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MCC continuous coalescent tree (n = 148) of Bufo gargarizans. High posterior probability nodes are represented in red circles. Green clade; Western China, red clade; Korea (+ 5 Chinese haplotypes), blue clade; all Chinese localities. DOI: 10.7717/peerj.4044/fig-4
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The effects of ice ages on speciation have been well documented for many European and North American taxa. In contrast, very few studies have addressed the consequences of such environmental and topographical changes in North East Asian species. More precisely, the Korean Peninsula offers a unique model to assess patterns and processes of speciatio...
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... Phylogenetic relationships between Old World bufonids (genera Bufo, Bufotes, Strauchbufo, and others) are under extremely intensive research. The results of these studies were summarized and discussed in fundamental publications [4,26,27], which hypothesize the dispersal of the Bufo genus path in the Palearctic. They integrate phylogeographic and taxonomic research, underlying that they are two rapidly evolving fields in an exciting era of new species discoveries. ...
... Within the genus, the "Western" Bufo complex and the east Asian Bufo gargarizans complex are distinguished [3,7]. The Bufo gargarizans species complex includes a number of species: the mainland Asiatic toad, B. gargarizans sensu lato (which includes B. tibetanus, B. andrewsi, and B. minshanicus), as well as the Taiwanese Bankor toad, B. bankorensis [3,7,8,[26][27][28][29][30]. Bufo gargarizans s.s., B. sachalinensis, and B. andrewsi host Pleistocene diversifications, but appear too young (<2 Myr) to underly additional speciation events [26,28,29]. ...
... The Bufo gargarizans species complex includes a number of species: the mainland Asiatic toad, B. gargarizans sensu lato (which includes B. tibetanus, B. andrewsi, and B. minshanicus), as well as the Taiwanese Bankor toad, B. bankorensis [3,7,8,[26][27][28][29][30]. Bufo gargarizans s.s., B. sachalinensis, and B. andrewsi host Pleistocene diversifications, but appear too young (<2 Myr) to underly additional speciation events [26,28,29]. Our phylogenetic tree agreed well with earlier phylogenies of the genus Bufo in East Asia [3,4,7,20,21]. ...
Based on a combination of molecular and morphological data, we herein report a new species within the bufonid Bufo gargarizans species complex. This is a widespread species complex with distribution from eastern Russia and the Korean Peninsula to China and the Ryukyu Islands of Japan. Records of this species have been documented in the Guangxi and Yunnan Provinces near the border with Vietnam and, for the first time from Vietnam, in Ha Giang Province. The new record of Bufo cf. gargarizans from Vietnam is from Ha Tinh Province. This species has never been reported from Vietnam so far south, about 550 km south from the previously known locality in Ha Giang Province. The female specimen was found in the Ha Tinh Province, Vu Quang National Park of central Vietnam and two specimens (male and female) were found Ha Giang Province. They are clearly distinguished from B. gargarizans and all the mentioned species by a specific color pattern on the belly and creamy-yellowish throat with large, bright red speckles. Genetic divergences of three Vietnam specimens from Ha Giang and Ha Tinh Provinces in the ND2 gene sequences between the В. sp. nov. and all other congeners ranged from 4.3% (with B. andrewsi) to 7.0% (with B. stejnegeri). We give a description of the morphological characters and coloration of the new record and provide an expanded diagnosis.
... This refers to the unusually similar distribution patterns between the current and the LGM scenarios. This has been further supported by studies of the phylogeography and demographic history of Southeast Asian species [96][97][98][99]. ...
The Mongolian racerunner, Eremias argus, is a small lizard endemic to Northeast Asia that can serve as an excellent model for investigating how geography and past climate change have jointly influenced the evolution of biodiversity in this region. To elucidate the processes underlying its diversification and demography, we reconstructed the range-wide phylogeographic pattern and evolutionary trajectory, using phylogenetic, population genetic, landscape genetic, Bayesian phylogeographic reconstruction and ecological niche modeling approaches. Phylogenetic analyses of the mtDNA cyt b gene revealed eight lineages that were unbounded by geographic region. The genetic structure of E. argus was mainly determined by geographic distance. Divergence dating indicated that E. argus and E. brenchleyi diverged during the Mid-Pliocene Warm Period. E. argus was estimated to have coalesced at~0.4351 Ma (Marine Isotope Stage 19). Bayesian phylogeographic diffusion analysis revealed out-of-Inner Mongolia and rapid colonization events from the end of the Last Interglacial to the Last Glacial Maximum, which is consistent with the expanded suitable range of the Last Glacial Maximum. Pre-Last Glacial Maximum growth of population is presented for most lineages of E. argus. The Glacial Maximum contraction model and the previous multiple glacial refugia hypotheses are rejected. This may be due to an increase in the amount of climatically favorable habitats in Northeast Asia. Furthermore, E. argus barbouri most likely represents an invalid taxon. The present study is the first to report a range-wide phylogeography of reptiles over such a large region in Northeast Asia. Our results make a significant contribution towards understanding the biogeography of the entire Northeast Asia.
... There are two major mountain ranges in South Korea, the Taebaek Mountains stretching along the east coast and running north to south, and the Sobaek Mountains dividing from the middle of the former range and running northeast to southwest direction (Kwon et al. 2016;NGII 2020). These mountain ranges have been known to play an important role in generating high species and genetic diversity in various organisms (Choi 2004;Choe et al. 2016;Chung et al. 2018), and such value as biodiversity reservoirs seems to be due to not only the role of the regions as glacial refugia during the Pleistocene (Chung et al. 2017;Borzée et al. 2017), but also the formation of the mountain ranges in the Korean Peninsula during the Miocene (Kim 1997;Kim & Hwang 2023). ...
Nine new species belonging to the Pholcus phungiformes-group are described from Korea, viz., Pholcus chugok Lee & Lee, sp. nov., Pholcus noeun Lee & Lee, sp. nov., Pholcus wonju Lee & Lee, sp. nov., Pholcus ungyo Lee & Lee, sp. nov., Pholcus hwangjeong Lee & Lee, sp. nov., Pholcus gimsatgat Lee & Lee, sp. nov., Pholcus solchi Lee & Lee, sp. nov., Pholcus mino Lee & Lee, sp. nov., and Pholcus maepo Lee & Lee, sp. nov. These new species were found from mountainous habitats in Gangwon-do and Chungcheongbuk-do, the northeastern regions of South Korea where the two major mountain ranges (Taebaeksan mountains and Sobaeksan mountains) are located together. Detailed descriptions, photographs, and a distribution map for the new species are provided with notes on their intraspecific variations of copulatory organs.
... The Bufonidae family has a global distribution, with two species complexes distributed in northeast Asia. Their regional distribution was shaped by several geological events, including glacial oscillations and the resulting sea level variations, several orogeneses, and the development of monsoon systems (Borzée et al., 2017;Fong et al., 2020;Othman et al., 2022aOthman et al., , 2022b. In addition, these natural phenomena were followed by a blurring of geographic boundaries due to anthropogenic activities. ...
There are two Bufonidae genera, Bufo and Strauchbufo, for a total of three and one species respectively, in continental northeast Asia as defined here. The two genera belong to two different species complexes with diverging evolutionary histories.
All four species have generally broad ranges, and they are distributed across a variety of habitats, except for Bufo stejnegeri, a species restricted to higher and cooler habitats. As a result, this species is breeding in cold fast-flowing streams after overwintering under the ice, while the other species breeds in lakes and other broad water bodies.
None of the species is currently highly threatened, although the decline in population size in B. stejnegeri needs attention, and addressing. The main threats are habitat and climate change, but also harvest in some parts of their range. The exact boundary between some of the species needs to be better understood.
... Many colonization events could have originated from the Paria Peninsula, which once extended into Trinidad as a more extensive peninsula. Peninsulas are known to have the capacity to harbor a significant number of reptile and amphibian taxa as they often represent distributional limits for species dispersal through changing conditions such as alternating glacial and interglacial periods (Hewitt 2000;Borzee et al. 2017). ...
The Trinidad and Tobago archipelago is in the southern Caribbean between the Caribbean Sea and the central Atlantic Ocean, northeast of Venezuela. The sister islands are often cited as the southernmost islands of the Lesser Antilles. However, Trinidad is a continental island with strong geologic and geomorphic
connections to mainland South America, having separated approximately 4 Ma. Tobago, on the other hand, is a far-traveled, Cretaceous oceanic arc that formed at the leading edge of the Caribbean plate and was accreted onto the northern coast of South America. Consequently, neither of the two islands are part of the Neogene Lesser Antilles Island arc.
... Many colonization events could have originated from the Paria Peninsula, which once extended into Trinidad as a more extensive peninsula. Peninsulas are known to have the capacity to harbor a significant number of reptile and amphibian taxa as they often represent distributional limits for species dispersal through changing conditions such as alternating glacial and interglacial periods (Hewitt 2000;Borzee et al. 2017). ...
The Trinidad-Tobago archipelago contains two major islands and about 25 satellite islands ranging in size from 3.6 km² to less than 1 ha. Despite different origins, both islands with satellites share a continental biota and have a rich snake fauna originating from vicariance and dispersal events. The authors used published molecular and morphological data to compare the two faunas and here discuss their origins and biogeography. Relatively high snake diversity and local endemics are present on both islands, which share some species with each other or with the mainland. Spatiotemporal distribution patterns illuminated by genetic clocks enhance understanding of evolution in the region and provide a more general example for the study of other biogeographic patterns. The authors also examined the literature for historical records. Trinidad shares many species with the Paria Peninsula in Eastern Venezuela, while Tobago species are more closely related to those from Venezuela’s more distant Central Caribbean Coastal Range.
... The origin and population history of B. g. miyakonis remain unknown. A previous morphological analysis suggested that B. g. miyakonis was closely related to a continental population around Shanghai (Matsui, 1984 (Liu et al., 2000;Fu et al., 2005;Cao et al., 2006;Hu et al., 2007;Yu et al., 2014;Tong et al., 2017;Borzée et al., 2017). Location names denoted by dots with numerals are further described in Appendix II. ...
... Although the obtained sequences reached a maximum of 348 bp, we used a 341 bp sequence that was shared by all sequences of the B. gargarizans species group. The sequences of B. g. miyakonis were aligned by ClustalW, as implemented in MEGA X (Kumar et al., 2018), with sequences from continental populations of B. gargarizans (Fu et al., 2005;Cao et al., 2006;Hu et al., 2007;Borzée et al., 2017;Tong et al., 2017) and Taiwanese populations of B. bankorensis (Liu et al., 2000;Yu et al., 2014) obtained from GenBank (NCBI; Fig. 1A, Appendix II). Similar to the previous studies, the common toad B. bufo (GenBank Accession No: MN122891), which had diverged with B. gargarizans species group in the Late Miocene (Zhan and Fu, 2011;Othman et al., 2022), was chosen as an outgroup. ...
... The continental populations of B. gargarizans were treated as four geographic groups as follows: western China (W), central China (C), eastern China (E), and northeastern China (NE), following Fu et al. (2005) and Hu et al. (2007). The previous studies suggested that the groups from eastern Taiwan (TE), western Taiwan (TW), and Korean peninsula (K) were all monophyletic (Yu et al., 2014;Borzée et al., 2017), so only one haplotype from each of these groups was used for our analyses. ...
The Miyako toad, Bufo gargarizans miyakonis, is a subspecies endemic to the Miyako Islands of the Ryukyu Archipelago. The distribution of B. g. miyakonis is curious since no toads are found on other islands in the remaining part of the Ryukyu Archipelago. In this study, we conducted phylogenetic estimation and population genetic analyses using sequence data of the mitochondrial control region to clarify the phylogenetic position of B. g. miyakonis in the B. gargarizans species group and its population history. Results of phylogenetic analyses suggested that B. g. miyakonis was closely related to eastern populations of B. gargarizans and that there is incomplete lineage sorting among these populations. The divergence time between B. g. miyakonis and its closest lineages was estimated to be 0.54–0.75 Ma, the Middle Pleistocene, which was a little younger than the estimated value provided by the previous study. It is probable that the migration of the ancestor of B. g. miyakonis was strongly affected by the Middle Pleistocene glacial cycle. Population genetics analyses suggested that this subspecies experienced a rapid population size expansion, and that there is moderate genetic differentiation of sub-populations. Although the B. g. miyakonis population has a relatively high-level genetic diversity, it is also unique, having a significant number of accumulated mutations following rapid demographic expansion.
... Zhan and Fu (2011) (Guo 2022). Several other works also contributed to the discussion, including Liu et al. (2000), Chen et al. (2013), Borzée et al. (2017), and Lee et al. (2021). ...
... Several molecular works are solely based on mitochondrial genes (e.g. Macey et al., 1998;Liu et al., 2000;Borzée et al., 2017), which presents specific problems associated with prevalent inter-specific hybridization within Bufonids (more discussion below). This "limited sample/data" problem is well recognized (e.g. Lee et al., 2021), and several studies resorted to combined data analysis, which increases both sample coverage and length of sequences (e.g. ...
... Othman et al. (2022) recently resurrected the name, which includes populations from the Korean Peninsula, the lower Amur river basin, and the Sakhalin Island. Borzée et al. (2017) first recognize a Korean clade based on one mitochondrial region (the control region). Lee et al. (2021) detected a clade (the Far North East clade), which includes populations from the Korean Peninsula and Russian Far East and the most northeastern corner of China based on one mitochondrial gene (ND2) and five nuclear loci. ...
... There are at least two independent historical colonization events from continental populations to the Japanese archipelago, and the land bridge during the LGM mediated the colonization and admixture (Komaki et al., 2015). Land bridge during glaciations may have served as corridor for the common toad (Bufo gargarizans, Borzée et al., 2017) as well. On the contrary, an analysis of the raccoon dog (Nyctereutes procyonoides) revealed no gene flow between the continental populations (China, Vietnam and Korea) and the Japanese archipelago population during glaciations (Kim et al., 2013). ...
... Siberian chipmunk (Tamias sibiricus) may have two refugia existed in South Korea and east Russia-northeast China during the Pleistocene(Lee et al., 2008). Many other species, including the Asiatic toads (Bufo gargarizans,Borzée et al., 2017), the short-tailed pit viper, and the Asian lesser white-toothed shrew (Crocidura shantungensis,Lee et al., 2018), had one or multiple refugia in the peninsula.Many species that are primarily distributed at the Korean peninsula and surrounding areas may also survived the Pleistocene glaciations in situ. The water toad (Bufo stejnegeri) is endemic to northeastern Asia. ...
Continental East Asia has a mild Pleistocene climate and a complex recent geological history. Phylogeographic studies of animals over the last 30 years have produced several distinctive patterns. Glaciation refugia are numerous and are not restricted to any particular regions. Most of them are localized and species-specific, although several large refugia, for example the mountains of SW China, are shared by multiple species and have refugia-within-refugia. Furthermore, postglaciation range expansion events vary greatly in time, scale and direction. Large-scale south-to-north post-LGM expansions are few and mostly occurred in the northern regions. Additionally, several unique geographic features, including the three-step terrain of China and the northern arid belt, have significant impacts on many species histories. Overall, the impacts of Pleistocene glaciations, particularly the LGM, on species history vary drastically from nondetectable to significant. The impacts are the least for species from the southwestern region and are most dominant for species from the north. Geological events play a more significant role in shaping species history than Pleistocene climatic changes. Phylogeographic patterns among animals species are highly consistent with those of plants. Future phylogeographic endeavour in East Asia should be hypothesis-driven and seek processes that underlie common patterns. The wide use of genomic data allow accurate estimates of historical population processes and exploration of older history beyond the Pleistocene.
... Nevertheless, recent clarifications regarding the distribution of S. raddei fossils, based on past habitat suitability models , provided much needed estimates for molecular dating using fossil calibrations. The use of fossil evidences as a primary calibration, combined with dating estimates of earlier works (Stöck et al. 2006;Dong et al. 2012;Borzée et al. 2017;Othman et al. 2020) and large samples sizes across the totality of the species' range helps reveal the patterns of phylogeography and landscape connectivity through time. ...
... This hypothesis is coherent with the biogeography of closely related toad species, as the climatic changes and sea level fluctuations during ice-ages have driven the phylogeography of Bufo sp. distributed in northeastern Asia, including for instance B. gargarizans (Borzée et al. 2017), B. sachalinensis sachalinensis (Othman et al. 2022) and B. stejnegeri (Fong et al. 2020). Despite the minimal impact of glaciations on Chinese S. raddei in the southern (Dong et al. 2012), the presence of glacial-driven refugia in Asia needs to be confirmed through a population dynamic lens, focusing on the northwestern limit of the current range of S. raddei. ...
... We highlighted the Pliocene and Pleistocene as the periods of active radiations, resulting in the simultaneous expansion of crown clades in both southern-and northeastern-originated lineages (Table 1; Fig. 3). Similarly, the extensive uplift of the QTP, drastic climatic variations such as the Quaternary climatic oscillations (Othman et al. 2022), sea level fluctuations (Borzée et al. 2017), and the development of the East Asian monsoon system impacted the dry seasons (Yuan et al. 2015), which have led to speciation, population dispersals and fragmentation of many animal taxa in the region (Cheng et al. 2017;Ding et al. 2020). ...
Landscape structures drive biogeographic patterns and population connectivity of animals distributed across diverse biotopes. Here, we provide a fresh insight on the impact of five landscape types in East Asia on the phylogeography and acoustic variability of the widespread Mongolian Toad, Strauchbufo raddei. For the first time, we reconstructed the biogeography of S. raddei over the species’ entire range throughout East Asia (N = 293; assembled up to 2,613 bp of concatenated CR-COI-12S rRNA-16S rRNA) using fossil-based molecular dating and genetic connectivity assessments. In addition, we addressed past population dynamics in relation to landscape types, and geographic variations in release calls for the clades occurring in the steppes of northern Mongolia and the Amur River basin (N = 147). Our results recovered two separate ancestors of S. raddei in East Asia, supporting a basal split between the northeastern and southern lineages in the Middle Miocene, c. 9.48 – 13.77 Mya. Ancestral range estimates suggested a Late Miocene radiation within the northeastern lineage, likely due to aridity-induced vicariance and dispersal from the central Asian steppes, c. 7.89 (5.25 – 11.50) Mya. The southern lineage emerged subsequently from glacial refugia, c. 6.84 (3.48 – 2.63) Mya, expanding northward and crossing the Gobi Desert and current-day Mongolia, c. 2.60 (1.15 – 3.72) Mya. At the exception of the pre-Tibetan Plateau clade, our reconstruction of migration trajectories highlighted the presence of effective gene flow across other landscapes, notably among the central and northeastern Chinese clades in the habitats defined as steppe, river basin and canyon. Significant variation in release calls between the clades in northern Mongolia and the Amur River Basin reflected the isolation between the two clades, and supported the presence of a northern refugium and post-glacial expansion of the southern lineage into northwestern Mongolia. In contrast with prior studies, our finding indicates that release calls can reflect phylogeographic patterns.
