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Index map of the Ryukyu Islands, Japanese Islands (Honshu, Hokkaido, Shikoku, and Kyushu), and Taiwan. Submarine contours are 200, 500, 1000, 2000, 3000, 4000, 5000 m. Base map from Vector Map (VMap) Level 0, National Geospatial-Intelligence Agency.
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In the Quaternary, the Ryukyu Islands evolved from a continental margin arc to an island arc by backarc spreading of the Okinawa Trough, accompanied by subsidence and isolation of the islands, a process that has continued to the present. Trough-parallel half grabens were filled with marine siltstone. Similar sediments filling orthogonal fault-contr...
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... Ryukyu island arc extends between Kyushu (southern Japanese islands) and Taiwan ( Figure 1). It consists of a chain of small islands that is convex (east) oceanwards in map view (Figure 1). ...
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... Ryukyu island arc extends between Kyushu (southern Japanese islands) and Taiwan ( Figure 1). It consists of a chain of small islands that is convex (east) oceanwards in map view (Figure 1). West of the arc, the Okinawa Trough is currently undergoing backarc spreading ( Letouzey and Kimura 1986;Park et al. 1998). ...
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... of the arc, the Okinawa Trough is currently undergoing backarc spreading ( Letouzey and Kimura 1986;Park et al. 1998). Because this opening is occurring in the absence of a buttress, the arc consists of small, subsided islands ( Figure 1). The Japan Sea ini- tially opened before 15 Ma, then stopped extending, and the uplift associated with continued subduction made the Japanese islands much larger than the Ryukyu Islands ( Figure 1; Osozawa 1997;Taira 2001). ...
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... this opening is occurring in the absence of a buttress, the arc consists of small, subsided islands ( Figure 1). The Japan Sea ini- tially opened before 15 Ma, then stopped extending, and the uplift associated with continued subduction made the Japanese islands much larger than the Ryukyu Islands ( Figure 1; Osozawa 1997;Taira 2001). Active backarc *Corresponding author. ...
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... Okinawa Trough has trapped a large amount of continentally derived clastic detritus, estimated to be up to 1000 m in thickness (Park et al. 1998), and mostly supplied by major streams, such as the Yangtze River (Figure 1). Yellow River detritus may flow in the northern Okinawa Trough through the Goto submarine canyon (Figure 1; e.g. ...
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... Okinawa Trough has trapped a large amount of continentally derived clastic detritus, estimated to be up to 1000 m in thickness (Park et al. 1998), and mostly supplied by major streams, such as the Yangtze River (Figure 1). Yellow River detritus may flow in the northern Okinawa Trough through the Goto submarine canyon (Figure 1; e.g. Oiwane et al. 2010). ...
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... marine transgression is recorded near the mouths of drainages that flowed in all directions, the westward-flowing drainages definitively record separa- tion of the landmass from the Chinese mainland to the west and creation of the island (Figure 2, 1 T a iw a n S tr a it OS: Osumi I. Figure 4. Shaded relief maps of Amami Oshima (A2), Okinawa-jima (O1), and Yaeyama (Y) islands (Ishigaki-jima and Iriomote-jima). Flat parts are covered by the Ryukyu Group. ...
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... most of the Ryukyu Islands, transgression continued, resulting in the deposition of Ryukyu limestone shortly after deposition of the earliest island-fringing sediments (Figure 2, 1.0 Ma). ...
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... summary of age constraints is given in graphical tabular form in Figure 5, and the marine siltstone age or basal limestone age is interpreted as the separation age based on the rationale given above. We also discuss the Tokara, Kerama, and Yonaguni gaps (Figures 1 and 3), which play a role in the isolation history (Figure 2), for they separate subgroups of the island chain from each other. ...
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... time when Kyushu, and therefore the rest of the Japanese islands were separated from Korean Peninsula and Asian mainland, corresponds to when the Tsushima warm current began to enter the Japan Sea. This geologic event is difficult to constrain from the geol- ogy of Kyushu because of the presence of two straits between Korea and Kyushu on either side of the island of Tsushima that has Palaeogene basement (Figure 1). The present Tsushima Current runs through both the Tsushima Strait (130 m depth) between Tsushima and Kyushu and the Korea Strait (230 m depth) between Tsushima and Korea. ...
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... Ma ( Sato et al. 1999; modified to 0.265 Ma by Sato et al. 2009). In addition, sediment core from the mouth of the Tsugaru Strait (Figure 1) records outflow of the Tsushima Current Figure 5. Graphical table of geochronologic constraints for the Shimajiri Group siltstone, Nakoshi Formation siltstone (time of isola- tion of island), reddish limestone, and Ryukyu limestone. Nannofossil datum planes correlated to the magnetic polarities are after Sato et al. (2009). ...
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... Watase line corresponds to the Tokara gap ( Figures 1 and 3), a narrow submarine valley connecting the Ryukyu trench to the Okinawa Trough that orthogonally truncates the Ryukyu arc chain and has a depth of 1500 m at the point of truncation. The Tokara gap divides the cor- respondingly named islands into northern and southern island subgroups. ...
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... Goto submarine canyon is also a fault-controlled valley (Figures 1 and 3; Oiwane et al. 2010), and together with the Tokara gap may represent part of the same sub- merged drainage system. If so, the Tokara gap may repre- sent the palaeo-mouth of the Yellow River (and the Huai River), reflecting when the Ryukyu area was a continental margin arc (cf., Figure 2). ...
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... island occupies the forearc high to the east of Amami Oshima (Figure 1). Its basement consists of marine siltstone of the Shimajiri Group, dated as planktonic foraminifera zone N22 (and older) ( Figure 5; Ujiié 1994), in agreement with the earlier work (Huang 1966). ...
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... Guga Formation fills these valleys. Upstream, these are terrestrial deposits, and downstream, they are shallow marine delta or fan deposits (Figure 2, 1.5 Ma), which were formed close to the present river mouth and shore- line. The maximum altitude of the marine Guga Formation is 100 m, and the Guga Formation is distributed both east- ern and western coast valleys of Okinawa-jima. ...
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... is because there was a large bay along the Kerama gap, with water deep enough to cover what became Kume-jima. Nakamura et al. (1999) postulated that the large-scale, cross-stratified sandstone (part of the Shimajiri Group) at Kume-jima represents deltaic sediments from the former Yangtze River (Figures 1-3). ...
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... present Kerama gap (Figures 1 and 3) is controlled by a graben (cf., Figure 2, 1.5 Ma). A series of normal faults trend NW-SE and dip SW in the area northeast of the axial part of the gap, cutting the Shimajiri Group and the Ryukyu limestone on both sides of the gap (Kimura 1996). ...
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... lies on the main Ryukyu Island chain (Figures 1 and 3), but lay originally at the eastern margin of the Chinese mainland based on the fact that the basement consists solely of the marine Shimajiri Group. Ryukyu limestone unconformably rests on the Shimajiri Group ( Figure 6E) and consists of basal conglomerate (part) and alternation of coral and detrital limestones (Honda et al. 1993). ...
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... isotopic compositions also reflect warm water (Lee 2000). There may be a branch of Kuroshio current passing through the Taiwan Strait (Figures 1 and 3). Owing to the shallow depth of the Taiwan Strait, it is likely that land bridges may have formed between Taiwan and the mainland during some glacial sea level lowstands. ...
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... the rifting of the Okinawa Trough began, a rift valley formed in the backarc of the Ryukyu arc (Figures 3 and 9, 1.5 Ma). The extension was con- trolled by NE-SW-striking normal faults, including the Nago fault (Osozawa and Watanabe 2011) on Okinawa- jima (O1). ...
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... small branch of extension (and also the Kuroshio current) might have existed between Tanega- shima (OS-1) and Yaku-shima (OS-2), as reflected by the fission track age of Tanega-shima (OS-1; Otsuka and Kuwayama 2000). The Tsushima gateway allowed inflow of the Kuroshio current to the Japan Sea at 1.552 ± 0.154 Ma ( Kitamura and Kimoto 2006), so the northern end of the rift valley likely extended to the Tsushima gate- way, through the Goto channel (Figure 9, 1.5 Ma). Marine deposits also record warm water through the Taiwan Strait, but such inflow would be unrelated to Okinawa Trough rifting. ...
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... deposits also record warm water through the Taiwan Strait, but such inflow would be unrelated to Okinawa Trough rifting. Japan, the Ryukyu Islands, and Taiwan separated from the continent at about the same time, 1.552 ± 0.154 Ma, through initial extension of the Okinawa Trough, for 1000 km along strike (Figure 9, 1.5 Ma). ...
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... the Tokara, Kerama, and Yonaguni gaps are NW-SE-trending normal fault valleys or probably N-S-trending fault valleys (Ujiié 1983;Kimura 1996;Lallemond et al. 2001) mostly orthogonal to the NE- SW-trending main rift valley, these gaps, particularly the former two, are also expected to be branches of the main rift, formed at 1.552 ± 0.154 Ma. The Kuroshio current entered through the Yonaguni gap, mixed with the western offshore warm current of Taiwan, and flowed out through the Tokara gap, with another branch separating to become the Tsushima Current through the Tsushima gateway (Figure 9, 1.5 Ma). ...
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... rifting was progressed, sea floor spreading followed to form the Okinawa Trough. Detrital sediments from the Yellow and Yangtze rivers were captured in the trough, and under the influence of the warm Kuroshio current, coral reefs formed around each island (Figure 2, 1.0 Ma). ...
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... Okinawa-jima (O1), how- ever, the subsidence rate accelerated at ca. 0.8 Ma, resulting in deposition of upper detrital limestone over reef lime- stone, before slowing. When the Ryukyu limestone was deposited, each island was much smaller than at present, and they could not have been connected to one another (Figure 2, 1.0 Ma). ...
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... the manner of a phylogenetic tree of biological species, we can graphically show the lineage/connection of each Ryukyu Island, including the Japanese Islands and Taiwan, compared to the mainland ( Figure 10). Our geological conclusion is that each island of Ryukyu formed since 1.552 ± 0.154 Ma, and as a consequence, new endemic species evolved on each island. ...
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Citations
... reveal mixing in various proportions with fine-grained sediment shed from other sources, including the Luzon and Ryukyu arcs and, subordinately, rivers of southeastern China (Fig. 15). A major role in the redistribution of clays in the western Philippine and northeastern South China seas is played by the northward flowing Kuroshio oceanic current (Fig. 1B;Liu et al., 2010b;Liu et al., 2011;Osozawa et al., 2012). ...
... Almost all islands are of continental origin. They formed on the eastern margin of the Asian continent in the past and underwent a complex geological history in the last two million years (Ota 1998;Osozawa et al. 2012; Government of Japan 2019), resulting in complex biogeography of lineages inhabiting the islands (e.g., Ota 1998; Kato and Yagi 2004;Kaito and Toda 2016;Hirano et al. 2019). The combination of all these aspects, i.e., the position of the islands, their climate, and their recent repeated connections and isolations from the continent and neighbouring islands, is the primary reason for the unique fauna and flora inhabiting the Ryukyu Archipelago. ...
We review the genus Anacaena Thomson, 1859 from the Ryukyu Archipelago, southern Japan. Three aquatic species are recognised: A. torikaii sp. nov. from Amami-ôshima Island, A. okinawana sp. nov. from Okinawa-jima Island and Kerama Islands, and A. kumejimana sp. nov. from Kumejima Island. All three species are very similar, with the morphology of the aedeagus being essential for a reliable identification. Dorsal colouration is also useful as a diagnostic character, despite some variation within species. We observe a possible geography-based variation between A. okinawana from Okinawa-jima I. and the neighbouring Kerama Is., but we treat both populations as conspecific based on genital morphology. Anacaena kumejimana and A. okinawana share many morphological characters possibly indicating their close relationship. We compare the endemism of aquatic Hydrophilidae in the Ryukyu Archipelago to that in other groups of aquatic beetles: the proportion of endemic species is higher in aquatic Hydrophilidae than in Dytiscidae, but much lower than in stream-inhabiting Hydraenidae and Elmidae. A list of Japanese species of Anacaena and a key to the Japanese species of the genus are provided.
... node 10 in Fig. 5 and Table 3], M. fruhstorferi on the Ryukyu Islands diversified. However, the past Ryukyu Islands stretched from north to south, and the separation of this archipelago began at approximately 1.55 Ma (Osozawa et al., 2012). The Meghimatium clades in this region may diversify prior to the separation of the islands. ...
East Asia, specifically the Japanese Archipelago, is a biodiversity hotspot of both vertebrates and invertebrates. Mollusks represent a burst of species diversity in this region due to the effects of biotic and abiotic factors on their morphological traits, such as shell shape and size. However, the evolutionary history of terrestrial slugs in East Asia remains unknown. In the present study, we investigated the molecular phylogeny of terrestrial slugs of the genus Meghimatium. This genus includes three described and eight undescribed species, and our study used all except for two. Based on phylogeny and the species delimitation tests, the genus Meghimatium was split into many putative species, suggesting higher species diversity than previously thought based on morphological and anatomical studies and that almost undescribed species may be inappropriate. Therefore, morphological traits, such as body size and colour, conventionally considered for classification may easily vary or be similar across geographic region. Moreover, the divergence time of this genus is almost concordant with the geographical time scale of the formation of the Japanese mainland. Our findings suggest that molecular phylogenetics helps classify Japanese Meghimatium slugs, but comprehensive taxonomic revisions using multi-locus analyses are needed.
... Ma) (Otsuka, 2002;Otsuka and Takahashi, 2000). However, dispersal of earthworms from the Ryukyus to the Japanese Islands may have been absent since the late Early Pleistocene-the connection between the Ryukyu Islands and the main Japanese Islands was lost after 1.55 Ma (Osozawa et al., 2011). ...
Megascolecid earthworms of the pheretimoid group are dominant detritivores of soil ecosystems in the Japanese Archipelago and East Asia. However, their diversity and phylogenetic relationships are poorly understood. We assembled whole mitogenome sequences for 197 megascolecid earthworms collected throughout Japan to study the phylogenetic relationships, phylogeography, divergence times, and diversification of important morphological characteristics among pheretimoid earthworms. Using 197 mitogenome sequences and 24 published mitogenome sequences from the East Asian mainland (221 sequences in total), we constructed a maximum likelihood tree and found that the pheretimoid earthworms currently assigned to Amynthas, Metaphire, Duplodicodrilus, and Manus are involved in the most senior genus Amynthas; thus, Amynthas can be treated as the sole genus encompassing all of the above genera. Within the Amynthas group, we identified three major lineages that led to four groups of endemic species in Japan. These lineages originated from different lineages on the East Asian mainland and Taiwan Island, indicating multiple colonization events from the East Asian mainland by different ancestral lineages, possibly after the Miocene. We also assembled nuclear ribosomal DNA sequences encompassing the 18S to 28S rRNA genes. The nuclear gene tree showed major groups consistent with the mitogenome tree except for different (and not well-resolved) relationships among major clades. Our molecular data covered 115-158 native and 7 non-native Amynthas group species in Japan in terms of DNA-based species delimitation. Our findings provide a basis for understanding the evolutionary relationships among diversified megascolecid earthworms in the Amynthas group in Japan and adjacent regions.
... Paleogeographic reconstruction of the islands' formation suggests that they were isolated from mainland Japan, mainland China, and Taiwan, ca. 1.55 million years ago (Osozawa et al., 2012). The unique flora of the Amami Islands is well recognized 17.3 (9-24) 6.4 (2)(3)(4)(5)(6)(7)(8)(9)(10)(11)(12) 53.6 59.7 16.0 76.8 51.7 TREES ATI SKS OKN 7.9 (5)(6)(7)(8)(9)(10)(11) 3.5 ...
Amami‐Oshima and Tokunoshima Islands (the Amami Islands) are part of an important area for biogeography in Japan. Although the uniqueness of the islands' flora is well recognized, their plant diversity, including non‐tree life forms, has yet to be compared with that of other regions. Here, we characterize the plant community structure of the Amami Islands, and analyze the α and β diversity patterns with respect to environmental factors as well as the frequency of threatened and endemic species compared with that in regions at higher and lower latitudes. We used 5 × 100 m plots at seven sites on the Amami Islands, three sites further north, and one site further south to record the abundance of all vascular plant species and tree sizes. We also used tree census data from five plots from the Monitoring Sites 1000 Project. Three principal components from five environmental variables were used as factors affecting diversity. We found that the broad‐leaved evergreen forests on the Amami Islands have higher α diversity resulting from species richness anomalies in shared families among regions. Additionally, generalized linear mixed models indicated that temperature and altitude affect α diversity positively. Furthermore, multiple regression models on distance matrices showed that temperature and regional differences in weather conditions affect β diversity. Tree species were found to affect species diversity and rareness, but herbaceous plants also affect these parameters substantially. These results indicate that the two studied islands are biodiversity hotspots and that evaluating herbaceous species is important for determining forest biodiversity.
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The study of population genetic structure and historical dynamics of species with disjunct distribution can reveal the mechanisms through with they were formed. Pinus densiflora is an essential tree species with ecological and economic value, and its natural distribution shows a disjunct pattern. Using transcriptome-level SNP data from 220 samples representing 32 naturally-distributed populations in East Asia, we investigated Pinus densiflora genetic diversity and structure, divergence time, and ancestral distribution. We identified five subpopulations which diverged approximately 2.02–1.49 million years ago, and found relatively low genetic differentiation among the three large subpopulations (SL, JH, and JK). Northeast China is the most likely origin, and its current distribution is the result of dispersal and vicariance events. It migrated southwest through the Liaodong Peninsula to the Shandong Peninsula and southeast through the Korean Peninsula to Japan. These results provide a basis for the conservation and management of P. densiflora in the future and the evolutionary study of species with similar life histories.
... In this section, we trace transport pathways of detritus generated on Taiwan Island from the river mouths to the deep-sea, where clay mineralogy (Fig. 6), elemental geochemistry (Fig. 13), and Sr and Nd isotopic ratios (Fig. 14) reveal mixing in various proportions with fine-grained sediment shed from other sources, including the Luzon and Ryukyu arcs and, subordinately, rivers of southeastern China (Fig. 15). A major role in the redistribution of clays in the western Philippine and northeastern South China seas is played by the northward flowing Kuroshio oceanic current ( Fig. 1B; Liu et al., 2010b;Liu et al., 2011;Osozawa et al., 2012). ...
... The restriction of melanic D. elegans and D. gunungcola to cooler conditions may be a case of niche conservatism (48-50). Taiwan and Okinawa islands, where the black morph of D. elegans is found, became isolated from the mainland around 1.5 million years ago (51). D. elegans might have colonized Taiwan from the mainland across the shelf area when the sea level dropped during the glacial period, which is believed to be the case in many taxa, such as damsel ies and ea beetles (52,53). ...
Background:
Variations in body size and body melanization are thought to be important features for local adaptation of environmental stresses in many insects and latitudinal clines of such variation have been found many taxa. When two species share similar resource, ecological divergence and niche partitioning may further evolve as the consequence of competition. Here, we examined the distribution, host plant usage, and body size variation of two closely-related species, Drosophila elegans, which has two discrete body color morphs, and D. gunungcola on three islands.
Results:
The brown morph of D. elegans has a similar distribution to D. gunungcola in Java and Sumatra, whereas the black morph of D. elegans is exclusively found in Taiwan. A significant correlation between latitudes and altitudes was found in sites where D. gunungcola was found in Sumatra south of equator. The brown morph of D. elegans was found to be smaller in body size and tends to live in warmer habitat compared to the black morph of D. elegans and D. gunungcola. A significant genetic correlation between body color and body size was found in recombinant inbred lines derived from hybrids of brown and black morph strains.
Conclusions:
The restricted distribution of D. gunungcola in Southern hemisphere coincides with the lack of highland habitat near the equator. Four plant species were found to be exclusively utilized by D. elegans only in Taiwan whereas the same flower species are shared by both fly species in Java and Sumatra, suggesting the presence of inter-specific competition for breeding sites in overlapping zones. Darker body coloration with larger body size appears to have evolved twice in this lineage, reflecting similar patterns of natural selection in Indonesia and postglacial Taiwan.
... Importantly, the three aforementioned archipelagos represent a gradient of geologic ages and degrees of geographic isolation. The Ryukyus are the oldest and most geographically connected, having originated as part of the Asian continent during the Permian Period (299-251 million years ago [Ma]; Hanzawa, 1935) and later separated into islands by tectonic activity approximately 1.5 Ma (Osozawa et al., 2012). Unlike hotspot archipelagos, the Ryukyus are continental in origin (Hanzawa, 1935) Myr old (Clague & Sherrod, 2014). ...
Islands make up a large proportion of Earth’s biodiversity, yet are also some of the most sensitive systems to environmental perturbation. Biogeographic theory predicts that geologic age, area, and isolation typically drive islands’ diversity patterns, and thus potentially impact non‐native spread and community homogenization across island systems. One limitation in testing such predictions has been the difficulty of performing comprehensive inventories of island biotas and distinguishing native from introduced taxa. Here, we use DNA metabarcoding and statistical modeling as a high throughput method to survey community‐wide arthropod richness, the proportion of native and non‐native species, and the incursion of non‐natives into primary habitats on three archipelagos in the Pacific ‐ the Ryukyus, the Marianas and Hawaii ‐ which vary in age, isolation and area. Diversity patterns largely match expectations based on island biogeography theory, with the oldest and most geographically connected archipelago, the Ryukyus, showing the highest taxonomic richness and lowest proportion of introduced species. Moreover, we find evidence that forest habitats are more resilient to incursions of non‐natives in the Ryukyus than in the less taxonomically rich archipelagos. Surprisingly, we do not find evidence for biotic homogenization across these three archipelagos: the assemblage of non‐native species on each island is highly distinct. Our study demonstrates the potential of DNA metabarcoding to facilitate rapid estimation of biogeographic patterns, the spread of non‐native species, and the resilience of ecosystems.
... The areas where T. jackii and two species of Amentotaxus occur are either presently overlapping or have experienced geological and floristic connection through time, particularly considering that Taiwan became separated from the Chinese mainland only ca. 1.5 Ma (Osozawa et al., 2012;Tang et al., 2013). Thus, T. jackii may have had the opportunity to hybridize with Amentotaxus. ...
Restriction site‐associated DNA sequencing (RAD‐seq) enables obtaining thousands of genetic markers for phylogenomic studies. However, RAD‐seq data are subject to allele dropout (ADO) due to polymorphisms at enzyme cutting sites. We developed a new pipeline, RADADOR, to mitigate the ADO in outgroups by recovering missing loci from previously published transcriptomes in our study of a gymnosperm genus Torreya. Using the supplemented RAD‐seq data in combination with plastome and mitochondrial gene sequences, morphology, and fossil records, we reconstructed the phylogenetic and biogeographic histories of the genus and test hypotheses on diversity anomaly in eastern Asian‐North American floristic disjunction. Our results showed that our pipeline recovered many loci missing from the outgroup, and the improved data yielded a more robust phylogeny for Torreya. Using the fossilized‐birth‐death model and divergence‐extinction‐cladogenesis method we resolved detailed biogeographic history of Torreya that suggested a Jurassic origin in the Laurasia and differential speciation and extinction among continents accounting for the modern diversity anomaly biased toward Eastern Asia (EA). The history also supported a vicariance origin of the modern Torreya from a widespread ancestor in EA and NA in the mid‐Eocene, cross‐Beringia exchange in the early Paleogene before the vicariant isolation, in contrast to the “Out of NA” pattern common to gymnosperms and in contrast to the “Out of EA” hypothesis previously proposed for the genus. Furthermore, we observed phylogenetic discordance between the nuclear and plastid phylogenies on T. jackii, suggesting differential lineage sorting of plastid genomes among Torreya species or plastid genome capture in T. jackii. This article is protected by copyright. All rights reserved.
