From threatened to widespread: a non-genuine conservation status update for a now widespread Asian Hylid - and the need for additional basic field research

From threatened to widespread: a non-genuine conservation status update for a now widespread Asian Hylid - and the need for additional basic field research | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article From threatened to widespread: a non-genuine conservation status update for a now widespread Asian Hylid - and the need for additional basic field research Amaël Borzée, Tae Eun Um, Vishal Kumar Prasad, Abhilasha Shrivastava, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7226956/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Context. Non-genuine conservation status updates threaten the trust in Red Listing, but they are necessary building blocks in science. In addition, they often highlight the critical need for basic field research. The Suweon treefrog, Dryophytes suweonensis , is a hylid species long thought to be range-restricted and assessed as Endangered by the IUCN Red List of Threatened Species. Aims. This study aimed at determining the species of individuals encountered during call surveys in 2024 in China, and determine the areas of suitable habitat for the species. Methods. We sequenced the Cyt b gene fragment to identify the species and compared the acoustic properties to identify the species. We also built ecological models with MaxEnt to predict suitable habitats for the species and guide future surveys. Key results. We confirmed the identity of the population found more than 985 km north of the northernmost known locality of D. suweonensis as the same species. The call properties also matched with those of D. suweonensis . The ecological niche models identified a very large area with suitable habitat spanning across the northern Chinese plains. Conclusions. The species is not endemic to the Korean Peninsula, occurring in a significantly broader range, and thus, the population size is higher than previously assumed. Implications. The species is unlikely to be threatened, and this huge range extension highlights the need for additional field surveys for all species in the area. Long term, the use of conservation prioritisation tools will highlight conservation needs in the region. Animal Science treefrogs conservation status northeast Asia species identification Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction All animal species can be assessed for their extinction risk, an exercise conducted by several agencies and globally led by the IUCN Red List of Threatened Species (IUCN 2025 ). These assessments result in species assessed as non-threatened, threatened (Vulnerable, Endangered or Critically Endangered) or Data Deficient. Assessments are considered out-of-date ten years after their release, and the aim is, therefore, to re-assess all species within this time frame (IUCN Standards and Petitions Committee 2024 ). As an example, this was done in 2004 for all amphibians through the first Global Amphibian Assessment (GAA; Stuart et al. 2004 ), although the second iteration of the global assessment could not be finished before 2023 (Luedtke et al. 2023 ). The assignment of species to specific extinction risk categories was updated between both global assessments, many species having to be uplisted towards higher threat levels and a few having to be downlisted, partially due to non-genuine changes (Luedtke, et al. 2023 ). Non-genuine changes consist of updates when the threat levels to a species are downlisted because of the discovery of new or larger populations. This was, for instance, the case of numerous species between the two GAAs because of the discovery of new populations (Luedtke, et al. 2023 ). Non-genuine changes are most generally the result of “basic” research, defined as the first stepping-stones of knowledge on species, but not necessarily basic in terms of efforts as they often require intense fieldwork (e.g. the Kalimantan Jungle Toad; Bickford et al. 2008 ). In comparison, “advanced” research can be conducted pipet in hand in the comfort of a laboratory (e.g. evolution of transcription factors in amphibians; Carron et al. 2024 ). In continental northeast Asia, many species were newly assessed during the second global amphibian assessment, highlighting treats focused on the western coastal areas of the Yellow Sea, along the Korean Peninsula and reaching marginally into Liaoning in the People’s Republic of China (hereafter China; Luedtke, et al. 2023 ). The assessments originated from a general increase in the data availability in the region (e.g. amphibianchina.org), and as a result, distribution data for most species became available and refined through the IUCN Red List portal ( https://www.iucnredlist.org/ ) during that period (e.g. Rana huanrenensis assessed in 2004 and in 2018, Amphibian Specialist Group 2019; Dryophytes japonicus assessed in 2004 and in 2017; Kuzmin et al. 2017a ). Among the species updated, Pelophylax nigromaculatus was downlisted due to a non-genuine change, as the level of threat was determined to have been overestimated by earlier assessments (Amphibian Specialist Group 2023 ). However, data availability is rarely universal. For instance, the downlisting of P. nigromaculatus is now questionable as two species were assessed under one name due to the absence of resources for taxonomic research as the nominative P. nigromaculatus includes two species with species-level divergence (Borzée 2024 ; Dufresnes et al. 2024 ). This highlights that when re-assessed, the conservation status of the species might change again, this time as well because of non-genuine changes. This back-and-forth is a clear indicator of a need for resources for additional field research and to fix the taxonomy, a critical foundation for all fields of biology (Godfray and Knapp 2004 ), including conservation (Thomson et al. 2018 ). The Suweon treefrog was described in 1980 from Suwon in the Republic of Korea (Kuramoto 1980 ) and it was at first expected to be restricted to the area generally centred on the city of Suwon (Roh et al. 2014 ). Further developments showed the species to be ranging further south (Borzée et al. 2017a ), where it borders the range of its sister species, D. flaviventris (Borzée et al. 2020a ). These two species, along with D. immaculatus , are part of the D. immaculatus species complex. The Suweon treefrog was then found to range north into the Democratic People’s Republic of Korea (hereafter DPR Korea), as far north as the Taeryong River (Borzée et al. 2021 ). The species was not found in the area in Liaoning, where the landscape seemed adequate (Borzée et al. 2024 ) and was therefore not thought to be present in China. Due to the very high level of habitat degradation in R Korea (Borzée, et al. 2017a ) and the perceived decline in population size (Borzée et al. 2018a ; Borzée 2020 ), the species was listed as Endangered (IUCN SSC Amphibian Specialist Group 2017 ). However, the surveys conducted here have shown the species to be present about 1000 km north of the northernmost site in DPR Korea, over an area at least twice the latitudinal width of the known indigenous range of the species (following the IUCN definition of the term for green status of species; Grace 2023 ). Materials and methods Field surveys In view of the lack of clear and supported geographic boundaries for many species in northeast China (Borzée 2024 ), we conducted general surveys consisting of a general rectangle across northeast China, with an angle in the west around Hingan in Inner Mongolia, an angle in the north around Da Hinggan in Heilongjiang, an angle in the southeast of the area around Hunchun in Jilin (with a bulge to include the area north of the Kankha Lake), and the last angle around Chaoyang in Liaoning (Fig. 1 ). The surveys were conducted in the form of rapid species assessments in June 2024 and June 2025 during the peak breeding season of the species, using aural surveys following the protocols developed in Borzée and Jang ( 2015 ) and completed in Borzée et al. ( 2017b ) and Borzée, et al. ( 2017a ). Broadly, sites with the highest probability of presence are visually assessed from satellite maps, and a route is plotted across the area with high apparent suitability between sunset and 3:00 am as the calling activity declines around that time of the night (or earlier, matching with sunrise at northern latitudes; Borzée et al. 2020b ). This protocol makes the detection of the focal species most likely, here an early summer breeder. We spent 5 minutes listening to the call activity of the species at each site, generally resulting in a 95% probability of detection of Asian Dryophytes , focusing on the presence of Kaloula borealis , Strauchbufo raddei , Pelophylax nigromaculatus , Pelophylax plancyi , Dryophytes japonicus , and here unexpectedly Dryophytes suweonensis . Molecular analyses Sampling this species does not require permits in China, and it was collected under the AICUC number 2024013 provided by the Ethical Committee of Nanjing Forestry University. Upon encountering the population of D. suweonensis, we orally swabbed five individuals in the vicinity of Qiqihar (47.59238°N; 124.27774°E) and stored them with a bead of desiccant for 10 days until storage at -20°C. We extracted DNA using the blood and tissue DNA extraction kit from Qiagen, using the manufacturer's protocols. Sequencing was performed for both the forward and reverse strands of a Cyt-b gene fragment (900 bp; primers: L0-ATGGCCCCTGTTTTACGCAA and1046- TAAATGGGTCTTCTACTG G; Dufresnes et al. 2016 ), by Tsingke Co., Ltd., Wuhan, People’s Republic of China. For species identification, we first trimmed and complemented the raw sequences for all five samples to 819-bp long fragments using Geneious Prime v. 2023.2.1 (Kearse et al. 2012 ). All sequences generated in this study were deposited to Genbank under the accession number PV231326 to PV231329 (Table 1 ). Table 1 Information for ingroup and outgroup taxa used to reconstruct the phylogeny and species assignments for the five Dryophytes suweonensis individuals collected in the vicinity of Qiqihar, China. We used a mtDNA Cyt b gene fragment of the analysis, for 77 Dryophytes individual vouchers. Species Isolate Localities Accession number (Cyt b ) Source Dryophytes suweonensis 1B China: Qiqihar PV231327 This study Dryophytes suweonensis 2B China: Qiqihar PV231326 This study Dryophytes suweonensis 3B China: Qiqihar PV231328 This study Dryophytes suweonensis 4B China: Qiqihar PV231325 This study Dryophytes suweonensis 5B China: Qiqihar PV231329 This study Dryophytes flaviventris mms8551 R Korea: Iksan, North Jeolla MK089227 This study Dryophytes flaviventris mms8552 R Korea: Iksan, North Jeolla MK089228 This study Dryophytes flaviventris mms8553 R Korea: Iksan, North Jeolla MK089229 This study Dryophytes flaviventris mms8548 R Korea: Iksan, North Jeolla MK089224 This study Dryophytes flaviventris mms8545 R Korea: Iksan, North Jeolla MK089221 This study Dryophytes versicolor MT759671 Bogart et al., 2020 Dryophytes versicolor MT759669 Bogart et al. 2020 Dryophytes versicolor MT759665 Bogart et al. 2020 Dryophytes immaculatus mms8671 China: Hefei, Anhui MK089239 Min (2008), direct submission Dryophytes immaculatus mms8670 China: Hefei, Anhui MK089238 Min (2008), direct submission Dryophytes immaculatus mms8669 China: Hefei, Anhui MK089237 Min (2008), direct submission Dryophytes immaculatus mms8668 China: Hefei, Anhui MK089236 Min (2008), direct submission Dryophytes immaculatus mms8667 China: Hefei, Anhui MK089235 Min (2008), direct submission Dryophytes immaculatus mms8666 China: Hefei, Anhui MK089234 Min (2008), direct submission Dryophytes immaculatus mms8665 China: Hefei, Anhui MK089233 Min (2008), direct submission Dryophytes immaculatus mms8664 China: Hefei, Anhui MK089232 Min (2008), direct submission Dryophytes suweonensis mms7859 R Korea: Siheung, Gyeonggi MK089210 Min (2008), direct submission Dryophytes suweonensis mms7857 R Korea: Siheung, Gyeonggi MK089208 Min (2008), direct submission Dryophytes suweonensis mms7852 R Korea: Asan, South Chungcheong MK089205 Min (2008), direct submission Dryophytes suweonensis mms7844 R Korea: Asan, South Chungcheong MK089203 Min (2008), direct submission Dryophytes suweonensis mms6871 R Korea: Cheongju, North Chungcheong MK089190 Min (2008), direct submission Dryophytes suweonensis mms6864 R Korea: Cheongju, North Chungcheong MK089184 Min (2008), direct submission Dryophytes suweonensis mms4470 R Korea: Paju, Gyeonggi MK089111 Min (2008), direct submission Dryophytes suweonensis mms4469 R Korea: Paju, Gyeonggi MK089110 Min (2008), direct submission Dryophytes suweonensis mms4462 R Korea: Pyeongtaek, Gyeonggi MK089107 Min (2008), direct submission Dryophytes suweonensis mms3715 DPR Korea: Moonduk, South Pyeongan MK089093 Min (2008), direct submission Dryophytes suweonensis mms0474 R Korea: Pyeongtaek, Gyeonggi MK089092 Min (2008), direct submission Dryophytes suweonensis mms0467 R Korea: Pyeongtaek, Gyeonggi MK089090 Min (2008), direct submission Dryophytes suweonensis mms0357 R Korea: Pyeongtaek, Gyeonggi MK089089 Min (2008), direct submission Dryophytes suweonensis mms0356 R Korea: Pyeongtaek, Gyeonggi MK089088 Min (2008), direct submission Dryophytes suweonensis mms0352 R Korea: Pyeongtaek, Gyeonggi MK089087 Min (2008), direct submission Dryophytes suweonensis mms0351 R Korea: Pyeongtaek, Gyeonggi MK089086 Min (2008), direct submission Dryophytes suweonensis mms0350 R Korea: Pyeongtaek, Gyeonggi MK089085 Min (2008), direct submission Dryophytes suweonensis KUZ:nn2H_Kv04 R Korea LC569676 Kuraishi et al., 2021 Dryophytes suweonensis 14SIH002_Hsuw R Korea KY055770 Dufresnes et al. 2011 Dryophytes suweonensis 14SIH001_Hsuw R Korea KY055769 Dufresnes et al. 2012 Dryophytes suweonensis 13GEN394_Hsuw R Korea KY055768 Dufresnes et al. 2013 Dryophytes suweonensis 13GEN718 R Korea KY055734 Dufresnes et al. 2014 Dryophytes japonicus 13GEN640 R Korea KY055733 Dufresnes et al. 2015 Dryophytes suweonensis 13GEN639 R Korea KY055732 Dufresnes et al. 2016 Dryophytes suweonensis 13GEN638 R Korea KY055731 Dufresnes et al. 2016 Dryophytes suweonensis 13GEN637 R Korea KY055730 Dufresnes et al. 2016 Dryophytes japonicus 13GEN636 R Korea KY055729 Dufresnes et al. 2016 Dryophytes japonicus 13GEN635 R Korea KY055728 Dufresnes et al. 2016 Dryophytes japonicus 13GEN135 R Korea KY055720 Dufresnes et al. 2016 Dryophytes japonicus Chi201204.47 R Korea KY055717 Dufresnes et al. 2016 Dryophytes japonicus Chi201204.46 R Korea KY055716 Dufresnes et al. 2016 Dryophytes japonicus Chi201204.45 R Korea KY055715 Dufresnes et al. 2016 Dryophytes japonicus Chi201204.42 R Korea KY055712 Dufresnes et al. 2016 Dryophytes japonicus HLi100277 R Korea KY055705 Dufresnes et al. 2016 Dryophytes japonicus HLi100141 R Korea KY055704 Dufresnes et al. 2016 Dryophytes japonicus HLi675 R Korea KY055701 Dufresnes et al. 2016 Dryophytes japonicus HLi599 R Korea KY055694 Dufresnes et al. 2016 Dryophytes japonicus HLi820 R Korea KY055683 Dufresnes et al. 2016 Dryophytes japonicus HLi819 R Korea KY055682 Dufresnes et al. 2016 Dryophytes japonicus HLi814 R Korea KY055677 Dufresnes et al. 2016 Dryophytes japonicus HLi813 R Korea KY055676 Dufresnes et al. 2016 Dryophytes japonicus HLi812 R Korea KY055675 Dufresnes et al. 2016 Dryophytes immaculatus Himm606002 China KY055673 Dufresnes et al. 2016 Dryophytes immaculatus Himm606003 China KY055672 Dufresnes et al. 2016 Dryophytes suweonensis Haplotype 10 R Korea: Pyeongtaek, Gyeonggi KF564864 Chun et al. (2013), direct submission Dryophytes suweonensis Haplotype 9 R Korea: Paju, Gyeonggi KF564863 Chun et al. (2013), direct submission Dryophytes suweonensis Haplotype 8 R Korea: Paju, Gyeonggi KF564862 Chun et al. (2013), direct submission Dryophytes suweonensis Haplotype 7 R Korea: Ganghwa, Incheon KF564861 Chun et al. (2013), direct submission Dryophytes suweonensis Haplotype 5 R Korea: Ganghwa, Incheon KF564859 Chun et al. (2013), direct submission Dryophytes suweonensis Haplotype 1 R Korea: Ganghwa, Incheon KF564855 Chun et al. (2013), direct submission Dryophytes suweonensis Haplotype 10 R Korea: Goseong, Gangwon-do KF564854 Chun et al. (2013), direct submission Dryophytes japonicus Haplotype 9 R Korea: Goseong, Gangwon-do KF564853 Chun et al. (2013), direct submission Dryophytes japonicus Haplotype 8 R Korea: Goseong, Gangwon KF564852 Chun et al. (2013), direct submission Dryophytes japonicus Haplotype 5 R Korea: Paju, Gyeonggi KF564849 Chun et al. (2013), direct submission Dryophytes japonicus Haplotype 4 R Korea: Eumseong, Chungcheong KF564848 Chun et al. (2013), direct submission Dryophytes japonicus Haplotype 3 R Korea: Gangneung, Gangwon KF564847 Chun et al. (2013), direct submission Table 2 Bayesian posterior distribution of migration rates among four clades of the Dryophytes immaculatus species complex, estimated using a partial Cyt b dataset (819 bp). The populations include D. immaculatus (Di) from eastern China, D. flaviventris (Df) isolated in Iksan, R Korea, and two clades of D. suweonensis : one from the Korean Peninsula (DsK) and a newly identified population from Qiqihar, China (DsQ). These estimates provide insights into gene flow and population connectivity with Θ indicating the effective population size (Ne) scaled by the mutation rate (µ), and M indicating the mutation-scaled migration rate. Parameter 0.03% 0.25% Mode 0.75% 0.98% Median Mean Θ Di 0.0028 0.0073 0.0136 0.0263 0.0756 0.0240 0.0305 Θ Df 0.0000 0.0007 0.0023 0.0037 0.0154 0.0032 0.0045 Θ DsK 0.0049 0.0103 0.0157 0.0263 0.0627 0.0236 0.0288 Θ DsQ 0.0000 0.0002 0.0019 0.0033 0.0182 0.0032 0.0048 M Df−>Di 6.70 6.70 22.30 205.30 205.30 275.70 335.00 M DsK−>Di 0.00 37.30 178.30 245.30 839.30 326.30 367.20 M DsQ−>Di 0.00 0.00 23.70 498.70 853.30 315.70 371.90 M Di−>Df 0.00 0.00 23.00 164.00 834.00 163.70 245.30 M DsK−>Df 0.00 4.00 23.00 260.70 592.00 271.00 329.70 M DsQ−>Df 0.00 1.30 103.00 501.30 756.70 384.30 421.10 M Di−>DsK 0.00 0.00 25.00 170.00 632.70 170.30 228.90 M Df−>DsK 214.70 639.30 884.30 988.00 999.30 620.30 582.70 M DsQ−>DsK 97.30 620.00 975.70 996.00 999.30 607.00 577.20 M Di−>DsQ 0.00 0.00 24.30 193.30 888.70 195.70 265.30 M Df−>DsQ 0.00 0.00 24.30 228.70 316.70 446.30 458.90 M DsK−>DsQ 0.00 0.00 23.70 276.00 699.30 289.70 351.60 Independently, we aligned the Cytb sequences of all five individuals with 72 homologous sequences available in Genbank ( https://www.ncbi.nlm.nih.gov/genbank/ ). We restricted the final alignment to closely related species: Dryophytes japonicus, Dryophytes suweonensis and D. immaculatus (n taxa = 77), as this analysis was conducted to confirm the species identification and not to conduct a population genetic study. These three species are closely related congeneric species distributed on the Korean Peninsula and China (see Table 1 for the details of the samples). We also included three samples of Dryophytes versicolor from the Nearctic as outgroups (Table 1 ). We first identified the best-fit partitioning scheme and substitution models for the Cytb sequences alignment. As the alignment included a partial intron (from base pair 1 to 12) and partial exon (from base pair 13 to 819), we included both the non-coding region and the three-codon strategies in our gene blocks parameter using Partition Finder v. 2.1.1 (Lanfear et al. 2012 ). We obtained HKY + I + G as the best substitution model for the four subsets of partition. We then integrated the information of the best substitution model in our concatenated alignment dataset (.Nexus) for the reconstruction of the Bayesian phylogenetic tree in Mr. Bayes v. 3.2.7 (Ronquist et al. 2012 ). We ran the analysis using the Monte Carlo Markov Chain (MCMC) algorithm, with four chains, 10,000,000 iterations, and a burn-in of 1,000. To ensure the convergence and robustness of the phylogenetic analysis, we optimised the value of split frequencies (< 0.05), the parameter space and the pattern of randomness of MrBayes at the end of its run. Ecological niche models Finally, we used ecological niche models to determine whether we are still missing some major populations. The data used for this analysis originated from the literature (Borzée, et al. 2017a ; Son et al. 2020 ; Borzée, et al. 2021 ) along with the new datapoint collected here (Supplementary File 1). To reduce the sampling bias, we thinned occurrence points within 4 km using the spThin package (Aiello-Lammens et al. 2015 ) and using the bias files created by the MASS package (Venables and Ripley 2002 ) in R version 4.2.1. We decided to rely on the 19 bioclimatic variables (Worldclim 2.1; Fick and Hijmans 2017 ), elevation (2.5 arc-minute resolution) and Landsat normalised difference vegetation index (NDVI, USGS) after the removal of highly correlated variables (Pearson’s r > 0.8; Bradie and Leung 2017 ). We used six variables for the final models: annual mean temperature (bio 1), temperature annual range (bio 7), annual precipitation (bio 12), precipitation seasonality (coefficient of variation) (bio 15), and elevation. We ran MaxEnt v. 3.4.4 (Elith et al. 2011 ; Phillips et al. 2017 ) with the default setting (Phillips and Dudík 2008 ) and used the cross-validate run type option with ten replicates (10k-fold). For the model evaluation, we calculated the area under the curve (AUC; Somodi et al. 2017 ) and true skill statistic (TSS; Allouche et al. 2006 ). To visualise the potential suitable habitat and to calculate the threshold area, we used ArcGIS pro v. 3.3.0 (ESRI, Redlands, CA, USA). Call description We recorded the advertisement calls of three individuals in the proximity of Qiqihar, Heilongjiang, China (47.592389°N; 124.277740°E) on 10 June 2024 at 20:25 (air temperature 24°C) in real-time using the 16-bit resolution and a sampling rate of 44.1 kHz. We used a linear PCM recorder (Tascam DR-40; California, USA) linked to a unidirectional microphone (unidirectional electret condenser microphone HT-81, HTDZ; Xi’an, China). The microphone was positioned straight, approximately 0.5 m from the mouth of the calling individual. We manually regulated the gain settings to avoid saturation and ensure the maximum amplitude peaks. The calls were deposited in the FonoZoo sound repository under accession numbers (FZ-SOUND-CODE 114785 − 14804). Prior to call data extraction, we removed background noise by filtering out the calls at 1 kHz and selected the cleanest section of the calls for each calling individual in Raven Pro 1.5 software (Bioacoustics Research Program 2014) for analysis. We kept the Hann window size at 256 sample windows, 128-sample hop size with 50% frame overlap and a 172 Hz frequency grid spacing. We analysed both temporal and spectral properties of the calls. We then extracted the data using Raven Pro v. 1.5 (Bioacoustics Research Program 2014). In total, we analysed 60 notes from three calling male individuals (20 notes from each individual) for the acoustic analysis, following the definitions of call properties from Park et al. ( 2013 ) and Borzée, et al. ( 2020a ). Further, we processed the calls in R (R Core Team 2024 ) using Seewave (Sueur et al. 2018 ) and tuneR (Ligges et al. 2018 ) packages to produce high-quality spectrograms, oscillograms and power spectrums. Conservation assessment Threat assessments over national boundaries are generally the most robust when following the IUCN Red List categories and criteria (Maes et al. 2015 ; IUCN Standards and Petitions Committee 2024 ), and they are suitable indicators to inform on conservation needs (Lamoreux et al. 2003 ; Rodrigues et al. 2006 ). Species are evaluated against a set of criteria, and when threatened, they can fall into either of three categories: Vulnerable, Endangered or Critically Endangered (IUCN Standards and Petitions Committee 2024 ). Assessments are conducted against quantitative thresholds for five criteria: population size reduction, geographic range size, small population size and decline, very small population and/or restricted distribution, and quantitative analysis of extinction risk. Complete datasets for the assessments are difficult to generate and can follow alternative protocols (Mace 1994 ; Hermoso et al. 2015 ). Here, the only data available is the one about the geographical range size, and we follow the IUCN Red List categories and criteria to suggest a threat assessment for the species (IUCN Standards and Petitions Committee 2024 ). Results Field surveys The northernmost new location of Dryophytes suweonensis is further north than Qiqihar, about 985 km north of the last northernmost known location, which is about 300 km north of the southern boundary of the range of the species, now bringing the species close to 1000 km of latitudinal extent (Fig. 1). At Harbin latitude, the longitudinal extant between the two extremes populations is greater than 880 km, in comparison with a past greater maximal longitudinal extend of 80 km. The westernmost population is now in Panjin in Liaoning, about 300 km further west than previously known. As a result, the distribution of D. suweonensis in R Korea is now comparatively smaller in terms of geographic coverage and potentially in terms of the number of individuals. Interestingly, all the individuals were encountered in rice paddies and showed the two dorsal stripes characteristic of D. suweonensis emerging from hibernation, only visible for a day or two after emergence in R Korea (Borzée 2024; Fig. 2). Molecular analyses The phylogenetic tree (Fig. 3) highlighted the clustering of the samples collected from Qiqihar with the D. suweonensis clade (Bayesian posterior probability [BPP] of 100%). Despite being distributed about 1000 km away, all five individuals specifically clustered within the clade of D. suweonensis and shared a paraphyletic relationship with D. immaculatus , endemic to China (Fig. 3). Ecological niche models The model was supported by AUC values of 0.958 ± 0.020 and a TSS value of 0.612 ± 0.108, and thus, both values indicated a perfect fit for the model (Phillips et al. 2006). The variable with the highest contribution to the distribution of D. suweonensis was bio 7 (temperature annual range), with 63.7% contribution and 52.9% permutation importance (Table 4). A very large area with suitable habitat was found through the models in the plains between Changchun, Harbin and Qiqihar in Jilin and Heilongjiang provinces, and on the Songliao plains in Liaoning (Fig. 1). The habitat suitability results were thresholded in the map using the maximum training sensitivity plus specificity (MTSS) value of 0.14 to classify MaxEnt output raster files into suitable and unsuitable habitats (Fig. 1; Liu et al. 2013). The calculated suitable range of the species within the area highlighted on Fig. 1 was 713 248 km 2 . Table 3 Call properties of Dryophytes suweonensis from Qiqihar, China, collected in June 2024. The data is based on 60 notes extracted from the calls of three individuals ( n = 20 per individual). Call properties Mean SD Min Max Note duration (s) 0.12 0.02 0.10 0.22 Note dominant frequency (Hz) 2962.97 398.41 0.13 3100.80 90% frequency bandwidth of note (Hz) 1521.68 696.44 516.8 2067.20 Duration of connected pulses (s) 0.07 0.02 0.04 0.18 Peak dominant frequency of connected pulses (Hz) 3031.88 85.12 2928.50 3100.80 90% frequency bandwidth of connected pulses (Hz) 1484.35 745.24 344.50 2067.20 Number of independent pulses 6.53 2.47 3 15 Table 4 Percentage of contribution and permutation importance for each of the five variables retained to build habitat suitability model with MaxEnt for Dryophytes suweonensis . Variables Contribution percentage Permutation importance Temperature annual range (bio 7) 63.7 52.9 Annual precipitation (bio 12) 24.8 32.3 Annual mean temperature (bio 1) 6.2 0.0 Precipitation seasonality (coefficient of variation; bio 15) 4.6 1.5 Elevation 0.7 13.4 Call properties In total, we analysed acoustic data for 60 notes from three D. suweonensis individuals from Qiqihar. The notes were produced at continuous intervals, and their temporal structure consisted of both independent and connected pulses. In the spectrogram, the sections of notes with independent pulses displayed double frequency bands for each pulse and four strong harmonics for the condensed pulsed parts (Fig. 4). The mean note duration was 0.12 ± 0.02 seconds, with a mean dominant frequency of 2962.97 ± 398.41 Hz. In comparison, the note duration for D. suweonensis in R Korea was 0.12 ± 0.02, while it was 0.11 for D. immaculatus and 0.15 for D. flaviventris (Borzée, et al. 2020a). The average number of independent pulses per note was 6.53 ± 2.47, while it is 5.34 for D. suweonensis in R Korea. The mean duration of connected pulses was 0.07 ± 0.02 seconds, and the mean peak dominant frequency of connected pulses was 3031.88 ± 85.12 Hz (Table 3). Conservation assessment The geographic range and the extent of occurrence of the species is well over the 20,000 km 2 limit to be listed as threatened (noting that the 713 248 km 2 are not the range of the species, which will only be a fraction of this value, but still estimated to be well above the threshold). The range is severely fragmented in the south only, and the continued decline cannot be inferred for the northern populations. In conclusion, globally, the species is not listed as threatened based on B1 and B2 criteria (IUCN Standards and Petitions Committee 2024). The earlier assessment as Endangered (IUCN SSC Amphibian Specialist Group 2017) can, however, be used as a national Red List assessment for R Korea. Discussion The discovery of new populations thousands of kilometres away from their known range is not unheard of, but it is unusual, and generally happens with cryptic species (e.g. Hydromantes strinatii , Lucente et al. 2016 ) or species with great dispersal capacities (e.g. Carduelis cucullate , Robbins et al. 2003 ). Here, a population of the Suweon treefrog ( Dryophytes suweonensis ) found about a thousand kilometres north of its type locality is entirely unexpected and raises questions about the lack of field research in the area and the threat assessment done on the species. However, there are decades old guidelines to remove infestation by the sister species, now known to be a misidentification, from silkworm farms within the range of D. suweonensis , also noting their resistance to common pesticide, and providing more information about their life cycle than other updated literature (Liu et al. 1989 ). Here, we highlight that the species is not endemic to the Korean Peninsula, the population is not as small as previously expected, although there are no population size estimates available, and more basic ecological research on distribution is highly needed. We also call for more research on the distribution of species in northeast China in view of the recent discovery of species thought to be restricted to the Korean Peninsula, including Pelophylax chosenicus (Borzée, et al. 2024 ), Rana coreana (Borzée et al. 2025 ) and the presence of one most-unlikely described Onychodactylus species. A potential shortcoming to the claim of a range extension would be the discovery of a new species, which is not the case here in view of the clustering within the D. suweonensis clade in the phylogenetic tree (Fig. 3). However, despite the species-specific assignment, cytochrome b alone is not sufficient to conduct phylogenetic analyses, a single gene fragment is not significant in terms of taxonomy, and the absence of introgression patterns and minimal incomplete lineage sorting is likely influenced by this limitation. For future studies, we recommended sampling along a latitudinal cline and the use of a genome-wide approach to achieve higher resolution and better insights into potential introgression and lineage sorting patterns of D. suweonensis . In addition, the absence of data from DPR Korea prevents concluding whether there are additional gaps in the distribution of D. suweonensis , and additional sampling would answer his question. Here, to follow the precautionary principle (Deville and Harding 1997 ) in environmental protection and conservation (Glasgow et al. 2020 , Kemp et al. 2024 ), the population in Qiqihar is assigned to D. suweonensis , but further genomic analyses will help better understand the relationship within the species. In addition, the call properties of the frogs from Qiqihar matched with that of D. suweonensis in R Korea (Borzée, et al. 2020a ), with some variations, likely linked to the inability to extract data in a uniform manner across datasets. Further analysis within the species, including populations from DPR Korea, could highlight the impact of repeated marine transgression on the evolution of traits in amphibians. The distribution of the suitable habitat is also along the same landscape as the one where D. suweonensis is present further south, with some gap in hilly areas. It is interesting to note that the area of suitable habitat in Jiangsu, China, does not match with the area where D. immaculatus is present (Borzée 2024 ). This abundance of species present on both the Korean Peninsula and in northeast China is, however, not unexpected as the area is a conservation-need hotspot for amphibians (Button and Borzée 2021 ). It is also a pattern that can be expected for numerous other species, and especially mammals, as most species found on the Korean Peninsula are also found further north (Jo et al. 2018 ). In addition, these broader ranges reinforce the conservation potential for these species (Borzée et al. 2019b ) as all extinct mammal species and their designated protected areas are also found in China (e.g. tigers, Lee and Miller-Rushing 2014 ). It is, therefore, critical to properly survey and reassess all species present on the Korean Peninsula and ranging into northeast China for their IUCN Red List status. In addition, the large number of threatened species in the area could result in the triggering of sites listed as Key Biodiversity Areas or under the Alliance for Zero Extinction (Mair et al. 2021 ; Re:wild et al. 2023 ). As a result, the extinction risk for species would become linked to conservation policies, decision-making, and funding availability, a critical need for amphibians (Guénard et al. 2025 ). The distribution of D. suweonensis is unlikely to go further north and west as surveys in Mongolia (Kuzmin et al. 2017b ; Borzée et al. 2019a ) and Russia (Othman et al. 2022 ) in the habitat that would be adequate for the species did not detect its presence. However, low plains in boundary areas in Russia need additional specific surveys. In addition, the species was only found in rice paddies, similar to the known sites in R Korea (Borzée and Jang 2015 ), and there is no extensive rice agriculture at these latitudes in nearby countries. It is, however, important to highlight that there was a bias in the selection of sites for the surveys as D. suweonensis was not one of the target species, and determining the presence of the target species is more easily done in riparian areas, where wetlands have been replaced by rice paddies. However, D. suweonensis was detected in rice paddies that were contiguous with reed beds, similar to the habitat in DPR Korea (Borzée, et al. 2021 ), and further surveys in both China and DPR Korea to identify natural sites are important for conservation plans in R Korea (Park et al. 2014 ). While these findings clearly show that the past threat assessment as Endangered for D. suweonensis is now outdated, it does not mean that these are wrong. At the time of the assessment, only the populations in R Korea were known, and it is therefore not a global assessment anymore, but a national assessment, in need of an update as the population has been declining since it was last done. To be able to properly assess the species following the IUCN Red List categories and criteria (IUCN Standards and Petitions Committee 2024 ), additional surveys and research on the species will be needed as the polygon that would be currently used to define the extent of occurrence would miss some of the populations (Fig. 1 ). Declarations Acknowledgements This species does not require special permits, and it was collected under the AICUC number 2024013 provided by the Ethical Committee of Nanjing Forestry University. We are grateful to Wang Yingwei and Wang Yatao for reaching out and sharing information about an additional locality for Dryophytes suweonensis in Liaoning. Declaration of Funding. This project was funded by the Foreign Youth Talent Program of the Ministry of Science and Technology of the People’s Republic of China (QN2023014004L), the Research Fund for International Scientists (RFIS) from the National Natural Science Foundation of China (NSFC; W2432021) and a grant from Forest City Studios to AB, and by the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR201803101) to MSM. Data Accessibility The data that support this study are available in the article and accompanying online supplementary material. Conflict of Interest Statement The authors declare no conflicts of interest Contribution: Data collection: AB, TEU, AS, MSM, SNO; Data analysis: TEU, VKP, AS, SNO, writing first draft: AB; writing revisions: TEU, VKP, AS, MSM, SNO References Aiello‐Lammens ME, Boria RA, Radosavljevic A, Vilela B, Anderson RP. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7226956","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":491516273,"identity":"2f4c2423-ca1a-4a0a-9128-0b6ab1b61dfc","order_by":0,"name":"Amaël Borzée","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5ElEQVRIiWNgGAWjYLCChw1givEBkODhI0pLIkQLswFICxspWtgkwCQh1fzTDj98kLjDJl++vflY5dccOxk2BuaHj27g0SJxO83YIPFMmmVjz7G027LbkoEOYzM2zsFnze0EM4nEtsMGzBI5ZrcltzEDtfCwSePTIn87/fsPkBY2+fffiiW31RPWYnA7x4wBpIVHgoeN8eO2w4S1GN7OKZYA+sVAgifNWJpx23EeNmYCfpG7nb7xw8cdNgby7Ycffvy5rdqen7354WO83kcGzDxgkljlIMD4gxTVo2AUjIJRMGIAAC9ZRXrl++PAAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0003-1093-677X","institution":"Nanjing Forestry university","correspondingAuthor":true,"prefix":"","firstName":"Amaël","middleName":"","lastName":"Borzée","suffix":""},{"id":491516418,"identity":"23de405e-a393-4f5d-8997-61bb031ac7e4","order_by":1,"name":"Tae Eun Um","email":"","orcid":"","institution":"Nanjing Forestry university","correspondingAuthor":false,"prefix":"","firstName":"Tae","middleName":"Eun","lastName":"Um","suffix":""},{"id":491516419,"identity":"d9d1d5be-c776-4f65-b141-b2578f3dc0c0","order_by":2,"name":"Vishal Kumar Prasad","email":"","orcid":"","institution":"Nanjing Forestry university","correspondingAuthor":false,"prefix":"","firstName":"Vishal","middleName":"Kumar","lastName":"Prasad","suffix":""},{"id":491516420,"identity":"f0adaaec-753a-4cae-8efd-d8338ae08eb7","order_by":3,"name":"Abhilasha Shrivastava","email":"","orcid":"","institution":"Nanjing Forestry university","correspondingAuthor":false,"prefix":"","firstName":"Abhilasha","middleName":"","lastName":"Shrivastava","suffix":""},{"id":491516421,"identity":"61ed971d-a692-4032-9558-e526e69a110f","order_by":4,"name":"Mi-Sook Min","email":"","orcid":"","institution":"Seoul National University","correspondingAuthor":false,"prefix":"","firstName":"Mi-Sook","middleName":"","lastName":"Min","suffix":""},{"id":491516422,"identity":"adc5a3d1-75f6-4583-9c79-ca46fcda5fca","order_by":5,"name":"Siti N. Othman","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA1UlEQVRIiWNgGAWjYDCCA0DM2AAk2EGEgQWRWg6CVPOAWAYSpGiRSABxidDCdyOB8fHHHdvkzSWfX93wo0CCgb+9OwGvFskbCcwGB8/cNtw5O6fsZg/QYRJnzm7Aq8XgRgKbxMG224wbbuek3eABajGQyCVOi/2Gm2fSbv4hRUvihhvsx24TZYvkmYfNBmfbbidvOJPDdlvGQIKHoF/4jicffFDZdtt2w/Hjz26++WMjx9/ei18Lg0BiA5TFYwAm8SsHAf4DMBb7A8KqR8EoGAWjYEQCAHwhUt+9gXlNAAAAAElFTkSuQmCC","orcid":"","institution":"Nanjing Forestry university","correspondingAuthor":true,"prefix":"","firstName":"Siti","middleName":"N.","lastName":"Othman","suffix":""}],"badges":[],"createdAt":"2025-07-27 14:57:21","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false},"doi":"10.21203/rs.3.rs-7226956/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7226956/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":87980203,"identity":"ef215a97-ebea-42a6-be4e-fe8f9bc40e6c","added_by":"auto","created_at":"2025-07-31 05:48:42","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":331023,"visible":true,"origin":"","legend":"\u003cp\u003eMap representative of the sites where the Suweon Treefrog, \u003cem\u003eDryophytes suweonensis\u003c/em\u003e, was encountered during surveys in July 2024 (yellow dots). The background colours show the results of the ecological niche models. The suitable habitat for \u003cem\u003eD. suweonensis\u003c/em\u003e (deep red) highlights the potential for additional populations in northern China.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-7226956/v1/4b54e663ff50211fe4f0121b.png"},{"id":87980206,"identity":"2007c439-f87b-4430-bfbe-08163aaa9ea5","added_by":"auto","created_at":"2025-07-31 05:48:42","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":566966,"visible":true,"origin":"","legend":"\u003cp\u003eRepresentative pictures of the Suweon treefrog, \u003cem\u003eDryophytes suweonensis\u003c/em\u003e, found in the proximity of Qiqihar, Heilongjiang, China (47.592389°N; 124.277740°E) on 10 June 2024 after 20:00. The spot aligned along the two dorsal stipes visible in the species only for a day or two are visible on frames A, B and C, although having almost entirely faded on B. The individual on frame C is the individual 24DsC001 of the genetic analyses. The individuals in the typical calling position for the species (E and F) were among the three individuals recorded.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-7226956/v1/9add053f1d8bdfb5c1658058.png"},{"id":87980205,"identity":"c1f8ff1d-8608-4998-bb71-d2383545ace7","added_by":"auto","created_at":"2025-07-31 05:48:42","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":196385,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree to confirm the identity of the \u003cem\u003eDryophytes suweonensis\u003c/em\u003e individuals sampled in Qiqihar, China. The phylogeny is reconstructed from 74 \u003cem\u003eDryophytes\u003c/em\u003e individuals (ingroup taxa), consisting of D\u003cem\u003e. japonicus\u003c/em\u003e, \u003cem\u003eD. suweonensis\u003c/em\u003e, and \u003cem\u003eD. immaculatus\u003c/em\u003e, and \u003cem\u003eD. versicolor\u003c/em\u003e as outgroup (n = 3 taxa). The number at each node represents the percentage of Bayesian Posterior probability (BPP) support for each clade. The five individuals sampled in Qiqihar are marked in salmon colour on the tree.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-7226956/v1/5f9c262fe3f4f526de70fc5f.png"},{"id":87980214,"identity":"51ec9558-3161-43f9-ae80-72aebe996fc7","added_by":"auto","created_at":"2025-07-31 05:48:43","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":306683,"visible":true,"origin":"","legend":"\u003cp\u003eOscillogram (A), spectrogram (B) and the power spectrum (C) of the advertisement calls of \u003cem\u003eDryophytes suweonensis \u003c/em\u003erecorded from Qiqihar, China. The graphs were created with the R package Seewave (Sueur et al. 2008).\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-7226956/v1/13ee6eddc9e2f18feeb88178.png"},{"id":87980967,"identity":"d42763e8-0120-4a2e-952b-3732fe83dda1","added_by":"auto","created_at":"2025-07-31 06:04:43","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2519627,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7226956/v1/d7ce761b-0936-4985-851e-484cd528cdcc.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003eFrom threatened to widespread: a non-genuine conservation status update for a now widespread Asian Hylid - and the need for additional basic field research\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAll animal species can be assessed for their extinction risk, an exercise conducted by several agencies and globally led by the IUCN Red List of Threatened Species (IUCN \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These assessments result in species assessed as non-threatened, threatened (Vulnerable, Endangered or Critically Endangered) or Data Deficient. Assessments are considered out-of-date ten years after their release, and the aim is, therefore, to re-assess all species within this time frame (IUCN Standards and Petitions Committee \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). As an example, this was done in 2004 for all amphibians through the first Global Amphibian Assessment (GAA; Stuart et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), although the second iteration of the global assessment could not be finished before 2023 (Luedtke et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The assignment of species to specific extinction risk categories was updated between both global assessments, many species having to be uplisted towards higher threat levels and a few having to be downlisted, partially due to non-genuine changes (Luedtke, et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eNon-genuine changes consist of updates when the threat levels to a species are downlisted because of the discovery of new or larger populations. This was, for instance, the case of numerous species between the two GAAs because of the discovery of new populations (Luedtke, et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Non-genuine changes are most generally the result of \u0026ldquo;basic\u0026rdquo; research, defined as the first stepping-stones of knowledge on species, but not necessarily basic in terms of efforts as they often require intense fieldwork (e.g. the Kalimantan Jungle Toad; Bickford et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). In comparison, \u0026ldquo;advanced\u0026rdquo; research can be conducted pipet in hand in the comfort of a laboratory (e.g. evolution of transcription factors in amphibians; Carron et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn continental northeast Asia, many species were newly assessed during the second global amphibian assessment, highlighting treats focused on the western coastal areas of the Yellow Sea, along the Korean Peninsula and reaching marginally into Liaoning in the People\u0026rsquo;s Republic of China (hereafter China; Luedtke, et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The assessments originated from a general increase in the data availability in the region (e.g. amphibianchina.org), and as a result, distribution data for most species became available and refined through the IUCN Red List portal (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.iucnredlist.org/\u003c/span\u003e\u003cspan address=\"https://www.iucnredlist.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) during that period (e.g. \u003cem\u003eRana huanrenensis\u003c/em\u003e assessed in 2004 and in 2018, Amphibian Specialist Group 2019; \u003cem\u003eDryophytes japonicus\u003c/em\u003e assessed in 2004 and in 2017; Kuzmin et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017a\u003c/span\u003e). Among the species updated, \u003cem\u003ePelophylax nigromaculatus\u003c/em\u003e was downlisted due to a non-genuine change, as the level of threat was determined to have been overestimated by earlier assessments (Amphibian Specialist Group \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eHowever, data availability is rarely universal. For instance, the downlisting of \u003cem\u003eP. nigromaculatus\u003c/em\u003e is now questionable as two species were assessed under one name due to the absence of resources for taxonomic research as the nominative \u003cem\u003eP. nigromaculatus\u003c/em\u003e includes two species with species-level divergence (Borz\u0026eacute;e \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Dufresnes et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This highlights that when re-assessed, the conservation status of the species might change again, this time as well because of non-genuine changes. This back-and-forth is a clear indicator of a need for resources for additional field research and to fix the taxonomy, a critical foundation for all fields of biology (Godfray and Knapp \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), including conservation (Thomson et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe Suweon treefrog was described in 1980 from Suwon in the Republic of Korea (Kuramoto \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1980\u003c/span\u003e) and it was at first expected to be restricted to the area generally centred on the city of Suwon (Roh et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Further developments showed the species to be ranging further south (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017a\u003c/span\u003e), where it borders the range of its sister species, \u003cem\u003eD. flaviventris\u003c/em\u003e (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e). These two species, along with \u003cem\u003eD. immaculatus\u003c/em\u003e, are part of the \u003cem\u003eD. immaculatus\u003c/em\u003e species complex. The Suweon treefrog was then found to range north into the Democratic People\u0026rsquo;s Republic of Korea (hereafter DPR Korea), as far north as the Taeryong River (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The species was not found in the area in Liaoning, where the landscape seemed adequate (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and was therefore not thought to be present in China. Due to the very high level of habitat degradation in R Korea (Borz\u0026eacute;e, et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017a\u003c/span\u003e) and the perceived decline in population size (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018a\u003c/span\u003e; Borz\u0026eacute;e \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), the species was listed as Endangered (IUCN SSC Amphibian Specialist Group \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, the surveys conducted here have shown the species to be present about 1000 km north of the northernmost site in DPR Korea, over an area at least twice the latitudinal width of the known indigenous range of the species (following the IUCN definition of the term for green status of species; Grace \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e\u003cem\u003eField surveys\u003c/em\u003e\u003c/p\u003e\u003cp\u003eIn view of the lack of clear and supported geographic boundaries for many species in northeast China (Borz\u0026eacute;e \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), we conducted general surveys consisting of a general rectangle across northeast China, with an angle in the west around Hingan in Inner Mongolia, an angle in the north around Da Hinggan in Heilongjiang, an angle in the southeast of the area around Hunchun in Jilin (with a bulge to include the area north of the Kankha Lake), and the last angle around Chaoyang in Liaoning (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The surveys were conducted in the form of rapid species assessments in June 2024 and June 2025 during the peak breeding season of the species, using aural surveys following the protocols developed in Borz\u0026eacute;e and Jang (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and completed in Borz\u0026eacute;e et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017b\u003c/span\u003e) and Borz\u0026eacute;e, et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017a\u003c/span\u003e). Broadly, sites with the highest probability of presence are visually assessed from satellite maps, and a route is plotted across the area with high apparent suitability between sunset and 3:00 am as the calling activity declines around that time of the night (or earlier, matching with sunrise at northern latitudes; Borz\u0026eacute;e et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e). This protocol makes the detection of the focal species most likely, here an early summer breeder. We spent 5 minutes listening to the call activity of the species at each site, generally resulting in a 95% probability of detection of Asian \u003cem\u003eDryophytes\u003c/em\u003e, focusing on the presence of \u003cem\u003eKaloula borealis\u003c/em\u003e, \u003cem\u003eStrauchbufo raddei\u003c/em\u003e, \u003cem\u003ePelophylax nigromaculatus\u003c/em\u003e, \u003cem\u003ePelophylax plancyi\u003c/em\u003e, \u003cem\u003eDryophytes japonicus\u003c/em\u003e, and here unexpectedly \u003cem\u003eDryophytes suweonensis\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eMolecular analyses\u003c/em\u003e\u003c/p\u003e\u003cp\u003eSampling this species does not require permits in China, and it was collected under the AICUC number 2024013 provided by the Ethical Committee of Nanjing Forestry University. Upon encountering the population of D. suweonensis, we orally swabbed five individuals in the vicinity of Qiqihar (47.59238\u0026deg;N; 124.27774\u0026deg;E) and stored them with a bead of desiccant for 10 days until storage at -20\u0026deg;C. We extracted DNA using the blood and tissue DNA extraction kit from Qiagen, using the manufacturer's protocols. Sequencing was performed for both the forward and reverse strands of a Cyt-b gene fragment (900 bp; primers: L0-ATGGCCCCTGTTTTACGCAA and1046- TAAATGGGTCTTCTACTG G; Dufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), by Tsingke Co., Ltd., Wuhan, People\u0026rsquo;s Republic of China.\u003c/p\u003e\u003cp\u003eFor species identification, we first trimmed and complemented the raw sequences for all five samples to 819-bp long fragments using Geneious Prime v. 2023.2.1 (Kearse et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). All sequences generated in this study were deposited to Genbank under the accession number PV231326 to PV231329 (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eInformation for ingroup and outgroup taxa used to reconstruct the phylogeny and species assignments for the five \u003cem\u003eDryophytes suweonensis\u003c/em\u003e individuals collected in the vicinity of Qiqihar, China. We used a mtDNA Cyt\u003cem\u003eb\u003c/em\u003e gene fragment of the analysis, for 77 \u003cem\u003eDryophytes\u003c/em\u003e individual vouchers.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eIsolate\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLocalities\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAccession number (Cyt\u003cem\u003eb\u003c/em\u003e)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSource\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1B\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Qiqihar\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePV231327\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2B\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Qiqihar\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePV231326\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3B\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Qiqihar\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePV231328\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4B\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Qiqihar\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePV231325\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5B\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Qiqihar\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePV231329\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes flaviventris\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8551\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Iksan, North Jeolla\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089227\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes flaviventris\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8552\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Iksan, North Jeolla\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089228\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes flaviventris\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8553\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Iksan, North Jeolla\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089229\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes flaviventris\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8548\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Iksan, North Jeolla\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089224\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes flaviventris\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8545\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Iksan, North Jeolla\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089221\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes versicolor\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMT759671\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBogart et al., 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes versicolor\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMT759669\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBogart et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes versicolor\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMT759665\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBogart et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8671\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089239\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8670\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089238\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8669\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089237\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8668\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089236\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8667\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089235\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8666\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089234\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8665\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089233\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms8664\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina: Hefei, Anhui\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089232\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms7859\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Siheung, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089210\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms7857\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Siheung, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089208\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms7852\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Asan, South Chungcheong\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089205\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms7844\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Asan, South Chungcheong\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089203\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms6871\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Cheongju, North Chungcheong\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089190\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms6864\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Cheongju, North Chungcheong\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089184\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms4470\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Paju, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089111\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms4469\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Paju, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089110\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms4462\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089107\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms3715\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eDPR Korea: Moonduk, South Pyeongan\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089093\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0474\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089092\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0467\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089090\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0357\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089089\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0356\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089088\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0352\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089087\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0351\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089086\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003emms0350\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMK089085\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eMin (2008), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eKUZ:nn2H_Kv04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eLC569676\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eKuraishi et al., 2021\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e14SIH002_Hsuw\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055770\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. 2011\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e14SIH001_Hsuw\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055769\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. 2012\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN394_Hsuw\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055768\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. 2013\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN718\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055734\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. 2014\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN640\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055733\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. 2015\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN639\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055732\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN638\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055731\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN637\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055730\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN636\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055729\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN635\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055728\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13GEN135\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055720\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChi201204.47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055717\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChi201204.46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055716\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChi201204.45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055715\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChi201204.42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055712\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi100277\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055705\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi100141\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055704\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi675\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055701\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi599\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055694\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi820\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055683\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi819\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055682\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi814\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055677\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi813\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055676\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHLi812\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055675\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHimm606002\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055673\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes immaculatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHimm606003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChina\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKY055672\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDufresnes et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Pyeongtaek, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564864\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Paju, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564863\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Paju, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564862\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Ganghwa, Incheon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564861\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Ganghwa, Incheon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564859\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Ganghwa, Incheon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564855\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes suweonensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Goseong, Gangwon-do\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564854\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Goseong, Gangwon-do\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564853\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Goseong, Gangwon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564852\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Paju, Gyeonggi\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564849\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Eumseong, Chungcheong\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564848\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eDryophytes japonicus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHaplotype 3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eR Korea: Gangneung, Gangwon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKF564847\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eChun et al. (2013), direct submission\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eBayesian posterior distribution of migration rates among four clades of the \u003cem\u003eDryophytes immaculatus\u003c/em\u003e species complex, estimated using a partial Cyt\u003cem\u003eb\u003c/em\u003e dataset (819 bp). The populations include \u003cem\u003eD. immaculatus\u003c/em\u003e (Di) from eastern China, \u003cem\u003eD. flaviventris\u003c/em\u003e (Df) isolated in Iksan, R Korea, and two clades of \u003cem\u003eD. suweonensis\u003c/em\u003e: one from the Korean Peninsula (DsK) and a newly identified population from Qiqihar, China (DsQ). These estimates provide insights into gene flow and population connectivity with Θ indicating the effective population size (Ne) scaled by the mutation rate (\u0026micro;), and M indicating the mutation-scaled migration rate.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.03%\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.25%\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMode\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.75%\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.98%\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eMedian\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eMean\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eΘ \u003csub\u003eDi\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.0028\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.0073\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.0136\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.0263\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.0756\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0240\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.0305\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eΘ \u003csub\u003eDf\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.0000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.0007\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.0023\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.0037\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.0154\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0032\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.0045\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eΘ \u003csub\u003eDsK\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.0049\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.0103\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.0157\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.0263\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.0627\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0236\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.0288\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eΘ \u003csub\u003eDsQ\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.0000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.0002\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0.0019\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.0033\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.0182\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0032\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0.0048\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDf\u0026minus;\u0026gt;Di\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e22.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e205.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e205.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e275.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e335.00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDsK\u0026minus;\u0026gt;Di\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e37.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e178.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e245.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e839.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e326.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e367.20\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDsQ\u0026minus;\u0026gt;Di\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e23.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e498.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e853.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e315.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e371.90\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDi\u0026minus;\u0026gt;Df\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e23.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e164.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e834.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e163.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e245.30\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDsK\u0026minus;\u0026gt;Df\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e23.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e260.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e592.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e271.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e329.70\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDsQ\u0026minus;\u0026gt;Df\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e103.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e501.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e756.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e384.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e421.10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDi\u0026minus;\u0026gt;DsK\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e25.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e170.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e632.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e170.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e228.90\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDf\u0026minus;\u0026gt;DsK\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e214.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e639.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e884.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e988.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e999.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e620.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e582.70\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDsQ\u0026minus;\u0026gt;DsK\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e97.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e620.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e975.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e996.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e999.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e607.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e577.20\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDi\u0026minus;\u0026gt;DsQ\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e24.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e193.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e888.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e195.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e265.30\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDf\u0026minus;\u0026gt;DsQ\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e24.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e228.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e316.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e446.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e458.90\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eM \u003csub\u003eDsK\u0026minus;\u0026gt;DsQ\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e23.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e276.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e699.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e289.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e351.60\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eIndependently, we aligned the Cytb sequences of all five individuals with 72 homologous sequences available in Genbank (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ncbi.nlm.nih.gov/genbank/\u003c/span\u003e\u003cspan address=\"https://www.ncbi.nlm.nih.gov/genbank/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). We restricted the final alignment to closely related species: Dryophytes japonicus, Dryophytes suweonensis and D. immaculatus (n taxa\u0026thinsp;=\u0026thinsp;77), as this analysis was conducted to confirm the species identification and not to conduct a population genetic study. These three species are closely related congeneric species distributed on the Korean Peninsula and China (see Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e for the details of the samples). We also included three samples of Dryophytes versicolor from the Nearctic as outgroups (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eWe first identified the best-fit partitioning scheme and substitution models for the Cytb sequences alignment. As the alignment included a partial intron (from base pair 1 to 12) and partial exon (from base pair 13 to 819), we included both the non-coding region and the three-codon strategies in our gene blocks parameter using Partition Finder v. 2.1.1 (Lanfear et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). We obtained HKY\u0026thinsp;+\u0026thinsp;I\u0026thinsp;+\u0026thinsp;G as the best substitution model for the four subsets of partition. We then integrated the information of the best substitution model in our concatenated alignment dataset (.Nexus) for the reconstruction of the Bayesian phylogenetic tree in Mr. Bayes v. 3.2.7 (Ronquist et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). We ran the analysis using the Monte Carlo Markov Chain (MCMC) algorithm, with four chains, 10,000,000 iterations, and a burn-in of 1,000. To ensure the convergence and robustness of the phylogenetic analysis, we optimised the value of split frequencies (\u0026lt;\u0026thinsp;0.05), the parameter space and the pattern of randomness of MrBayes at the end of its run.\u003c/p\u003e\u003cp\u003e\u003cem\u003eEcological niche models\u003c/em\u003e\u003c/p\u003e\u003cp\u003eFinally, we used ecological niche models to determine whether we are still missing some major populations. The data used for this analysis originated from the literature (Borz\u0026eacute;e, et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2017a\u003c/span\u003e; Son et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Borz\u0026eacute;e, et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) along with the new datapoint collected here (Supplementary File 1). To reduce the sampling bias, we thinned occurrence points within 4 km using the spThin package (Aiello-Lammens et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and using the bias files created by the MASS package (Venables and Ripley \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) in R version 4.2.1. We decided to rely on the 19 bioclimatic variables (Worldclim 2.1; Fick and Hijmans \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), elevation (2.5 arc-minute resolution) and Landsat normalised difference vegetation index (NDVI, USGS) after the removal of highly correlated variables (Pearson\u0026rsquo;s \u003cem\u003er\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.8; Bradie and Leung \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). We used six variables for the final models: annual mean temperature (bio 1), temperature annual range (bio 7), annual precipitation (bio 12), precipitation seasonality (coefficient of variation) (bio 15), and elevation. We ran MaxEnt v. 3.4.4 (Elith et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Phillips et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) with the default setting (Phillips and Dud\u0026iacute;k \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) and used the cross-validate run type option with ten replicates (10k-fold). For the model evaluation, we calculated the area under the curve (AUC; Somodi et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and true skill statistic (TSS; Allouche et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). To visualise the potential suitable habitat and to calculate the threshold area, we used ArcGIS pro v. 3.3.0 (ESRI, Redlands, CA, USA).\u003c/p\u003e\u003cp\u003e\u003cem\u003eCall description\u003c/em\u003e\u003c/p\u003e\u003cp\u003eWe recorded the advertisement calls of three individuals in the proximity of Qiqihar, Heilongjiang, China (47.592389\u0026deg;N; 124.277740\u0026deg;E) on 10 June 2024 at 20:25 (air temperature 24\u0026deg;C) in real-time using the 16-bit resolution and a sampling rate of 44.1 kHz. We used a linear PCM recorder (Tascam DR-40; California, USA) linked to a unidirectional microphone (unidirectional electret condenser microphone HT-81, HTDZ; Xi\u0026rsquo;an, China). The microphone was positioned straight, approximately 0.5 m from the mouth of the calling individual. We manually regulated the gain settings to avoid saturation and ensure the maximum amplitude peaks. The calls were deposited in the FonoZoo sound repository under accession numbers (FZ-SOUND-CODE 114785\u0026thinsp;\u0026minus;\u0026thinsp;14804).\u003c/p\u003e\u003cp\u003ePrior to call data extraction, we removed background noise by filtering out the calls at 1 kHz and selected the cleanest section of the calls for each calling individual in Raven Pro 1.5 software (Bioacoustics Research Program 2014) for analysis. We kept the Hann window size at 256 sample windows, 128-sample hop size with 50% frame overlap and a 172 Hz frequency grid spacing. We analysed both temporal and spectral properties of the calls.\u003c/p\u003e\u003cp\u003eWe then extracted the data using Raven Pro v. 1.5 (Bioacoustics Research Program 2014). In total, we analysed 60 notes from three calling male individuals (20 notes from each individual) for the acoustic analysis, following the definitions of call properties from Park et al. (\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and Borz\u0026eacute;e, et al. (\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e). Further, we processed the calls in R (R Core Team \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) using Seewave (Sueur et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and tuneR (Ligges et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) packages to produce high-quality spectrograms, oscillograms and power spectrums.\u003c/p\u003e\u003cp\u003e\u003cem\u003eConservation assessment\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThreat assessments over national boundaries are generally the most robust when following the IUCN Red List categories and criteria (Maes et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; IUCN Standards and Petitions Committee \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), and they are suitable indicators to inform on conservation needs (Lamoreux et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Rodrigues et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Species are evaluated against a set of criteria, and when threatened, they can fall into either of three categories: Vulnerable, Endangered or Critically Endangered (IUCN Standards and Petitions Committee \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Assessments are conducted against quantitative thresholds for five criteria: population size reduction, geographic range size, small population size and decline, very small population and/or restricted distribution, and quantitative analysis of extinction risk. Complete datasets for the assessments are difficult to generate and can follow alternative protocols (Mace \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Hermoso et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Here, the only data available is the one about the geographical range size, and we follow the IUCN Red List categories and criteria to suggest a threat assessment for the species (IUCN Standards and Petitions Committee \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003eField surveys\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe northernmost new location of \u003cem\u003eDryophytes suweonensis\u003c/em\u003e is further north than Qiqihar, about 985 km north of the last northernmost known location, which is about 300 km north of the southern boundary of the range of the species, now bringing the species close to 1000 km of latitudinal extent (Fig.\u0026nbsp;1). At Harbin latitude, the longitudinal extant between the two extremes populations is greater than 880 km, in comparison with a past greater maximal longitudinal extend of 80 km. The westernmost population is now in Panjin in Liaoning, about 300 km further west than previously known. As a result, the distribution of \u003cem\u003eD. suweonensis\u003c/em\u003e in R Korea is now comparatively smaller in terms of geographic coverage and potentially in terms of the number of individuals. Interestingly, all the individuals were encountered in rice paddies and showed the two dorsal stripes characteristic of \u003cem\u003eD. suweonensis\u003c/em\u003e emerging from hibernation, only visible for a day or two after emergence in R Korea (Borzée 2024; Fig.\u0026nbsp;2).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMolecular analyses\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe phylogenetic tree (Fig.\u0026nbsp;3) highlighted the clustering of the samples collected from Qiqihar with the \u003cem\u003eD. suweonensis\u003c/em\u003e clade (Bayesian posterior probability [BPP] of 100%). Despite being distributed about 1000 km away, all five individuals specifically clustered within the clade of \u003cem\u003eD. suweonensis\u003c/em\u003e and shared a paraphyletic relationship with \u003cem\u003eD. immaculatus\u003c/em\u003e, endemic to China (Fig.\u0026nbsp;3).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eEcological niche models\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe model was supported by AUC values of 0.958 ± 0.020 and a TSS value of 0.612 ± 0.108, and thus, both values indicated a perfect fit for the model (Phillips et al. 2006). The variable with the highest contribution to the distribution of \u003cem\u003eD. suweonensis\u003c/em\u003e was bio 7 (temperature annual range), with 63.7% contribution and 52.9% permutation importance (Table\u0026nbsp;4). A very large area with suitable habitat was found through the models in the plains between Changchun, Harbin and Qiqihar in Jilin and Heilongjiang provinces, and on the Songliao plains in Liaoning (Fig.\u0026nbsp;1). The habitat suitability results were thresholded in the map using the maximum training sensitivity plus specificity (MTSS) value of 0.14 to classify MaxEnt output raster files into suitable and unsuitable habitats (Fig.\u0026nbsp;1; Liu et al. 2013). The calculated suitable range of the species within the area highlighted on Fig.\u0026nbsp;1 was 713 248 km\u003csup\u003e2\u003c/sup\u003e.\u003c/p\u003e\n\u003cdiv\u003e\n \u003ctable border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003eTable 3\u003c/div\u003e\n \u003cdiv\u003e\n \u003cp\u003eCall properties of \u003cem\u003eDryophytes suweonensis\u003c/em\u003e from Qiqihar, China, collected in June 2024. The data is based on 60 notes extracted from the calls of three individuals (\u003cem\u003en\u003c/em\u003e = 20 per individual).\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCall properties\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eMean\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eSD\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eMin\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eMax\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eNote duration (s)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.02\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.22\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eNote dominant frequency (Hz)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2962.97\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e398.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3100.80\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e90% frequency bandwidth of note (Hz)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1521.68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e696.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e516.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2067.20\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eDuration of connected pulses (s)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.02\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.04\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.18\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePeak dominant frequency of connected pulses (Hz)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3031.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e85.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2928.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3100.80\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e90% frequency bandwidth of connected pulses (Hz)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1484.35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e745.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e344.50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2067.20\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eNumber of independent pulses\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.53\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2.47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003ctable id=\"Tab3\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv\u003eTable 4\u003c/div\u003e\n \u003cdiv\u003e\n \u003cp\u003ePercentage of contribution and permutation importance for each of the five variables retained to build habitat suitability model with MaxEnt for \u003cem\u003eDryophytes suweonensis\u003c/em\u003e.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eVariables\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eContribution percentage\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePermutation importance\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eTemperature annual range (bio 7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e63.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e52.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eAnnual precipitation (bio 12)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e24.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e32.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eAnnual mean temperature (bio 1)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePrecipitation seasonality (coefficient of variation; bio 15)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eElevation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e13.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003cem\u003eCall properties\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eIn total, we analysed acoustic data for 60 notes from three \u003cem\u003eD. suweonensis\u003c/em\u003e individuals from Qiqihar. The notes were produced at continuous intervals, and their temporal structure consisted of both independent and connected pulses. In the spectrogram, the sections of notes with independent pulses displayed double frequency bands for each pulse and four strong harmonics for the condensed pulsed parts (Fig.\u0026nbsp;4). The mean note duration was 0.12 ± 0.02 seconds, with a mean dominant frequency of 2962.97 ± 398.41 Hz. In comparison, the note duration for \u003cem\u003eD. suweonensis\u003c/em\u003e in R Korea was 0.12 ± 0.02, while it was 0.11 for \u003cem\u003eD. immaculatus\u003c/em\u003e and 0.15 for \u003cem\u003eD. flaviventris\u003c/em\u003e (Borzée, et al. 2020a). The average number of independent pulses per note was 6.53 ± 2.47, while it is 5.34 for \u003cem\u003eD. suweonensis\u003c/em\u003e in R Korea. The mean duration of connected pulses was 0.07 ± 0.02 seconds, and the mean peak dominant frequency of connected pulses was 3031.88 ± 85.12 Hz (Table\u0026nbsp;3).\u003c/p\u003e\n\u003cdiv\u003e\n \u003cbr\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003cem\u003eConservation assessment\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe geographic range and the extent of occurrence of the species is well over the 20,000 km\u003csup\u003e2\u003c/sup\u003e limit to be listed as threatened (noting that the 713 248 km\u003csup\u003e2\u003c/sup\u003e are not the range of the species, which will only be a fraction of this value, but still estimated to be well above the threshold). The range is severely fragmented in the south only, and the continued decline cannot be inferred for the northern populations. In conclusion, globally, the species is not listed as threatened based on B1 and B2 criteria (IUCN Standards and Petitions Committee 2024). The earlier assessment as Endangered (IUCN SSC Amphibian Specialist Group 2017) can, however, be used as a national Red List assessment for R Korea.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe discovery of new populations thousands of kilometres away from their known range is not unheard of, but it is unusual, and generally happens with cryptic species (e.g. \u003cem\u003eHydromantes strinatii\u003c/em\u003e, Lucente et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) or species with great dispersal capacities (e.g. \u003cem\u003eCarduelis cucullate\u003c/em\u003e, Robbins et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Here, a population of the Suweon treefrog (\u003cem\u003eDryophytes suweonensis\u003c/em\u003e) found about a thousand kilometres north of its type locality is entirely unexpected and raises questions about the lack of field research in the area and the threat assessment done on the species. However, there are decades old guidelines to remove infestation by the sister species, now known to be a misidentification, from silkworm farms within the range of \u003cem\u003eD. suweonensis\u003c/em\u003e, also noting their resistance to common pesticide, and providing more information about their life cycle than other updated literature (Liu et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1989\u003c/span\u003e). Here, we highlight that the species is not endemic to the Korean Peninsula, the population is not as small as previously expected, although there are no population size estimates available, and more basic ecological research on distribution is highly needed. We also call for more research on the distribution of species in northeast China in view of the recent discovery of species thought to be restricted to the Korean Peninsula, including \u003cem\u003ePelophylax chosenicus\u003c/em\u003e (Borz\u0026eacute;e, et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), \u003cem\u003eRana coreana\u003c/em\u003e (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and the presence of one most-unlikely described \u003cem\u003eOnychodactylus\u003c/em\u003e species.\u003c/p\u003e\u003cp\u003eA potential shortcoming to the claim of a range extension would be the discovery of a new species, which is not the case here in view of the clustering within the \u003cem\u003eD. suweonensis\u003c/em\u003e clade in the phylogenetic tree (Fig.\u0026nbsp;3). However, despite the species-specific assignment, cytochrome b alone is not sufficient to conduct phylogenetic analyses, a single gene fragment is not significant in terms of taxonomy, and the absence of introgression patterns and minimal incomplete lineage sorting is likely influenced by this limitation. For future studies, we recommended sampling along a latitudinal cline and the use of a genome-wide approach to achieve higher resolution and better insights into potential introgression and lineage sorting patterns of \u003cem\u003eD. suweonensis\u003c/em\u003e. In addition, the absence of data from DPR Korea prevents concluding whether there are additional gaps in the distribution of \u003cem\u003eD. suweonensis\u003c/em\u003e, and additional sampling would answer his question. Here, to follow the precautionary principle (Deville and Harding \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) in environmental protection and conservation (Glasgow et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, Kemp et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), the population in Qiqihar is assigned to \u003cem\u003eD. suweonensis\u003c/em\u003e, but further genomic analyses will help better understand the relationship within the species.\u003c/p\u003e\u003cp\u003eIn addition, the call properties of the frogs from Qiqihar matched with that of \u003cem\u003eD. suweonensis\u003c/em\u003e in R Korea (Borz\u0026eacute;e, et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e), with some variations, likely linked to the inability to extract data in a uniform manner across datasets. Further analysis within the species, including populations from DPR Korea, could highlight the impact of repeated marine transgression on the evolution of traits in amphibians. The distribution of the suitable habitat is also along the same landscape as the one where \u003cem\u003eD. suweonensis\u003c/em\u003e is present further south, with some gap in hilly areas. It is interesting to note that the area of suitable habitat in Jiangsu, China, does not match with the area where \u003cem\u003eD. immaculatus\u003c/em\u003e is present (Borz\u0026eacute;e \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThis abundance of species present on both the Korean Peninsula and in northeast China is, however, not unexpected as the area is a conservation-need hotspot for amphibians (Button and Borz\u0026eacute;e \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). It is also a pattern that can be expected for numerous other species, and especially mammals, as most species found on the Korean Peninsula are also found further north (Jo et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In addition, these broader ranges reinforce the conservation potential for these species (Borz\u0026eacute;e et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e) as all extinct mammal species and their designated protected areas are also found in China (e.g. tigers, Lee and Miller-Rushing \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). It is, therefore, critical to properly survey and reassess all species present on the Korean Peninsula and ranging into northeast China for their IUCN Red List status. In addition, the large number of threatened species in the area could result in the triggering of sites listed as Key Biodiversity Areas or under the Alliance for Zero Extinction (Mair et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Re:wild et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). As a result, the extinction risk for species would become linked to conservation policies, decision-making, and funding availability, a critical need for amphibians (Gu\u0026eacute;nard et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe distribution of \u003cem\u003eD. suweonensis\u003c/em\u003e is unlikely to go further north and west as surveys in Mongolia (Kuzmin et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2017b\u003c/span\u003e; Borz\u0026eacute;e et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2019a\u003c/span\u003e) and Russia (Othman et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) in the habitat that would be adequate for the species did not detect its presence. However, low plains in boundary areas in Russia need additional specific surveys. In addition, the species was only found in rice paddies, similar to the known sites in R Korea (Borz\u0026eacute;e and Jang \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), and there is no extensive rice agriculture at these latitudes in nearby countries. It is, however, important to highlight that there was a bias in the selection of sites for the surveys as \u003cem\u003eD. suweonensis\u003c/em\u003e was not one of the target species, and determining the presence of the target species is more easily done in riparian areas, where wetlands have been replaced by rice paddies. However, \u003cem\u003eD. suweonensis\u003c/em\u003e was detected in rice paddies that were contiguous with reed beds, similar to the habitat in DPR Korea (Borz\u0026eacute;e, et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), and further surveys in both China and DPR Korea to identify natural sites are important for conservation plans in R Korea (Park et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eWhile these findings clearly show that the past threat assessment as Endangered for \u003cem\u003eD. suweonensis\u003c/em\u003e is now outdated, it does not mean that these are wrong. At the time of the assessment, only the populations in R Korea were known, and it is therefore not a global assessment anymore, but a national assessment, in need of an update as the population has been declining since it was last done. To be able to properly assess the species following the IUCN Red List categories and criteria (IUCN Standards and Petitions Committee \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), additional surveys and research on the species will be needed as the polygon that would be currently used to define the extent of occurrence would miss some of the populations (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAcknowledgements\u003c/p\u003e\n\u003cp\u003eThis species does not require special permits, and it was collected under the AICUC number 2024013 provided by the Ethical Committee of Nanjing Forestry University. We are grateful to Wang Yingwei and Wang Yatao for reaching out and sharing information about an additional locality for\u0026nbsp;\u003cem\u003eDryophytes suweonensis\u003c/em\u003e in Liaoning.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of Funding.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis project was funded by the Foreign Youth Talent Program of the Ministry of Science and Technology of the People\u0026rsquo;s Republic of China (QN2023014004L), the Research Fund for International Scientists (RFIS) from the National Natural Science Foundation of China (NSFC; W2432021) and a grant from Forest City Studios to AB, and by the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR201803101) to MSM.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eData Accessibility\u003c/p\u003e\n\u003cp\u003eThe data that support this study are available in the article and accompanying online supplementary material.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of Interest Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflicts of interest\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eContribution:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData collection: AB, TEU, AS, MSM, SNO; Data analysis: TEU, VKP, AS, SNO, writing first draft: AB; writing revisions: TEU, VKP, AS, MSM, SNO\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAiello‐Lammens ME, Boria RA, Radosavljevic A, Vilela B, Anderson RP. 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New York, USA: Springer.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Supplementary Material","content":"\u003cp\u003eSupplementary file 1 is not available with this version.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"Nanjing Forestry University","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"treefrogs, conservation status, northeast Asia, species identification","lastPublishedDoi":"10.21203/rs.3.rs-7226956/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7226956/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eContext.\u003c/h2\u003e\u003cp\u003eNon-genuine conservation status updates threaten the trust in Red Listing, but they are necessary building blocks in science. In addition, they often highlight the critical need for basic field research. The Suweon treefrog, \u003cem\u003eDryophytes suweonensis\u003c/em\u003e, is a hylid species long thought to be range-restricted and assessed as Endangered by the IUCN Red List of Threatened Species.\u003c/p\u003e\u003ch2\u003eAims.\u003c/h2\u003e\u003cp\u003eThis study aimed at determining the species of individuals encountered during call surveys in 2024 in China, and determine the areas of suitable habitat for the species.\u003c/p\u003e\u003ch2\u003eMethods.\u003c/h2\u003e\u003cp\u003eWe sequenced the Cyt\u003cem\u003eb\u003c/em\u003e gene fragment to identify the species and compared the acoustic properties to identify the species. We also built ecological models with MaxEnt to predict suitable habitats for the species and guide future surveys.\u003c/p\u003e\u003ch2\u003eKey results.\u003c/h2\u003e\u003cp\u003eWe confirmed the identity of the population found more than 985 km north of the northernmost known locality of \u003cem\u003eD. suweonensis\u003c/em\u003e as the same species. The call properties also matched with those of \u003cem\u003eD. suweonensis\u003c/em\u003e. The ecological niche models identified a very large area with suitable habitat spanning across the northern Chinese plains.\u003c/p\u003e\u003ch2\u003eConclusions.\u003c/h2\u003e\u003cp\u003eThe species is not endemic to the Korean Peninsula, occurring in a significantly broader range, and thus, the population size is higher than previously assumed.\u003c/p\u003e\u003ch2\u003eImplications.\u003c/h2\u003e\u003cp\u003eThe species is unlikely to be threatened, and this huge range extension highlights the need for additional field surveys for all species in the area. Long term, the use of conservation prioritisation tools will highlight conservation needs in the region.\u003c/p\u003e","manuscriptTitle":"From threatened to widespread: a non-genuine conservation status update for a now widespread Asian Hylid - and the need for additional basic field research","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-31 05:48:38","doi":"10.21203/rs.3.rs-7226956/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"fb607584-d3b5-4332-94c5-c3d9c55e75b4","owner":[],"postedDate":"July 31st, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":52342881,"name":"Animal Science"}],"tags":[],"updatedAt":"2025-07-31T05:48:38+00:00","versionOfRecord":[],"versionCreatedAt":"2025-07-31 05:48:38","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7226956","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7226956","identity":"rs-7226956","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}