenetic analyses indicated that Philippine Gekko species are a diverse clade with a long history in the archipelago. Ancestral range reconstruction and divergence time analyses suggest a Palawan microcontinental origin for this clade, coinciding with Palawan’s separation from Asia beginning 30 Ma, with subsequent diversification in the oceanic Philippine islands. The widespread species G. mindorensis and G. monarchus diversified in the late Miocene/early Pliocene and are potentially complexes of numerous undescribed species. Main conclusions The view of the Philippine islands as a ‘fringing archipelago’ does not explain the pattern of species diversity in the genus Gekko. Philippine Gekko species have diversified within the archipelago over millions of years of isolation, forming a large diverse group of endemic species. Furthermore, the Philippine radiation of gekkonid lizards demonstrates biogeographical patterns most consistent with stochastic colonization followed by in situ diversification. Our results reveal the need to consider deeper time geological processes and their potential role in the evolution of some Philippine terrestrial organisms.     

禄丰古猿化石地点的鱼化石

在禄丰古猿化石产地采获许多鲤的咽喉齿化石,由形态特征看,相近于现代广泛分布的鲤(Cyprinus carpio Linn.)的咽喉齿,当代表其早期记录。现今云南一带生存有多种鲤,可能为其繁衍的后代,且相当繁盛。这一发现为鲤属(Cyprinus)起源于亚洲提供了化石依据。     

Origin of the pork tapeworm Taenia solium in Bali and Papua,Indonesia

Global distributions of zoonotic pathogens have been strongly affected by the history of human dispersal and domestication of livestock. The pork tapeworm Taenia solium is distributed worldwide as the cause of neurocysticercosis, one of the most serious neglected tropical diseases. T. solium has been reported in Indonesia but only endemic to restricted areas such as Bali and Papua. Previous studies indicated the distinctiveness of a mitochondrial haplotype confirmed in Papua, but only one isolate has been examined to date. In this study, genetic characterization of T. solium and pigs in Bali and Papua was conducted to clarify the distributional history of the parasite. Mitochondrial haplotype network analysis clearly showed that Indonesian T. solium comprises a unique haplogroup which was the first to diverge among Asian genotypes, indicating its single origin and the fact that it was not introduced in the recent past from other area in Asia in which it is endemic. Although phylogenetic analysis based on the mitochondrial D-loop revealed multiple origins of pigs in Bali and Papua, the majority of pigs belonged to the Pacific Clade, which is widely dispersed throughout the Island Southeast Asia (ISEA) and Oceania due to Neolithic human dispersal. Given the results of our network analysis, it is likely that the Pacific Clade pigs played a key role in the dispersal of T. solium. The data suggest that T. solium was introduced from mainland Asia into Western Indonesia, including Bali, by modern humans in the late Pleistocene, or in the early to middle Holocene along with the Pacific Clade pigs. Introduction into New Guinea most likely occurred in the late Holocene through the spread of Pacific Clade pigs. Over time, T. solium has been eradicated from most of Indonesia through the middle to modern ages owing to religious and cultural practices.     

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.     

The Leporid Datum: a late Miocene biotic marker

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The legacy of a vanished sea: a high level of diversification within a European freshwater amphipod species complex driven by 15 My of Paratethys regression

The formation of continental Europe in the Neogene was due to the regression of the Tethys Ocean and of the Paratethys Sea. The dynamic geology of the area and repetitious transitions between marine and freshwater conditions presented opportunities for the colonization of newly emerging hydrological networks and diversification of aquatic biota. Implementing mitochondrial and nuclear markers in conjunction with a large‐scale sampling strategy, we investigated the impact of this spatiotemporal framework on the evolutionary history of a freshwater crustacean morphospecies. The Gammarus balcanicus species complex is widely distributed in the area previously occupied by the Paratethys Sea. Our results revealed its high diversification and polyphyly in relation to a number of other morphospecies. The distribution of the studied amphipod is generally characterized by very high local endemism and divergence. The Bayesian time‐calibrated reconstruction of phylogeny and geographical distribution of ancestral nodes indicates that this species complex started to diversify in the Early Miocene in the central Balkans, partially in the shallow epicontinental sea. It is possible that there were several episodes of inland water colonization by local brackish water lineages. Subsequent diversification within clades and spread to new areas could have been induced by Alpine orogeny in the Miocene/Pliocene and, finally, by Pleistocene glaciations. The present distribution of clades, in many cases, still reflects Miocene palaeogeography of the area. Our results point out that investigations of the historical aspect of cryptic diversity in other taxa may help in a general understanding of the origins of freshwater invertebrate fauna of Europe.     

Evolutionary history of Spalacidae inferred from fossil occurrences and molecular phylogeny

1. The Spalacidae is a family of strictly subterranean rodents with a long evolutionary history. It is unclear how ecological changes have influenced the evolutionary history of these mammals, and the phylogenetic relationship of the subfamilies within Spalacidae is controversial.
2. Through compiling fossil records, reconstructing molecular phylogeny from molecular data, determining the date of divergence, and analysing their geographical evolution based on molecular data and fossil taxa, we explore the origin and evolutionary process of Spalacidae in detail. Diversification within Spalacidae dates to the Late Oligocene, approximately 25 million years ago, based on molecular data.
3. This family originated in South and East Asia in the Late Oligocene, and then split into four clades. The first clade includes Rhizomyinae, which was highly diversified in South Asia in the Early-to-Middle Miocene. Then Rhizomyinae from Asia migrated to northern Africa in multiple waves through the Afro-Eurasian land bridge. Its range largely contracted in the Late Miocene, notably in Central Asia. The second clade includes the extinct Tachyoryctoidinae, which was confined to East and Central Asia, and survived from the Late Oligocene to the Late Miocene. The third clade includes Spalacinae, which have remained around the Mediterranean region since the Late Oligocene with slight trend of northward expansion. The fourth clade is Myospalacinae. Ancient genera of this subfamily in East Asia dispersed eastward during the Late Miocene and reached northern China and south-east Russia.
4. The general distribution pattern of Spalacidae has persisted since the Late Miocene. Extinction of Tachyoryctoidinae and clear range contraction of Rhizomyinae in Central and East Asia are likely to have resulted from increased aridification, while the slight northward expansion of Myospalacinae and Spalacinae since the Quaternary was probably a response to a similar northward expansion of suitable vegetation for these animals.

     

Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago

Gigantism and dwarfism evolve in vertebrates restricted to islands. We describe four new species in the Rhinolophus hildebrandtii species-complex of horseshoe bats, whose evolution has entailed adaptive shifts in body size. We postulate that vicissitudes of palaeoenvironments resulted in gigantism and dwarfism in habitat islands fragmented across eastern and southern Africa. Mitochondrial and nuclear DNA sequences recovered two clades of R. hildebrandtii senso lato which are paraphyletic with respect to a third lineage (R. eloquens). Lineages differ by 7.7 to 9.0% in cytochrome b sequences. Clade 1 includes R. hildebrandtii sensu stricto from the east African highlands and three additional vicariants that speciated across an Afromontane archipelago through the Plio-Pleistocene, extending from the Kenyan Highlands through the Eastern Arc, northern Mozambique and the Zambezi Escarpment to the eastern Great Escarpment of South Africa. Clade 2 comprises one species confined to lowland savanna habitats (Mozambique and Zimbabwe). A third clade comprises R. eloquens from East Africa. Speciation within Clade 1 is associated with fixed differences in echolocation call frequency, and cranial shape and size in populations isolated since the late Pliocene (ca 3.74 Mya). Relative to the intermediate-sized savanna population (Clade 2), these island-populations within Clade 1 are characterised by either gigantism (South African eastern Great Escarpment and Mts Mabu and Inago in Mozambique) or dwarfism (Lutope-Ngolangola Gorge, Zimbabwe and Soutpansberg Mountains, South Africa). Sympatry between divergent clades (Clade 1 and Clade 2) at Lutope-Ngolangola Gorge (NW Zimbabwe) is attributed to recent range expansions. We propose an “Allometric Speciation Hypothesis”, which attributes the evolution of this species complex of bats to divergence in constant frequency (CF) sonar calls. The origin of species-specific peak frequencies (overall range = 32 to 46 kHz) represents the allometric effect of adaptive divergence in skull size, represented in the evolution of gigantism and dwarfism in habitat islands.     

Hidden Diversity in Sardines: Genetic and Morphological Evidence for Cryptic Species in the Goldstripe Sardinella,Sardinella gibbosa (Bleeker, 1849)

Cryptic species continue to be uncovered in many fish taxa, posing challenges for fisheries conservation and management. In Sardinella gibbosa, previous investigations revealed subtle intra-species variations, resulting in numerous synonyms and a controversial taxonomy for this sardine. Here, we tested for cryptic diversity within S. gibbosa using genetic data from two mitochondrial and one nuclear gene regions of 248 individuals of S. gibbosa, collected from eight locations across the Philippine archipelago. Deep genetic divergence and subsequent clustering was consistent across both mitochondrial and nuclear markers. Clade distribution is geographically limited: Clade 1 is widely distributed in the central Philippines, while Clade 2 is limited to the northernmost sampling site. In addition, morphometric analyses revealed a unique head shape that characterized each genetic clade. Hence, both genetic and morphological evidence strongly suggests a hidden diversity within this common and commercially-important sardine.     

Pseudo-nitzschia pungens (Bacillariophyceae): A cosmopolitan diatom species?

Genetic, reproductive and morphological variation were studied in 193 global strains of the marine diatom species Pseudo-nitzschia pungens (Grunow ex Cleve) Hasle to assess potential intraspecific variation and biogeographic distribution patterns. Genetic differentiation between allo- and sympatric strains was investigated using the ITS1–5.8S–ITS2 rDNA region. Three ITS clades were found. Clones of opposite mating type were sexually compatible within clades I or II, and viable F1 hybrid offspring were produced in crosses between them. The molecular differences between these clades were correlated with slight but consistent morphological differences. At present, nothing can be said about morphology and mating behavior for clade III clones because only ITS data were available. The three ITS clades showed different geographic distributions. Clade II was restricted to the NE Pacific, whereas clones belonging to clade III originated from geographically widely separated areas (Vietnam, China and Mexico). ITS clade I was recovered in all locations studied: the North Sea (Belgium, The Netherlands, France), the eastern and western N Atlantic (Spain, Canada), the NW and S Pacific (Japan, New Zealand) and the NE Pacific (Washington State). Clade I thus appears to be globally distributed in temperate coastal areas and provides the first strong evidence to date for the global distribution of a biologically, genetically and morphologically defined diatom species.     

Phylogenetic reconstruction of the genus Triptilion (Asteraceae,Nassauvieae) based on nuclear and chloroplast DNA sequences

The genus Triptilion is endemic to central Chile, the Mendoza Province and western Patagonia in Argentina. It is currently composed of seven species: T. achilleae, T. benaventii, T. berteroi, T. capillatum, T. cordifolium, T. gibbosum, and T. spinosum. The main objectives of this paper were to determine the phylogenetic relationships of species of Triptilion. We also traced the evolution of annual and perennial life-forms. Historically a close relationship has been described between genera Triptilion and Nassauvia. Phylogenetic analysis of the genus Triptilion and more closely related genera was undertaken using two nuclear (ITS, ETS) and two chloroplast (trnL-F, rpl32-trnL) markers. The topology of the Bayesian inference tree shows that the genus Triptilion is paraphyletic, because Nassauvia lagascae, the only representative of Nassauvia section Caloptilium grouped with T. achilleae, Clade I. The other species of Triptilion form two clades: Clade II composed of T. cordifolium and T. gibbosum and Clade III that includes T. benaventii, T. berteroi, T. capillatum, and T. spinosum. The genus Triptilion originated and diverged during the Miocene. The results of the life history reconstructions indicate that the common ancestor of Triptilion and Nassauvia was perennial. The annual habit appears to be derived in Triptilion. The life-form of the common ancestor of Triptilion was ambiguous; it may have been annual or perennial.     

采自吉林省的中国丝盖伞属新记录种

报道了采自吉林长白山的中国丝盖伞属丝盖伞亚属Inocybe subgen. Inocybe的3个新记录种：黄囊丝盖伞I.?muricellata、新卷毛丝盖伞I. neoflocculosa和灰白丝盖伞I. senkawaensis,并提供详尽的形态描述和线条图。研究标本存放于吉林农业大学菌物标本馆（HMJAU）。     

Evolution of bird communities in the Neogene of Central Asia,with a review of the Neogene fossil record of Asian birds

A complete taxonomic review of Neogene birds of continental Asia is provided. To date, avifauna from the latter half of the Miocene and Pliocene of Central Asia (Mongolia and adjacent regions of Inner Asia) are most thoroughly investigated. Available data enable a reconstruction of successive replacement of Early and Middle Miocene avifaunas by communities of the Recent type. Middle Miocene avifaunas of Mongolia include a great number of extinct genera and species, many of which were widespread in Eurasia. Extant genera became dominant in the Late Miocene and taxa close to living species appear in the Late Pliocene fossil record. Late Pliocene communities of birds of Central Asia were complex in genesis, composed of Miocene relicts (Struthio), immigrants from the European regions of the Palearctic (phasianid Plioperdix), North American immigrants (Calcarius), and also autochthonous elements, the origin of which is apparently connected with the arid belt of Central Asia (diverse passerines).     

Colliding fragment islands transport independent lineages of endemic rock-crawlers (Grylloblattodea: Grylloblattidae) in the Japanese archipelago

Fragment islands, viewed from the paradigm of island biogeographic theory, depend on continual immigration from continental sources to maintain levels of species diversity, or otherwise undergo a period of relaxation where species diversity declines to a lower equilibrium. Japan is a recently derived fragment island with a rich endemic flora and fauna. These endemic species have been described as paleoendemics, and conversely as recently derived Pleistocene colonists. Geological events in the Miocene period, notably the fragmentation and collision of islands, and the subsequent uplift of mountains in central Japan, provided opportunities for genetic isolation. More recently, cyclical climatic change during the Pliocene and Pleistocene periods led to intermittent land bridge connections to continental Asia. Here we investigate the pattern and timing of diversification in a diverse endemic lineage in order to test whether ongoing migration has sustained species diversity, whether there is evidence of relaxation, and how geological and climatic events are associated with lineage diversification. Using multi-locus genetic data, we test these hypotheses in a poorly dispersing, cold-adapted terrestrial insect lineage (Grylloblattodea: Grylloblattidae) sampled from Japan, Korea, and Russia. In phylogenetic analyses of concatenated data and a species tree approach, we find evidence of three deeply divergent lineages of rock-crawlers in Japan consistent with the pattern of island fragmentation from continental Asia. Tests of lineage diversification rates suggest that relaxation has not occurred and instead endemism has increased in the Japanese Grylloblattidae following mountain-building events in the Miocene. Although the importance of climate change in generating species diversity is a commonly held paradigm in Japanese biogeography, our analyses, including analyses of demographic change and phylogeographic range shifts in putative species, suggests that Pleistocene climatic change has had a limited effect on the diversification of rock-crawlers.     

Multigene molecular phylogenetics reveals true morels (Morchella) are especially species-rich in China

The phylogenetic diversity of true morels (Morchella) in China was estimated by initially analyzing nuclear ribosomal internal transcribed spacer (ITS) rDNA sequences from 361 specimens collected in 21 provinces during the 2003-2011 growing seasons, together with six collections obtained on loan from three Chinese herbaria. Based on the results of this preliminary screen, 40 Esculenta Clade (yellow morels) and 30 Elata Clade (black morels) were chosen to represent the full range of phylogenetic diversity sampled. To investigate their species limits, we generated DNA sequences from portions of three protein-coding genes (RPB1, RPB2 and EF-1α) and domains D1 and D2 of the nuclear large subunit (LSU) rDNA for all 70 collections. To fully assess evolutionary relationships, previously published multilocus DNA sequence data representing all known Morchella species was included in this study. Phylogenetic analyses employing maximum parsimony and maximum likelihood frameworks resolved 30 species in China compared with 22 in Europe and 19 within North America. Eleven novel phylogenetically distinct species were discovered in China, including two species within the Elata Clade and nine within the Esculenta Clade. Of the 30 species in China, 20 appear to be endemic, nine were also represented in Europe, and four putatively fire-adapted species have disjunct distributions in China, Europe and western North America. Although the diversification time estimates place the Esculenta Clade in China as early as the late Cretaceous and the Elata Clade by the early Oligocene, 27 of the 30 species evolved between the middle Miocene 12Mya and present.